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ZOOLOGICA

SCIENTIFIC CONTRIBUTIONS
OF THE

NEW YORK ZOOLOGICAL SOCIETY

PUBLISHED BY THE SOCIETY
THE ZOOLOGICAL PARK, NEW YORK

NEW YORK ZOOLOGICAL SOCIETY

General Office: 630 Fifth Avenue, New York City

OFFICERS

President, Fairfield Osborn
First Vice-President, Alfred Ely

Chairman, Executive Committee & Second Vice-President, Laurence S. Rockefeller
Treasurer, Cornelius R. Agnew

General Director, Zoological Park and Aquarium, Allyn R. Jennings
Assistant General Director, Harry Sweeny, Jr.

SCIENTIFIC STAFF

Zoological Park

Raymond L. Ditmars, Curator of Reptiles and Insects
Lee S. Crandall, Curator of Birds
Claude W. Leister, Curator of Mammals and Educational Activities
Leonard J. Goss, Curator of Health
William Bridges, Editor and Curator of Publications

Aquarium

Charles M. Breder, Jr., Director
Christopher W. Coates, Aquarist
Ross F. Nigrelli, Pathologist
G. M. Smith, Research Associate in Pathology
Homer W. Smith, Research Associate in Physiology

Department of Tropical Research

William Beebe, Director
John Tee-Van, General Associate
Gloria Hollister, Research Associate
Jocelyn Crane, Technical Associate

Editorial Committee

Fairfield Osborn, Chairman

Allyn R. Jennings Charles M. Breder, Jr.

William Beebe William Bridges

CONTENTS

Part 1. March 18, 1940.

1. The Breeding Behavior of the Common Shiner, Notropis

cornutus (Mitchill). By Edward C. Raney. (Plates I-IV ;
Text-figure 1) 1

2. Divergence and Probability in Taxonomy. By ISAAC GlNS-

BURG 15

3. Miscellaneous Notes on the Eggs and Young of Reptiles. By

Roger Conant & Alexander Downs, Jr 33

4. Occlusion of the Venom Duct of Crotalidae by Electro-coagu-

lation: an Innovation in Operative Technique. By Duval
B. Jaros. (Text-figure 1) 49

5. Eastern Pacific Expeditions of the New York Zoological So-

ciety. XVII. A Review of the American Fishes of the Family
Cirrhitidae. By John Tee-Van. (Plate I; Text-figures 1-4) 53

6. Eastern Pacific Expeditions of the New York Zoological So-

ciety. XVIII. On the Post-embryonic Development of
Brachyuran Crabs of the Genus Ocypode. By Jocelyn
Crane. (Text-figures 1-8) 65

7. New Species of British Guiana Heterocera. By W. Schaus . . 83

8. A Papillary Cystic Disease Affecting the Barbels of Amei-

umis nebulosus (Le Sueur), Caused by the Myxosporidian
Henneguya ameiurensis sp. nov. By R. F. NlGRELLi & G. M.
Smith. (Plates I- VIII; Text-figure 1) 89

9. Caudal Skeleton of Bermuda Shallow Water Fishes. IV.
Order Cyprinodontes : Cyprinodontidae, Poecilidae. By
Gloria Hollister. (Text-figures 1-17) 97

10. The Histology of the Eye of the Cave Characin, Anoptichthys.

By E. B. GRESSER & C. M. Breder, Jr. (Plates I-III) 113

Part 2. July 3, 1940.

11. Plankton of the Bermuda Oceanographic Expeditions. IX.

The Bathypelagic Caridean Crustacea. By Fenner A.
Chace, Jr. (Text-figures 1-64) 117

12. Eastern Pacific Expeditions of the New York Zoological So-

ciety. XIX. Actiniaria from the Gulf of California. By
Oskar Carlgren. (Text-figures 1-8) 211

13. Morphological and Embryological Studies on Two Species

of Marine Catfish, Bagre marinus and Galeichthys felis.

By Daniel Merriman. (Plates I-V; Text-figures i-9) 221

PAGE

14. Propagation of the Electric Impulse Along the Organs of the

Electric Eel, Electrophorus electricus (Linnaeus) . By C. W.
Coates, R. T. Cox, W. A. Rosenblith & M. Vertner
Brown. (Plate I; Text-figures 1-3) 249

15. Notes on the Display Forms of Wahne’s Six-plumed Bird of

Paradise. By Lee S. Crandall. (Text-figures 1-3) 257

16. Acute Hemorrhagic Gastro-enteritis in a Giant Panda. By

Leonard J. Goss 261

17. Two New Species of Trematodes from the Deep Sea Scorpion

Fish, Scor])aena maclurensis Cuv. & Val. By Ross F.
Nigrelli. (Plate I; Text-figures 1 & 2) 263

18. Report of the Hospital and Laboratory of the New York

Zoological Park, 1939. Mortality Statistics of the Society’s
Collection. By Leonard J. Goss 269

Part 3. November 14, 1940.

19. Eastern Pacific Expeditions of the New York Zoological So-

ciety. XX. Medusae of the Templeton Crocker and Eastern
Pacific Zaca Expeditions, 1936-1938. By Henry B. Bigelow.
(Text-figures 1-20) 281

20. Two New Species of Trematodes ( Apharyngostrigea bilobata :

Strigeidae, and Cathaemasia nycticoracis : Echinostomidae)
from Herons, with a Note on the Occurrence of Clinos-
tomum companulatum (Rud.). By 0. Wilfred Olsen.
(Plate I) 323

21. Nesting of the Sunfish, Lepomis auritus (Linnaeus), in Tidal

Waters. By Neil D. Richmond. (Plate I) 329

22. Eastern Pacific Expeditions of the New York Zoological So-

ciety. XXL Notes on Echinoderms from the West Coast of
Central America. By Hubert Lyman Clark. (Plates I &

II; Text-figures 1-4) 331

23. The Nesting Behavior of Eupomotis gibbosus (Linnaeus), In

a Small Pool. By C. M. Breder, Jr. (Plates I & II; Text-
figures 1 & 2) 353

24. Reproductive Activities of a Hybrid Minnow, Notropis cor-

nutus X Notropis rubellus. By Edward C. Raney 361

Part 4. December 31, 1940.

25. Eastern Pacific Expeditions of the New York Zoological So-

ciety. XXII. Mollusks from the West Coast of Mexico and
Central America. Part I. By Leo George Hertlein & A. M.
Strong. (Plates I & II) 369

26. On the Electric Powers and Sex Ratios of Foetal Narcine

brasiliensis (Olfers). By C. M. Breder, Jr., & Stewart
Springer 431

VI

PAGE

27. A Study of the Activities of a Pair of Galago senegalensis

moholi in Captivity, Including the Birth and Postnatal
Development of Twins. By Florence De L. Lowther.
(Plates I-VI) 433

28. Diets for a Zoological Garden: Some Results During a Test

Period of Five Years. By Herbert L. Ratcliffe 463

29. The Biology of the Smoky Shrew ( Sorex fumeus fumeus

Miller). By W. J. Hamilton, Jr. (Plates I-IV ; Text-figure
1) 473

30. Social and Respiratory Behavior of Small Tarpon. By Arthur

Shlaifer & C. M. Breder, Jr. (Plates I & II ; Text-figure 1)493

31. New Observations on the Blood Group Factors in Simiidi

and Cercopithecidae. By P. B. Candela, A. S. Wiener

& L. J. Goss 513

32. Muscle Dystrophy in Tree Kangaroos Associated with Feeding

of Cod Liver Oil and. Its Response to Alpha-Tocopherol.

By Leonard J. Goss 523

33. Mortality Statistics for Specimens in the New York Aquarium,

1939. By Ross F. Nigrelli. (Plates I-III) 525

y- 34. A Comparison of Some Electrical and Anatomical Character-
istics of the Electric Eel, Electrophorus electricus (Lin-
naeus). By R. T. Cox, W. A. Rosenblith, Janice A. Cut-
ler, R. S. Mathews & C. W. Coates. (Text-figures 1-7) . .553

Index to Volume XXV 563

V

ZOOLOGICA

SCIENTIFIC CONTRIBUTIONS
OF THE

NEW YORK ZOOLOGICAL SOCIETY

VOLUME XXV
Part 1

Numbers 1-10

PUBLISHED BY THE SOCIETY
THE ZOOLOGICAL PARK, NEW YORK

March 18, 1940

CONTENTS

PAGE

1. The Breeding Behavior of the Common Shiner, Notropis

cornutus (Mitchill). By Edward C. Raney. (Plates
I-IV; Text-figure 1) 1

2. Divergence and Probability in Taxonomy. By Isaac

Ginsburg 15

3. Miscellaneous Notes on the Eggs and Young of Reptiles.

By Roger Con ant & Alexander Downs, Jr. 33

4. Occlusion of the Venom Duct of Crotalidae by Electro-

coagulation: an Innovation in Operative Technique.

By Duval B. Jaros. (Text-figure 1) 49

5. Eastern Pacific Expeditions of the New York Zoological

Society. XVII. A Review of the American Fishes of the
Family Cirrhitidae. By John Tee-Van. (Plate I; Text-
figures 1-4) 53

6. Eastern Pacific Expeditions of the New York Zoological

Society. XVIII. On the Post-embryonic Development of
Brachyuran Crabs of the Genus Ocypode. By Jocelyn
Crane. (Text-figures 1-8) 65

7. New Species of British Guiana Heterocera. By W.

Schaus 83

8. A Papillary Cystic Disease Affecting the Barbels of
Ameimus nebulosus (Le Sueur), Caused by the Myxo-
sporidian Henneguya ameiurensis sp. nov. By R. F.
Nigrelli & G. M. Smith. (Plates I- VIII; Text-fig-

ure 1) 89

9. Caudal Skeleton of Bermuda Shallow Water Fishes. IV.
Order Cyprinodontes: Cyprinodontidae, Poecilidae.

By Gloria Hollister. (Text-figures 1-17) ... 97

10. The Histology of the Eye of the Cave Characin, Anop-
tichthys. By E. B. Gresser & C. M. Breder, Jr. (Plates
I-III) 113

ZOOLOGICA

SCIENTIFIC CONTRIBUTIONS
OF THE

NEW YORK ZOOLOGICAL SOCIETY

1.

The Breeding Behavior of the Common Shiner, Notropis
cornutus (Mitchill) .

Edward C. Raney

Zoology Department, Cornell University,

Ithaca, New York

(Plates I-IV; Text-figure 1).

Although several authors have written concerning the habits of the
common shiner, Notropis cornutus (Mitchill), there is little detailed infor-
mation available on the breeding behavior of this widely distributed
cyprinid. The more pertinent papers giving data on spawning are those
by Adams & Hankinson (1928), Greeley (1927 and 1929) and Hankinson
(1932). The author has made an intensive study of the breeding habits of
the common shiner, Notropis cornutus cornutus, during the past spring
(1939) in the region about Ithaca, New York, and these data, together
with observations made elsewhere mostly during the previous four years,
are presented here. The breeding act has been observed many times at vari-
ous places and under different conditions. However, the details were
obscured, as they are in many fishes, because of the rapidity of consumation
until photographs were taken under nearly ideal conditions in nature (see
Plates I-IV). The method used in securing the photographs was similar to
that used by some ornithologists. A nest of common shiners was located at
a spot where the water was shallow and clear. If the shiners were at a
high pitch in their spawning activity they would return after a period rang-
ing from one-half to several hours and carry on normally with the camera
mounted on a tripod and operated by an observer only 4 to 5 feet away. By
utilizing exposures of 1/100 of a second or less most action could be stopped
and the spawning process pictured. This method will probably be helpful in
studying the habits of other nest building species among our fresh water
fishes.

Migration.

An inshore migration in lakes and, at least in some localities, an up-
stream movement is made by the common shiner. At the south end of
Cayuga Lake, near Ithaca, New York, large schools are found moving toward

2 2 mo

2

Zoologica: New York Zoological Society

[XXV :1

shore during the first two weeks in May when the water has reached a
temperature of 55° to 60° F. Their coming is eagerly awaited by bait
dealers since, at this time, the common shiner is in demand as bait for the
pike, Esox lucius. Nurnberger (1931, p. 215) has noted the presence of
large schools of the common shiner appearing along the shore of Big Sandy
Lake, Minnesota, on June 1, 1928, when the water temperature was 68° F.
In the absence of actual reported observations of spawning in ponds and
lakes it is assumed that the common shiner resorts to the gravel riffles of
tributary streams, but Hubbs & Cooper (1936, p. 65) have pertinently
stated that “its abundance in some of the inland lakes (Michigan) suggests
that it may also spawn successfully over the gravel shoals of these lakes.”

In streams there appears to be at least a slight upstream migration from
the deeper pools where the adults have wintered. On May 8, 1939, adult
common shiners were found moving upstream into the trap of a weir placed
across Grout Brook, the inlet of Skaneateles Lake, Cortland County, New
York. Constant surveillance of the waters below the weir had failed to note
any shiners before this date. The nearest satisfactory wintering place was
300 or more yards away and it is not at all improbable that they wintered
in the lake 600 yards distant. Just how far individual shiners may migrate
is, in the absence of tagging experiments, unknown.

Sexual Dimorphism.

Sexual dimorphism is pronounced in the common shiner, as in most
other nest building or territory guarding cyprinids, as Semotilus, Cam-
postoma and Nocomis. Breeding males of the common shiner attain a larger
size, have well developed pearl organs or breeding tubercles and are more
highly colored than females. It is obvious that the largest females in a
stream exceed in size the smallest breeding males (Text-figure 1). Neverthe-
less the difference in size is one of the best characters for distinguishing the
sexes at a given nest and in many instances of spawning I have yet to see a
male spawn with a larger female. It is difficult to generalize regarding the
actual sex ratio but with adult shiners on the spawning ground invariably
there are more females than males. The maximum size for male common
shiners is about 8 inches in total length. A male Notropis cornutus chryso-
cephalus in breeding color is figured in Forbes & Richardson (1920: oppo-
site page 147) and a male Notropis cornutus frontalis in Greeley (1927:
plate number 5).

The following color notes were made one hour after preservation in
10% formalin, from specimens of Notropis cornutus cornutus taken from
over a nest in Salmon Creek at Ludlowville, New York, on June 6, 1939.
The sides of the breeding males become suffused with red, the amount
varying considerably with the individual. The posterior border of all fins
tends to have a red band which in turn is edged by a much narrower clear
area. There is a pronounced tendency for the pectoral and ventral fins to
become suffused throughout with red except for a light posterior border and
a milky white anterior border. The rays of the dorsal, caudal and pectoral
fins tend to become dusky just anterior to the red band while the base of the
anal fin usually remains milky white. The base of the ventral fins shows a
slight tendency to become dark in some individuals. The branchiostegals
are red and the lower part of the cheek and opercles are at times a dull red.
The upper part of the opercles and cheek are a slate blue as well as a narrow
band immediately behind the opercle. There is a longitudinal mid-dorsal
streak and a streak on each side of the back which vary from a silver to a
greenish color but at all times are highlights on a dark background which
make excellent field recognition characters (see Plate II).

Breeding females are dusky above and silvery along the sides. In a few
cases a slight amount of red has been seen, especially at the posterior border
of the caudal and anal fins. The rays of the dorsal, pectoral and caudal fins

1940]

Raney: Breeding Behavior of Notropis cornutus

3

tend to become dusky while the ventral and anal fins are usually white.
While females have a faint longitudinal streak down the mid-line of the
back and one along each side of the back they are not as pronounced as in
the males. The swollen abdominal region of the ripe or near ripe female
is also a helpful field character.

The males begin to show color about a month before spawning, a sub-
dued red tint appearing first on the lower fins. The red which is to be seen
along the sides in the ripe males develops last and appears only a few days
before spawning takes place. After breeding the color fades quickly and is
usually gone in two weeks, the red on the sides being the first to go. How-
ever, an occasional male is taken even in late summer with a slight amount
of color in the fins.

The breeding male has well developed pearl organs scattered on the
back in front of the dorsal fin, on the nape, on the top of head and on the
snout; a few under the anterior part of the eye, in the region of the pre-
maxillaries and maxillaries and a single row along the lower jaw. The
tubercles on the head are better developed than the more posterior ones.
A few minute pearl organs occur on the upper sides, below and in front
of the dorsal fin; on the posterior border of the opercle; on a few of the an-
terior scales just dorsad of the base of the pectoral and on the belly between
the pectoral and ventral fins. An irregular row of fairly large pearl organs
is to be seen on the anterior edge of the first ray of the dorsal fin and
smaller ones occur on the sides of the first few dorsal rays. The dorsal
side of the pectoral fins are well supplied with fairly large tubercles which
undoubtedly are of use in enabling the male to hold the female in spawning.
Minute tubercles occur on the ventral side of the pectoral and on both sides
of the ventral fins near the anterior border. A careful examination of a
dozen ripe females disclosed no tubercles. However, Fowler (1912, p. 473)
reports that “occasionally a tuberculate female is found.”

As with the color, the tubercles of the male grow gradually, starting
about a month before spawning, and are usually gone within two weeks
after spawning, although the scars of the larger pearl organs may oftem
be seen up until one month after breeding.

Functions of Breeding Tubercles.

The well developed breeding tubercles on the snout, chin, head and ori
the back, in front of the dorsal fin, of the males are of unquestionable value
in the fighting which takes place in attempting to hold a territory and in
driving away predators which attempt to eat the eggs. These tubercles are
also a protection to the male in the sporadic digging activities in which,
they engage. Probably the only tubercles which are of value in the spawn-
ing act are those on the dorsal side of the pectoral fin which functions as
a grasping organ and those on the ventral side between the pectoral and
ventral fins which come in direct contact with the side of a female (Plate II).

Time of Spawning.

In Big Sandy Lake, Minnesota, Nurnberger (1931, p. 215) reports that
spawning occurred in 1929 between June 24 and 29 when the water reached
66°-70° F. In Illinois, according to Forbes & Richardson (1920, p. 148)
spawning occurs from May 1 to the last of June. Hubbs & Cooper (1936, p.
65) give the spawning season in Michigan as extending from the latter part
of May into June and mention that “spawning probably rarely occurs at
water temperatures lower than 60° to 65° F.” In western Pennsylvania
spawning takes place in May and June while in New York the season may
extend from the last two weeks in May until the middle of July in some of
the colder Adirondack streams. About Philadelphia, Pennsylvania, Fowler

4

Zoologica: New York Zoological Society

[XXV :1

(1909, p. 540) reports that the shiner spawns from late April to early
summer. Tracy (1910, p. 68) gives spring and early summer as the season
in Rhode Island. In the Connecticut Lakes in Maine Kendall & Goldsborough
(1908, p. 31) give the breeding time as spring or early summer, depending
on the temperature of the water.

The lowest water temperature at which I have actually observed shiners
spawn is 64° F. but they very likely begin to spawn at a temperature of
60 F. or thereabouts. However, in certain cold streams spawning probably
occurs at a slightly lower temperature. Spawning has personally been ob-
served when the water temperature was as high as 78° F. and Greeley (1929,
p. 172) saw Notropis cornutus frontalis spawning in Silver Creek, a tribu-
tary of Lake Erie, Chautauqua County, New York, as late as July 9 at a
water temperature of 83° F. At any one locality spawning is usually over in
ten days although ripe males are often seen for longer periods.

No breeding was observed taking place at night and at this time few
adult fishes remain in close proximity to the nest. Most of them retire to
the deeper pools nearby where they are relatively inactive. The spawning
act was seen as early as 9 A.M. on May 27, 1939, in Willseyville Creek at
Willseyville, Tioga County, New York. Spawning reached a high point about
12 noon and gradually tapered off after 4 P.M., none being noted later in
the day than 6 P.M. Spawning had reached a peak of activity at this par-
ticular locality on May 27. Earlier and later in the season there was but
little breeding activity at this place except in the afternoon.

Spawning Site.

A very important factor in explaining the wide distribution and the
local abundance of the common shiner is its capacity to spawn successfully
in many different habitats. Shiners may (1) spawn over gravel beds in
running water, (2) excavate small depressions in gravel or sand in running
water or, (3) utilize the nests built by other species (Plate I), even when
these nests occur in pools as is frequently the case ivith Exoglossum. They
appear to prefer the nests of other species as nesting sites when they are
available, although in 100 yards of Catatonk Creek, at Candor, New York,
on May 31, 1939, shiners were observed utilizing all three of the above-men-
tioned nesting places. Their preference for Exoglossum nests, especially in
small streams, often finds them spawning over these nests in the relatively
quiet waters of shalloAV pools rather than using the gravel riffles which are
so common nearby. The depth of the water over a spawning site is usually
less than 8 inches except when nests of other fishes are used. Four
large male shiners were once observed spawning over an Exoglossum nest
situated at the edge of a deep pool in 2 feet of water.

Gravel Beds as a Nesting Site.

Shiners often utilize gravel in or at the head of riffles as a spawning
site. These areas can hardly be considered nests although they can easily
be distinguished by the cleared nature of the bottom. The clearing away
of the silt is accomplished in two ways; first, by the constant moving of
the male fish over the gravel during which the lower fins make some contact
with the bottom, and, second, by the occasional sporadic digging motions
practiced by the male. He inserts his head between two pebbles and, by a
quick sidewise motion, dislodges one or both of them. Only rarely do they
pick up small stones in their mouths.

Fowler (1909, p. 540) has observed breeding schools of several hundred
brilliantly colored males packed close together over clear gravel or sandy
shallows in the region about Philadelphia, Pennsylvania. Several splendid
opportunities have been had recently for observing spawning (subspecies
cornutus ) over gravel in Fall Creek, Ithaca, New York, and in Salmon Creek,

1940]

Raney: Breeding Behavior of Notropis cornutus

5

at Ludlowville, New York. In western Pennsylvania Notropis cornutus
chrysocephalus has been observed spawning over gravel in riffles. Of the
above-mentioned localities when gravel is utilized many more shiners spawn
together over a given area than when the nests of other species are used.
At Ludlowville on June 6, 1938, from 80 to 100 males (Text-figure 1) had
taken a position in a riffle over a gravel bottom where the water varied in
depth from three to five inches. Under such conditions each male attempts
to hold a small territory which, of course, is quite limited because of the
large number of males in such a restricted area and, at most, consists of a
distance of three inches or so to either side. Much fighting occurs among the
males but after one retreats from the rush of another male he usually comes
back to his original position and the rushes may be repeated many times
between spawnings. The offensive activity of a male consists in quickly

50 60 70 80 00 100 110 120 130

STANDARD LENGTH IN MM.

Text-figure 1.

Frequency distributions of breeding males and females of Notropis cornutus
cornutus taken from over one spawning area in Salmon Creek, at Ludlowville,
Tompkins County, New York, on June 6 and 9, 1939. Measurements of standard
length were taken on 130 males and 232 females preserved in alcohol.

6

Zoologica: New York Zoological Society

[XXV :1

circling either to right or left and attempting to strike with the tubercled
snout. Few direct hits are made as opposing males are usually alert and
beat a quick retreat at the moment an antagonist starts circling. The water
■“boils” over such groups of males as they break water with their dorsal and
caudal fins during this fighting and maneuvering for position. Occasionally
“deferred combat” (Reighard, 1910, p. 1129) occurs as two males of nearly
the same size (Plate III) tensely swim off parallel to each other, often for
a distance of 3 to 5 feet. If they are of equal size they soon return to or
close to their original position on the gravel. There was a tendency at
times for this large breeding group to break into two, about half the males
moving upstream several feet and there repeating the same combat for
territory.

Approximately 150 females could be seen on the gravel below the hold-
ings of the males. When they were ready to breed the females would come
upstream and take a position over a male and spawning would take place in
a manner to be described later. This large spawning group was somewhat
unusual in that it was located in a shaded spot under a huge American elm
Wee and received no direct sun during the afternoon. In such a situation
where spawning occurs over coarse gravel the eggs may be seen and re-
covered by picking up or scuffing the pebbles. Some of the eggs are free
between pebbles, singly or in groups, while others are found adhering to
the stones.

At times, especially toward the end of the spawning season, a large
depression is made on the bottom by a large group of spawning males.
Whether a nest-like excavation is apparent depends somewhat on the type
of bottom. At Ludlowville the pebbles were fairly large, averaging about
one inch in diameter with few under one-half inch in diameter. Under con-
ditions such as this the digging activities of the males have little effect other
than to give a clean look to the area. In lower Fall Creek, at Ithaca, New
York, a similar breeding group will excavate a shallow depression in the
bottom consisting of sand and small gravel, which is often 3 to 4 feet in
diameter. Such nests are hardly distinguishable from the type which is
discussed next.

Nests Excavated by the Common Shiner.

Both subspecies cornutus and frontalis have been observed digging and
guarding their own circular nests, which are usually 8 to 12 inches in
diameter. In most cases a pit partly or wholly excavated by Semotilus
(Plate I) or Campostoma is taken over by a number of shiners, the number
varying usually from 3 to 20 males although occasionally but one male oc-
cupies a nest. These nests are most often built at the head of a riffle but are
also often seen in fairly fast flowing riffles. They are probably never built
in quiet water. The males make the nests by digging activities in which
they insert their heads between two stones and by jerking the head to the
side dislodge the stones laterally. Several males will engage in this digging
at one time although there are frequent interruptions as the males dash at
each other, vieing for the position furthest upstream in the nest. This
digging usually occurs irregularly between spawning acts.

Utilization of Nests of Other Fishes.

Several authors including Greeley (1929, p. 172), Hankinson (1920, p.
8, and 1932, p. 415), Hubbs & Cooper (1936, p. 65) and Van Duzer (1939,
p. 73) have pointed out instances of the common shiner utilizing the nests
of other species, such as Nocomis micropogon, Nocomis biguttatus, Semo-
tilus a. atromaculatus , Leucosomus corporalis, Exoglossum maxillingua and
Campostoma anomalum. Shiners appear to prefer the pebble nests of
Nocomis and Exoglossum when available. Their second choice under most

1940]

Raney: Breeding Behavior of Notropis cornutus

7

conditions appears to be the pit and ridge nest dug by Semotilus and the
author has observed them taking over and enlarging the circular pits exca-
vated by male Campostoma anomalum pullum. The breeding pattern is the
same, however, whether the nest of another species is utilized or a nest of
its own is excavated.

On several occasions Semotilus and Nocomis have been working on their
nests at the same time that shiners were spawning there. The male shiners
seldom attacked a male Semotilus and the Semotilus in turn continued re-
moving stones from his pit and piling them in a ridge upstream (see
Reighard, 1910, p. 1125). At times he would turn on and strike with his
tuberculate head a shiner that became too bold. As Hankinson (1932, p.
418) has pointed out a male Nocomis will frequently continue bringing
stones to add to his ever-growing nest even though many shiners are present
and spawning is occurring. An amazing example of the complacency of a
male Nocomis micropogon was noted in Catatonk Creek at Candor, New
York, on May 31, 1939. Twenty male and thirty female cornutus were
swarming over a Nocomis nest. These were accompanied by a dozen breed-
ing Notropis rubellus. A number of specimens of Exoglossum maxillingua
and Rhinichthys cataractae were darting in from their position on the
periphery to eat what eggs they could secure. During this great activity
which made the water “boil” over the nest, the male Nocomis calmly added
stones to his nest for a period slightly more than one hour without at any
time attacking any of the breeding fishes that were so close. At the same
time the male shiners carefully respected the Nocomis. However, female
Semotilus or Nocomis seldom appear when shiners swarm the nests, and
actual spawning has not been seen under these conditions.

With spawning Exoglossum, as Van Duzer (1939, p. 73) has pointed
out, “the mating of the cut-lips was always definitely lessened and sometimes
stopped by their ( cornutus ) presence and activity at the nest.” It is quite
probable that the male cornutus have learned to respect the sizeable male
Semotilus and Nocomis, armed as they are with well developed pearl organs
on the head, while they have become equally familiar with the abortive
rushes of the non-tuberculate and usually smaller male Exoglossum.

An interesting side light is the considerable number of natural hybrids
which occur largely as a result of this habit of spawning over the nests of
other species. Two species have often been seen spawning at the same time
over a Nocomis nest.

Hybrids of the combination Notropis cornutus X Notropis rubellus are
frequently collected and both of these species have been seen spawning over
a Nocomis nest at the same time (see Hankinson, 1932, p. 417). The hybrid
Notropis cornutus X Nocomis micropogon has been recorded by Greeley
(1938, p. 51). Clinostomus elongatus spawns over the nests of both Notropis
cornutus and Semotilus atromaculatus atromaculatus and both Greene (1935,
p. 89) and Greeley (1938, p. 51-52) have recorded the hybrids Notropis
cornutus X Clinostomus elongatus and Semotilus atromaculatus atromacul-
atus X Clinostomus elongatus. Greeley (1938, p. 52) has also reported
hybrids Notropis cornutus X Semotilus atromaculatus atromaculatus and
Notropis cornutus X Leucosomus corporalis which likely result from
cornutus spawning over the nests of these species. Thompson (1935, p.
492) has mentioned that in Illinois hybrids of Notropis cornutus are com-
mon presumably because cornutus spawns in the nests of other species.

Sex Recognition.

The female that successfully spawns, invariably approaches the nest
from the downstream side and assumes a position dorsad and slightly down-
stream from the male, who faces upstream (Plate I). Should she continue
upstream to a position in front of a male, as occasionally happens, she would
be driven away by the rush of a male, usually the male holding the dominant

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position in the nest. The male would usually dart at and cause to retreat
downstream any female getting into the nest at his body level, that is,
near the bottom of the nest. In case a female was chased she would beat
a retreat downstream, assume a position from one to several feet below the
nest and then gradually come upstream again over a male. The constant
turning of his head from side to side enables him to spot any fish which
comes above him. The behavior of a male when a female has assumed a
position above him is evidence that he recognizes her by the position she
takes. The male at this time will, while facing upstream with body nearly
straight, incline his body from side to side so that his reddish sides flash
alternately (Plate I). While the male may react to the mere presence of a fish
body above and behind him, there is a possibility that the male is attracted
by the milky white area about the slightly protruding external opening of
the oviduct of the female. In the case of one ripe female, but two inches long,
this white area was so pronounced it might easily be seen by an observer
standing five feet away. It must surely be an attractive mark when viewed
from the position of the male. A female will take a position above any male
who is in or slightly below the nest. Often a small male on the outside edge
of the nest (Plate II) will be successful in spawning more often than the
large male dominating the nest since the latter is forced to spend a large
part of his time attempting to chase away other fishes. However, one excep-
tionally large and vigorous male was once seen holding a Semotilus pit so
effectively by his wild dashes at intruding male cornutus that he successfully
spawned with a half-dozen females in about one and one-half minutes before
any other males seriously challenged and finally displaced him.

There is one bit of evidence that female cornutus react to fish of larger
size whether of the same species or not. At one instance a male Exoglossum
returned to his nest, and took a position facing upstream. He moved
slightly to right and left over the nest much as does a male cornutus, and
several female cornutus took a position above and dipped down to take a
spawning position beside him. The male Exoglossum reacted by turning and
rushing at them and finally chased the female shiners away from the nest.

Spawning Act.

With a female above a male and the male alternately inclining himself
to a semi-recumbent position first to one side then to the other, conditions
are set for the spawning act (Plate I). The female dips downward beside
that side of a male which is inclined toward the bottom or making the more
acute angle. The anterior end of the head of the female is just beyond the
anterior border of the pectoral fin of the male. The pectoral fin of. the
male is inserted under or slightly behind the head of the female, the distal
end of his pectoral being curved upward around the female. At the same
time his caudal peduncle is swung up over the caudal peduncle of the female
and then the caudal peduncle of the male moves downward. As a result of
this pressure the female is lying on her side at right angles to the male
with her head toward shore and usually with her ventral side upstream
(Plate II).

The male continues to bend his body in such a way as to enclose the
female within a curve (Plate III). It appears that the eggs are laid at this
moment, the eggs being forced out by the downward pressure exerted by
the male. The female then straightens out (Plate IV) and shoots head
upwards to, or almost to, the surface just as the male begins to straighten
himself. When observing a large group of spawning shiners, females may
constantly be seen breaking water as they come out of the spawning em-
brace. The speed with which this spawning act takes place is a fraction of
a second or just slightly longer than the time necessary to snap ones fingers
quickly, and the details are about impossible to see clearly with the unaided
eye, even when observed at close range. The spawning act is much like that

1940] Raney: Breeding Behavior of Notropis cornutus 9

of Semotilus atromaculatus atromaculatus as described by Reighard (1910,
p. 1130).

The male shiner immediately takes his position facing upstream and
may spawn again with another female in a few seconds. The female swims
slowly downstream a few feet, takes a position below the nest and soon
comes back to spawn again, at times with the same but usually with a
different male. One female was observed to spawn twice with the same male
in a period of one minute. A small male about three inches in total length
who at the time was the lone male on the nest, was counted spawning 20
times within 10 minutes. There were about 30 females below and over this
nest. Although a male will spawn with the female to either side, many
more righthanded matings were noted. An exceptionally vigorous male may
hold the dominant position in the nest, and spawn many times, for as long
as 20 minutes. Most males, however, drop downstream and rest in less than
half this time; thus the spawning individuals over a nest are changing
constantly.

Male shiners recognize the spawning act when they observe another
male spawn. When a breeding group has been purposely frightened away
from a nest it is usually one of the smaller male shiners that returns to the
nest first, drives away the egg predators and spawns with the first female to
come upstream. On the completion of the spawning act from 5 to 10 males
would immediately rush to the area and the never-ending battles for the
leading places in the nest would resume.

One group of breeding shiners had become conditioned so that when a
daily train rushed by at high speed only 20 yards away, spawning con-
tinued without interruption. A horned dace that was present in the nest at
the same time, however, beat a hasty retreat toward the deep pool above.
These shiners were easily frightened by any quick movement such as the
passage of a bird close above the water or the movement over the surface
of a ripple caused by a sudden gust of wind. However, they did not move
when loud vocal notes were made within six feet of them.

A common water snake, Natrix sipedon sipedon, approximately two feet
in length, once quietly moved downstream into a nest. The snake stopped
with its head near the surface and its body dragging on the bottom six
inches below. Neither the spawning shiners nor other fishes such as Rhin-
ichthys a. atratulus and Campostoma anomalum pullum appeared at all
disturbed. Several shiners swam leisurely up and casually examined the
intruder and one even investigated by touching the snake’s body with its
mouth as if testing its edibility.

An Exoglossum nest that was used by a spawning male shiner was
completely covered by the author with large flat stones. The male shiner
returned shortly and made a few dashes at adult Rhinichthys atratulus
atratulus that were eating what eggs they could get in the crevices. The
male shiner remained in the immediate vicinity for three minutes and then
dropped downstream twenty feet and attempted to drive away another male
shiner with a holding over an Exoglossum nest.

No exact observations ai'e available but it is thought that but few eggs
are laid at one time, probably not more than fifty. Occasionally a very ripe
female will lose her entire supply of eggs when she is handled and it is thus
probable that at times many more eggs are laid than at others. As has
been mentioned above the eggs fall to the bottom and lodge on and among
the pebbles of the nest. Apparently few are washed downstream. The flow
of water over a depression such as cornutus may make, or over the pit of a
Semotilus nest, is such that eddys are formed, bringing the water in the
bottom of the pit to a relatively quiet condition even in fairly fast riffles.
Thus very few eggs are washed downstream. When Exoglossum and Nocomis
nests are used the eggs are usually driven into the anterior slope of the nest
by the current.

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Other species of fishes, chiefly darters and other minnows, are quick
to dart in to eat the eggs after a spawning act is consumated. Indeed some
Rhinichthys atratulus atratulus seem to be conditioned so that they make a
mad dash to the spot at the moment spawning occurs. The male shiners are
usually successful in keeping these predators at bay.

Although some species of fishes desist from taking food during spawn-
ing, both male and female Notropis cornutus cornutus will eat during this
period. Ball (as reported by Marshall, 1939, p. 153) “found no cessation of
feeding during the breeding season” in Notropis cornutus chrysocephalus.
Ripe females lying in the middle layers of water just below the nest will
readily rise to the surface to take insects which are floating by. Males, when
not too much occupied with their territory guarding or spawning, will rise
to eat these insects.

After the spawning period many of the larger males are found dead.
Invariably they are badly fungused. However, most have probably reached
the end of the normal life span of the species at this time. Numbers of
dead specimens of other species of minnows such as Campostoma and
Nocomis are often seen just after the spawning season is over.

Eggs and Young.

The demersal eggs become adhesive when water-hardened about two
minutes after they are laid. At first they are orange but fade considerably
in a short time. The average diameter is 1.5 mm. The eggs drop between
and are washed under pebbles and some are subsequently buried by the
digging activity of the males to a depth of several millimeters in the sand.
Some single eggs are found and numbers up to twenty have also been found
in loose clumps.

The incubation period is unknown, as every effort to hatch eggs in
aquaria resulted in failure. Little definite data is available on the behavior
of the newly hatched larvae, but later, after the loss of the yolk sac, the
post-larvae may be seen in small schools at and slightly under the surface.
At this time they often school with the young of other species as Catostomus
commersonnii commersonnii, Hybognathus regius and Notropis hudsonius
hudsonius in the sluggish waters about Ithaca, New York. In smaller
streams in the same region the young cornutus are to be found in the com-
pany of the young of Rhinichthys atratulus atratulus, Clinostomus elongatus
and Catostomus commersonnii commersonnii and other species. After they
have reached a length of around 15 mm. they are still to be found in small
schools but now they are often seen in fast water as well as in pools.

Throughout the summer the young of the year tend to refrain from
associating with juveniles and adults of the same species although by Sep-
tember a few mixed collections of young and small adults have been taken.
The schools of juveniles and adults, with but few exceptions, resort to the
deeper pools during winter. However, the young are often to be found in
great numbers, associated with the young of other species, _ in shallow, ice
covered backwaters along streams. For a detailed description of the eggs
and young of Notropis cornutus chrysocephalus, the reader is referred to
Fish (1932, p. 339).

Acknowledgment is due James Kezer who read the manuscript critically
and to Ernest A. Lachner and Dwight A. Webster for their assistance in
the field.

Summary.

1. In May an inshore migration in lakes and, in some cases at least, an
upstream movement is made by Notropis cornutus.

2. Sexual dimorphism is pronounced. Breeding males have well devel-

1940]

Raney: Breeding Behavior of Notropis cornutus

11

oped pearl organs or breeding tubercles, are highly colored and reach a
larger size than breeding females. The breeding tubercles on the various
parts of the body are of value in the fighting which occurs among the males,
in driving away predators and in the breeding act.

3. Spawning occurs from May 1 through the middle of July in the north-
eastern states, beginning usually when the water has reached a temperature
of 60° to 65° F. At any one locality spawning lasts about ten days and is
limited to the daylight hours.

4. The common shiner may (1) spawn over gravel beds in running
water, (2) excavate small depressions in gravel or sand in running water
or, (3) utilize the nests built by other species such as Nocomis micropogon,
Nocomis biguttatus, Leucosomus corporalis, Semotilus atromaculatus atro-
maculatus, Exoglossum maxillingua and Campostoma anomalum whether
these are built in running or the still waters of shallow pools. They prefer
to spawn over the nests of other species when these are available.

5. Hybridization often occurs with Notropis cornutus as one parent
largely as a result of their spawning over the nests of other fishes.

6. The number of males that will spawn over one nest varies con-
siderably, there being from one hundred or more over a gravel bed to as few
as one male over a small depression. More females are present at the nests
than males.

7. Males fight continually for the leading position, the position furthest
upstream in the nest.

8. A male shiner recognizes a female and takes a semi-recumbent posi-
tion alternately from one side to the other when the female approaches
from the downstream side of the nest and comes to a position above and
slightly downstream from the male.

9. The female reacts to a fish whether it be a male common shiner or
another species providing he has taken a position over a nest, head facing
upstream and moving slightly from side to side.

10. The female takes the initiative in the breeding act by dipping down-
ward, to lie beside the male. The male throws his caudal peduncle over that
of the female, curves his body by bringing his head and tail in close
proximity with his pectoral fin underneath the head of the female. The eggs
are forced from the female at this moment while she lies, usually on her
side, her ventral surface facing upstream and her head pointing toward
shore. The entire breeding act is over in a fraction of a second and the
details cannot be clearly seen with the unaided eye.

11. Probably fewer than fifty eggs are laid at once. The demersal eggs
become adhesive after water-hardening in about two minutes after being
laid and drop between the pebbles on the bottom of the nest to which they
adhere. When first laid they are orange and average 1.5 mm. in diameter.

Bibliography.

Adams, Charles C. & Hankinson, T. L.

1928. The ecology and economics of Oneida Lake Fish. Bull. New York
State College of Forestry, Syracuse University, Vol. 1, No. 4a ( Roose-
velt Wild Life Annals, Vol. 1, Nos. 3 and 4) : 235-548.

Fish, Marie Poland

1932. Contributions to the early life histories of sixty-two species of fishes
from Lake Erie and its tributary waters. Bull. U. S. Bur. of Fisheries,
Vol. 47: 293-398.

Forbes, Stephen Alfred & Richardson, Robert Earl

1920. The fishes of Illinois. Second Edition. Nat. Hist. Surv. 111., Vol. 3:
pp. 1-cxxxi; 1-357.

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Zoologica: New York Zoological Society

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Fowler, H. W.

1909. A synopsis of the Cyprinidae of Pennsylvania. Proc. Acad, of Nat.
Sci. of Phil, for 1908 (1909) : 517-553.

1912. Some features of ornamentation in fresh water fishes. Am. Naturalist,
Vol. 46: 470-476.

Greeley, J. R.

1927. Fishes of the Genesee Region with Annotated List. In: A Biological
Survey of the Genesee River System. Suppl. 1 6th Ann. Rept. N. Y.
Cons. Dept., 1926: 47-66.

1929. Fishes of the Erie-Niagara Watershed. In: A Biological Survey of
Erie-Niagara System. Suppl. 18 th Ann. Rept. N. Y. Cons. Dept., 1928:
150-179.

1931. Fishes of the Area with Annotated List. In: A Biological Survey of
the St. Lawrence Watershed. Suppl. 20 th Ann. Rept. N. Y. Cons.
Dept., 1930: 44-94.

1932. Fishes of the Area with Annotated List. In: A Biological Survey of
the Oswegatchie and Black River Systems. Suppl. '21st Ann. Rept.
N. Y. Cons. Dept., 1931: 54-92.

1938. Fishes of the Area with Annotated List. In: A Biological Survey
of the Allegheny and Chemung Watersheds. Suppl. 21th Ann. Rept.
N. Y. Cons. Dept., 1937 : 48-73.

Greene, C. Willard

1935. The distribution of Wisconsin fishes. State of Wisconsin, Conserva-
tion Commission, Madison: 1-235.

Hankinson, T. L.

1908. A biological survey of Walnut Lake, Michigan. Mich. State Board
Geol. Surv., Rept. for 1907 (1908) : 157-288.

1920. Report on investigations of the fish of the Galien River, Berrien
County, Michigan. Occ. Papers Univ. of Mich., Mus. Zoology, No. 89:
1-14.

1932. Observations on the breeding behavior and habits of fishes in southern
Michigan. Papers Mich. Acad. Sci., Arts and Letters, Vol. 14, 1931
(1932) : 411-425.

Hubbs, Carl L. & Cooper, Gerald P.

1936. Minnows of Michigan. Bull, of Cranbrook Inst, of Science, No. 8: 1-95.

Kendall, W. C. & Goldsborough, E. L.

1908. The fishes of the Connecticut Lakes and neighboring waters with
notes on the plankton environment. U. S. Bur. Fisheries, Doc. 633:
1-77.

Marshall, Nelson

1939. Annulus formation in scales of the common shiner, Notropis cornutus
chrysocephalus (Rafinesque). Copeia, (3): 148-154.

Nurnberger, Patience Kidd

1931. Observations on the spawning temperature of Luxilus cornutus. Trans.
Am. Fish. Soc., Vol. 61: 215.

Reighard, J. E.

1910. Methods of studying the habits of fishes with an account of the
breeding habits of the horned dace. U. S. Bur. Fisheries Bull., Vol.
28, for 1908 (1910) : 1111-1136.

1915. An ecological reconnaissance of the fishes of Douglas Lake, Cheboygan
County, Michigan, in Midsummer. U. S. Bur. Fisheries Bull., Vol.
33: 215-249.

Thompson, David H.

1935. Hybridization and racial differentiation among Illinois fishes. Natural
Hist. Surv. 111. in Appendix. Vol. 20: 492-494.

1940]

Raney: Breeding Behavior of Notropis cornutus

13

Tracy, H. C.

1910. Annotated list of the fishes known to inhabit the waters of Rhode
Island. Fortieth Ann. Rep. Comm, of Inland Fisheries of Rhode
Island: 35-176.

Van Duzer, Evelyn M.

1939. Observations on the breeding habits of the cut-lips minnow, Exoglos-
sum maxillingua. Copeia, (2) : 67-75.

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Zoologica: Neiv York Zoological Society

EXPLANATION OF THE PLATES.

Plate I.

A typical breeding group of Notropis cornutus cornutus over a nest begun
by a male Semotilus atromaculatus atromaculatus at Willseyville, New York.
The large male at the right, occupying the leading position in the nest, is as-
suming a semi-recumbent position in anticipation of spawning with the small
female now dipping toward his left side. He is followed by eight males which
are easily recognized by the prominent light colored streak along the upper side.
Three small females are to be seen to the left and one is downstream from this
group of eight males. Another female appears above the third from the last
male in the nest. Several other females, that do not appear in the photograph,
were scattered further downstream. All fishes are facing upstream.

Plate II.

The beginning of a spawning act in which a small male cornutus, to the right
of the larger leading male in the nest, has thrown his body over that of the
female who now lies on her side. The breeding male is just starting to bring his
caudal peduncle toward the head in the typical breeding curve. The right
pectoral fin of the male is under the anterior end of the female. This much of
the spawning act had happened so rapidly that the other three males to his
left have apparently not noted it. The upstream male will probably turn on the
spawning male and drive him off when he finally sees him. The open mouth is
typical of a spawning male. Two female cornutus are to the left and one is
behind the male on the left side of the spawning group. Two males on the down-
stream side of the nest appear at the extreme left of the figure. Note the adult
Rhinichthys atratulus atratulus in the right foreground on the periphery of the
nest in a search for eggs.

Plate III.

The climax of the breeding act has been reached with the male cornutus in
the foreground curved over the female. Although the pectoral fin of the male is
usually under the female it is above in this case and the position of the female is
somewhat abnormal. The eggs are forced out of the female at this point. Another
female may be seen above and to the left of the leading male in the nest. The
two males near the center of the figure are in a position males assume before
attempting to dash at each other. Note that their heads and caudal fins are apart
while their bodies almost touch in the middle. The light colored tubercles of the
males stand out against the dark background of the skin. Three males may be
seen at the extreme left of the figure where the water breaks over the riffle. An
adult Rhvnichthys atratulus atratulus is present in the right foreground.

Plate IV.

The breeding act has been completed. The male cornutus is straightening
his body and the female has started to move upward. The large dominating male
to his right is turning to rush at the male that has just spawned. Three other
males are near the middle of the nest and four more appear at the left of the
figure. One small female is present just in back of the largest male near the
center of the photograph. Two other females appear in the upper right hand
corner. An adult Rhinichthys atratulus atratulus is present in the foreground.

RANEY.

PLATE I.

THE BREEDING BEHAVIOR OF THE COMMON SHINER, NOTROPIS CORNUTUS (mITCHILl).

RANEY.

PLATE II.

THE BREEDING BEHAVIOR OF THE COMMON SHINER, NOTROPIS CORNUTUS (mITCHILl)

RANEY.

PLATE III.

THE BREEDING BEHAVIOR OF THE COMMON SHINER, NOTROPIS CORNUTUS (MITCHILL

RANEY.

PLATE IV.

THE BREEDING BEHAVIOR OF THE COMMON SHINER, NOTROPIS CORNUTUS (mITCHILl).

Ginsburg : Divergence and Probability in Taxonomy

15

2.

Divergence and Probability in Taxonomy.

Isaac Ginsburg

U. S. Bureau of Fisheries, Washington, D. C.

Taxonomists of past generations have generally been content with de-
scribing and establishing species based on one or but a very few specimens.
The business of distinguishing species by this method is a comparatively
easy matter. Using very few specimens as a basis of comparison, closely
related species, in their large majority, appear to be sharply differentiated.
In occasional instances a sharp distinction on the basis of even a few speci-
mens proved troublesome, and such specimens were generally assumed to
represent “varieties,” “races,” etc., of the same species.

This easy method proved to be inadequate, as it was bound to. Later
investigators found that such distinctions all too frequently did not accord
with their material. This is due to the fact that related species often ap-
proach closely or even intergrade in their differentiating characters. From
a taxonomic point of view we do not know all we should about a species
until we know its range and manner of variability, at least in the few
crucial characters by which it is distinguished from closely related ones.
This is, of course, true of races and other subdivisions of the species. Tax-
onomists come more and more to realize this and act accordingly. In dealing
with mass data obtained in the study of variability, it is desirable to reduce
them, when it is consistent to do so, to single figures, statistical constants,
for convenience in comparison, discussion and interpretation. This paper
considers one such class of constants, that concerned with the measure of
divergence, which is of the utmost importance to taxonomists, as related
to another class, that concerned with the expression of probability, which
is often used in place of the first.

Probability in its numerical expression is often referred to as the
“test of significance.” As it is my hope that this paper will prove to be of
interest to taxonomists to whom the latter term is not a household phrase,
it may be well to consider briefly here its precise meaning.

When a taxonomist compares the likenesses of and the differences be-
tween two closely related populations — be they species, subspecies, races,
etc. — he does not study the variability of the entire population, but his
comparison is based on a relatively restricted number of specimens ; in other
words, on two samples drawn one each from the two populations. The
degree of difference or divergence shown by the two samples determines
his conclusion regarding the taxonomic rank of the two populations,
whether they are to be considered as species or as belonging to a category
of the next or second next lower rank. However, we know that different
samples drawn even from the same variable population will generally not
be the same, but, on the contrary, due solely to chance, will exhibit dif-
ferences of greater or lesser degree. The question then comes up, is the
difference shown by the two samples compared in taxonomic research a real
population difference, or is it due to the fortuities of sampling? It may
be assumed that due solely to chance it may happen sometimes that two

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samples drawn even from the same population will exhibit a difference as
great or greater than that between the two samples of the two populations
compared, and in that case it would, of course, be inappropriate to separate
the two samples taxonomically. An answer to the above question, in part,
as discussed below, is given by certain mathematical formulae developed
in connection with the theory of probability. By the use of such formulae
— based on the difference between the averages of the two samples, the
squares of the deviations of the individual specimens from the averages,
and the number of specimens in the samples — it may be determined, for
any one given character, how often such a difference, or a greater difference,
is likely to be obtained at random, by mere chance, from two samples of
the same population. If such an eventuality is likely to occur but rarely,
say, two times or less per 100 trials, we may state that the difference is
“significant” and that it may be concluded with comparative assurance that
the two samples compared in our taxonomic research belong to two distinct
populations.

The above is a bare outline of the “test of significance,” but it is hoped
that it presents its essential idea. The “test of significance” then results
in a number that represents the numerical expression of probability, indi-
cating the probable value of a difference determined in biologic research, or
the probable reliability of the two samples compared, for the purpose of
drawing pertinent conclusions. In the practical application of the formulae
an arbitrary limit is postulated and a figure obtained as a result of the
test of significance, which equals or is higher than the accepted limit, is
taken to denote “significance.” It should be noted in particular that this
test merely establishes that a determined difference is “significant.” It does
not indicate definitely whether the difference is of specific, subspecific or
racial magnitude. The taxonomic rank of the two populations compared is
determinable definitely only by some appropriate measure of divergence.

It is very important and can not be too strongly emphasized that it is
necessary to draw a sharp distinction of the fundamental difference between
the two concepts, measure of divergence and expression of probability, from
both a theoretical and a practical standpoint. This fundamental idea has
been formulated by Fisher (1936, p. 59) as follows: “It must be stressed
that the test of significance calculates a probability; it does not calculate a
racial difference.” Although it is, or should be, generally realized that a
test of probability is not the same as a measure of divergence, yet, some-
how, the two concepts become inextricably mixed in deliberation and discus-
sion. Somehow or other there appears to be a lingering idea with some
biologists that the greater the numerical value of the figures showing “sig-
nificance” obtained by the use of current formulae that express probability,
the greater the divergence between the pair of populations compared. Often
this is true; but it is only a partial truth, and like all partial truths it is
bound to lead us sadly astray. This confusion of concepts appears to be a
stumbling block not only with biologists who are not given much to the
employment of statistical formulae, but even with some who employ them
extensively.

If two separate comparisons be made of two pairs of populations, and
the test of significance have a much greater numerical value as between one
pair of populations than between the other, it does not always mean that the
former pair diverges to a greater extent; although in many cases a greater
numerical value for the test of significance does coincide with a greater de-
gree of divergence. The real meaning is that for the comparison showing a
greater numerical value, one or both samples are too large for our purpose,
for that particular pair of populations with their spread and relative regu-
larity of the frequency distributions and their difference between the means.
Smaller samples would have been sufficient to prove what we set out to dis-
cover, if our purpose was the determination of the probable mathematical
significance of the difference between the means. More specifically, when

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17

two values of significance obtained in two comparisons are above its accepted
limit, but differ widely— say, one is 10, the other 50 — the meaning is that
the samples in the latter comparison are larger than necessary for the pur-
pose of determining significance. It is evident, therefore, that figures of
different magnitudes which express mathematical probability or signifi-
cance, cannot consistently be employed for the purpose of expressing rela-
tive divergence.

A notable example of a substitution of such constants is furnished by
the “coefficient of racial likeness” which is extensively used by some physical
anthropologists as a measure of population divergence. Regarding its proper
use Morant (1923, pp. 205-207) states as follows:

“It [the coefficient of racial likeness] is not a true measure of absolute
divergence, and must not for a moment be considered as such, but neverthe-
less we shall speak of it, for convenience, as if it were an absolute measure
of racial affinity. When it is said that a low coefficient between two races
A and B indicates a closer relationship than a higher coefficient between,
say, A and C, what is meant always is that it is more probable that A and B
are random samples from the same population than that A and C are.”

This is a lucid statement of the underlying idea. The coefficient of
racial likeness is essentially an expression of probability and not a measure
of divergence. Only as an expedient make-shift is it used as a measure of
divergence. It has been extensively used as such by Morant and others.
However, a make-shift should be used only when it does not lead to false
conclusions ; but the coefficient of racial likeness often does lead to absurdly
inaccurate biological conclusions, as shown by Seltzer (1937). It would seem
to be best to abandon altogether the use of this coefficient as a measure of
divergence, and if it is still desired to employ it as an expression of prob-
ability, to change its misleading designation. (The coefficient of racial like-
ness is used primarily to combine two or more characters for the purpose of
measuring divergence. For any single character the misleading results ob-
tained by using as a measure of divergence a certain formula that funda-
mentally expresses probability, is discussed by me in another place (1938,
pp. 279-282). The problem of measuring divergence for a multiplicity of
characters I have considered in another paper (1939).)

Physical anthropologists of the school of the London Biometric Labora-
tory having become inured to the use of the coefficient of racial likeness —
which, as stated, is essentially an expression of probability — as a measure
of racial divergence, we find a similar substitution of constants employed
in still another connection. In a later paper, Morant (1936, p. 32) states as
follows: “Different characters will arrange the series in very different

orders, and it is not clear, at first, why more importance should be attached
to one of these orders than to any other ... A grading of the characters
in order of importance for the purpose in view can be obtained by noting
the number of significant differences found for each in a particular set of
comparisons.” He then lists the percentage of times, of the total number of
comparisons made, in which each one of a number of characters showed a
“significant” value for a (alpha is the chief, compound factor in the formula
for determining the coefficient of racial likeness; Morant postulates that if a
is greater than 10, it shows significance).

Now, what do we understand by an “important” character? Obviously
a character is important in distinguishing populations when it manifests a
comparatively high degree of divergence, and the opposite is true of an
unimportant character. With respect to populations of specific or lower
rank, the degree of divergence it shows is the criterion by which the im-
portance of a character may be judged. The importance of characters in
such populations may be considered from two points of view.

First, often a character may be said to be important in the sense that
it may be employed to divide a number of related populations of similar
taxonomic rank, such as a number of races within a species or a number

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of species within a genus, into two major groups. Its importance then con-
stitutes a group divergence. Such a character will show a comparatively
high divergence when a population of one group is compared with a popula-
tion of the other group, and a relatively low divergence when a pair of
populations within either group is compared, although even within the limits
of each group it will generally show different degrees of divergence to a
certain extent. In such species or genera, when the populations are divided
into pairs in all possible combinations and the pairs compared, the char-
acter, in general, will appear important in approximately half the number
of comparisons and unimportant in the remainder; providing the number of
populations in one group approaches equality to that of the other group, as
they often do.

Second, more generally, the terms “important” and “unimportant” as
applied to any given character is applicable only in connection with a given
pair of populations, and they have no universal application. A character that
may be important, that is, manifest a comparatively high degree of diver-
gence, with respect to one pair of populations may be unimportant with
respect to another pair, and vice versa. This is often true of a series of
closely related populations. In a species containing many races, or in a genus
comprising many species, that are divisible by important group characters
into primary, secondary and tertiary groups, certain other important char-
acters may crop up independently in some of the groups, and the same im-
portant character may appear in groups that are otherwise not immediately
related. Often important characters thus appear in such kaleidoscopic fash-
ion that they cannot be used consistently for the major division of the
species in a genus or the races in a species. In general, therefore, a char-
acter may be said to be important only with respect to the comparison of a
particular pair of populations.

Morant’s attempt to determine certain characters as of general impor-
tance is, therefore, rather irrelevant. However, this is beside the point.
What I am after is to point out that here also Morant uses a test of prob-
ability to express what is fundamentally a divergence.

In comparative biological research, the essential thing we are after, in
general, is to determine a difference or a divergence. This is true of both
morphological and physiological comparisons, using the latter term in a broad
sense to comprise all life processes including the complicated chain of events
connected with the reproductive process. Whether we compare the morph-
ology of pairs of related populations in taxonomic work, the comparative
yield of milk for a given breed of cattle in feeding experiments, the percept-
ible effects of a particular drug on guinea pigs or human beings in phar-
macological research as compared with controls, etc., we are trying to deter-
mine the precise divergence between two variable quantities or populations
(in cases similar to the latter, between treated and untreated individuals, or
between the same individuals before and after treatment) . This is our prime
object. A secondary consideration is the mathematical determination of the
probable reliability of the samples from a study of which the data are drawn
that form the basis of our conclusions.

This being so, it is remarkable that hitherto most attention has been
directed to the secondary consideration, the determination of probability,
while the primary object, the determination of an adequate measure of
divergence, has been rather neglected. A measure of divergence that is
universally employed is the difference between the means of the two sets of
data compared, but this is evidently not always adequate. It is certainly
altogether inadequate in taxonomic research. A fundamental defect of
measures of divergence in taxonomic research based on such values as the
mean, median, or mode, is that they represent denominate numbers which
are altogether unlike, their absolute values differing widely, in pairs of
populations that differ by widely unlike characters. Consequently, the figures
expressing the measures of divergence for different pairs of populations,

1940] Ginsburg : Divergence and Probability in Taxonomy 19

when based on denominate numbers, are not fairly comparable. That the
figures are not comparable for characters the measures of which are ex-
pressed in different units is self evident; but even when expressed in the
same unit they are often not fairly comparable, if the characters are unlike.
For instance, if the divergence of a pair of closely related populations of
mice be expressed by the difference of the means of the tail length measure-
ments, and that of a pair of populations of fishes by the same difference of
the head length measurements in the same unit, the relative divergence of
the two pairs may not be fairly comparable. Furthermore, measures of
divergence based even on the same character expressed in the same unit,
are not fairly comparable for different pairs of populations if the spread
of their distributions differ widely. An ideal measure of divergence, one
that could be used as a universal yardstick, should be an abstract number
based on the degree of overlap, positive or negative, of the two frequency
distributions, such as the measure employed by me (1938). That measure
appears to be fairly adequate for taxonomic work. Whether that measure,
or a modification of it, will be found applicable to research problems similar
to the other two mentioned above, I am not prepared to discuss. I am here
speaking chiefly from the point of view of the taxonomist. (In another paper
considering the measure of divergence with respect to a multiplicity of
characters, I (1939) concluded that a measure of divergence based on the
principal character is fairly adequate, and that if the other characters are
to be considered at all in its determination, they are to be afforded minor
weights. In any further attempt at the combination of several characters,
the figures for the different characters used should be such abstract num-
bers that measure the divergence of every one separately, rather than de-
nominate numbers that express their absolute values.)

Current formulae that are generally suitable for the determination of
the probable reliability of samples investigated in taxonomic research, have
as their fundamental bases the difference between the means of the two
samples and their probable or standard errors, the size of the samples, and
their variability as expressed by the standard deviation. The practical use
of this determination in taxonomic research is rather limited. Fisher (1936,
p. 59) states succinctly the proper, general application of the determination
of significance as follows:

“It will be seen that the test of significance does no more, and attempts
no more, than to answer the straightforward question, ‘Could these samples
have been drawn at random from the same population?’ It calculates a
probability. If the probability is very small the answer is ‘No.’ If it is not
so small as to reach the level of significance required, the answer is ‘Yes,
they could.’ The answer never is ‘Yes, they must have been.’ ”

To this I may add that in taxonomic practice, in the majority of cases,
the actual arithmetical determination is rather unnecessary. If the two
frequency distributions are fairly regular (that is, the frequencies in, the
successive classes diminish successively at both sides of the mode, even
though the distribution be skewed) as they usually are when based, respec-
tively, on homogenous material and the sampling is adequate; and further,
if the modes are at different even though closely a'djacent classes, as they
usually are when the two populations represented by the distribution really
differ and the degree of divergence is rather considerable, the arithmetical
determination of probability will usually result in a “significant” figure.
Most of the cases covered by Fisher’s first contingency may then be judged
for practical purposes by a mere inspection of the data arranged in the
form of frequency distributions.

In regard to pairs of distributions falling under Fisher’s second con-
tingency, that is, distributions showing a divergence of relatively low mag-
nitude, and the differences of which do not reach the level of mathematical
significance as determined by the samples examined ; this class of examples
will no doubt include many in which the differences are biologically signifi-

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cant. In nature, differences in taxonomic characters between pairs of pop-
ulations form a gradual series from small to large values, with virtually all
possible intermediate values. (The series may be visualized as represented
by a straight line of the equation, mx—y = 0.) Small values near the
extreme of the series must have a biological significance, although mathe-
matically their significance appears doubtful. For such populations the
arithmetical determination of probability is of no practical value by itself,
because it fails to give a definite answer to the question in which we are
interested, namely, is the difference real, even though small, or is it due to
the vicissitudes of sampling? The mathematical answer to this question,
to adapt Fisher’s style in the preceding citation, virtually is “no” or “yes,”
which is no direct answer at all. When the test for significance results in a
low numerical value, lower than the accepted limit, it may mean either one
of two things: (1) The difference is not real. (2) The difference is real
but its magnitude is such that the samples are not large enough to prove
its reality mathematically. Larger samples are necessary for a mathematical
test of significance. The meaning of too low a figure then may be similar
but opposite to what was noted above that too high a figure for significance
shows that the samples are too large; but when the figure expressing signifi-
cance is high the answer is direct and positive, and when it is too low the
answer is indirect and limited.

While some of the small but real differences that are not too extreme
will show mathematical significance when the size of the samples are greatly
increased — and theoretically any real difference, no matter how small, will
show significance by taking samples that become infinite as the differences,
in a series of pairs of populations compared, approach zero — in actual prac-
tice the size of the samples necessarily must be more or less limited. In work-
aday biological practice, therefore, it can hardly be doubted that small differ-
ences of biological significance will appear mathematically insignificant.
In passing, it may be mentioned that instances may occur in which it would
be impossible to obtain very large samples. Supposing we compare two
populations of which the actual number of living individuals is very lim-
ited, and find a small difference which, based on the entire number of living
individuals, does not show any mathematical significance. That does not
mean that such a small difference does not have a biological significance.
In general, in cases coming under Fisher’s second contingency, our conclu-
sions must be based on the biological evidence rather than on mathematical
deduction.

From the standpoint of the comparative practical unimportance of the
determination of probability in taxonomic research, it has received an undue
share of attention from certain biologists whose work is essentially taxo-
nomic, such as those investigations dealing with population or “racial”
differences in various groups of living things. In general, this is also true
of some statistical constants now in use in taxonomic work as noted below.
From the point of view of the taxonomist at least, a great deal of what is
being done along this line may be said to represent mathematical, rather
than biologic research, employing biological data for the purpose of solving
mathematical problems or formulating mathematical propositions. Of course,
mathematics represents one of the important disciplines in the sum total of
human culture, and there can be no objection for workers who are inter-
ested in mathematical research to illustrate their problems and propositions
by the use of biological data, if they wish to do so. But it should be remem-
bered that a great part of such research is of little importance in solving
taxonomic problems. In taxonomic problems, what we are greatly interested
in is to determine divergence as precisely as possible, while the determina-
tion of probability is of secondary importance. The thing to be regretted
is not so much that a great deal of attention is being paid the latter, but
that it is apparently done at the expense of the former. A few examples of
published reports will illustrate this idea.

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An outstanding, valuable and well known taxonomic work to which
reference is often made in biological discussions is that by Crampton (1916,
1925, 1932) dealing with the terrestrial gastropods of the genus Partula.
The investigation forming the basis of Crampton’s reports is unusual as
compared with taxonomic studies in general, by the number of specimens
examined and the detail with which they were examined. The information
furnished by Crampton makes it evident that Partula is at the present time
level in an early and active stage of speciation, and as such, its detailed
study is of special importance for an understanding of the process of
evolution.

The study of speciation or raciation in Partula does not lend itself
altogether readily to statistical treatment, because some of the important
distinguishing characters are rather of a qualitative nature and are not
readily expressible in terms of definite figures, although it is not altogether
impossible to do so. One important character, the direction of the spiral of
the shell, dextral or sinistral, can be expressed in terms of exact figures.
In some species it is always either sinistral or dextral ; while in other species,
or populations of lesser rank, the direction of the spiral varies with the
individual. In the latter, Crampton very helpfully gives the precise numbers
or proportion of the sinistral and dextral individuals in his samples. For
certain other characters that are measurable with more or less precision,
Crampton furnishes a wealth of statistical data and constants in tabular
form.

However, the tables published by Crampton furnish only a part of the
information that his data were in position to furnish, and that not the most
important part. For each character he generally gives the range of varia-
tion, the mean, the standard deviation and the error of the last two figures.
For some species he also gives their coefficient of variation. Now, in the
distinction of the species, or the different populations within a given species,
and in the interpretation of the relationship of the various populations, of
specific or lower rank, of what material difference, in general, is a knowl-
edge that the standard deviation, or the coefficient of variation, in one is
larger than in another population? Also, at their best such data are only
approximate, and of what material difference, in general, is a knowledge of
the small value of the error of the mean for the foregoing purposes? These
figures are interesting, but they are largely of academic interest. Of course,
there can be no objection if an author wishes to furnish such figures. What
is regretable is that more pertinent information is omitted. From a taxo-
nomic point of view we are intensely interested in how far or to what degree
the different populations diverge with respect to the various characters.
For that purpose we are presented only with the ranges and the means of
the various characters, and these are altogether inadequate. To determine
the precise extent of divergence, by some such method employed by me
(1938), frequency distributions for the different characters for the sep-
arate populations are needed and these are omitted for the characters based
on measurements.

For three characters Crampton does give frequency distributions,
namely, the direction of the spiral of the shell, the degree of tooth develop-
ment, and the color pattern. Crampton’s presentation of the data for the
last two characters is especially interesting, because they are rather quali-
tative in their nature. As such, their determination in terms of definite
figures is only approximate and dependent to some extent on a subjective
estimate. Qualitative characters are generally described by authors in
adjectival words or phrases that necessarily must be indefinite to a certain
extent, and not in terms of definite figures. Crampton shows that such
characters also can be expressed, approximately, in the form of frequency
distributions. Similarly qualitative characters in other groups as well may
be expressed in figures, and although such figures necessarily must be only
approximate at their best, they should yet prove to be of importance in

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determining divergence between closely related populations. An interesting
example of this kind is furnished also by the work of Sumner on mice of the
genus Peromyscus. Characters based on color differences, in general, are
qualitative, yet Sumner (1929) has found it possible to express quantita-
tively seven such characters, presenting his results for two of them in the
form of histograms (p. Ill), and giving the averages for the other five.

I have used Crampton’s reports as an example because they constitute
a work of unusual value and interest as compared with the ordinary run of
taxonomic papers, but the foregoing statements apply to many other pub-
lished papers m which taxonomists employ statistical methods. I may here
cite three recent papers in my own specialty, in fishes, that happened to
come to my attention, namely, by Schultz (1937), by Matsubara (1938) and
by Storey (1938). These papers are much less extensive in scope than
Crampton’s reports, in that they deal with much fewer populations. They
also differ more or less in the manner of the statistical presentation of the
data; but they illustrate in different ways some of the points raised above.

Schultz compares the Pacific with the Atlantic population of the cape-
lin. He compares a larger number of characters than usual in such cases,
but for each character he publishes only the range of variation and the mean
with its error. These figures are altogether inadequate for determining the
precise divergence between the two populations, the thing in which Schultz
as well as other taxonomists are chiefly interested. Had Schultz’s data been
published in the form of frequency distributions, they would constitute a
valuable example showing the differing degrees of intergradation of the
several characters in two closely related populations that differ by more
than one character, in addition to forming a basis for the determination of
the precise divergence between the two populations. (Schultz’s method of
combining several characters for the purpose of determining divergence
I consider in another paper (1939).)

Matsubara, working with Japanese lizardfishes, does not employ statis-
tical formulae or constants and does not calculate probabilities. Neverthe-
less, his method is essentially statistical in its nature, as it properly should
be in a problem such as the author was confronted with. However, the data
for the variability of the characters that are employed in comparing and
distinguishing his populations (which happen to be of specific rank), are
presented in graphic form and are not altogether suitable for the purpose
of calculating the extent of divergence in terms of precise figures. Of
course, the frequency distributions of the several characters may be approxi-
mately determined from the graphs, but it is very difficult or impossible to
get the exact figures. For the precise determination of divergence, it is im-
portant to have the actual frequency distributions obtained during the
investigation.

Storey, repoi’ting on an investigation of the Atlantic populations of
Harengula, also presents her data in graphic form and the same remarks
apply to hers as well as to Matsubara’s method of presentation. Further-
more, her data for characters having a continuous variation, namely, pro-
portional measurements, are presented in the form of curves “smoothed by
threes three times.” “Smoothing” has the slight advantage of producing
somewhat more regular curves which are rather more pleasing to the eye,
but it has an important disadvantage in that the curves tend to mask hetero-
geneity in the material studied. That the material of Harengula pensacolae,
for instance, possibly was heterogeneous is shown by her comparison (p. 35)
of the specimens from Sanibel with those from other localities. Storey sug-
gests that the differences in the measurements may be due to the different
preservative used, formaldehyde instead of alcohol. This may be so to a
certain extent, but part of the differences quite possibly represent a popula-
tion divergence. The difference in the gill raker count of the Sanibel speci-
mens would certainly seem to represent a population divergence. However,
in clupeid species in general, the gill raker count differs greatly with the

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23

size of the specimens and Storey does not appear to have segregated her
data in sufficiently restricted size groups to reveal any possible intraspecific
population differences in this character.

Detailed studies of other clupeid species have shown that they tend to
diversification into distinct, statistically measureable, local populations of
lesser rank, subspecies or races. It is highly probable that this is also true
of the four Atlantic species distinguished by Storey. In view of the close
approach or even general intergradation between these four species in the
characters determined, it is quite possible that if such a detailed study be
made, the relationship of the various populations will receive a modified
interpretation than that obtained by the data available to the author. The
size of the samples studied by the author were rather restricted (Storey,
1938, pp. 16-17), and in order to apply to them current statistical formulae,
the grouping of the data adopted necessarily had to be comprehensive.
A study of larger samples and of the same characters determined by the
author, measurements, gill raker count and ventral scute count, with the
data segregated by locality, and those of the measurements and gill raker
count by smaller size groups, would possibly present a somewhat different
picture of the relationship of the various populations, than that obtained
by the grouping of the data as adopted by the author. It is evident that
not only is it important— in order to determine precise divergence, distin-
guish properly the different populations and determine their relationship —
to have frequency distribution tables published, but to subdivide the data
where necessary by size, sometimes also by sex, and also by locality where
heterogeneity is suspected.

Instead of presenting detailed frequency distribution tables, Storey
gives derivatives of her data (table 3, pp. 16-17) in the form of certain
constants, the most important of which are: the standard deviation, the
mean, the difference between the means of the two populations compared
and its standard error. These are not of much value in determining diver-
gence as stated above. She also gives the relative deviate and the value of p,
which express probabilities, as they are intended by the author to do, but
are not suitable to determine divergence.

A paper based on the study of populations of flies that are of much in-
terest in connection with some phases of the species problem, was very re-
cently published by Mather & Dobzhansky (1939). It deals with the two
well known “races” of Drosophila pseudoobscura, generally designated in
the literature, following Lancefield’s suggestion in his original report (1929)
announcing their distinction, as race A and race B. The two populations
occupy different but overlapping geographic ranges; they are also incom-
pletely segregated ecologically (Dobzhansky 1937a, pp. 406-408).

The apparent principal character proving that the two populations are
distinct is a physiological one and refers to the sterility of hybrid offspring
when they are crossed. The sterility is partial, being confined to the males.
Hybrid females are fertile, at least in part. A backcross of Fi females to
males of either parent population gives rise to both sterile and fertile males.
Besides this principal character, Mather & Dobzhansky review and enumer-
ate other, minor diverging physiological characters, and differences based on
gene arrangement in the chromosones.

Morphological differences between the two populations that have so far
been discovered show certain degrees of intergradation. The object of the
paper mentioned is to deal with the morphological characters, and it takes
up five such characters, namely, the number of teeth in both the proximal
and distal sex comb on the leg of the male, the length and width of the wing
and the length of the tibia. The former two are sex characters. The latter
three characters were determined for males and females separately and
they were found to differ by sex as well as by population. All five characters
differed also according to minor populations or “strains” within each one of
the two major populations.

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Now, this problem is fundamentally taxonomic in its nature. Our con-
cern is the determination of the relative rank of the taxonomic category in
which the populations are to be placed. This determination, in its turn,
must have as its basis a determination of the relative divergence of the
populations. Given the known facts regarding the populations, let us con-
sider this particular case from the taxonomist’s viewpoint. This case may
not be as remarkable as it appears. Every careful taxonomist of wide ex-
perience no doubt can cite similar instances in which distinct populations
show relatively low and varying degrees of divergence with respect to mor-
phological differences. Its apparent remarkableness rests on the partial
sterility of the hybrid offspring correlated with a relatively low morpho-
logical divergence, and very likely is due to the fact that relative sterility
and fertility of hybrids has been definitely determined only in a rather
negligible number of very closely related populations.

In appraising the case under consideration taxonomically, it is well to
consider the relative importance of physiological and morphological criteria
in classification. There is no fundamental reason why the former should
not be used for this purpose the same as the latter. Morphological criteria
are generally used in taxonomy because they are determinable more readily
and with greater precision. In the relatively few known cases in which a
physiological character shows a greater divergence than any known mor-
phological character, the former may be employed as the principal character
in determining the taxonomic rank of the pair of populations compared.

In the case considered, the sterility criterion is evidently important,
and we may confine ourselves to a consideration of this physiological cri-
terion. The precise value of this criterion in classfieation in general cannot
be said to be as yet firmly established, and it cannot well be appraised, be-
cause it is known for relatively few populations as compared with their
untold multitude. However, in general, it is evident that this criterion is
not absolute, but, on the contrary, it is fully expressible in terms of degrees
of magnitude only. Even in regard to the classical example of hybrid
sterility, the mule, one now and then finds in the literature apparently au-
thentic records of fertile individuals. Possibly, if sterility in the mule be
investigated extensively and systematically, the percentage of fertile indi-
viduals may be found to be greater than such haphazard observations would
seem to indicate. At any rate, judged by what we already know in regard
to hybrid sterility in general, it is evident that this criterion shows all de-
grees of differences, from perfect or almost perfect sterility through dif-
ferent degrees of partial sterility to comparatively unhampered fertility,
depending on the populations crossed.

There is an incomplete correlation between relative sterility and the
relative degree of divergence of morphological characters. When an at-
tempt is made to cross two closely related populations that have reached a
sufficiently high degree of divergence, as determined by morphological cri-
teria, to be generally regarded as species, one of several things may happen:
(1) They may not be crossable. (2) They may produce zygotes showing
various degrees of inviability, that is, they die at various stages of develop-
ment, depending on the populations.' (3) When viable offspring are pro-
duced, they may be sterile or show infertility of varying and rather pro-
nounced degrees. On the other hand, when a cross is made between two
populations the divergence of which is of such a rather low degree, as de-
termined by morphological criteria, that they are generally regarded as of
a taxonomic rank below that of species, fertile offspring generally seem to
be produced. However, even with our present rather meager knowledge re-
garding sterility of hybrid offspring, it is evident that there are frequent
exceptions to the above generalizations. A pair of closely related popula-
tions, which, judged by morphological criteria, are generally regarded as
species sometimes, perhaps often, on being crossed give rise to a progeny
that is fertile.

1940]

Ginsburg: Divergence and Probability in Taxonomy

25

Since the magnitude of sterility is relative, being merely a matter of
degree, it follows that if it be used as a criterion for grading populations into
taxonomic categories, it would be necessary to draw arbitrary lines between
the species and various categories of lower rank, the same as when morpho-
logical characters, especially quantitative ones, are used for that purpose. It
is then necessary to devise a measure for expressing the degrees of sterility,
and the most obvious measure that suggests itself is the relative percentage
of sterile and fertile individuals in the hybrid progeny of the pair of popula-
tions compared. For instance, we may decide arbitrarily that if a pair of
closely related populations on being crossed produce, on the average, a
progeny 90% or more of the individuals of which are sterile, the populations
are to be designated as species; they are to be designated as subspecies
when the percentage of sterile individuals is 75 — 85%, other things being
equal; they are to be designated as races when the same percentage is
60 — 70. These are the tentative arbitrary lines which I (1938) suggested
to draw for morphological characters. For the sterility criterion even less
data are extant than for morphological criteria, to enable us to draw the
most pertinent arbitrary lines ; but wherever drawn the lines evidently must
be arbitrary. It may perhaps be found desirable to draw arbitrary lines
for the sterility criterion that differ in numerical value from those employed
for morphological criteria.

There being no absolute correlation between morphological divergence
and relative sterility, the sterility criterion, if employed in taxonomy, evi-
dently is to be used on a par with morphological criteria and coordinate with
them. Whichever is the most divergent, it is to be used as the principal
character for determining the taxonomic rank of the pair of populations
compared. If any morphological character shows a divergence of specific
magnitude the two populations are to be designated as species even though
a cross between them produces offspring that are 100% fertile. Conversely,
if the degree of sterility is greater than any morphological character that
has been discovered, the former is the chief factor to be used in determining
the taxonomic rank of the pair of populations compared.

Bearing the preceding propositions in mind, let us turn to the question
of the taxonomic rank of the two major populations of Drosophila pseu-
doobscura. It is evident that for a pertinent decision of the question we need
to know the degree of divergence of the various characters, morphological
as well as physiological. For this purpose the paper by Mather & Dobzhan-
sky furnishes only the averages of the morphological characters, which are
entirely inadequate for measuring divergence, as discussed above. From a
taxonomist’s viewpoint, the data presented educe further questions. The
authors very properly subdivide each one of the two major populations,
races A and B, into minor populations which they designate as “strains.”
In the morphological characters the differences between the extreme strains
within each race is nearly as great or greater than the differences between the
major, composite populations. The question then is, when is a population to
be designated as a “strain” and when is it to be designated as a “race”?
More specifically, from the data presented it is evident that we have two
(or more) “strains,” one in each “race,” that morphologically are approxi-
mately alike. What criterion then do they use for placing one strain in one
race, and the other, morphologically similar strain, in the other race? Al-
though I searched the paper for a definite statement in answer to this ques-
tion, it could not be found. Apparently their basic criterion is sterility, for
Dobzhansky (1937b, p. 285) states: “. . . the Fi hybrid males from crosses
between race A and race B are always sterile . . .” But other questions
present themselves: On how extensive a body of data is the above quoted
statement based? Especially, on how many crosses between different strains
is it based? Is sterility of Fi hybrid males absolute also between strains that
are alike morphologically?, since we have seen that there is a rough corre-
lation between morphological divergence and sterility. I did not deepi it

26

Zoologica: New York Zoological Society

[XXV :2

necessary for the present purpose to enter into a complete analysis and re-
view of the recorded investigations that have a bearing on an answer, if
any, to the preceeding questions. Even should sterility of the hybrid male
eventually prove not to be absolute between all the strains, it nevertheless
seems evident from the above quoted statement that its degree, in general,
is comparatively high.

Let us see now to what taxonomic conclusion we may come on the basis
of the data that have been recorded.

A consideration of the condensed data presented by Mather & Dobzhan-
sky makes it apparent that the morphological character showing the greatest
divergence between the two major populations, taken in their entirety, re-
fers to the number of teeth in the proximal sex comb of the male (with that
based on the number in the distal comb a close second). While the precise
degree of divergence between the two primary populations cannot be de-
termined from the condensed data, it seems apparent that divergence is
rather considerable but intergradation must also be pronounced. Arrang-
ing the averages in the authors’ table 2 in their order of magnitude, five
of the extreme minor populations, “strains,” of race B have averages of
5.88, 5.92, 5.96 6.00 and 6.08 respectively, nearly the same as three extreme
minor populations of race A with averages of 5.92, 5.96, and 6.00. The total
number of minor populations compared are 20 of race B and 19 of race A;
those having the averages nearly alike are, consequently, 25% of the total
“strains” of B and 16% of A, or an average of about 20%. Had the authors
given their data in the form of frequency distributions, the individuals com-
prised in the samples enumerated would apparently be seen to represent
intergrades to a large extent. In addition, it seems apparent that a con-
siderable number of individuals of the other minor populations, especially
of those populations the averages of which are next in magnitude of those
enumerated above, would also prove to be intergrades. It may be reasonably
expected then that the two major populations taken in their entirety would
show an intergradation of 25% or more in the morphological character of
greatest divergence, according to the measure of divergence suggested by
me (1938). According to the suggested arbitrary lines in the foregoing
paper, this represents a divergence of racial or nearly subspecific magnitude.
Judged by this criterion, on the basis of the incomplete data, the two popu-
lations perhaps are to be taxonomically designated as races, or they may
possibly be near the borderline between the subspecies and the race.

The other important matter to consider is sterility. At stated, for the
present it is difficult to form a judgment regarding the precise treatment of
this criterion in taxonomy in general. We must also bear in mind the fact
that in this particular case sterility is confined to one sex, the male. While
by the use of morphological criteria species are sometimes based on char-
acters of one sex, and properly so, the question remains whether the same
course is to be followed with respect to the sterility criterion. Nevertheless,
the fact of hybrid male sterility appears to be of tremendous biological
significance, especially if we assume that it is 100%, or nearly so, as the
above quotation from Dobzhansky would seem to indicate. Therefore, tak-
ing into consideration the sterility criterion, the rather considerable diver-
gence in two morphological characters, and the other differences mentioned
by Mather & Dobzhansky, the divergence would seem to be approximately of
subspecific magnitude. A taxonomist’s best and most reasonable judgment
based on the extant incomplete evidence, would then seem to dictate the
course, at least tentatively, of recognizing the two major populations as
subspecies. According to a common taxonomic practice, it may be desirable
to formally distinguish them by name, as, for instance, to designate them
Drosophila pseudoobscura2 and Drosophila lancefieldi2, employing the nota-
tion for subspecies as suggested in my (1938) paper.

The course here suggested will give taxonomic expression to the relative
divergence of some of the major populations of the genus Drosophila with

1940]

Ginsburg : Divergence and Probability in Taxonomy

27

respect to morphological criteria and the sterility criterion. Some of the
other major populations diverge in varying but pronounced degree morpho-
logically, and when crossed produce inviable or altogether sterile offspring;
they are, therefore, recognized as species. The two populations under con-
sideration diverge morphologically in lesser degree and produce partly fer-
tile offspring, and are consequently designated as subspecies.

The sterility criterion may be broadened to include the various graded
results that occur when a cross is made between two populations. The
grades are: incrossability, inviability of zygotes graded, in its turn, accord-
ing to stage of development at which they die, offspring viable but sterile
(even such populations may be only partly crossable), progeny partly fertile.
It is interesting to note that in the one genus Drosophila there appears to
be a correlation, at least partial, between hybridization results and morpho-
logy. Species that are readily separable morphologically appear to be either
not crossable or to produce inviable zygotes. D. melanogaster and D. simu-
lans, a cross of which results in viable but sterile offspring (Sturtevant
1929), also diverge morphologically in a lesser degree than some other
species, so much so that they were not distinguished until comparatively
recently (see Sturtevant, 1921, pp. 91-92). Finally, the two major popula-
tions of D. pseudoobscura1 show a comparatively low morphological diver-
gence correlated with partial fertility of hybrids.

The differences between D. miranda and D. pseudoobscura1 ( Dobzhan-
sky, 1935) seem to be rather intermediate between that of the latter two
pairs of populations mentioned. They produce viable but altogether sterile
offspring the same as the cross between D. simulans and D. melanogaster.
But morphologically the difference between them is evidently not greater
than that between “races” A and B of pseudoobscura, whereas D. simulans
and D. melanogaster are more greatly divergent morphologically, showing
one apparently discontinuous difference, that relating to the structure of
the male genitalia (compare Sturtevant, 1921, especially his figures 13-14.
p. 34, with Dobzhansky, 1935).

Dobzhansky (1935) and Dobzhansky & Tan (1937) describe important
differences in the chromosome structure between D. miranda and D. pseu-
doobscura; but, except in the peculiar distribution and number of sex
chromosomes, such differences are evidently also nothing more than a matter
of degree, since similar differences appear to exist not only between the two
major populations of pseudoobscura, but also between the “strains” or minor
populations (see Dobzhansky, 1937b, pp. 92-95). In general, hardly the sur-
face has been scratched in elucidation of differences in chromosome structure
between very closely related populations, and it cannot be used at present
as a criterion in classification with any degree of assurance.

D. azteca and D. athabascae evidently constitute another example of
two closely related populations of Drosphila, “. . . that are very similar ex-
ternally but that produce sterile Ft hybrids . . .” (Dobzhansky, 1937b, p.
113).

At least some of the major populations of the genus Drosophila have
evidently reached at the present time level a fairly advanced stage of spe-
ciation, so that they are rather easily separable and are designated as
species. Yet, even in this genus, when all major populations are considered,
there is a certain gradual transition in degrees of divergence as determined
by both morphological criteria, and physiological criteria based on hybridiz-
tion. As for the minor, intraspecific populations, it may well be expected
that various degrees of divergence will be discovered when the several
species are subjected to such taxonomic analysis as was carried out by
Mather & Dobzhansky on D. pseudoobscura x. The same state of affairs very
likely will be found to exist when we know more about the variability of
some of the numerous poorly known species, or what are now recognized as
species.

28

Zoologica: Netv York Zoological Society

[XXV :2

The taxonomic course suggested above indicates how a taxonomist
would or should form a judgment in this particular case. A decision could
be made with greater assurance, had the authors presented frequency dis-
tributions of at least the morphological character showing the greatest di-
vergence. That would have given us a basis for a determination of the
precise degrees of divergence between the minor populations and of the
divergence of the two major populations by combining the data of each one
of the two groups of minor populations, by some such method as was sug-
gested by me (1938). Instead, the authors present only derivatives of their
data in the form of means and squares of deviations for each character, and
for a combination of characters, “a score,” the latter by a method developed
by Fisher. The main results of these figures is that they lead to other, final
derivative figures expressing mathematical tests of significance of the dif-
ferences between averages by standard methods. Now, just what are the
values of the final figures in taxonomy? Do they convey any special mean-
ing or ideas to the biologist, which would help him to come to a better
decision, one made with greater assurance, in regard to the taxonomic status
of the two populations, than that based on their morphological divergence
and on the sterility criterion? The test of significance is interesting, but
it is only of minor interest in taxonomy. In this instance especially, the
definite determination of mathematical significance would seem to be of no
more than academic interest. Arranging the averages given by the authors
in their tables 2 and 4 in order of magnitude, one with some experience in
applying current statistical formulae may see at a glance, without going
through the actual arithmetical calculation, that the figures for the averages
would result in values for the test of significance that would reach its ac-
cepted limits for the principal character. Any difference in the magnitude
of the values beyond that limit does not have any special taxonomic meaning,
as noted above. To an experienced and careful taxonomist, the figures for
the averages, even condensed summaries though they are, speak much more
eloquently, they constitute a much better basis for the formation of con-
structive decisions than the figures resulting from the test of significance.
In sum, what the paper virtually accomplishes is to determine the figures
for the test of significance. But this is only a secondary part of the problem.
The main thing that we need to determine is the precise degree of diver-
gence. This question is very inadequately answered.

While the greater part of the paper deals with the test of significance
which is a matter of but minor interest in taxonomy, the subject of our
primary concern, the determination of precise divergence to serve as a
basis for forming a decision with assurance is incidentally considered, only
insofar as the given averages form a very inadequate measure for such
a determination. The valuable data determined by the authors in their in-
vestigation is not presented in such manner that the precise degree of di-
vergence could be determined. Here then is an example of an investigation
that bears the earmarks of care and reliability in its execution and carried
out by reputable investigators, but the report of which fails to furnish the
data in such form as will be of most help in deciding the question of our
chief concern. To use a favorite expression of biological statisticians, the
authors failed to extract all the information which their data were capable
of furnishing, and the most important part is omitted. They, fail to give
even the ranges of variability of the different characters. This investigation
is especially interesting in that, in a sense, it represents a study in ex-
perimental taxonomy. The groups of individuals, designated by the authors
as “strains,” upon which the data were determined, were bred in the labora-
tory from parents of known origin. It would be very interesting to compare
divergence in such populations with that of the same populations as they
occur wild ; but the data as presented permit only an inadequate comparison.

I wish to emphasize here that I am not criticizing the paper as such.
What I am after is to discuss its value in taxonomy, and the problem
with which the paper deals is primarily a taxonomic one.

1940J

Ginsburg: Divergence and Probability in Taxonomy

29

Physical anthropologists have been very assiduous in determining series
of measurements of their material in many characters, as may be gathered
by going through the volumes of Biometrica, for instance. Now, although
they apparently have different standards than other systematists by which
they determine the taxonomic rank of the populations studied by them,
physical anthropology is nothing more than a highly specialized branch of
taxonomy, dealing chiefly with one »genus, Homo, and the same methods
are applicable to this specialized branch as to taxonomy in general. In going
through the published reports on physical anthropology in Biometrica, one
finds that, omitting correlation studies, the figures on which the discus-
sions and conclusions are based generally consist of the mean, the standai'd
deviation, the coefficient of racial likeness, and their errors; sometimes the
coefficient of variation is presented. However, what we are chiefly interested
in, is the precise divergence between the populations, and those figures are
not of much value in determining that, as stated.

The published reports that are used to illustrate the foregoing dis-
cussion, represent taxonomic investigations carried out by the use of appro-
priate modern methods, statistical methods. No fault can be found with the
methods of investigation adopted by the authors. These are the methods
that are to be recommended in taxonomic research. One or another of the
investigations reported may be incomplete in one or another direction, but
the methods adopted are correct as far as they go. Nevertheless they fail
to determine the thing that is most important taxonomically, namely, the
precise degree of divergence. This is generally true of published reports of
taxonomic investigations in which modern statistical methods are employed.
It seems that, in general, workers labor under the spell of statistical for-
mulae developed through generations, and having to do chiefly with the
theory of probability, something which is generally of but secondary im-
portance in taxonomy.

What is to be especially regretted is that the class of reports under con-
sideration, with frequent fortunate exceptions, omit the necessary data,
complete frequency distribution tables, by which precise divergence could
be determined by anybody who is interested. In general, in any investiga-
tion it is the data determined that are of primary importance. The manner
of treatment of the data, their interpretation and the conclusions drawn
from them, as given by the investigator, may not be the only possible ones.
It is possible that some biologist at a later date, considering the subject
from another point of view, may wish to treat the data in a different man-
ner, and such treatment may even necessitate a different interpretation and
lead to different conclusions. However, this would be impossible unless
frequency distribution tables of the original data are presented. As it is,
valuable data forming the quintessence of any investigation, representing a
great deal of time and patient and concentrated effort, is thus practically
lost to future workers. For instance, I (1938) employed a certain measure
for determining precise divergence. This measure seems adequate in taxo-
nomic work. However, some future worker may propose a measure that is
more adequate, better expressive of the essential facts. But as most reports
are presented now, it is not possible to determine divergence by the method
employed by me, and will probably prove to be indeterminable by any future
method that may be proposed. For the method mentioned at least, frequency
distribution tables of the data are necessary, and this will likely prove to be
so in any case.

In this connection a few words may not be amiss in regard to the
economic aspect of the subject, having the editor’s point of view in mind.
Authors are sometimes confronted with the editor’s desire to abbreviate
manuscript reports by eliminating parts that appear not very essential. (It
is unfortunate that this happens even with reports of outstanding merit.)
In such cases, if elimination of some parts becomes necessary, and the ques-
tion comes up whether to eliminate frequency distribution tables or graphic

30 Zoologica: New York Zoological Society [XXV:2

representations of such tables, it is best to dispense with the latter. For,
graphic representations merely constitute a device to “catch the eye” and
clinch the author’s conclusions, and it is usually possible to represent data
graphically in more than one way, while the data on which the report is
based are quintessential, as stated. If absolutely necessary, frequency dis-
tributions should be included even at the expense of parts of the discussion.
In some cases, a report may be abbreviated even without urging from the
editor, and yet be more informative than in the form in which it finally
appears. For instance, some reports include tables giving detailed meas-
urements of individual specimens, and yet they fail to include frequency
distribution tables. But in mass data individual measurements are not of
much consequence. (They may be of some interest in the case of very ex-
treme or palpably aberrant specimens.) Such tables are rather superfluous
and not likely to be carefully perused even by specialists directly concerned.
In mass data what we are chiefly interested in, and what our conclusions
are likely to be based on, is the frequency distribution of a population with
respect to a given character under consideration.

It is fortunate for the cause of science that taxonomists are more and
more abandoning the idea that taxonomy consists chiefly of the publication
of local lists and catalogs, and descriptions of new species. This was all
right for taxonomy in its pioneering stages, and if carefully and skillfully
done, such papers still serve a useful purpose. However, gradually it is com-
ing to be realized that the backbone of taxonomy is to be found in the careful
and adequate comparison of related populations, whether they be of specific,
subspecific or racial rank, by statistical methods, for the purpose of deter-
mining the intrapopulational variability, and interpopulationally, their precise
degree of intergradation, or divergence, with each other. This forms a proper
basis for an understanding of the relationship of groups of closely related
populations. Our chief interest is to determine the precise divergence be-
tween pairs of closely related populations, or concomittantly their precise
intergradation, these two values being complementary. The determination
of probability is of but secondary importance in taxonomy, although most
attention has hitherto been given to it. In issuing reports of taxonomic in-
vestigations in which statistical methods are employed, it is essential to in-
clude frequency distribution tables of the data, so that precise divergence,
or intergradation, may be determined, by existing methods or by methods
that may be discovered in the future.

Literature Cited.

Crampton, Henry E.

1916. Studies on the variation, distribution and evolution of the genus
Partula. The species inhabiting Tahiti. Publ. Carnegie Inst. Wash-
ington, no. 228, 313 pp.

1925. The species of the Mariana Islands, Guam and Saipan. Ibid., no.
228A, 116 pp.

1932. The species inhabiting Moorea. Ibid., no. 410, 335 pp.

Dobzhansky, Theodosius

1935. Drosophila miranda, a new species. Genetics, vol. 20, pp. 377-391.

1937a. Genetic nature of species differences. Amer. Nat., vol. 71, pp. 404-420.

1937b. Genetics and the origin of species. Columbia University Press, 364 pp.

Dobzhansky, T. & Tan, C. C.

1937. Studies on hybrid sterility. III. A comparison of the gene arrange-
ment in two species, Drosophila pseudoobscura and Drosophila mir-
anda. Zeitsch. Induk. Abstam. Vererbl., bd. 72, pp. 88-114.

1940]

Ginsburg: Divergence and Probability in Taxonomy

31

Fisher, R. A.

1936. “The coefficient of racial likeness” and the futui’e of craniometry.
Jour. R. Anthrop. Inst. Great Britain and Ireland, vol. 66, pp. 57-63.

Ginsburg, Isaac

1938. Arithmetical definition of the species, subspecies and race concept,
with a proposal for a modified nomenclature. Zoologica, vol. 23,
pp. 253-286.

1939. The measure of population divergence and multiplicity of characters.
Jour. Washington Acad. Sc.i., vol. 29, pp. 317-330.

Lancefield, D. E.

1929. A genetic study of crosses of two races or physiological species of
Drosophila obscura. Zeitsch. Induk. Abstam. Vererbl. bd. 52, pp.
287-317.

Mather, K. & Dobzhansky, T.

1939. Morphological differences between the “races” of Drosophila pseudo-
obscura. Amer. Nat., vol. 73, pp. 5-25.

Matsubara, Kiyomatsu

1938. A review of the lizard-fishes of the genus Synodus found in Japan.
Jour. Imp. Fish. Inst. Tokyo, vol. 33, no. 1, pp. 1-36.

Morant, G. A.

1923. A first study of the Tibetan skull. Biometrica, vol. 14, pp. 193-260.

1936. A contribution to the physical anthropology of the Swat and Hunza
Valleys based on records collected by Sir Aurel Stein. Jour. 'R.
Anthrop. Inst. Great Britain and Ireland, vol. 66, pp. 19-42.

Schultz, Leonard P.

1937. Redescription of the capelin Mallotus catervarius (Pennant) of the
North Pacific. Proc. U. S. Nat. Mus., vol. 85, pp. 13-20.

Seltzer, Carl C.

1937. A critique of the coefficient of racial likeness. Amer. Jour. Phys.
Anthrop., vol. 23, pp. 101-109

Storey, Margaret

1938. West Indian clupeid fishes of the genus Harengula. Stanford Ichthy.
Bull., vol. 1, pp. 3-56.

Sturtevant, A. H.

1921. The North American species of Drosophila. Carnegie Inst., Wash-
ington, Publ. no. 301, 150 pp.

1929. The genetics of Drosophila simulans. Ibid., no. 399, pp. 1-62.
Sumner, Francis B.

1929. The analysis of a concrete case of intergradation between two sub-
species. Proc. Nat. Acad. Sci. U. S., vol. 15, pp. 110-120, and 481-493.

J

'

■

(

\

Conant & Doums: Miscellaneous Notes on Reptiles

33

3.

Miscellaneous Notes on the Eggs and Young of Reptiles.

Roger Conant & Alexander Downs, Jr.

Philadelphia Zoological Garden.

Everyone who is in close daily contact with an extensive collection of
living reptiles soon accumulates an assortment of notes, which, while inter-
esting in themselves, fail to fit the pattern of his contemplated papers.
Thus we have assembled miscellaneous data on some 38 clutches of eggs and
broods of young which, except as noted, were laid or hatched in the reptile
house in the Philadelphia Zoological Garden. These data, while fragmentary
in spots, we append below with the hope that like Klauber’s (1938) “odd
items of general interest,” they may prove of use. At least we are fortified
by the thought that with startlingly few exceptions the life histories of
most reptiles are so poorly known that even scraps of information about
them are welcome additions to herpetological knowledge.

The data available in the literature on this subject have come chiefly
from captive specimens. The chances of observing egg-laying or hatching
in the field are slight, and far more can be learned by maintaining gravid
females in captivity under conditions as closely approximating natural ones
as possible. Professional curators have contributed much information but
here is a big field for the amateur herpetologist to make useful observa-
tions. All he needs is an ability to record data accurately, an inexpensive
meter stick, and a friendly corner druggist who does not mind having his
scales used in the interests of science.

Our weights and measurements were recorded as soon after laying or
hatching as was convenient — usually on the same or the following day. Eggs
were incubated in aquaria or glass jars nearly filled with moist, decayed
wood. A flat piece of glass placed on top of each vessel served as a lid and
permitted control of the moisture content within each one. When the under-
surface of the glass “sweated” appreciably the lid was moved slightly to
one side to allow evaporation. Conversely, when the hatching media became
too dry it was sprinkled with water and the lid was replaced. Ordinary
room temperatures prevailed during the incubation periods.

The weights and measurements of all female specimens were recorded
after their eggs were laid. All weights are recorded in grams and all
measurements in millimeters.

The letters “H” and “L” indicate the highest and lowest numbers, re-
spectively, in each column of figures, thus representing the extremes of
variation in each group. Unusually abnormal specimens are excepted. In
cases where the highest and lowest figures appear more than once, only the
first is designated by a letter.

For help in taking weights and measurements and in securing other
data we are indebted to Mark Mooney, Jr., and to Edmond Malnate, succes-
sive superintendents of the reptile collection in the Philadelphia Zoological
Garden. Several members of the Junior Zoological Society of Philadelphia
also have given valuable assistance.

[XXV :3

'34 ’ Zoological: NeiO York Zoological Society

Alligator mississipiensis (Daudin).

A number of alligators, apparently recently hatched, were discovered
along the swampy margins of a large pool on St. Simon’s Island, Georgia,
about August 20, 1937, by Robert S. Ingersoll, Jr. There were about 36
specimens in the group, but not all of them were captured. Measurements
and weights were recorded when the alligators arrived at the Philadelphia
Zoological Garden a few days later.

No.

Length

Weight

No.

Length Weight

1

241

53.2

11

243

46.5

2

233

50.9

12

244

52.6

3

249

51.9

13

251

52.5

4

252

56.4— H

14

247

51.7

5

231— L

42.9

15

257— H

52.6

6

247

53.2

16

247

52.5

7

247

47.0

17

244

51.2

8

249

53.2

18

236

42.7 — L

9

237

48.7

19

232

47.6

10

241

43.0

Average

243.58

50.02

Basiliscus vittatus (Gray).

Eggs found scattered about on the floor of the

cage on June 29, 1937;

all separate from one another. Shells buff, ovoid and rather soft.

Female

and male

in captivity since March 20, 1935. Both

were confined

in same

cage but

eggs were infertile. The shell of egg No. 10 was badly dented.

No.

Length

Width

Weight

1

18.7— H

10.9

1.40

2

17.7

11.1

1.31

3

18.3

11.7

1.60— H

4

17.4

12.6— H

1.50

5

18.3

10.5

1.25

6

16.9

10.8

1.40

7

17.0

10.4

1.09

8

16.2

11.3

1.05

9

17.8

10.5

1.25

10

15.0

10.1

.65

11

15.6— L

11.5

1.10

12

16.2

10.0— L

.91

13

16.4

12.0

1.10

14

15.9

11.1

.90— L

Average

16.96

11.04

1.18

Sceloporus undulatus fasciatus (Green).

Group of eggs brought to the Philadelphia Zoo about August 1, 1938.
Donor could not furnish the locality where they were found, but stated it
was in southern New Jersey.

Total Length

No.

Length

( snout to vent)

Weight

1

45— L

22— L

0.4

2

46

22

0.4

3

49— H

23— H

0.3

4

45

22

0.3

Average

46.25

22.25

0.35

Egg No. 1 hatched on August 8, the others on August 11.

1940]

Coimnt & Downs: Miscellaneous Notes on Reptiles

35

Ophisaurus apoclus (Pallas).

Eggs found on July 20, 1937, in a cage occupied by several large, adult
specimens received on June 9, 1937, from the Zoological Society of London.
Probably the two eggs did not constitute the entire complement from any
one female, but no others were laid.

No.

Length

Width

Weight

1

33.4

18.7

6.45

2

33.2

18.4

6.00

Average

33.3

18.55

6.23

Farancia abacura abacura (Holbrook),

A large female of this subspecies was found about August 20, 1937,
coiled around her eggs under a large log on St. Simon’s Island, Georgia,
by Robert S. Ingersoll, Jr. There were approximately 15 or 20 eggs but un-
fortunately some of them were destroyed in transit to Philadelphia and the
exact number could not be determined. Two of the eggs were hatched on

October 9, 1937.

, .

No.

Length

Weight

1

204 (

4.8

2

231

6.2

Average

217.5 !

5.5

Specimen No. 1 was deformed, its tail being kinked iii an S-shape.

Opheodrys aestivus (Linnaeus).

A female specimen. (collected at Royal Oak, Talbot County, Maryland,
on July 2, 1938, was kept, alive at Treasure Island Boy Scout Camp, where it
deposited 9 eggs on July 13. The , eggs subsequently were given to the
Philadelphia Zoological Garden, where they began hatching on September
9, 1938.

A To.

Length

Weight '

1

183 v

1,3

■2

201 — II

L4 — H

'3

190

1.2

,■4

182 =

,, .. 1.1— L

5

191

1.3

6

180— L

1.4

7

192

1.3

8

187

1.2

9

190

1.2

Average

188.4

1.27

When the eggs first were discovered to be hatching one snake had
completely escaped from the shell, another had its head half-way out of the
egg and a third had the tip of its nose exposed. Six young had escaped by
the morning of September 10; the numbers of slits made by these snakes
in their respective egg shells were 5, 3, 2, 2, 2 and 1. All the young were very
timid, drawing their heads back into the shells when an observer approached,
or hiding in the hatching media after they had left the eggs. The last
snakes escaped from their eggs on September 11. On this date the eyes of
all were overcast, preparatory to shedding. The young were uniformly
greenish-gray in coloration at first, but changed to uniform light green
upon shedding their skins.

36

[XXV :3

Zoologica: New York Zoological Society

Coluber constrictor constrictor Linnaeus.

A clutch of eggs of this species was found buried in the earth by H.
Leschke at Haddonfield, N. J., on June 19, 1936. They were stained brown,
and salt-like protuberances on the shells made them rough to the touch.

No.

Length

Width

Weight

1

28.0

22.4

8.1

2

26.8— L

22.9

7.8

O

O

32.3

20.4— L

7.4

4

31.0

22.0

8.3

5

29.8

22.1

8.0

6

27.3

23.2

7.8

7

27.5

21.7

7.1— L

8

29.5

21.9

8.3

9

27.0

23.7— H

8.3

10

29.5

22.0

7 8

11

29.0

22.4

8.1

12

28.2

21.8

7.3

13

29.7

20.8

7.9

14

31.0

21.7

8.4

15

27.7

22.8

7.2

16

28.5

22.4

7.7

17

33.7— H

21.4

8.5

18

29.8

20.5

7.3

19

31.9

22.0

8.7— H

20

30.0

21.5

7.6

Average

29.41

21.98

7.88

Twelve of these eggs hatched between August 7 and August 12, 1936.

No.

1

2

3

4

5

6

7

8
9

10

11

12

Length

278
288
275
272

279
282
256

253— L
265

305— H

294

284

Weight

5.2

5.2

6.1

4.6

5.8

6.0

4.5— L

4.5
5.0

62— H

5.6
5.8

277.5

5.38

Average

1940]

Conant & Downs: Miscellaneous Notes on Reptiles

37

A group of 15 eggs of the same species was laid on June 6, 1938, in a
c,age containing several females, all caught a short time before near Phila-
delphia. Determination of which adult laid the eggs was impossible, but
it was presumed that the entire lot was deposited by one female. All the
eggs were white, the shells had salt-like protuberances on them and all
except No. 11 were approximately the same size.

No.

Length

Width

Weight

1

35.6

18.2

7.5

2

36.3

18.6

8.0

3

32.1

20.0— H

7.9

4

33.7

19.4

7.7

5

36.7

19.2

7.8

6

33.6

19.6

8.3

7

34.1

19.8

7.8

8

37.6

18.4

7.3— L

9

31.7— L

19.4

7.5

10

34.9

18.6

8.0

11

46.5— H

17.4— L

8.3

12

35.0

19.4

8.1

13

35.5

19.7

8.6— H

14

37.4

18.9

7.9

15

33.6

19.5

—

Average

35.62

19.07

7.9 (of 14)

Eggs No. 14 and No. 15 each had one end prolonged into a tip, and
No. 15 was broken and leaking when found.

Four of the above eggs hatched in August, 1938, three on the 13th and
the fourth on the 15th.

No.

Length

Weight

1

309

6.3

2

294— L

5.95— L

3

336— H

6.8— H

4

303

6.4

Average

310.5

6.36

Elaphe obsoleta obsoleta (.Say).

A female, probably fx-om near Philadelphia, deposited 11 eggs in cap-
tivity on July 24, 1937. Five of these were separate but the others were
adherent in pairs. Very probably the snake was disturbed during ovoposi-
tion ; several specimens of other species were confined in the same cage
with it.

No.

Length

Width

Weight

1

50.8

20.2

13.7

2

55.5— H

19.7

14.95

3

43.8

22.4

12.23

4

48.0

18.2

10.15

5

44.9

17.4— L

9.15

6

47.2)1

21.7 ) 1

27.8

7

46.3)

23.7)— H

8

47.7 (

22.5 \

26.7

9

45.5)

22.3)

10

45.6 1

22.5 (

25.8

11

43.5)— L

22.6)

Average

47.16

21.2

12.77

Egg No. 4 had a marked constriction around its middle.

i Eggs which were adherent to each other. The measurements of eggs in such condition are
grouped by brackets and their combined weights are expressed by one number, a policy followed
throughout the text.

38

Zoologica: New York Zoological Society

[XXV :3

Two of the above eggs hatched on October 9, 1937.

No. Length

Weight

1 361

10.3

2 360

11.2

Average 360.5

10.75

Elaphe vulpina (Baird &

Girard)

Three large females collected by Robert H. Mattlin in May, 1938, at

Little Cedar Point, Lucas

County,

Ohio, all deposited eggs during July,

1938. In two instances, many of the eggs were

adherent, making it im-

possible to measure them and necessitating the weighing of them in groups

as indicated in the tables below. In only one case

was it possible to deter-

mine which female laid which eggs.

Her weight and length are recorded in

their proper place.

I. Eggs laid on July 20, 1938.

No.

Length

Width

Weight

1

431

241

2

43|

251

3

39 1

25 1

4

42 1'

25 1

5

42 1

|

6

43)

l

161.7

7

39|

— [

8

46 1

24 1

9

40

1

10

41 i

241

11

42 J

24 J

12

36— L

24

13.8

13

44

24

16.1

14

47— H

27

14.9

Average

41.93

24.6 (of 10) 14.75

II. Eggs laid on July

21, 1938.

Length of female, 1229 mm.; weight,

409.5 g.

No.

Length

Width

Weight

1

47.3

25.7

18.9

2

47.0

24.1

17.3

3

48.2

24.0

18.7

4

50.4

24.5

18.5

5

48.0

25.8— H

19.3

6

42.9

23.7

14.8

7

38.11— L

22.31— L

8

48.9|

' 24.01

9

51.2)

24.1)

84.8

10

44.81

24.2|

11

58.5 1— H

23.3 J

Average

47.75

24.15

17.48

Five of the eggs in this group hatched in September— three on the 1st
and two on the 5th.

No. Length Weight

1 306 14.4

2 334— H 15.4— H

3 333 14.0

4 300— L 11.9— L

5 331 14.3

Average

320.8

14.0

1940]

Conant & Downs: Miscellaneous Notes on Reptiles

39

III. Eggs laid on July 23, 1938. All were separate, probably because
the female moved about while depositing them. The moist substance with
which they were covered when laid was sufficiently adhesive to make pebbles
from the cage floor stick to several of the eggs.

No.

Length

Width

Weight

1

39.9

26.9

16.65

2

40.7

24.5

14.4

3

40.7

26.1

15.4

4

37.8

27.3—

-H 16.2

5

44.4

23.5

14.4

6

29.1— L

25.0

14.7

7

45.0

22.9—

-L 15.0

8

47.2— H

25.0

16.5

9

41.8

24.4

14.8 ■ i

10

40.4

24.4

14.3

11

40.6

24.6

14.9

12

38.4

24.3

• 14.1

13

42.9

27.3

18.5

14

45.7

26.9

19.8— H

15

42.8

24.3

14.5

16

39.9

26.1

15.8

17

43.1

27.2

13.9

18

41.9

26.3

18.1

19

35.7

25.5

13.6

20

38.1

24.6

13.4— L

21

38.6

26.3

15.1

Average 40.7

25.38

15.43

Pituophis melanoleucus melanoleucus (Daudin) .

Two females from undetermined localities

in southern New Jersey laid

eggs in July, 1938.

Many of the eggs in each

group were adherent to one

another, preventing complete measurements and separate weighings.

I. Eggs laid on

July 12, 1938.

Length of female, 1514 mm.; weight,

952.3 g.

No.

Length

Width

Weight

1

531

341

2

49 1

35 1

3

— 1

33 1

4

5

48|

49|

35 1

34 r

285.85

6

53|

7

-1

34 1

8

61 1

321

9

49

33

32.3

Average 51.71 (of 7) 33.75 (of 8) 35.35

Two of the above eggs hatched on September 19, 1938.

No.

Length

W eight

1

412

35.8

2

376

26.8

Average

394.0

31.3

40

Zoologica: New York Zoological Society

[XXV :3

II. Eggs laid on July 13, 1938.

No.

Length

Width

Weight

1

571

331

2

3

— l
58 f

321
33 f

155.0

4

59)

341— H

5

60)

32)

76.4

6

57)

33)

7

66

31— L

39.9

Average

59.5 (of 6)

32.57

38.76

The eggs in this group started to hatch on September 18, and the last
of the 7 emerged on the 20th. There were as many as 5 and 6 slits in some
of the eggs and in one or two of them 2 or more slits crossed, to form loose
flaps.

No.

Length

Weight

1

433

36.5— H

2

452— H

36.5

3

416— L

34.5— L

4

433

35.4

5

416

35.0

6

429

35.0

7

437

36.1

Average

430.86

35.57

The young snakes might be described as follows:

Ground color (above) yellowish-cream, becoming orange or brownish-
cream on the sides. Dorsal blotches very dark brown (almost black), the
most anterior dorsal ones enclosing two rather small, light areas. The
orange or brownish coloration on the sides is caused by the anterior parts
of the scales being of these colors. Some scales also have a longitudinal
streak of the same colors on them.

Top of head olive-gray, marked with a few dark spots. A dark line
across in front of the eyes, situated on the anterior edges of the frontal and
supraoculars and the posterior edges of the prefrontals. A short, dark dash,
obliquely downward from the back of the eye; another straight down from
the eye. Some of the sutures between the upper labials dark. Belly whitish,
washed with orange-pink.

Lampropeltis getulus florklana Blanchard.

Female collected 20 miles west of Miami, Florida, and sent to the Phila-
delphia Zoological Garden through the courtesy of Dr. Thomas Barbour.
Eggs were deposited on May 19, 1936. Length of female, 1047 mm.; weight,

282.2 g.

No.

Length

Width

Weight

1

47.7

22.0— L

13.4

2

50.0— H

22.6

13.8— H

3

36.8

23.8— H

11.2

4

38.9

22.9

11.0

5

43.2

23.0

13.2

6

33.0— L

23.4

9.8— L

7

38.9

23.2

12.3

8

40.4

22.9

12.4

9

38.6

23.6

12.0

10

47.0

23.0

13.6

11

44.3

22.8

13.4

Average

41.70

23.01

12.37

1940]

Conant & Downs: Miscellaneous Notes on Reptiles

41

Egg No. 2 had a constricted nipple at one end and No. 6 had a flesh-
colored area covering about one-third of the egg. All other eggs were plain
cream white, with smooth, parchment-like shells. Numbers 7 to 11, inclu-
sive, were adherent in a single cluster but wei’e separated for weighing and
measuring. Several young had escaped from their shells when the clutch
was examined on July 26, 1936, and the last emerged on July 27. Two eggs

were not fertile.

No.

Length

Weight

1

290

11.1

2

285

9.4—

3

289

10.5

4

277— L

10.8

5

300— H

11.1

6

282

11.0

7

295

12.3—

Average (of 7)

288.2

10.89

8

232

7.8

9

216

5.5

Average (of 9)

274

9.93

The last two specimens obviously were runts. In addition their sides
were somewhat shrunken except in the region of the belly, which was
noticeably swollen. They lived only a few days. The other specimens began
eating DeKay’s snakes, Storeria dekayi, and one another almost at once;
eventually only one specimen remained.

Rhinocheilus lecontei Baird & Girard.

Eggs were found in the cage on July 1, 1935. The female was purchased
from a dealer and its origin is unknown.

No.

Length

Width

Weight

1

37.0

15.9

7.1

2

36.2

16.0— H

6.7

3

37.2

16.0

6.6

4

41.0— H

15.4

7.2—

5

35.6

15.8

6.5

6

30.1— L

15.3— L

5.3—'

Average

36.18

15.73

6.57

42

Zoologica: New York Zoological Society

[XXV :3

Natrix cyclop-ion floridana Goff.

Two specimens from Marion County, Florida, bore young in captivity.
I. Young born on July 27, 1936.

No.

Length

Weight

1

249— H

8.3— H

2

248

7.0

3

229

6.3

4

230

5.9— L

5

240

7.5

6

229

6.2

7

245

7.0

8

248

7.0

9

234

6.9

10

233

7.1

11

225

6.6

12

243

8.1

13

244

7.9

14

222— L

6.0

Average

237.07

6.99

This litter probably consisted of more than 14 specimens since some
had escaped from the cage and were found on the floor. Number 4 had its
hemipenes everted; No. 14 was dead. The pattern of the young snakes was
more conspicuous than that of adults; it consisted of black reticulations on
an olive-brown ground. Their bellies were plain yellow except for the antero-
lateral edges of the ventrals, which were black.

II. Young born on August 7, 1936. Length of female, 1389 mm.;
weight, 686.6 g.

No.

Length

Weight

1

245

9.0

2

259— H

9.2

3

241

7.6— L

4

257

9.1

5

250

8.9

6

244

8.4

7

245

8.0

8

240— L

8.7

9

246

8.7

10

251

9.6— H

Average

247.8

8.72

1940]

Conant & Downs: Miscellaneous Notes on Reptiles

48

Matrix erythrog aster transversa (Hallowell) .

A litter was born on September 5, 1933, to a female collected in Kansas
and presented to the Toledo Zoological Park by Dr. Reeve M. Bailey. The
parent measured 1071 mm. and weighed 335.1 g.

No.

Length •

Weight

1

255

6.0

2

267

5.9

3

268

6.4— B

4

256

S.l

5

250— L

5.6

6

267

6.0

7

262

8.1

8

268

6.4

9

257

6.2

10

2 70

6.2

11

264

6.1

12

261

6.1

13

266

6.3

14

267

6.3

15

258

5.8

16

255

6.0

17

265

5.9

18

271— H

6.3

19

271

5.7

20

265

5.5— L

Average

263.15

6.05

An additional specimen (not recorded) was born dead. The living young
might be described as follows :

Pattern (above) consists of a dorsal and lateral series of blotches
alternating clear forward to the head. In the center of each light area, on
the sides between the black blotches, is a large area of rose-brown, narrowly
edged with white. Belly uniform white except for the antero-lateral edges
of the ventrals, which are black.

These small snakes began eating chopped fish almost at once. As they
grew they were measured and weighed at intervals. On December 9, 1933,
they averaged 259.3 mm. and 6.61 g. ; on April 5, 1934, 19 of them (one
having died) averaged 306.8 mm. and 10.90 g. ; on June 13, 1934, 331.6 mm.
and 13.33 g. All but 3 had died by May 2, 1935, when the survivors aver-
aged 420 mm. and 26.70 g. These snakes did not hibernate; they were kept
warm and active over both winters.

Several young of this subspecies were born at the Toledo Zoological
Park during September, 1930, to a female from New Braunfels, Texas. They
grew rapidly in captivity on a diet of fish, but as a result of cannibalism the
group eventually was reduced to one. This, a female, passed a large, red, in-
fertile ovum on September 2, 1933, and two others on September 4, 1933.
Thus it would appear that the snake was sexually mature at the age of three
years. At that time it weighed 429.7 g. and measured 960 mm. in length.
While it was still light in color, its pattern had become rather indistinct
and the contrast between blotches and ground color was considerably less-
ened. The light crossbands between the dorsal blotches were the most
prominent feature of the pattern.

44

Zoologica: New York Zoological Society

[XXV: 3

Natrix septemvittata (Say).

Two females of this species from Detroit, Michigan, gave birth to young
on August 23, 1938. Since both females were in the same cage it was im-
possible to determine how many each had borne.

No.

Length

Weight

1

212

2.8

2

221

2.6

3

222

3.3— H

4

215

2.6

5

222

3.2

6

223

3.3

7

227

3.1

8

226

3.2

9

225

2.8

10

218

2.7

11

216

2.6

12

206— L

2.3— L

13

230— H

3.1

Average

220.2

2.89

Another female from Delaware County, Pennsylvania, bore 2 young
and passed 2 dead embryos and 2 infertile ova on August 22, 1938. The
female weighed 52.9 g. and measured 688 mm. Data on the 2 living young
are:

No. Length Weight

1 168 1.4

2 206 2.5

Average 187 1.95

No. 1 was deformed.

1940]

Conant & Downs: Miscellaneous Notes on Reptiles

45

Natrix sipedon pictiventris Cope.

Young were born on August 2, 1936, to a specimen from near Palmetto,
Florida, collected by the late C. C. Goff. The female weighed 368.9 g. and
measured 771 mm. in length. Most of its tail was missing.

No.

Length

Weight

1

211

3.3

2

213

3.4

3

194

3.2

4

215

3.5

5

206

3.5

6

208

4.0

7

217

3.9

8

213

4.2

9

215

3.7

10

180— L

2.9— L

11

210

4.3

12

211

4.0

13

199

3.7

14

205

3.4

15

223— H

3.8

16

217

4.2

17

201

4.0

18

200

3.9

19

206

4.4— H

20

199

3.7

21

221

3.7

22

192

3.2

23

206

3.7

24

204

3.5

25

218

4.1

Average

207.36

3.73

No. 24 and No. 25 were dead.

[XXV: 3

46 Zoologica: New York Zoological Society

Thamnophis sirtalis sirt'alis (Linnaeus).

A specimen from Philadelphia, Pennsylvania. Young born on August 4,
1936. Length of female. 706 mm.; weight, 92.2 g.

No.

Length

Weight

1

143

1.1

2

142

0 9—1.

3

156

1.1

4

155

1.0

5

156

1.2— H

6

155

1.0

7

162

1.2

8

164

1.2

9

155

1.1

10

157

0.9

11

147

1.1

12

149

1.0

13

162

1.1

14

153

1.1

15

164

1.2

16

158

1.0

17

150

1.1

18

165

1.2

19

161

1.1

20

160

1.1

21

168

1.0

22

170— H

1.1

23

158

1.0

24

166

1.1

25

152

1.0

26

160

1.2

27

158

1.1

28

157

1.1

29

149

1.0

30

155

1.0

31

140— L

0.9

Average

156.35

1.07

No. 12 had a deformed back. No. 31 was dead.

Crotalus viridis viridis Rafinesque.

Young born to two females received from South Dakota, collected by
A. M. Jackley in August, 1938.

I. Born on September 7, 1938. Parent weighed 297.4 g. and measured

mm. in length, to the base of

the rattle.

No.

Length

Weight

1

281

14.9

2

282

13.8— L

3

286

15.1

4

290— H

15.2

5

273

14.2

6

274

14.3

X

251— L

14.1

8

259

14.2

9

270

14.8

10

279

15.6— H

11

277

15.5

Average

274.73

14.7

The eyes of all were overcast, indicative of an approaching moult.

1940] Conani & Downs: Miscellaneous Notes on Reptiles 47

II. Young- born on September 10, 1938; two dead embryos also passed.
Female undertermined; several others in cage.

No.

Length

Weight

1

256

14.8

2

271— H

15.6— H

9

O

237— L

11.6— L

4

249

12.8

5

253

12.8

Average

253.2

13.52

The “button” was included in measuring the young of both litters.

Sternotherus odoratus (Latreille).

A turtle of this species was collected early in June, 1936, by Byron
Gardner, Jr., who found it in South Carolina directly across the Savannah
River from Augusta, Georgia. It deposited 2 eggs on June 22.

No.

Length

Width

Weight

1

25

15.1

3.6

2

23.2

15.4

3.5

Average

24.1

15.25

3.55

On October 2, 1936, a baby was found walking about in the hatching
media. That it may have emerged at some time previous is indicated by the
fact that the other egg also had hatched, but the young had died and was
badly dessicated. Data on the living one are as follows:

■'} r Carapace Depth of Plastron

Length Width Shell Length Weight

22 18.9 12 15.8 2.2

Chelydra serpentina serpentina (Linnaeus).

A large snapper collected 3 miles north of New Castle, Delaware, on
April 24, 1936, deposited 3 eggs in its tank on June 22, 1936.

No. Diameter Weight

1 27.2 10.8— H

2 26.8 — L i. 9.7— L

3 29.5— H 10.7

Average 27.83 10.4

Since the nature of the tank prevented this turtle from nesting, the 3
eggs doubtless represent only a fraction of the entire complement. The
specimen was transferred to a spacious, out-door enclosure but whether
other eggs were laid was not determined.

48

Zoolog tea: New York Zoological Society

Terrapene Carolina Carolina (Linnaeus).

A set of eggs probably found near Philadelphia, was given to the Phila-
delphia Zoological Garden by an anonymous donor. Young hatched on Sep-
tember 1 and 2, 1938.

Carapace

No.

Length

Width

Depth of Shell

Weight

1

34.6

33.3

16.6—1,

7.9

2

33.3— L

32.1— L

16.7

7.8

3

34.9— H

33.5

17.0— H

8.4—1

4

33.7

33. 7— H

16.8

7.6—

Average

34.13

33.15

16.78

7.93

Pseudemys floridana ssp.

A female from central Florida laid 2 eggs in its tank on June 7, 1936.

No.

Length

Width

Weigh t

1

33.0

24.3

9.2

2

35.7

24.5

8.7

Average

34.35

24.4

8.95

As with the specimen of Chelydra s. serpentina above, conditions were
not optimum for laying and the 2 eggs doubtless are but a part of the
number the turtle might have been expected to lay.

Testudo tornieri Siebenrock.

Four of these tortoises were secured from a dealer on August 7, 1935.
One of them, weighing 375.9 g., carapace length 144 mm., carapace width
106 mm., and depth of shell 38 mm., deposited an egg on January 9, 1937.
This weighed 33.5 g. and measured 48 mm. in length by 28 mm. in width.
As Loveridge (1928, p. '51 ) has noted, this remarkable turtle has a com-
plement of only a single egg.

References.

Klauber, L. M.

1938. Notes from a Herpetological Diary, I. Copeia, No. 4, pp. 191-7.
Loveridge, Arthur

1928. Field Notes on Vertebrates Collected by the Smithsonian-Chrysler
East African Expedition of 1926. Proc. U. S. Nat. Mus., Vol. 73, Art.
17, pp. 1-69, pis. 1-4.

Jaros: Occlusion of Venom Duct

49

4,

Occlusion of the Venom Duct of Crotalidae by Electrocoagulation:
an Innovation in Operative Technique.1

Duval B. Jaros

Riverside, Illinois.

(Text-figure 1).

An article by Tait2 on the surgical removal of the venom glands of
rattlesnakes prompted Mr. Emil Rokosky of the Chicago Zoological Park to
suggest the use of electrocoagulation on the venom gland. It was thought,
however, that occlusion of the duct would be a less drastic procedure than
removal of the gland. In addition, conservation of the gland would permit
interesting experiments and histologic studies. The immediate problem pre-
sented was the development of the technical details of a rapid operation
which would occlude the duct, preserve the gland and result in minimum
injury to the snake. In the procedure here outlined, electrocoagulation is
employed.

Electrocoagulation produces destruction of tissue by heat, and corres-
ponds to the solidifying of the white of an egg during boiling. The heat of
electrocoagulation, however, is not conducted but is induced within the tissue
itself, which insures coagulation without even surface carbonization. The
coagulum is sterile and simulates a scab, nature’s own wound covering.
Sterility, firmness and pliability of the coagulum permit healing without
scarring.

Essentials of an electrocoagulating device are two electrodes and a high
frequency current. The electrodes may be in the form of a single duo-
terminal instrument, a body-plate and a point, or as in the present experi-
ment, two separate uniterminal instruments with one serving also as the
probe for securing the duct.

The high frequency current alternates at about one million cycles per
second. Suitable currents can be obtained from simple spark-gap or ther-
mionic tube apparatuses sold by all physiotherapy equipment dealers. The
coagulating current of 800 kilocycles, 150 milliamperes and 2000 volts, seals
the severed ends of the duct, and is therefore preferable to the cutting
current.

In the operation, several innovations employed made general anaesthesia
unnecessary and facilitated single-handed performance with relative safety
to the operator and the snake. The first device, an operating board (Text-
fig. 1) was designed to hold the snake firmly and prevent it from thrashing.
The board was fitted with a neck pad, strap and a thin rubber restraining
flap for the body. When in use, this flap was drawn over the snake and at-
tached along the opposite edge of the board by a zipper running its entire

1 Presented at a meeting of the Amateur Herpetologist Group of the Chicago Academy of
Sciences, June 15, 1939, and at the annual meeting of the American Society of Ichthyologists and
Herpetologists, Chicago, September 14, 1939.

2 Copeia, 1938:1.

50

Zoologica: New York Zoological Society

[XXV :4

length. Pressed down by the restraining sheet which yields to its struggles,
the snake cannot crawl. This operative convenience can be augmented by
having the surface of the board on which the snake rests very smooth or
covered with yielding rubber similar to the overlying restraining sheet.

A second aid was the surgical skin clip or Michel clip. These clips, small
strips of metal clawed at each end and preferably bent into a V shape, when
pressed point to point, firmly grasp any interposed tissue. Applied to the
snake’s lips, they hold the snake’s mouth shut and make handling rela-
tively safe without the use of general anaesthesia.

Operating board designed to hold a snake firmly while the venom duct is
occluded.

Another innovation was an improvised scalpel which proved, for my
purpose, to be superior to any other obtainable. These scalpels were made
from slivers of the cutting edges of Gillette “Thin Blades,” broken off with
pliers and clamped in ordinary artery forceps. With the forceps serving
as handles, these tiny bayonettes afforded a generous supply of extremely
sharp, delicate and inexpensive knives.

The Operation.

The snake, grasped behind the head, was fastened to the operating
board, and Michel clips were clamped on the lips ; one on each side below the
pit. Care was taken not to include the mandible.

At this time, if desired, a local anaesthetic may be injected at the site
of the incision. A 2% procain borate solution was used. Almost imme-
diately after the injection a quarter-inch incision was made just below the
eye and on a level with the pit. Through this small incision the duct was
secured and drawn out with a blunt curved probe. When clear of the sur-
rounding tissue, it was coagulated sufficiently to sever it. The severed ends
were dropped back into place and the wound was touched with tincture of
merthiolate. The wound was so small, and there was so little trauma, that
sutures were unnecessary and healing was remarkably rapid. In many cases
it was difficult to locate the site of incision after one week.

As the snake is not deprived of glands or fangs, the outward appearance
of the head remains unaltered. In the specimens under observation no
change in disposition was noticed; feeding remained regular in those which
ate regularly before treatment. Each specimen was tested weekly by allow-
ing it to bite a rabbit. In every case the result was negative. A cotton-
mouth moccasin, ( Agkistrodon piscivorus ) in good health forty-five weeks
after treatment and eating well, was nevertheless unable to inflict other than
mechanical injuries when biting. Deaths resulted from causes other than the
operation — from mouth rot and intestinal disorder.

1940]

Jar os: Occlusion of Venom, Duct

51

The obstructed glands taken from snakes three, seven and eight weeks,
and ten months, after treatment were sectioned for microscopic examination.
The sections were studied by Dr. George J. Rukstinat, Associate Professor
of Pathology at Rush Medical College. He found large acinar spaces filled
with secretion, and lined by cells which evidently became cuboidal as a
result of the retained venom; but even the compressed cells showed little
signs of atrophy. In portions of the sections normal columnar cells survived.
Nuclei and cytoplasm were demarcated clearly. Confluence of acini in some
regions seemed apparent from the remnants of acinar walls which projected
into the lumens of the cyst-like spaces. Many cells contained clear or slightly
stippled vacuoles, the character of which is being investigated.

The histological evidences of cell viability were corroborated by the
sub-cutaneous injection into mice of secretion from the glands of one snake
which was killed ten months after occlusion of the duct. Minute quantities
(0.02 cc.) of this secretion proved fatal to mice in four hours, and induced
the usual changes of a general haemolysis and of local necrosis.

The potentialities for study of the venom gland through this method are
numerous. Perhaps some internal absorption of secretion occurs; or perhaps
some specialized cells elaborate components of venom as do specialized cells
in organs such as the pancreas.

Summary.

A method of rendering venomous serpents harmless is presented. Elec-
trocoagulation is used to prevent, by the destruction of a portion of the
duct, the escape of venom from the gland and its passage to the fang. The
operator can manage, unassisted and with safety, the entire procedure. The
snake is but slightly injured and is not disfigured. The desired results are
apparently permanent. The habits of none of the thirty-six snakes treated
seemed in any way effected by the operation, even after forty-five weeks.
The operation was successfully performed on crotalids ranging in size from
eight inches to four feet.

In the performance of the operation three innovations were introduced:
a practical device for holding the snake, a cheap and efficient operating
knife, and the use of Michel surgical clips to eliminate all danger of being
bitten.

I wish to express my grateful acknowledgement to Dr. George J. Ruks-
tinat, to Mr. Walter L. Necker of the Chicago Academy of Sciences, and
to Mr. Emil J. Rokosky, for advice and encouragement; and to my father,
Dr. Joseph F. Jaros, for his help with the technical aspects of the work.

Tee-Van: Review of American Cirrhitidae

53

5.

Eastern Pacific Expeditions of the New York
Zoological Society. XVII.

A Review of the American Fishes of the Family Cirrhitidae.'

John Tee-Van

Department of Tropical Research, New York Zoological Society.

(Plate I; Text-figures 1-4).

[This is the seventeenth of a series of papers dealing with the collections
of the Eastern Pacific Expeditions of the New York Zoological Society made
under the direction of Dr. William Beebe. The present paper is concerned
with specimens taken on the Templeton Crocker Expedition (1936), the
Eastern Pacific Zaca (1937-1938) and the Arcturus Oceanographic (1925)
Expeditions. For data on localities, dates, dredges, etc., of these expeditions,
refer to Zoologica, Vol. VIII, No. 1, pp. 1-45 ( Arcturus ) ; Zoologica, Vol.
XXII, No. 2, pp. 33-46 (Templeton Crocker) ; and Zoologica, Vol. XXIII,
No. 14, pp. 278-298 (Eastern Pacific Zaca ).]

Contents.

Page

Introduction 53

Key for the differentiation of the American
genera (Family Cirrhitidae) 54

Cirrhitus 54

Page

Cirrhitus rivulatus (Valenciennes) 54

Cirrhitichthys 58

Cirrhitichthys corallicola new species.... 58

Pseudocirrhites 61

Pseudocirrhites pinos Mowbray 61

Introduction.

For many years the only representative of the family Cirrhitidae known
from the Americas was the west coast Cirrhitus rivulatus (Valenciennes).
In 1927, however, Mowbray described a species from the Isle of Pines, Cuba,
establishing a new genus, Pseudocirrhites, for his fish, and marking the
first West Indian record for the family. During the 1937-1938 expedition
of the Department of Tropical Research of the New York Zoological Society
along the west coast of Mexico and Central America, still another form
was found; it is described as a new species in this paper. Specimens of all
three species have been examined and are reviewed and described herewith.

In the light of the recent discovery of a cirrhitoid fish in West Indian
waters, the finding of a closely related form on the west coast of America
tends to explain the apparent isolation of the Atlantic fish and provides an-
other link in the chain of evidence that demonstrates the close relationship
of the fishes of the Atlantic and Pacific sides of tropical American waters.

1 Contribution No. 590, Department of Tropical Research, New York Zoological Society.

54

Zoologica: Neiv York Zoological Society

[XXV :5

For an explanation of the position of localities mentioned in this paper,
see Text-fig. 4.

I am indebted to Dr. Leonard P. Schultz of the U. S. National Museum
and Prof. Albert E. Parr of the Peabody Museum, Yale University, for the
loan of specimens, and to Miss Janet B. Wilson, who made the drawings
used in this paper.

Family Cirrhitidae.

Key for the differentiation of the American genera.

la. Scales on the cheek very small, 16 to 20 rows from eye to edge of pre-

opercle (Pacific) Cirrhitus.

lb. Scales on the cheek large, 4 to 5 rows from eye to edge of preopercle.
2a. Scales absent on nape and interorbital space ; a small patch of scales

behind and between the anterior nosti’ils (Pacific) ... Cirrhitichthys.

2b. Scales present on nape and interorbital space. (Atlantic)

Pseudocirrhites.

Cirrhitus Lacepede, 1803.

Genotype by monotypy, Cirrhitus maculatus Lacepede, Lacepede, 1803,
Hist. Nat. des Poiss,, 5: 2.

Cirrhitus rivulatus (Valenciennes).

(Plate I, Text-fig. 1).

Proportions in Percentage of the Standard Length, and Counts: Depth
34-36 (2.4-3) 2 ; head 38-40 (2.46-2.75); eye 7.7 in fish of 119 to 182 mm.,
9.2 in 65 mm. fish, 11-11.4 in 30 to 35 mm. fish (3-5.7) ; snout 11-14.7
(2. 3-3.1) ; maxillary 13.6-17.6 (2. 1-2. 8) ; interorbital space 5-6.5 (5.4-7.66) ;
pectoral length 25.4-28.6 in fish of 119 to 182 mm., 32.2-34 in 31 to 65 mm.
fish; pelvic length 20-23; snout to 1st dorsal fin 38-41; snout to 2nd dorsal
fin 64-68; snout to anal fin 68-73; snout to pectoral fin 34-38; snout to pelvic
fin 45-51; 1st dorsal height 11.7-14.6; 2nd dorsal height 12.2-15.4; anal
height 17.4-20.6. Dorsal fin X, 11 or 12; anal fin III, 6 or 7 ; pectoral fin with
14 rays, the uppermost and the lower 7 (rarely 8) simple; scales: 5% to
6V2 from origin of 1st dorsal fin to lateral line, 46 to 49 in a longitudinal
series, 12 to 14 scales from lateral line to origin of anal fin, 7 to 8 predorsal
scales; gill-rakers. 5 to 0 on upper limb and 10 to 11 on the lower limb of the
first arch.

Body compressed, sturdily and heavily built, the caudal peduncle deep.
Anterior profile strongly convex, especially in larger individuals, the eye
entering the profile very slightly. Scales present on the body, absent on the
head except for the preopercle, which is very finely scaled, and the opercle
which has scales on its anterior part similar to those of the body; somewhat
smaller scales on the posterior flap of the opercle. Lateral line continuous,
the openings of the canals tilted upward.

Interorbital space concave; a low longitudinal crest on the nape; pre-
opercle broadly rounded, its upper portion finely serrate, the serrations most
conspicuous in small specimens; opercle with an obtuse flap extending back-
ward over the pectoral base. Branchiostegal rays 6, the innermost one very
small; the branchiostegal membranes broadly united but free from the
isthmus; mouth low, the lips thick in large specimens; maxillary extending
backward to the vertical of the posterior border of the orbit in large fish and
to the center of the orbit in small fish. Upper jaw with small, rather widely
spaced canines, the posterior tooth and three or four near the symphysis en-

2 Figures in parentheses are proportions stated in terms of times in the head or standard length.

1940]

Tee-Van: Review of American Cirrhitidae

55

larged; inside of these teeth is a band of smaller teeth, the band widest
anteriorly, becoming narrower laterally and vanishing posteriorly. Lower
jaw with similar teeth but with two or three of the outer canine-like teeth
near the center of each side of the jaw enlarged. Tongue large, wide and
free anteriorly. Gill-rakers short, covered with fine asperities.

Spinous dorsal fin rather low, the spines increasing in size to the 5th
and 6th, then decreasing, the ultimate spine longer than the penultimate;
soft dorsal fin higher than the spinous ; caudal fin truncate; or slightly con-
cave ; anal fin with the second spine largest and heaviest, the rays long, twice
the length of the spines; pectoral fin with its lower rays considerably
swollen, especially in large individuals, the 5th and 6th from the bottom
longest, posterior edge of the branched rays oblique, the uppermost simple
ray shortest of all; pelvic fins originating under the center of the adpressed
pectoral fin, the tip of the pelvics reaching to the vent or slightly beyond.

Color: During growth there is considerable change in the coloration of
this species, the principal alterations being caused, first, by the breaking up
of the simple vertical bands of the young into groups of spots bordered with
darker and lighter, the spots often acquiring a different basal color; second,
the shift from the pearly-gray ground color of the very young to the browns
or greenish-browns of the adults; third, the loss of scarlet on the upper
anterior surface markings of the head and the disappearance of the scarlet,
black and gray spinous dorsal fin.

The original description of the young, (nominal Cirrhitus betaurus),
given by Gill in 1862, agrees excellently with our similar-sized specimens
(35 mm.). Gill’s description is appended here: “The color is ivhitish on the
body, blackish on the shoulders and from the dorsal fin to the eyes, and with
four complete, oblique, blackish bands; the first under the middle of the
spinous dorsal; the second under the last spine; the third under the middle
of the soft dorsal, and the fourth encircling the caudal peduncle. The head
has three lateral bands, one on the preorbital region, a second on the cheek,
and a third on the posterior margin of the preoperculum. The operculum
has a longitudinal oblong spot. The chin has four spots forming the angles
of a rhomb, and there is another one behind, on the branchiostegal membrane
near the margin. The spinous dorsal is margined with blackish, and the
two bands beneath more or less ascend on it; anal blackish. The caudal has
a blackish B-shaped mark and a band at its base divided by the lateral line.
The pectoral is dusky, with a black spot at its base nearly surrounded by a
clear area, and separated from a spot in front of the base. The ventrals are
blackish, with nearly transparent sides and margin.”

A color plate made in the field from a living 35 mm. specimen shows the
following slight differences from this color description : The ground color is
pearly gray, the spinous dorsal fin is scarlet with a blackish narrow upper
border, the vertical body bands being continued upward onto the fin. All of
the dark markings on the head and nape have a central core of reddish-
brown. Jordan (Fishes of Sinaloa, Proc. Cal. Acad. Sciences, (2) 5: 473)
states of similar-sized small individuals: “First dorsal fin bright orange red
in life; second reddish; cross bands on body black.”

In a 65 mm. standard length fish the ground color is much duller and
the bands on the sides are starting to break up into spots. This is especially
noticeable on the dorsal surface of the body near the dorsal fin.

In our series the next largest specimen (119 mm.) has the pattern and
coloration of all of the remaining larger specimens.

The adults may be described as follows, the description being combined
from other descriptions and our own observations. Color of body dark green-
ish- or yellowish-brown, occasionally more ochraceous brown on the lower
head and belly (dark gray: Fowler). Dark brown, spots and bands, occasion-
ally becoming yellowish-brown, usually bordered first with darker brown or

56

Zoologica: New York Zoological Society

[XXV :5

Text-figure 1.

Cirrhitus rivulatus Valenciennes. Specimens of 35, 65, 142 and 182 mm.
standard length, showing the changes of pattern with age.

black and then with pale blue, distributed as follows: three narrow bands
crossing the interorbital space; two to three bands from the eye to the
maxillary or premaxillary; a saddle-like transverse band on the nape pos-
terior to the eye, in the center of which, posteriorly, is a small circular spot;
on the nape are three longitudinal short bars, one on the mid-dorsal line
and one on either side of this; a band from eye to supra-scapular ; one from
eye to upper border of the preopercle; a vertical band on the preopercle
anteriorly and another on the posterior border ; an upwardly ascending band
from the upper edge of the preopercle toward the upper tip of the opercle;
a short vertical band on the base of the pectoral fin and another similar one

1940]

Tee-Van: Review of American Cirrhitidae

57

on the pre-pectoral region. Body with five upright oblique bands of similarly
colored and bordered broken spots, the bands extending onto the spinous
and soft dorsal fins; in addition there is a similar band on the posterior end
of the caudal peduncle. Caudal fin with sub-circular brown spots bordered
with darker brown and light blue, the spots roughly forming vertical and
semicircular vertical lines; this arrangement is not always apparent, in
large specimens. In some descriptions the tail is described as being dark with
a network of pale blue reticulated lines. Dorsal fin dark brown, somewhat
mottled with continuations of the bars of the sides of the body; pectoral fins
dusky; anal fin dark brown, sometimes green basally, with two to five prom-
inent spots similar to those on the body on its posterior portion; pelvic fins
dusky, especially toward the tip, sometimes olive basally, gray-black termin-
ally.

Iris in life (327 mm. fish) olive green above and below with a broad
silvery zone extending longitudinally, within which are two rounded spots
in front and two more behind the pupil. A narrow green area immediately
surrounding the pupil.

Range: Pacific mainland from Lower California and the Gulf of Cali-
fornia southward to Panama (Mexico: Cape San Lucas, Gulf of California,
Mazatlan, Sihuatanejo, Acapulco; Nicaragua: Corinto; Costa Rica: Piedra
Blanca Bay, Uvita Bay; Panama: Panama) ; Revillagigedo Islands (Clarion
Is., Socorro Is.) ; Galapagos Islands (Hood Is., James Is., Tower Is.) ;
Malpelo Island.

Local Distribution : A rocky reef and tidepool species, hiding in crevices
and darting out for prey.

Method of Capture: Hook and line baited with bait or with a shiny piece
of metal, traps, poisoning in tidepools.

Size: Grows to ITV2 inches. A 327 mm. (I2V2 inches) fish weighed 3
pounds and a 450 mm. ( 171/2 inches) fish weighed 5 pounds.

Shidy Materials: 8 specimens from 31 to 450 millimeters, from the
following localities: Nicaragua: Corinto; Costa Rica: Piedra Blanca Bay,
Uvita Bay; Galapagos Islands: Tower Island. In addition, water-glass
sight records were made of this species at the following places: Mexico:
Sihuatanejo, Acapulco; Clarion Island.

References: Cirrhites rivulatus Valenciennes, Voyage autour du Monde, sur
la fregatte “La Venus,” tome 5, Ichthyologie, 1855: 309, plate 3, fig. 1 (Descrip-
tion, color, poor plate: type locality, Galapagos Islands .1 ; Gunther, Catalogue of
the Acanthopterygian Fishes in the Collection of the British Museum, 2, 1860:
519 (Short description) ; Jordan, D. S. and Gilbert, C. H., List of Fishes col-
lected at Mazatlan, Mexico, by Charles H. Gilbert, Proc. U. S. Nat. Mus., 2,
1882 (1883) : 108 (Check-list, name only) ; Jordan, D. S., A list of the fishes
known from the Pacific coast of tropical America, from the Tropic of Cancer to
Panama, Proc. U. S. Nat. Mus., 8, 1885 (1886) : 381 (Check-list; Cape San Lucas,
Galapagos Islands) ; Jordan, D. S. and Evermann, B. W., The Fishes of North
and Middle America, Bull. U. S. Nat. Mus., 47 (2), 1898: 1491 (Description,
color, range, short synonymy) ; Jordan, D. S. and McGregor, R. C., List of
fishes collected at the Revillagigedo Archipelago and neighbouring islands, Rept.
U. S. Fish Comm, for 1898 (1899) : 283 (Clarion and Socorro Islands) ; Jordan,
D. S. and Evermann, B. W., The Fishes of North and Middle America, Bull. U. S.
Nat. Mus., 47 (4), 1900: plate 227, fig. 576 (figure); Pellegrin, J., Poissons
recueillis par M. Leon Diguet dans le Golfe de Californie, Bull. Mus. Hist. Nat.
(Paris), 7, 1901: 163 (Gulf of California); Gilbert, C. H. and Starks, E. C.,
The Fishes of Panama Bay, Mem. Calif. Acad. ScL, 4, 1904: 139 (Panama;
restates Gunther’s 1868 record) ; Beebe, W., Galapagos, World’s End, G. P.
Putnam’s Sons, New York and London, 1924: plate 5 (colored figure) ; Ulrey,
A. B., A check -list of the fishes of Southern California and Lower California,
Journ. Pan-Pacific Res. hist., 4 (4), 1929: 18 (Check-list only; Cape San Lucas) ;
Terron, C. C., Lista de los peces de los costas de la Baja California, Ann. Inst.
Biol., Univ. Nac. Auton. Mexico, 3, 1932: 79 (Check-list, name only, Cape San

58

Zoological New York Zoological Society

[XXV :5

Lucas) ; Breder, C. M., Jr., Heterosomata to Pediculata from Panama to Lower
California, Bull. Bingham Oceanogr. Coll., 2 (3), 1936: 37 (Unknown locality).

Cirrhitus rivulatus : Gill, T., Synopsis of the family of Cirrhitoids, Proc.
Acad. Nat. Sci. Phila., 1862: 107 (Name, synonymy, range); Gill, T., Catalogue
of the fishes of Lower California, in the Smithsonion Institution, collected by
Mr. John Xantus, Proc. Acad. Nat. Sci. Phila., 1862: 259 (Cape San Lucas,
LoWer California) ; Snodgrass, R. E. and Heller, E., Shore fishes of the Revil-
lagigedo, Clipperton, Cocos and Galapagos Islands, Proc. Wash. Acad. Sci., 6,
1905: 385 (Galapagos Islands, range, few proportions) ; Beebe, W., The Arcturus
Adventure, G. P. Putnam’s Sons, New York and London, 1926: 150 (Hood
Island, Galapagos; method of capture). See p. 434 for specific determination;
Jordan, D. S., Evermann, B. W. and Clark, H. W., Check list of the fishes and
fishlike vertebrates of North and Middle America north of the northern boundary
of Venezuela and Colombia, Rep. U. S. Comm, of Fish, for 1928 (1930) : 358
(Check list, range) ; Fowler, H. W., The Fishes of the George Vanderbilt South
Pacific Expedition, 1937, Acad. Nat. Sci. Phila., Monograph No. 2, 1938: 15
(Description, color; Malpelo Island), 53, (Note on color, proportions; James
Island, Galapagos), 257, (check list).

Cirrhitus betaurus : Gill, T. N., Catalogue of the fishes of Lower California
in the Smithsonian Institution, collected by Mr. John Xantus, Proc. Acad. Nat.
Sci. Phila., 1862: 259 (Original description, color; type locality: Cape San Lucas,
Lower California) ; Jordan, D. S. and Gilbert, C. H., Catalogue of the fishes
collected by Mr. John Xantus at Cape San Lucas, which are now in the United
States National Museum, with descriptions of eight new species, Proc. U. S.
Nat. Mus., 5, 1882 (1883): 371 (Synonymized with Cirrhitus rivulatus).

Cirrhitichthys rivulatus: Gunther, A., An account of the Fishes of the
States of Central America, based on collections made by Captain J. M. Dow,
F. Godman, Esq., and 0. Salvin, Esq., Trans. Zool. Soc. London, 6 (7), 1868:
387 (Check list); Galapagos Islands, Panama), 421, plate 86, fig. 4 (Panama,
description, figure).

Cirrhites betaurus: Jordan, D. S., The fishes of Sinaloa, Proc. Calif. Acad.
Sci., (2) 5, 1895: 472 (Relationship of betaurus and rivulatus discussed, color,
Mazatlan) ; Jordan, D. S. and Evermann, B. W., The Fishes of North and Middle
America, Bull. U. S. Nat. Mus., 47, 1898: 1492 (Description, color, range);
Ulrey, A. B., A check-list of the fishes of Southern California and Lower
California, Journ. Pan-Pacific Res. Inst., 4 (4), 1929: 18 (Check list only. Cape
San Lucas) ; Terron, C. C., Lista de los peces de las costas de la Baja Cali-
fornia, Ann. Inst. Biol., Univ. Nac. Anton. Mexico, 3. 1932: 79 (Check list,,
name only; Cape San Lucas).

Cirrhitichthys Bleeker, 1856.

Genotype by original designation Cirrhites graphidopterus— Cirrhites
aprinus Cuv. and Val. ; Bleeker, Naturk. Tijdschr. N ederl. -Indie, Deel X
(new series, Deel VII) 1856: 474. The generic description was published
in 1857 by Bleeker, Vischfauna van Ambonia, Acta Soc. Sci. Indo-Nederl.,
2. 1857: 39.

Cirrhitichthys corallicola sp. nov.

(Text-figure 2) .

Type : Holotype, No. 28,710a, Eastern Pacific Zaca Expedition of the
Department of Tropical Research, New York Zoological Society; Gorgona
Island, off the Pacific coast of Colombia, South America (Lat. 2° 58' N.,
Long. 78° 11' W.), in coral, March 30, 1938; standard length 58 mm. Para-
types: 32 specimens, No. 28,710, same locality and date as the holotype, 22
to 59 mm. standard length. Types in the collections of the Department of
Tropical Research, New York Zoological Society.

Measurements in Percentage of the Standard Length, and Counts:
Measurements of the holotype: Depth 36.6 (2.7) ; head 34.5 (2.9) ; eye 8.4

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59

(4.1 in head;; snout 11.2 (3.1 in head); interorbital space 5.7 (6.1 in
head); maxillary 13.3 (2.6); pectoral fin length 33.5 pelvic fin length 24;
snout to origin of 1st dorsal fin 33.5; snout to 2nd dorsal fin 64; snout to
origin of anal fin 68; snout to pectoral fin 32; snout to pelvic fin 45; 1st
dorsal height 17.6; 2nd dorsal fin height 26.8; anal fin spine height 21.6.

Measurements of 10 individuals, 34 to 58 mm. standard length, including
those of the holotype: Depth 33-39.5 (2. 6-2. 9) ; head 34.2-39 (2. 5-2.9) ; eye
7.8-10.9 (4.1-4.5); snout 10.2-12.4 (3. 1-3.5) ; maxillary 12.3-14.2 (2.6-2.9) ;
interorbital space 4.4-6. 1 (6.1) ; pectoral fin length 31-37; pelvic fin length
23.7-26.3; snout to origin 1st dorsal fin 31.5-37; snout to 2nd dorsal fin
62-65; snout to origin of anal fin 65-72; snout to pectoral fin 32-36; snout to
pelvic fin 42-49; 1st dorsal height 16-19; 2nd dorsal height 23-29; anal fin
height (spine) 20-22. Counts: dorsal fin X, 12; anal fin III, 6; pectoral fin
with 14 days, the uppermost ray and the lowermost 6 or 7 simple ; scales 43-45
in a lateral series, 4 from origin of dorsal fin to lateral line, 9 or 10 from
lateral line to origin of anal fin; gill-rakers 3 to 5 on upper limb, 9 to 11 on
the lower limb of the first gill-arch, the lowermost two rudimentary; verte-
brae 10 plus 15 plus 1.

Body compressed, the head obtusely pointed, the greatest depth at %
the length of the pectoral, depth of caudal peduncle 13.3% of the length.
Anterior profile at a 45 degree with the axis of the body, with a slight notch
just above the orbit, the latter entering the profile. Upper profile from
origin of dorsal fin to the 7th spine straight, the profile then gently curving
toward the peduncle. Lower profile from snout to beneath base of pelvic fins
at a considerable angle to the line of the abdomen.

Scales cycloid, absent on the head with the exception of the opercles and
a small patch between the posterior nostrils; lips naked. Four oblique rows
of scales on the preopercle from the eye to the rounded angle of the pre-
opercle. Scales present on the opercles and on the branchiostegal membranes
immediately beneath the isthmus; the latter scales are very small. Scales
extending on the bases of the spinous dorsal fin and on the membranes of
the soft dorsal and anal fins. Lateral line continuous, the tubes short and
opening upward ; nape with a large number of tubes.

Head obtusely pointed, the eye entering dorsal profile ; interorbital space
concave, the supraorbital ridges prominent, especially posteriorly; anterior
nostril circular, with a short tube, nearer eye than snout and with a 5-fin-
gered fleshy tentacle on its posterior border; posterior nostril with a raised
border internally and anteriorly, placed close to the orbit; preopercle
broadly rounded with 17 to 18 short but strong serrae, the uppermost slightly
longer than the lower; opercle ending in an obtuse flap and with two flat
spines, the lower much larger and more evident than the upper; branchio-
stegal rays 6, the innermost one on each side very small; branchiostegal
membranes broadly connected but free from the isthmus; mouth small, the
lips fleshy; maxillary extending to below the center of the orbit, the max-
illary almost completely hidden beneath the suborbital. Teeth of the upper
jaw with an outer row (20 to 21 on each side) of small, recurved canines,
two on each side of the isthmus considerably larger and stronger than the
others, the posterior teeth of the jaw slightly larger than their fellows;
behind this outer row is a villiform band of small teeth, widest anteriorly,
becoming narrower as it progresses backward. Lower teeth similar to those
of the upper jaw, except that on the middle of the side of each jaw there is
a group of enlarged canines; anterior to these the teeth are similar in size
to those of the upper jaw, posterior to the enlarged group the teeth are quite
small. The interna] band of villiform teeth on the lower jaw extends back-
ward only as far as the anterior tooth of the lateral enlarged canines.
Vomer and palatines with villiform teeth, the teeth of the vomer in a broadly
arched band. Gill-rakers small and short.

Dorsal fin with the 3rd, 4th and 5th spines highest, the upper anterior

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edge of each interspinal membrane with a series of 8 to 10 dermal tentacles;
soft dorsal high, the anterior two rays elongate and forming a slight lobe.
Anal fin high, the second spine longer and heavier than the others, its length
1.5 in the head, length of the 1st spine 1.9 in the second, length of the third
spine 1.3 in the second. Caudal fin truncate. Pelvic fins in close juxta-
position, their origin beneath the anterior 2/5ths of the pectoral fin, the tips
of the pelvics extending to the vent. Pectoral fin with 14 rays, the upper-
most one unbranched, the six lowermost (rarely the 7th) simple and un-
branched, slightly swollen and with their tips free from membrane; 5th
from bottom ray longest (1.15 in the head in the type), the 4th and 6th next
longest.

Text-figure 2.

Cirrhitichthys corallicola Tee- Van. Drawing of the type, 58 mm. standard
length.

Color: In life the general body color is orange yellow with a pinkish or
lavender cast, heavily covered with dark orange or red spots placed somewhat
in alternate oblique upright rows of larger and smaller spots, the spots ex-
tending onto the lower portion of the dorsal fin; similarly colored spots on
the head, snout and base of the pectorals are much smaller than those on
the body. Dorsally and posteriorly the spots become brighter red in color.
Snout reddish. Dorsal fin transparent yellow, mottled with brownish, the
tips of the spines and the tentacles on the interspinous membranes scarlet;
caudal fin yellow mottled with dusky; pectoral, pelvic and anal fins trans-
parent yellow, the latter somewhat dusky. Iris golden red.

In the Gorgona Island specimens the spots are exceptionally well-marked
and clean cut. In many of the specimens from more northerly localities,
the spots tend to merge and thus produce irregular vertical bands. This
condition is especially noticeable in a 30 mm. fish from Acapulco, less so in
some others.

In preservative our specimens have become yellow with brownish and
gray spots, the pattern of the body being retained. In two specimens from
the Pearl Islands, Panama (Vanderbilt Collection) there remain only vague
traces of the spots on the body.

Range: Found by us from Sihuatanejo, Mexico, southward along the
coast to Gorgona Island, Colombia. (Mexico: Sihuatanejo Bay, Acapulco;
Costa Rica: Port Parker, Port Culebra; Panama: Bahia Honda, Pearl

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Tee-Van: Review of American Cirrhitidae

61

Islands (Vanderbilt Alva 1938 Expedition); Colombia: Gorgona Island).
The majority of the specimens seen and taken by us were found in the
interstices of coral. The fishes were rarely seen from above, but they were
exceptionally conspicuous when viewed from a diving helmet while sub-
merged.

Remarks : This species is blenny-like in the rapidity of its movements
and general habits and also resembles in this respect, some of the smaller
serranids. Our field names of “lavender blenny” or “lavender serranid,”
given before we had been able to catch an example, are evidence of the ap-
pearance of the fish in life.

Specimens Examined : 45, including the types and paratypes, from the
localities mentioned above under Range.

Pseudocirrhites Mowbray, 1927.

Genotype by monotypy, Pseudocirrhites pinos Mowbray, in Breder, Bull.
Bingham Oceanogr. Coll., 1 (1), 1927: 48.

This genus is close to the Pacific Cirrhitichthys, differing principally
in the possession of scales on the nape and interorbital space. In Cirrhitich-
thys, as represented by C. corallicola and a Japanese specimen of C. aureus,
scales are absent on the regions mentioned above, but present in a small
subcircular patch between and posterior to the anterior nostrils. I have not
checked on other species of Cirrhitichthys to determine whether this condi-
tion is true of all of the species of the genus.

Considering the geographical isolation of the Atlantic species plus the
difference in scalation, Pseudocirrhites is maintained as a valid genus, as
opposed to synonymizing it with Cirrhitichthys, to which it is closely related.

In the original description of Pseudocirrhites , Mowbray mentions the
following as the principal character of his new genus: “The above new
genus is based on the broadly united gill-membranes.” This conception of
the new genus was probably based on a misstatement in Jordan and Ever-
mann, Bull. U. S. Nat. Mus., 47 : 1490, which gained emphasis by being re-
peated in Jordan and Evermann, The Shore Fishes of the Hawaiian Islands
(Bull. U. S. Fish Comm., 23 (1), 1905: 446), and in Jordan and Herre’s A
Review of the Cirrhitoid Fishes of Japan ( Proc . U. S. Nat. Mus., 33, 1907:
157). This error described the gill-membranes in Cirrhitidae as separate
and free from the isthmus. Unfortunately this is not true, as the possession
of broadly united gill-membranes is characteristic of all members of the
family that have been examined and is so mentioned in other descriptions
beyond those listed here. (See also: Regan, On the Cirrhitoid Percoids, Ann.
Mag. Nat. Hist., (8) 7: 259-262).

The definition of Pseudocirrhites may be rewritten as follows: Cirr-
hitoid fishes with rather large cycloid scales, approximately 42 rows in a
lateral series; 4 to 5 rows of scales on the cheek; nape and interorbital
space as far forward as the anterior nostrils fully scaled. Teeth present on
the vomer and palatines; jaws with an outer row of small canines inside of
which are villiform bands of smaller teeth, a few strong, backwardly-turned
canines on center of side of each lower jaw, upper jaw with a few slightly
enlarged teeth near the symphysis and toward the posterior end of each jaw.
Preopercle serrate. Branchiostegals 6.

Pseudocirrhites pinos Mowbray.

(Text-figure 3).

Proportions in Percentage of the Standard Length, and Counts : Depth
33-38 (2.6-3) ; head 37 (2.7) ; eye 9.2-9.6 (3.84-4) ; snout 11.3-11.7 (3.1-3.3) ;

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[XXV :5

maxillary 13-13.4 (2.74-2.86) ; interorbital space 4.8-5.76 (4.15-6.4) ; pectoral
fin length 36.5-39; pelvic fin length 21.6-25; snout to 1st dorsal fin 36; snout
to 2nd dorsal fin 62-64.5; snout to anal fin 70; snout to pectoral fin 34-36;
snout to pelvic fin 45-49. 1st dorsal fin height 15.7 ; 2nd dorsal fin height
18.5-21.5; anal fin height 20.8. Counts : dorsal fin X,ll; anal fin III, 6; pec-
toral fin, 14 rays, the top 1, and lower 5 simple; pelvic fin I, 5; scales: 6 from
origin of dorsal fin to lateral line, 42 in a lateral series, 8 from the lateral
line to the origin of the anal fin; gill-rakers 6 on upper limb of first arch,
8 to 9 on the lower limb of the first arch.

Text-figure 3.

Pseudocirrhites pinos Mowbray. Drawing of the type, 54 mm. standard
length.

Body oblong, compressed; depth of the caudal peduncle 2.8 in the head.
Body and head covered with large cycloid scales, with the exception of the
snout and preorbital, and the maxillary and premaxillary; branchiostegal
membranes scaled. Lateral line continuous, nearly straight, extending onto
the base of the caudal ; pores turned obliquely upward. A series of small
dermal tentacles on the anterior upper edge of each interspinal membrane
of the dorsal fin.

Eye placed high and slightly entering the dorsal profile; interorbital
space concave; anterior nostril with a fringed tentacle; preopercle rounded,
finely serrate; opercle with a single flat spine; branchiostegal membranes
united (accidentally bi'oken during re-examination of the type) and free from
the isthmus, covered with small scales; mouth small, placed low; maxillary
narrow, about % entirely covered by the preorbital; premaxillaries little
protractile; villiform teeth in bands in each jaw, on the outer border of
which are a band of small canines, 3 or 4 slightly enlarged canines near the
symphysis of the upper jaw, a few enlarged teeth at the posterior end of the
jaw; a few strong canines on the middle of the sides of the lower jaw; vomer
and palatines with small teeth. Caudal fin truncate, its rays about equal in
length to the second anal spine, the first spine of the anal fin about 2 in the
second, the second spine longest and strongest, the third intermediate in
length and strength to the other two; pectoral fin with 14 rays, the upper-
most 1, and lowermost 5 simple, the remainder branched; lowermost simple
rays considerably longer than the other rays; tip of pectoral extending to
the 1st soft ray of the anal fin; pelvic fins with their tips extending to the
1st anal spine.

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63

A 26 mm. fish differs from the type in being less deep, and in having
the two upper rays of the pectoral fin simple on the right side, and the upper
three simple on the left side.

Coloration : Head and body light brown; three vertical light-colored
bands on body, broadest interiorly, in the center of each band is a narrow
brown line ; a broad band of dark brown covering the entire caudal peduncle ; a
brown spot larger than the eye, two-thirds of which is on the body, at the
base of the posterior dorsal rays. The bands of the body, both dark and
light, extend onto the dorsal fin. Pectoral, ventral and caudal fins pale. Head
and nape and dorsal fin with bright red spots.

In a color plate made from life of the 26 mm. fish, the narrow brown
lines in the center of the light colored bands are faint; the spot at the base
of the soft dorsal and the band on the caudal peduncle are black ; the pectoral
and anal rays and the basal portions of the upper rays of the caudal fin
are reddish.

Text-figure 4.

Map of the Eastern Pacific Zaca (1937-1938) Expedition of the Department
of Tropical Research of the New York Zoological Society, showing the localities
mentioned in the text. The Isle of Pines, Cuba, the type locality of Pseudo-
cirrhites pinos Mowbray, is the small island immediately south of the western
end of Cuba.

Range : Known from the Isle of Pines, Cuba, and from Saba Bank, 6
miles S. W. of Saba Island, West Indies (17° 35' N., 63° 21' W.)

Local Distribution: The two known specimens of this species were taken
in coral, in shallow water (type) and at 25 fathoms.

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Zoologica: New York Zoological Society

Study Materials: The type and a specimen taken by the Arcturus
Oceanographic Expedition on Saba Bank, West Indies, March 15, 1925
(Dept. Trop. Res., N. Y. Zoological Society, No. 5061, color plate A. 822.)

References: Pseudocirrhites pinos. Mowbray, in Breder, Bull. Bingham
Oceanogr. Coll. 1 (1), 1927; 48, fig. 23 (Description, color; figure; type locality.
Isle of Pines, Cuba; Type: No. 382, Bingham Oceanogr. Coll., Peabody Museum,
Yale University).

Note on Cirrhitus rivulatus (Valenciennes).

A 505 mm. fish of this species from the Galapagos Islands (U. S. Nat.
Mus., 38,302) examined after this paper was in page proof, has the color
bars and bands, especially those of the head, much wider than in smaller
specimens and occupying relatively a greater amount of space when
compared with the interspaces. The teeth of the upper jaw in this speci-
men show a few enlarged canines. In the lower jaw the enlarged canines
of the middle of the side of each jaw (3 on the right side and 5 on the
left) are very large and conspicuous and are wider apart than in smaller
fish. Anterior to these there are two or three small canines and posterior
to the enlarged group on each side are a group of smaller canines. The
villiform teeth in the lower jaw occupy a small patch in the front of each
jaw anterior to the large lateral teeth.

EXPLANATION OF THE PLATE.

Plate I.

Cirrhitus rivulatus (Valenciennes). Seven individuals from Corinto,
Nicaragua, 65 mm. to 240 mm. standard length, showing the alteration in pat-
tern correlated with growth and size.

TEE-VAN.

PLATE I.

A REVIEW OF THE AMERICAN FISHES OF THE FAMILY CIRRHITIDAE.

Crane: Post-embryonic Development of Ocypode

65

6.

Eastern Pacific Expeditions of the New York
Zoological Society. XVIII.

On the Post-embryonic Development of Brachyuran Crabs
of the Genus Ocypode.1

Jocelyn Crane

Technical Associate, Department of Tropical
Research, New York Zoological Society.

(Text-figures 1-8).

[This is the nineteenth of a series of papers dealing with the collections
of the Eastern Pacific Expeditions of the New York Zoological Society made
under the direction of William Beebe. The present paper is concerned
principally with specimens taken on the Eastern Pacific Zaca (1937-1938)
and Arcturus Oceanographic (1925) Expeditions. For data on localities,
dates, dredges, etc., of these expeditions, refer to Zoologica, Vol. VIII, No.
1, pp. 1-45 {Arcturus), and Zoologica, Vol. XXIII, No. 14, pp. 287-298
(Eastern Pacific Zaca) f]

Contents.

Page

I. Introduction 65

II. Summary of Important Points 66

III. Previous Knowledge of Ocypodid Devel-
opment 66

A. Zoea 66

B. Megalops 66

IV. The First Zoea of Ocypode gaudichaudii 67

A. Material and Methods 67

B. Diagnosis 67

C. Description 67

D. Comparison 69

Page

V. The Megalops of Ocypode 69

A. Material and Methods 69

B. Taxonomy and Identification 70

C. Comparison of Uca and Ocypode. . 72

D. Generic Characters 72

Diagnosis 72

Description 72

E. Specific Characters 80

F. Ecology 81

VI. Bibliography 81

I. Introduction.

The present paper is the first of a series dealing with the brachyuran
crabs of the Eastern Pacific Zaca Expedition. It concerns the first zoea of
Ocypode gaudichaudii and the megalopa of the two Pacific and single
Atlantic species of the genus.

Previous reports on the Brachyura of the various Eastern Pacific Ex-
peditions of the New York Zoological Society are the following: Rathbun,
1924, “Brachyuran Crabs Collected by the Williams Galapagos Expedition,
1923” ( Zoologica , Vol. V, No. 14) ; Boone, 1927, “Galapagos Brachyura”
{Zoologica, Vol. VIII, No. 4) ; Glassell, 1936, “Templeton Crocker Expedi-
tion. I. Six New Brachyuran Crabs from the Gulf of California” {Zoo-

1 Contribution No. 591, Department of Tropical Research, New York Zoological Society.

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[XXV :6

logica, Vol. XXI, No. 17) ; Crane, 1937, “The Templeton Crocker Expedition.
III. Brachygnathous Crabs from the Gulf of California and the West Coast
of Lower California” ( Zoologica , Vol. XXII, No. 3) ; Crane, 1937, “The
Templeton Crocker Expedition. VI. Oxystomatous and Dromiaceous Crabs
from the Gulf of California and the West Coast of Lower California”
( Zoologica , Vol. XXII, No. 7).

The drawings in the present paper are the work of Miss Harriet
Bennett.

To Mr. Templeton Crocker I wish to express my appreciation for the
opportunity of collecting material on a cruise of his yacht Zaca; to Dr.
William Beebe, Director of the Department of Tropical Research, for his
supervision and advice in the preparation of this paper; and to Dr. Waldo
L. Schmitt of the United States National Museum for the loan of speci-
mens, and for his generous permission to dissect an Atlantic megalops and
include the results of the study in this report.

II. Summary of Important Points.

1. The zoea of a species of Ocypode (O. gaudichaudii from the eastern
Pacific) is described for the first time. It differs from that of the most
closely related genus, Uca, principally in having lateral spines.

2. The megalopa of both eastern Pacific species of the genus, O.
gaudichaudii and O. occidentalis, are described for the first time. They
differ only in small details (such as the number of setae on the last pleopods)
from the western Atlantic megalops, O. albicans, but are totally distinct from
the known megalops of Uca.

3. The megalops described by Rathbun (1924, p. 155) as being perhaps
referrable to O. gaudichaudii is shown to belong instead to the genus
Plagusia, probably to P. depressa tuberculata.

4. A probable explanation of the peculiar characteristics of Ocypode
megalopa is presented: Since a previous investigator (Smith, 1880) found
evidence that Ocypode moults into the first crab stage in shallow holes on
the exposed beach, the megalops must be washed in by the waves. Hence its
obesity, its thick, hard cuticle, and its ability, through the development of
special grooves, to fold all its appendages tightly against its body, serve
as protections against the buffeting of the waves, scraping of the sand, and
exposure to the drying effect of air in landing on the beach.

III. Previous Knowledge of Ocypodid Development.

A. Zoea: The zoeal stages of Ocypode have been hitherto unknown.
Zoeae of a number of other ocypodids, however, have been identified : Those
of Uca pugilator (Bose), U. pugnax (Smith) and U. minax (Le Conte)
have been described by Hyman (1920, pp. 485 ff.). Aikawa (1929, pp. 49-51)
has described the first zoea of each of the following species : Macrophthalmus
dilatatus de Haan, M. japonicus de Haan, M. depressus Ruppell, Scopimera
globosa de Haan and Ilyoplax pusillus (de Haan). The same author (1937,
p. 152) gives in tabular form a resume of the principal characteristics of
all the above species.

B. Megalops: Apparently only in the genera Ocypode and Uca have
megalopa been described. In 1817 Say (p. 155 ff.) described as the type of
a new genus and species Monolepis inermis, a strange creature which he
referred to the Macroura near Porcellana, although he recognized and was
puzzled by its resemblances to the Brachyura. Since the larval stages of
crabs were, of course, quite unknown in those days, Say’s conclusions were
perfectly natural. In this, the first notice of its habits, he says: “Of this
interesting animal I found several specimens on the eastern shore of Mary-

1940]

Crane: Post-embryonic Development of Ocypode

67

land, which had been cast on the beach by the refluent tide. They appeared
desirous to protect themselves from the dashing of the surf, and the influ-
ence of the sun, by burrowing in the sand, in order to wait the return of the
tide; but their efforts had no further effect on the compact sand, than to
raise a small portion of the surface, which, by the action of the waves was
spread over them so as to be distinguishable from the general surface by a
small elevation.”

Dana, in 1852 (p. 491) published the only figure of M. inermis up to the
present time; with it, he described and figured a second species of Monolepis,
M. orientalis, from the Philippines. It was not until 1873 that Monolepis was
properly identified, when Smith (p. 67) set forth the opinion that M. inermis
was the young of M. albicans; in 1880 (p. 25), he gave evidence, based on
watching the actual moulting of a captured specimen taken in Vineyard
Sound, that this theory was indubitably correct.

In 19‘24 Rathbun fp. 155) described a megalops from the Galapagos
which she referred, with a question mark, to O. gaudichandii. Boone in 1927
(p. 168) reproduced this description, and a photograph, of the same specimen.
As I shall show later (p. 70), this Galapagos megalops should be referred
instead to the genus Plagusia.

Lebour in 1932 (pp. 18-21) questionably referred two megalopa from the
Philippines to this family; at least one of the specimens is certainly an
Ocypode and the other doubtless belongs in the family if not in that genus
(see p. 70).

Hyman (1920, pp. 496-497) has described the megalops of western
Atlantic species of Uca.

IV. The First Zoea of Ocypode gaudichaudii Milne Edwards & Lucas.

(Text-figure 1).

A. Material and Methods: A single ovigerous female was taken during
the Eastern Pacific Zaca Expedition, at Bahia Honda, Panama, on March
15, 1938. The crab was kept in an aquarium with shallow water at one end
and sand banked up at the other; it was fed bits of shrimp, mussel and fish.
Most of its time was spent half submerged in the water. The eggs were
well developed at time of capture, dark orange in color; they did not hatch,
however, until 11 days later, on March 26. Unfortunately, the prezoea was
not secured. The zoeae lived from 24 to 30 hours — no special attempt hav-
ing been made to rear them — and were preserved in 70% alcohol. There
was no difference between those killed as soon as it was discovered they had
hatched, and those that died naturally the next day. They are catalogued in
the collections of the Department of Tropical Research, New York Zoological
Society, as Nos. 38,798 and 38,818.

In the following description, Aikawa’s system of classification of
antenna and telson, and his general methods of description, are used, though
in amplified form (see Aikawa, 1929, 1933 and 1937).

B. Diagnosis: Dorsal, rostral and lateral spines present. Second an-
tenna B-4 type. Telson B-type; hair formula: endopodite of first maxilla,
4-0; of second maxilla 2-1 (3) with a slight, unequal bifurcation; of second
maxilliped, 5-0-0. Spines present on second and third abdominal somites.

C. Description: No primary chromatophores apparent (after 18
months in alcohol) ; almost all specimens lack even secondary chromato-
phores, pigment being completely absent except in the eyes.

The following measurements were made from specimens killed and pre-
served in 70% alcohol: Total length, from front to telson tip, ca. 1.6 mm;
dorsal spine .54 mm.; rostral spine .43 mm.; perpendicular distance between
their tips 1.47 mm.; lateral spine .23 mm.; abdomen 1.13 mm. long, .2 mm.
wide (at fourth segment) ; telson .34 mm. long.

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Ocypode gaudichaudii : first zoea, total length (preserved in alcohol) 1.6 mm.
A. lateral view; B, abdomen, dorsal view; C, first maxilla; D, second maxilla; E,
mandible; F. antennule and antenna; G, first maxilliped; H, second maxilliped.

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Postero-lateral region of carapace finely punctate. Dorsal and rostral
spines moderately long, the dorsal the longer; dorsal straight, rostral
slightly concave anteriorly; lateral spine a little more than half as long as
rostral, curved strongly downward.

First antenna with 2 aesthetes and 2 unequal hairs on tip. Second
antenna closest to Aikawa’s B-4 type (1933, p. 126) ; much shorter than
rostral spine but slightly longer than first antenna excluding terminal hairs;
spinous process of peduncle thick with 2 longitudinal rows of serrations ex-
tending most of its length, but stopping short of the tip; about 17 serrations
in each row; exopodite about one-fourth length of peduncle with a somewhat
squared off tip giving rise to 2 unequal terminal hairs, the longer not reach-
ing end of spinous process; flagellum rudimentary or absent.

Coxopodite of first maxilla with 4 hairs, basipodite with 5; endopodite
2-jointed, with 4 hairs on second, none on first segment. Second maxilla with
7 hairs on coxopodite, 9 on basipodite; endopodite with a slight, unequal
bifurcation, 2 hairs on larger, distal part, and 1 on smaller, proximal part;
scaphognathite with 4 hairs. Protopodite of first maxilliped with 2-2-2-2
hairs, of second with l-l-l-l ; endopodite of first maxilliped with 5-2-1-2-2
hairs, of second with 5-0-0.

Abdomen with six segments ; a pair of small spines directed anteriorly on
second segment, a similar pair directed posteriorly on third segment; fourth
segment swollen laterally in posterior portion, the postero-lateral corners
projecting backward over succeeding segment, but not actually spinous;
corners of fifth segment similarly produced to a lesser extent. Telson B-type
(i.e., with a normal, moderate fork, the horns beinjy_ without spines) ; fork
shallow, slightly more than half length of telson.

D. Comparison: The zoea of Ocypode differs from that of Uca as
described by Hyman (1920) as follows: it is larger (at least 1.6 mm. long
instead of not more than 1.0 mm.) ; lateral spines are present; the exopodite
of the second antenna is shorter, with 2 unequal terminal hairs, instead of
being long, with a short hair arising part way down its length2; the telson
is less deeply forked; the endopodite of the second maxilla is slightly but
noticeably bifurcated, the smaller, most basal lobe having 1 hair, the larger
distal lobe 2 hairs, instead of there being 3 hairs on an unbifurcated tip.
Chromatophore differences must be determined from fresh material.

If Aikawa’s convincing suggestions as to the evolutionary value of the
various characters be accepted, these differences lend support to his remarks
on the relationships of the genera of Ocypodidae (1937, pp. 154-5), showing
Ocypode as a more nearly direct link than Uca between the Macrophthal-
minae and Mictyrinae: While preserving the less specialized form of antenna
and telsons found in the Macrophthalminae, it has lateral spines hitherto
known in the family only in the Mictyrinae. However, certain knowledge
of the relationships of this most interesting family must await a great deal
of future study.

V. The Megalops of Ocypode.

(Text-figures 2-8).

A. Material and Methods: The present study of the megalops is
based on the three American species, O. albicans Bose, O. yaudichaudii Milne
Edwards & Lucas and O. occidentalis Stimpson. The first ranges from Rhode
Island to Brazil, the second from the Gulf of Fonseca, Salvador, to Chile,
and the third from Lower California to Peru.

2 This difference may prove to be slighter than it now appears, since Hyman was not certain
whether one terminal hair continued the segment of the exopod : “The outer half of the tip of the
basal portion bears the distal segment, which is small and cylindrical. From its tip arise two setae,
one long, which seems to be a continuation of the segment, and a short, outer one.’* (Hyman, 1920,
p. 491). If it is not a continuation, the antenna would be a B-4 instead of a B-2 type, according to
Aikawa’s classification (1933, pp. 126-7). There seems no doubt that the present specimens
have two, distinct, terminal setae.

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Megalopa of both Pacific species (here identified and described for the
first time) are included in the collections of the Eastern Pacific Expeditions,
as follows:

O. gaudichaudii, 8 specimens :

No. 25,205; Sta. 63 T-l; Arcturus Oceanographic Expedition; 20 mi. W. of
Mariato Point, Panama (6° 58' N. Lat., 81° 08' W. Long.); night surface haul;
May 11, 1925; 2 megalopa; carapace length’ 4.72, 4.86 mm.; breadth 4.58 mm.;
total length 5.43, 5.58 mm.

No. 38,112; Sta. 205 L-l; Eastern Pacific Zaca Expedition; Potrero Grande
Bay, Costa Rica; night light; Jan. 23, 1938; 1 megalops ; ‘ carapace length 5.58
mm.; breadth 4.72 mm.; total length 9.0 mm.

No. 38,823; Sta. 232 L-l to L-4; Eastern Pacific Zaca Expedition; Gorgona
Island, Colombia; night light; March 27-30, 1938; 5 megalopa; carapace lengths
4.86 to 5.72 mm.; breadths 4.3 to 4.86 mm.; total lengths 8.44 to 9.4 mm.

0. occidentalis , 2 specimens:

No. 37,160; Sta. 182 L-l; Eastern Pacific Zaca Expedition; Chamela Bay,
Mexico; night light; Nov. 17, 1937; 1 megalops; carapace length 4.15 mm.;
breadth 4.15 mm.; total length 7.43 mm.

No. 37,547a; Sta. 195 L-l; Eastern Pacific Zaca Expedition; Port Guatulco,
Mexico; night light; Dec. 3, 1937; 1 megalops; carapace length 5 mm.; breadth
4.29 mm.; total length 7.7 mm.

Three megalopa of O. albicans from Woods Hole, Mass., were studied
through the kindness of Dr. Waldo L. Schmitt (U- S. N. M. Nos. 10,995 and
11,175; carapace lengths 5.43 to 6.43 mm.; breadths 4.58 to 5.3 mm.; total
lengths 8.15 to 9.58 mm.). Although the megalops of this species has long
been known (see p. 000), it has never been adequately described and
figured. Hence, illustrations of this form are included in the present report.
Fortunately, one of these specimens has all the appendages tightly folded
against the body, giving the clue for many of the peculiarities of form in the
megalops of Ocypode (see below).

From a comparison of these three species, the well-marked generic
chax-acters of this stage of Ocypode become clearly evident. In spite of the
similarity of the forms, drawings of the parts of all three are reproduced,
as a basis for future work.

B. Taxonomy and Identification: As has already been remarked,
(p. 67), Monolepis inermis has been identified without question as the
megalops of Ocypode albicans. It is by analogy with this identification that
Dana’s very similar M. orientalis (1852, p. 491) and our own specimens are
referred without question to Ocypode. Megalopa of the closely related genus
JJca are, fortunately, very distinct, so that there is no question of confusion
(see below, and Hyman, 1920, pp. 496-497 and figs).

The megalopa questionably referred to Ocypodidae by Lebour (1932,
pp. 18-21) are undoubtedly members of that family, although the genera
cannot at the present time be settled. Her megalops A 1, 12 mm. long, with
only seven hairs on the last pleopod and no hairs at the bases of the third
and fourth legs, almost certainly does not belong to Ocypode; megalops A 2,
on the other hand, will very likely prove to be a member of that genus.

A reexamination of the Galapagos megalops identified questionably as
O. gaudichaudii by Rathbun in 1924, p. 155, shows that this specimen
(U.S.N.M. No. 57,735) should be referred instead to the grapsid genus
Plagusia, probably to P. depressa tuberculata, for the following reasons:

1. The spines on the tarsi are characteristic of grapsid, not ocypodid,
megalopa.

2. The antennular fossae characteristic of the Plagusiinae are clearly
marked, being more deeply incised in a straight dorsal view than the illus-
tration indicates (ibid., pi. VII, p. 154).

;>> Measured in median line.

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Text-figure 2.

Ocypode albicans, megalops, showing position of appendages when folded
against body. A, lateral view; B. ventral view. (From U. S. National Museum
specimen No. 10,995; length of carapace in median line, 6.43 mm.).

72 Zoologica: New York Zoological Society [XXV :6

3. The form of the maxillipeds is more similar to that of the adults of
Plagusa than of Ocypode.

4. There are no hairs between the bases of the third and fourth legs.
These are clearly apparent in true Ocypode megalopa.

5. Minor points such as the arrangement of hairs on the manus of the
cheliped and on the carapace are typical of Plagusia, and especially of P.
depressa tuberculata.

6. Finally, comparison with Aikawa’s description and figures of
Plagusia dentipes (1937, p. 136, fig. 36), and with Percnon megalopa in our
own collections, show the close relationship in many similarities, while the
thick cuticle, the exaggerated body depth and the sternal sculpturing which
are so characteristic of true Ocypode are lacking.

The Pacific megalopa of the present collections, listed above, are divided
without hesitation into the two Pacific species, O. gaudichaudii and O.
occidentalis, since both specimens of the latter form were taken well north
of the northern boundary of O. gaudichaudii’ s range, and exhibit distinct
specific characters in the number of setae on the last pleopods and pattern
of the sternal sculpture.

C. Comparison of Uca and Ocypode: The megalopa of Ocypode and Uca
have in common the following characters, which seem to be of the most
diagnostic value in determining families (see especially Lebour, 1928, pp. 488-
491) : Rostrum pointed, sharply bent, with a single median spine; no spines
on carapace ; 3 curved “feelers” on dactyls of fifth legs ; more than 6 hairs on
exopodites of last pleopods; hooks on basal segments of legs rudimentary or
absent. In addition they have small, 10-11-segmented antennae; small
chelipeds; external maxillipeds and other mouthparts of a general shape
typical of adult Ocypodinae; and a tuft of hairs between the bases of the
third and fourth legs, also as in the adults. There are no spines on the
dactyls of the second, third and fourth legs.

Although present material is inadequate for a complete comparison,
megalopa of these two genera may be distinguished by the following key.
The characteristics of Uca, as given in the above paragraph and in the key,
are taken from Hyman (1920, pp. 496-497).

A. More than 20 hairs on exopodite of last pleopod ; anterior part of
sternum conspicuously sculptured ; no distinct bulge on inner mar-
gin of merus of third maxilliped; cuticle thick and hard; body ex-
cessively obese, almost cylindrical; carapace grooved for reception
of appendages when folded Ocypode

AA. Less than 15 hairs on exopodite of last pleopod; anterior part of
sternum smooth; a distinct bulge on inner margin of merus of third
maxilliped; cuticle not noticeably thick and hard; body not un-
usually obese, but instead flattened dorso-ventrally as is usual in
megalopa; carapace not grooved for reception of appendages when
folded Uca

D. Generic Characters : The following description of Ocypode is based
only on the three American species.

Diagnosis. Megalops large (total length up to 9.4 mm.), extremely
obese4, heavy, deeply pigmented, with thick, hard cuticle; rostrum small, ver-
tical ; sternum sculptured anteriorly, the pattern varying with the species ;
deeply grooved posteriorly for reception of the short abdomen; a prominent
transverse, truncate, suborbital projection; chelipeds small; three curved
“feelers” on tip of fifth leg; hooks on ischium of legs rudimentary or
absent; a tuft of hairs present between bases of third and fourth legs; no
spines on dactyls of walking legs ; 22 to 28 setae on last pleopod.

Description. General color yellowish to dark brown with individual
black chromatophores large and numerous or small and sparse, varying with
the individual; carapace large, rounded dorsally, almost dome-shaped,

4 Maximum body depth almost or quite equalling carapace width.

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straight in front with small, three-pointed rostrum, the central spine
longest, triangular, sharply bent, vertical in large specimens; two frontal
eminences; frontal, gastric,, cardiac and intestinal regions sometimes with
pores, variously numbered and arranged, present or absent in individuals of
the same species, possibly connected with moulting; a groove in the cardiac
region, curving posteriorly, and a corresponding one in the intestinal
curving anteriorly; both grooves varying slightly with the individual within
the species; in the posterolateral region is a depression for the reception of
the folded fifth leg.

Lateral region of carapace very deep, vertical, provided with shallow
grooves for reception of the bent second, third and fourth legs. A promi-
nent, rounded, horn-like projection extends sidewise from the suborbital
region. Anterior sternal region sculptured with a prominent central knob,
and various adjacent smaller projections, depressions and excavations, the
pattern depending upon the species, and varying slightly with the individual ;
into and around this area fit the folded dactyls of the chelipeds and of the
second and third legs. Sternal segments well marked, a projection on the
one at the base of the second leg. Sternal groove deeply excavated for
reception of the short, thick abdomen.

Antennule (Text-fig. 7 A-C) almost concealed, with a large, swollen
basal segment containing a statocyst already well developed, and a five-
jointed terminal process composed of the second to the sixth segments. A
fringe of hairs makes a continuous line across the lower part of the basal
segment and the upper part of the second, which arises from the inner,
lower portion of the base. Anteriorly the base gives rise to two hairs.
From the third segment arises a small, single-jointed external flagellum
tipped with three or four hairs. The fourth, fifth and sixth segments bear
about 6 to 10 long, sensory hairs each; in addition the sixth segment is
tipped distally with a single stout hair. The antennule differs from that
of the megalops of Uca (Hyman, 1920, p. 496, fig. 26) chiefly in the presence
of an external flagellum, of three, not two segments in the terminal process,
and of a fringe of hairs across the basal and second segments. In the
adult Ocypode the basal segment is proportionally much larger, the external
flagellum has two, not one, joints, and the terminal process is degenerate,
almost lacking hairs and with only one segment apparent.

Antenna (Text-fig. 7 D-F) very short, similar to antenna of Uca in seg-
mentation {ibid., fig. 34), composed of three basal segments and a seven-
jointed flagellum of which the fifth segment is longest (later to be divided
into two, giving the 11 joints of the adult) ; a few sensory hairs at base
of penultimate segment and a single distal hair; one to several hairs
usually present on first two basal segments and third and fourth segments
of flagellum. Because of the fragility of the antenna, an exact hair count
and comparison of the three species studied is not possible with the present
material.

Eyes moderately large, little projecting. Mandible (Text-fig. 7 G-I)
similar to that of adult, with the cutting edge worn variously by use;
palp three-jointed, as in adult and in Uca. Maxillae (Text-fig. 7 J, K) and
maxillipeds (Text-fig. 8) similar to those of adult and of Uca {ibid., figs.
51, 60, 68, 70, 74). First maxillae with very strong bristles on basal lobes,
and a two-jointed palp. Epipodites large in first and third maxillipeds,
moderate in second. No prominent bulge on inner edge of ischium of third
maxillipeds, as there is in Uca, in both megalops and adult.

Chelipeds small, being short and only moderately broad and heavy.

Second, third and fourth legs small, fifth leg very small. All have
various grooves, excavations and bulges on the segments which enable them
to fit with extreme precision and tightness against the body and each other
when folded : The chelipeds are held against the pterygostomium region,
the second and third legs against the sides, each other and the sternum,

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B

Text-figure 3.

A. Ocypode albicans ; B. Ocypode occidentalis. Dorsal view of megalopa, total
lengths 8.15 and 7.43 mm. respectively. Short hairs on legs omitted.

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Text-figure 4.

A, Ocypode gaudichaudii. Dorsal view of megalops, total length 8.6 mm.
Short hairs on legs omitted. B, C, D, fifth legs of megalops of O. albicans, O. occi-
dentalis and O. gaudichaudii, respectively, all drawn to same scale.

the fourth leg, which is the longest, straight forward with the dactyls
hooking over the eye-stalk, and the tiny fifth legs exactly fitting into the
posterolateral grooves. The few interstices left on the ventral surface of

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B

Text-figui'e 5.

A, Ocypode albicans; B, O. occidentalis. Ventral views of megalopa in Text-fig.
3. Short hairs on legs omitted.

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Text-figure 6.

A, Ocypode gaudichaudii. Ventral view of megalops in Text-fig. 4. Short
hairs on legs omitted. B, C. D, exopodites of last pleopods of O. albicans, O. occi-
dent.alis and O. gaudichaudii, respectively, all drawn to same scale.

the megalops by the folded legs are exactly filled with the suborbital,
epistomal and sternal projections. Hooks on ischia or meri rudimentary
or absent (except in chelipeds of a Philippine specimen: see Lebour, 1934,
p. 20, fig. 18).

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Appendages of megalopa of Ocypode. A, B, C, antennule of O. albicans, 0.
occidentalis and O. gaudichaudii, respectively; D, E, F, antenna of 0. albicans, 0.
occidentalis and 0. gaudichaudii, respectively; G, H, 1, mandible of 0. albicans,
0. occidentalis and 0. gaudichaudii, respectively; J, K, second maxilla of O. albi-
cans, and O. gaudichaudii, respectively; L, M, N, first maxilla of 0. albicans, 0.
occidentalis and O. gaudichaudii, respectively. The corresponding parts of the
three species are, in each group of drawings, drawn to the same scale.

A tuft of hairs present, as in the adult, between the bases of the third
and fourth legs.

No spine^ on dactyls of second, third and fourth legs.

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Text-figure 8.

Maxillipeds of megalopa of Ocypode. A, B, C, first maxilliped of O. albicans,
0. occidentalis and O. gaudichaudii, respectively; D, E, F, second maxilliped of
O. albicans, O. occidentalis and O. gaudichaudii, respectively; G, H, I, third maxilli-
ped of O. albicans, O. occidentalis and 0. gaudichaudii, respectively. All drawn
to same scale. Number of hairs on epipodites approximate.

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Fifth legs (Text-fig. 4 B-d) with three curved feelers on last segment.
Last pleopods with 22 to 28 setae (Text-fig. 6 B-D). Telson rounded.

From the present material it is impossible to tell whether there are
one or two megalopal stages, since none has been reared. Smaller examples
within the species differ considerably from larger in minor details of cara-
pace modeling, as well as in size, and it seems likely that unlike Uca (Hyman,
1920, pp. 496-497) there are two, not one. If this is so, then both speci-
mens of O. occidentals are doubtless in the first megalops stage.

E. Specific Characters: Although, during this period, the three

American species are so similar, there are two characters by which the
forms may be easily differentiated: the number of setae on the exopodites
of the last pleopods and the form of sculpturing on the anterior part of the
sternum. These differences are best shown as follows :

O. occidentals :

Anterior part of sternum as in fig. ;
22 to 23 setae on last pleopods.

O. albicans:

Anterior part of sternum as in fig. ;
26 to 28 setae on last pleopods.

O. gaudichaudii:

Anterior part of sternum as in fig. ;
27 to 28 setae on last pleopods.

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81

Minor differences are shown in the illustrations. The mouthparts are
larger and hairier in gaudichaudii than in the other two species, as in the
adults.

F. Ecology: The outstanding characteristics of the megalops of
Ocypode are its obesity, the thickness of its cuticle and its provision for
folding the appendages tightly against the body. These peculiarities are
explained when the habits of the megalops are taken into account. The
adult lives on beaches which are usually fully exposed to a heavy surf. The
megalops described by Say (1817, the type of Monolepis inermis) and some
of the specimens taken subsequently at Woods Hole by Smith (1873 and
1880) were taken on the beach, sometimes in the act of digging a rudi-
mentary hole, apparently preparatory to moulting, or at least as protection
from the waves. Thus it seems almost certain that the moult to the first
crab stage takes place on the beach. If this is so, all the megalopal peculiari-
ties named above are easily explained by the following theory: The thick-
ness of the cuticle, the rotundity of form and the complicated arrangements
for the folding of the appendages are all protections against the pounding
of the surf and the scraping of sand in the shallows, as the megalops
literally rides to shore on the crest of a wave. Were it fragile, with the
appendages outspread, it would obviously lose most of the limbs and be bat-
tered in landing; also, it would run the risk of having the breathing
apparatus clogged by the sand swirling at the edge of the tide. Afterwards,
when the megalops has been left stranded, the thick cuticle helps conserve
vital body moisture, and, if the megalops does actually, under normal cir-
cumstances, dig at this time, such hardness is obviously essential.

All of the specimens in the present collection were taken at night
lights, usually just off shore; one, however, was captured twenty miles from
the nearest land; in all probability such megalopa never reach a beach,
and, of course, countless others must be washed up on rocky or otherwise
uninhabitable coasts. Still others, as with New England examples of 0.
albicans, are carried by currents far out of their range, where the climate
is so unsuitable that ultimate survival is impossible.

It will be interesting to learn whether other beach-dwelling ocypodids
have similar adaptations, when their megalopa are discovered. Species of
Uca which live in protected marshes and lagoons are not exposed to the same
dangers in landing at the end of the megalopal stage; the lack of these
specializations in Hyman’s (ibid.) examples of Uca are apparent.

The stomachs of five megalopa (one of occidentalis, four of gaudi-
chaudii) were examined and found to be completely empty except, in one
case, for the remains of two minute crustaceans, apparently copepods.

VI. Bibliography.

Aikawa, H.:

1929. On larval forms of some Brachyura. Rec. Ocean. Works in Japan,
Vol. II, No. 1.

1933. On larval forms of some Brachyura, Paper II: A note on indetermin-
able zoeas. Rec. Ocean. Works in Japan, Vol. V, No. 2.

1937. Further notes on brachyuran larvae. Rec. Ocean. Works in Japan,
Vol. IX, No. 1.

Beebe, W.:

1926. The Arcturus Oceanographic Expedition. Zoologica, Vol. VIII, No. 1.

1938. Eastern Pacific Expeditions of the New York Zoological Society, XIV.
Introduction, itinerary, list of stations, nets and dredges of the Eastern
Pacific Zaca Expedition, 1937-1938. Zoologica, Vol. XXIII, No. 14.

Boone, L.:

1927. Galapagos Brachyura. Zoologica, Vol. VIII, No. 4.

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Zoologica: New York Zoological Society

Dana, J. D.:

1852. Crustacea. United States Exploring Expedition during the years
1838, 1839, 18i0, 18J)1, 18U2, under the command of Charles Wilkes,
U.S.N., Vol. VIII, Part I.

Hyman, O. W.:

1920. The development of Gelasimus after hatching. Journ. Morphology ,
Vol. XXXIII, No. 2.

1922. Adventures in the life of a fiddler crab. Smithsonian Rep’t. for 1920
(1922), pp. 443-460.

Lebour, M. V.: .

1928. The larval stages of the Plymouth Brachyura. Proc. Zool. Soc. London,
1928, Part 2.

1934. Larval Crustacea (Decapoda and Stomatopoda) . Expedition S. A. R.
Prince Leopold of Belgium, Duke of Brabant, to the Extreme East
(1932). Bull Mus. Roy. d’Hist. Nat. de Belgique, Vol. X, No. 8.

Rathbun, M. J.:

1918. The Grapsoid crabs of America. Smith. Inst. Nat. Mus. Bull. 97, 1917
(1918).

1924. Brachyuran crabs collected by the Williams Galapagos Expedition,
1923. Zoologica, Vol. V, No. 14.

Say, T.:

1817. An account of the Crustacea of the United States. Jour. Acad. Nat.
Sci. Philadelphia, Vol. I, pp. 155-160.

Smith, S. J.:

1873. The megalops stage of Ocypoda. Am. Jour. Sci. & Arts, Vol. VI (3),
No. 1.

1880. Occasional occurrence of tropical and subtropical species of decapod
Crustacea on the coast of New England. Trans. Conn. Acad. Arts &
Sci., Vol. IV, No. 6.

Schaus: New Species of Heterocera

83

7.

New Species of British Guiana Heterocera.1

W. Schaus

United States National Museum,
Washington, D. C.

Introduction.

During the eight years of occupancy, by the Department of Tropical
Research, of the New York Zoological Society’s Station at Kartabo, British
Guiana, extensive collections of insects were made. These were all taken
in the quarter-mile area under intensive study — what may be called the
Guiana Jungle-zone. The first collection of Lepidoptera was reported on by
Dr. H. G. Dyer, in Zoologica, Vol. I, No. 4, 1909, pp. 125-138, where 5 new
genera and 24 new species were described.

Dr. W. Schaus has recently examined other Kartabo collections and has
written the following paper, with descriptions of 14 new species of Hetero-
cera. The types are all deposited in the collection of the United States Na-
tional Museum at Washington, D. C. — William Beebe.

New species described in this paper are the following:

Noctuidae: Pseudbarydia elipha, Argidia alonia, Argidia azania.

Lasiocampidae : Euglyphis marbodia, Tolype loisa.

Saturniidae: Hylesia derica.

Dalceridae: Ca restricta.

Limacodidae: Venadicadia cetiona.

Geometridae: Macaria barticaria, Microgonia hildonia, Dithadama ?
domarita.

Lithosiinae: Leucotmenis kaietura, Prepiella deicoluria.

Hepialidae: Phassus guianensis.

Noctuidae.

Pseudbarydia elipha sp. nov.

Female: Palpi grayish-white above, black underneath; head white; col-
lar dark gray; thorax grayish-white; abdomen above citrine drab with
grayish-white segmental lines faintly indicated toward angle segment, under-
neath whitish with darker lines. Fore wing: some whitish-gray mottling
at base of inner margin; antemedial line black, wavily inbent from costa,
and excurved on inner margin; medial space dark olive buff limited by a
black and white lunular line, almost vertical from costa; a postmedial black
line inbent on costa then outbent, lunular, and incurved beyond cell to inner
margin, inwardly edged with white; the space toward costa has the lunular
line double, also edged with white and containing some small brown spots ;
a curved double black line on discocellular ; terminal space brownish-olive
with some white scaling subterminally above inner margin; apex brown

1 Contribution No. 592, Department of Tropical Research, New York Zoological Society.

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edged below with broad black scaling and an incurved series of thick black
spots; termen with a double black line alternately broken on interspaces;
cilia whitish. Hind wing glossy deep colonial buff with purplish-brown
suffusions along inner margin; a short black streak across end of cell; a
dark brown subterminal fascia widest at costa and diminishing to anal
angle; termen with small white spots and white cilia. Wings below purplish-
gray, the veins finely darker; short dark steaks at end of cells; a subterminal
fuscous fascia crossed by black streaks; termen pinkish-gray with a ter-
minal wavy black line.

Expanse : 55 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34787, U. S. N. M.

Argidia alonia sp. nov.

Male: Palpus and throat coral red, the palpi inwardly white; head and
collar dull gray, the latter with a black medial line; thorax mostly dull
purplish with black lines; abdomen dorsally vinaceous fawn, laterally and
underneath pale rhodonite pink. Fore wing vinaceous fawn, the base of
costa darkly shaded, the base of inner margin paler, both limited by an
antemedial dentate black and white line followed by a bluish-gray, black-
edged round spot; a medial short black line across costal margin; at end of
cell a broken, finely edged black, round spot, containing some bluish-green
scales which are continued to a larger more outward spot somewhat kidney-
shaped, bluish-violet edged by a fine white line; costal edge to apex yellow-
ish, interrupted postmedially by some black scales, below it a fine white-
edged black line straight from vein to inner margin, termen partly avel-
laneous with a series from apex of small white lunular spots. Hind wing
dull purplish-lilac, the costal margin pale bluish-white; below end of cell an
irregular black-edged spot containing white scales and four black points;
a medial series of small black and white lunular and similar postmedial spots
on interspaces; along termen bluish-white spots, the cilia black. Wings
below cream buff, the markings fine, dark brown. Fore wing: a point in
cell, a faint inangled medial line, an incurved line on discocellular ; an in-
curved postmedial line fine and pinkish to end of cell, then darker and inbent
to inner margin; from it above vein 6 a line to termen, the latter Hay’s
lilac with fine dark edges. Hind wing: a faint antemedial line and a more
intense postmedial straight line, the termen as on fore wing.

The species is near Argidia hilaris Walker.

Expanse: 43 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34738, U. S. N. M.

Argidia azania sp. nov.

Female: Palpus black with some grayish scales at tip; head buckthorn
brown; thorax wood brown, the patagia with tufts of cinnamon hairs; ab-
domen dorsally at base dark olive buff, becoming cinnamon buff at anal
segment; a lateral black line; underneath, dull tawny olive; hind tibiae
with white spots.

Fore wing: basal half yellow ocher with pinkish suffusions; an ante-
medial lunular line followed in cell by a small round white spot circled with
a black line; two smaller similar spots medially at end of cell and well
separated; a broad vinaceous rufous fascia postmedially from costa at apex
to inner margin, crossed by a fine black and white line, excurved below
costa, then inbent, straight to inner margin followed by a subterminal series
of small black spots pi'eceded by white scales above inner margin ; a terminal

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Schaus: New Species of Heterocera

85

fine black line partly cut by veins; cilia red. Hind wing dull purplish-
ocher at base; a white point in cell; a narrow fascia, vinaceous rufous
medially, limited by a straight black and white line, followed by clusters of
black and white scales, then by an irregular series of small black spots;
termen dull red. The veins with long black lines. Fore wing below cinna-
mon, somewhat darker on termen; a black point in cell and a small black
and white spot on discocellular ; a fine postmedial black line outwardly
edged with white; costal edge dark purplish; a terminal black line. Hind
wing below slightly paler, the inner margin grayish; a fine white and black
line on discocellular and a fine dark medial line.

Expanse: 62 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34750, U. S. N. M.; Department of Tropical Research
No. 2084.

Lasiocampidae.

Euglyphis marbodia sp. nov.

Female: Head wood brown, also the thorax; abdomen above light cin-
namon drab with dark transverse lines edged with gray and silver scales,
underneath cinnamon drab. Fore wing grayish-pink; a black line above
inner margin connected with a fine black outcurved medial line touching a
dentate black spot at end of cell; an outangled fine black postmedial line
inbent to spot at end of cell, then somewhat wavy to inner margin; a sub-
terminal dark line outcurved from costa, then sinuous to inner margin,
inwardly bordered with the ground color; termen wood brown. Hind wing
mostly wood brown; a medial wavy black line, outwardly edged with white
at costa; a fine wavy subterminal black line; termen narrowly wood brown.
Underside of wings buffy brown without any markings.

Expanse: 60 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34739, U. S. N. M.

Tolype I oisa sp. nov.

Male: Antennae light vinaceous fawn. Head, collar and thorax white,
the patagia with pinkish-white hairs. Abdomen above pinkish-white with
grayish mottling, underneath pallid gray. Fore wing partly bluish-white;
costal margin light cinnamon drab slightly paler toward apex ; a double fine
antemedial line across cell, angled close below cell, the veins crossing the
medial space well defined; a fine line across end of cell; a double dark,
sinuous, postmedial line followed subterminally by three fuscous spots and
a dark spot at tornus; cilia white; the inner margin with some long, fine
black and cinnamon hairs. Hind wing very pale vinaceous lilac, the termen
faintly darker, the cilia white. Wings underneath whiter, the fore wing,
with indications of the dark costa and the subterminal spots.

Expanse: 20 mm.

Habitat: Kartabo, British Guiana.

Type : Cat. No. 34740, U. S. N. M.

Saturn iidae.

Hylesia derica sp. nov.

Female : Head and thorax chaetura drab. Abdomen above, benzo brown
with light brownish-drab transverse lines, underneath chaetura drab. Fore

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wing dark purple drab; costa medially pale brownish drab, and a similar
transverse fascia, within it a dark streak across end of cell; a similar sub-
terminal wavy fascia. Hind wing light vinaceous drab, the veins defined
by fine dark lines; indications of a broad postmedial fascia, the termen
narrowly dark purple drab. Wings below light purple drab, the hind wing
with a broad postmedial fascia cut by fine dark lines on veins.

Expanse: 60 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34741, U. S. N. M.

Dalceridae.

Ca restricta sp. nov.

Male: Antennae silvery olivaceous yellow; palpi yellow; thorax and base
of abdomen silvery white. Fore wing pale grayish-vinaceous ; costal margin
silvery white; faint darker shading at tornus; a black point at end of cell;
termen with short black streaks between the veins; cilia and hind wing
silvery white; a very faint terminal line; wings below silvery white.

Expanse: 21 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34742, U. S. N. M.

Limacodidae.

Venadicadia cetiona sp. nov.

Male: Palpi reddish brown; vertex with white scales; collar black;
thorax reddish with a pinkish-white lateral line; metathorax with roseate
tufts; abdomen above reddish-brown with black segmental lines, under-
neath silvery white with dark gray segmental lines. Fore wing: costa and
two longitudinal veins, also the inner margin, broadly grayish-red; the
three interspaces on anterior half of wing dark purple brown, all limited by
an outbent pinkish line from costa to termen above vein 2; apex narrowly
white, the space below it postmedially with a curved white line from costa
and some dark brown shading between the veins; a subterminal series of
small black spots on interspaces; termen narrowly reddish, the cilia roseate.
Hind wing mostly black with reddish suffusions on interspaces according to
light; cilia white or dark also according to light. Fore wing below fuscous
brown, the costal edge narrowly pinkish-white, the inner margin broadly
white toward outer margin, the termen pinkish on interspaces, the cilia
narrowly brownish. Hind wing below dull grayish thickly irrorated with
darker scales, the costa with some white shading, the cilia white. All colors
variable according to light.

Expanse: 20 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34743, U. S. N. M.

Geometridae.

Mac aria barticaria sp. nov.

Male: Antennae dark brown; palpi and frons roseate; vertex gray;
collar pale pinkish; thorax white; abdomen white with some black points.
Fore wing mostly white irrorated with short black and reddish streaks;
paired black streaks on base of costa; a fine black antemedial line, slightly
curved, more intense on inner margin; a fine wavy reddish line medially

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Schaus: New Species of Heterocera

87

starting from a red and black streak on costa; a postmedial straight black
line outwardly edged with dark violet from costa to anal angle; termen
whitish with minute black points, and subterminal dark spots, not reaching
apex; a double black terminal line. Hind wing: basal half white with a
fine wavy antemedial pinkish line, and a black point beyond cell; outer half
of wing light purplish-vinaceous; a terminal black line. Wings underneath
white, the markings of upper side more intense, the lines broader, mostly
brick red, the termen of hind wing harmonizing with the fore wing.

Expanse: 35 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34744, U. S. N. M,

Microgonia hildonia sp. nov.

Male: Palpus below roseate; head gray; the thorax pinkish-white;
abdomen dorsally with some black scales at base, otherwise pale pinkish-
white with fine darker striae and an antemedial fine reddish line followed
by a black point; a dark spot on costa at apex, from it a dark angled line
continued as a fine reddish line to inner margin postmedially ; termen yel-
lowish-white with fine darker streaks and some faint brownish spots sub-
terminally. Hind wing: the costal margin white, the wing otherwise vina-
ceous with numerous darker streaks; a black point at cell before a reddish
antemedial line; some postmedial faint dark spots and striae; a fine, dark,
terminal line. Wings below white with scattered dark scales and black
points at cells; the termen broadly yellow ocher, not reaching anal angle
on hind wing.

Expanse: 30 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34745, U. S. N. M.

Dithadama ? domarita sp. nov.

Female: Palpi reddish; vertex light gray, thorax gray, laterally black;
abdomen above partly roseate with narrow and broad fuscous lines; anal
segment grayish-white, underneath the same with fine dark segmental lines.
Fore wing largely purplish-gray; costal edge black not reaching apex with
a whitish gray streak below it to subterminal line; the basal color crossed
by a medial wavily sinuous dark brown line from close below costa to inner
margin, followed by a dark shade parallel to it; a subterminal wavy dark
brown line from costa slightly outbent, well incurved from vein 3 to inner
margin; termen lilacine gray crossed by dark veins with a terminal fine and
wavy black line. Hind wing from base to subterminal line of the color of
fore wing crossed by a short reddish-brown line near base; a short curved
line at cell, the postmedial line a continuation of the subterminal line of
fore wing; the terminal black line cut by veins. Wings below iridescent,
greenish or purplish with the lines reduced to small points.

Expanse: 22 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34746, U. S. N. M.

LlTHOSIINAE.

Leucotmenis kaietura sp. nov.

Female : Palpi black above, orange underneath ; head black with a lateral
fine orange line ; abdomen above rather broadly black with a broad lateral

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orange line, underneath dark plumbago slate. Fore wing black; a bluish-
white streak in cell and a small white streak, somewhat triangular, post-
medially. Hind wing with yellow hairs on inner margin, and a small hyaline
spot beyond cell, cut by two veins; cilia on both wings dark bluish-gray.
Fore wing below with the white streak in cell broader and more distinct.

Expanse: 25 mm.

Habitat: Kaietur, British Guiana.

Type: Cat. No. 34747, U. S. N. M.

Prepiella deieoluria sp. nov.

Female: Head white; thorax roseate gray; the metathorax with a
curved white line; abdomen above pale vinaeeous pink with traces of fine
dark segmental lines. Fore wing: basal third pallid purple drab outwardly
edged by a lunular black line with projecting lines based and a black' point
at base of costa; medial space bittersweet pink; subterminal space pinkish-
white limited by an irregular double black line connected by transverse
black lines ; termen narrowly pink with white cilia. Hind wing white with
some pale pinkish suffusions and a few black spots at apex; fore wing
below broadly black on basal third of costa; a medial curved pink space, the
subterminal curved space black without the irregular lines of the upper
side. Hind wing below pinkish-white with a black spot at apex.

Expanse: 21 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34748, U. S. N. M.

Hepialidae.

Phassus g uianensis sp. nov.

Male: Antennae fuscous. Head, collar and patagia vinaeeous rufous,
the thorax only visible as a black point. Abdomen dorsally etruscan red
with narrow darker segmental lines, underneath paler with broader seg-
mental lines. Fore wing with pale purplish-vinaceous stripes on a light
cinnamon drab background; costa narrowly fuscous; a pale triangular patch
below base of costa limited by a fine, dark inbent line to inner margin; a
pale medial space, its base on costa crossed by three wavy, fine reddish
lines; the medial dark space to inner margin crossed by black and pinkish
lines, the dark veins partly pinkish; the postmedial dark space narrow on
costa, cut by white veins, expanding toward inner margin. The black and
white line edging the triangular pale space continued to inner margin with
some black spots anteriorly, the posterior half with a black and pink line;
the apical pale space with subterminal double pale and darker lunules, toward
inner margin with irregular pinkish and black lines; termen with short,
irregular, spots and lines; cilia dark brown. Hind wing light russet vina-
ceous, paler on termen. Fore wing below dark at base becoming pinkish-
gray on terminal half with a faint medial line and a wavy postmedial line.
Hind wing below slightly paler than on upper surface.

Expanse: 55 mm.

Habitat: Kartabo, British Guiana.

Type: Cat. No. 34749, U. S. N. M.

Nigrelli & Smith: Papillary Cystic Disease of Ameiurus

89

8.

A Papillary Cystic Disease Affecting the Barbels of Ameiurus
nebulosus (Le Sueur), Caused by the Myxosporidian
Henneguya ameiurensis sp. nov.

R. F. Nigrelli & G. M. Smith

New York Aquarium; Yale Medical School and New York Aquarium
(Plates I- VIII; Text-figure 1).

Introduction.

Many of the myxosporidians are histozoic parasites of fishes and cause
only slight host tissue responses. Occasionally, lesions are produced that
eventually develop into large tumor masses (see Nigrelli & Smith, 1938).
The tissue response in such cases manifests itself by the proliferation of
fibroblastic material which forms a supportive frame-work for the develop-
ing spore masses. The end result in such growths, however, is the degenera-
tion of muscle, bone and other tissues in the path of the growing fibroblasts
and spores. It is interesting to note that in many of these myxosporidian
infections the inflammatory reaction is invariably mild and only when a
secondary bacterial or fungal infection sets in does it become severe, the
latter occurring perhaps at the time the tumors rupture to release the
mature spores.

As a general rule, histozoic myxosporidians are definitely localized by
circumscribing connective tissue. Their effects on host tissues are usually
limited to these circumscribed areas and only in certain instances do they
cause pathological changes to adjacent tissues and other organs. The present
contribution deals with an instance where not only a local phenomenon
occurs, but also immediate neighboring tissue responses develop. In the
disease to be described here, the myxosporidian infection is localized in the
connective tissue layer of the skin of the barbels of the common bullhead.
Although the parasites are encapsulated by host tissue and although they
elicit degenerative changes within these enclosed areas, nevertheless they
also induce pathological changes adjacent to the encapsulated collections of
organisms, such as hyperplasia of the overlying epithelium. The details of
these two sets of changes will be described below.

The parasites causing this papillary cystic disease belong to the genus
Henneguya Thelohan. Surprising as it may seem, it is the first myxospori-
dian species to be described from the common bullhead, Ameiurus nebulosus
(Le Sueur). This parasite, however, differs sufficiently from other species
of Henneguya to be considered new and for which the name Henneguya
ameiurensis is proposed.

Material and Methods.

The infected catfish was taken from a lake in New Hampshire during
the summer of 1938. The disease was found in a single fish and attempts to

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obtain more material were without success, although a great many fish were
examined for this purpose.

The fish was an adult, measuring 7% inches in total length. As seen
in the accompanying drawing (Fig. 1) and photograph (Fig. 2), the disease
affected the barbels, especially the two dorsal ones, and the barbel at the
angle of the mouth on the right side of the body. Externally there were
no other manifestations of the disease and the fish appeared normal in all
other respects.

The bases of the affected barbels were completely surrounded by tumor
masses (Figs. 1, 2). On closer inspection these masses had a very distinct,
irregular papillary appearance. The parts of the barbels distal to the lesions,
however, were normal in appearance.

The material was fixed in 10% formalin and numerous blocks were
prepared for histological studies. They were sectioned at 5-10 microns and
stained with hematoxylin-eosin, Giemsa’s, Mallory’s and Masson’s.

Description of Henneguya ameiurensis sp. NOV.

Vegetative Stages : The cysts containing the masses of organisms were
found in the dermal connective tissue of the barbels (Figs. 3, 4, 5). The
cysts varied considerably as to shape and in size. In certain cross sections
of the barbels as many as fifteen cysts were found, of which a few were
spherical but the majority of which were oval or irregular in shape. They
measured from .190 x .342 to .760 x 1.22 mm. In practically all instances,
the cysts were matured, containing for the most part fully developed spores.
However, pansporoblasts were present at the periphery of the cysts, but in
the majority of the cases they were fairly well advanced in their develop-
ment. In younger cysts all stages of sporogenesis were recognized (Figs.
6, 7). The pansporoblasts are disporous, although monosporous forms were
evident. Henneguya ameiurensis is polysporous.

The young sporonts are spindle-shaped organisms. Polar capsule nuclei
are present and the sporoplasm contains but a single nucleus (Text-fig. 1, d).
As the sporont becomes mature, the definite capsules are completely devel-
oped and the shell with the posterior process begins to elongate (Figs. 6,
7). The two nuclei found in the adult spore appear to be the result of a
subsequent division, for many spores were found with dividing nuclei in
the telophase stage (Text-fig. 1, a, b). According to Kudo (1926), the
sporont of many Myxsosporidia contains six nuclei, two of which form the
capsule, two for the shell and two remain as the nuclei of the sporoplasm.

It is interesting to report here that many uninucleate, irregular shaped
organisms were encountered within the cysts, scattered among the matured
spores. These have been interpreted by us as newly released sporoplasms
which will perhaps migrate from the cysts into adjacent tissues and set up
new foci of infections. Such possible migrations might also explain certain
connective and epithelial reactions encountered in these preparations.

Spores: The spores are more or less lanceolate in shape, being approxi-
mately six times as long as they are broad (Text-fig. 1; Figs. 8, 9, 10, 11).
The mature forms average about 23.3 microns in length, 4.1 microns in
width, 3.0 microns in thickness. The posterior process varies considerably in
length and thickness. The more elongated the process, the thinner it is.
However, the length measurements for this process vary from 15-41.5
microns. The process is an extension of the shell material and therefore
composed of two halves. Occasionally they may be separated along the pos-
terior tip, giving it a bifurcated appearance (Text-fig. la).

The sporoplasm is granular and in the majority of forms binucleated.
Uninucleate stages were rare in our material. A small iodinophilous vacuole
was occasionally found anterior to the nuclei but most always it was found

1940] Nigrelli & Smith: Papillary Cystic Disease of Ameiurus

91

posterior to them. At the posterior end of the sporoplasm, at about the point
where the posterior process begins, one or two deeply staining basophilic
granules are always present, (Text-fig. 1; Fig. 11). The nature of these
granules is unknown, although they have been seen and described by other
investigators (e. g., Gurley, 1888).

Text-fig. 1.

Henneguya ameiurensis sp. nov. from the barbels of the common bullhead,
Ameiurus nebulosus. a, b, c, mature spores. Note dividing nuclei in a and b.
c, uninucleate spore, d, young spore with uninucleate sporoplasm and two polar
capsule nuclei. X 1500.

Although the general shape of the shell is lanceolate, there is a char-
acteristic indentation at the level of the posterior end of the polar capsules
(Text-fig. 1; Fig. 10). The sutural ridge is faintly evident. The polar cap-
sules measure on the average about 5.4 microns in length and 1.6 microns in
width at their greatest diameter. They do not, however, fill the entire an-
terior end of the shell. The anterior end of the shell tapers slightly at the
tip, giving it a blunt conical appearance.

The staining reactions of the spores to the various techniques used may
be mentioned here. With Masson’s stain the shell, capsules and vacuole are
colored a very dark green; polar filaments and sporoplasm take on a red
color; while the nuclei are stained a deeper red. With Mallory’s stain the
shell and capsules are colored yellowish; sporoplasm is orange; nuclei and
posterior sporoplasmic granules take on a light blue color. With Giemsa’s

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stain the polar filaments take on a deep blue-green color; shell and exten-
sion are a lighter blue-green color; sporoplasm and vacuole take on a
pinkish color.

In several of the cysts, spores with three and four polar filaments and
nuclei were encountered. Similar spore abnormalities were described by
Schaferna & Jirovec (1931) for Hemneguya acerinae.

Comparisons : Henneguya ameiurensis appears to be closely related to
H. gurleyi Kudo (1910 (= Myxobolus cf. linearis Gurley, 1894), a species
found in lesions at the base of the spines of the second dorsal fin of Ameiurus
melas. However, H. ameiurensis differs from H. gurleyi in the shape of the
shell and in size measurements, two important diagnostic characters.
Ameiurus nebulosus, the common bullhead, insofar as is known to the
authors, is a new host species for which a myxosporidian infection has
been reported.

Description of the Myxosporidian Lesion in Ameiurus nebulosus.

The cross section of the barbel, including the tumor, reveals the fact
that the tumor mass arises from the skin at the base of the barbel (Figs. 3,
4). Part of the tumor is composed of numerous cystic cavities separated by
connective tissue septa (Fig. 5). These cavities contain the organisms re-
ferred to above. Each cyst is surrounded by a dense fibrous capsule vary-
ing in thickness from 6 to 16 microns. The septum separating two adjacent
cysts is formed also from fibrous tissue varying somewhat in density. This
enclosing fibrous tissue is composed of fusiform connective tissue cells, in
places hyalin, with here and there a few scattered lymphocytes. The inner
lining of the cysts is formed of very flattened fibroblastic cells. In contact
with the inner lining may be found globular or slightly flattened masses
which are the developing schizonts of Henneguya ameiurensis (Figs. 5, 6,
7). The remainder of each cavity is loosely packed with large numbers of
young and matured spores.

It is obvious that the cysts occupy the region directly below the epithe-
lium; that is, they lie in the corium or just below this layer.

All of the cysts seem to contain varying amounts of a stringy mucoid
substance in which may be found embedded small numbers of spores. Other
cysts contain, beside the mucoid material, lymphocytes, some fibroblasts and
occasional melanophores (Fig. 12). The presence of these cellular elements
invading the cavity may indicate an early healing response. Later stages of
repair were not encountered in our material. It would be interesting to
know the final fate of the cysts and their contents, for it might shed light
on the subsequent stages of the developmental cycle of these organisms.
Our recent studies on lymphocystic disease in its later stages has indicated
the probable fate of these large hypertrophied cells characteristic of the
disease by a rupture of the hypertrophied lymphocystis cell, permitting the
escape of their contents into the surrounding water (Nigrelli & Smith,
1939). Here, again, the same outcome may be effected, particularly as we
find in greatly distended cysts occasional areas of thin overlying epithelium.

As stated earlier in this paper, the finding of released sporoplasms may
indicate that reinfection of adjacent tissues occurs when these organisms
migrate from the cysts, without actual rupture of the cyst wall.

As a rule, however, the epithelium is considerably thickened over the
infected area, giving the lesion a papillary or warty appearance. The epi-
thelium is composed of many layers and contains mucus cells and giant
dermal gland cells, the latter referred to in German literature as “Kolben
Zellen” (Figs. 13, 14). Reed (1924) has called these last-named giant cells
dermal gland cells and according to him they arise from elements which he
termed “clavate” cells. Dermal gland cells are demonstrated very clearly in
our material, as can be seen in Fig. 15. They are often binucleate and stain

1940] Nigrelli & Smith: Papillary Cystic Disease of Ameiurus

93

a faint pink or yellow color with eosin and Masson’s respectively. Their
nuclei are intensely basophilic. Such cells seem to be plentiful in the normal
skin of the catfish barbel. In the infected barbel, however, they appear to be
increased in numbers, just as are the mucus and squamous cells of the over-
lying skin. Their exact function is not known.

The sensory end organs are retained in the hyperplastic skin overlying
the tumors without evidence of degeneration. They probably are not in-
creased in numbers (Fig. 16).

Melanophores are of very frequent occurrence in the papillary epithe-
lium and in places suggest a definite hyperplasia of this type of cell (Fig. 17).

Histological examination of sections of other organs and tissues of the
catfish showed their structure apparently normal in all respects.

Discussion.

Tissue responses to myxosporidian infections have been reported by
other investigators (Gurley, 1893; Hahn, 1915, Kudo, 1926, 1929; Bond,
1938; etc.). Kudo (1926, 1929) has noted and described the histo-pathology
in lesions due to Myxosporidia, and in 1929 he reported the development of
a hyperplastic growth of the epithelium of the gills in channel catfish, Ictal-
urus punctatus, infected with Henneguya exilis. In these catfish, the para-
sites are encysted on the gill lamellae, and the surrounding tissue responded
to the infection by an enormous hyperplastic growth of the adjacent epi-
thelium together with a severe inflammatory reaction. In the present case,
the inflammatory reaction is a mild one but there is a definite hyperplasia
of the epithelium overlying the subcutaneous infected regions, and such
specialized cells as melanophores, mucus and dermal gland cells found in
the epithelium are involved to a certain extent in this hyperplasia.

Summary.

1. A papillary cystic disease is described affecting the barbels of
Ameiurus nebulosus (Le Sueur).

2. The parasite producing this disease is a new species of Henneguya
Thelohan, for which the name H. ameiurensis is proposed.

3. The spores are more or less lanceolate in shape, measuring 23.3 x 4.1
x 3.0 microns. The posterior shell extension, characteristic of the genus,
varies considerably in length and thickness.

4. The binucleate condition of the mature sporoplasm appears to be the
result of division of the nucleus in a slightly earlier stage of the spore.

5. The pansporoblasts are disporous, although monosporous forms do
occur. This myxosporidian species is polysporous.

6. H. ameiurensis is the first myxosporidian parasite to be described
from the common bullhead.

7. The various tissue responses of the host to this form of myxospori-
dian infection is described.

References.

Bond, F. F.

1938. Cnidospordia from Fundulus heteroclitus Lin. Trans. Amer. Micro.
Soc. 57: 107-122.

Gurley, R. R.

1893. The Myxosporidia, or Psorosperms of Fishes, and the Epidemics Pro-
duced by Them. U. S. Comm, of Fish and Fisheries. Report for 1892.
pp. 65-304.

94

Zoologica: New York Zoological Society

[XXV :8

Hahn, C. W.

1913. Sporozoan parasites of Certain Fishes in Vicinity of Woods Hole,
Mass. Bull. U. S. Bureau Fish. 33:193-214.

Kudo, R.

1919. Studies on Myxosporidia. Illinois Biol. Monogr. 5: 1-265.

1926. On Myxosoma catostomi Kudo 1923, a Mvxosporidian Parasite of the
Sucker, Catostomus commersonii. Arch. f. Protist. 56: 90-115.

1929. Histozoic Myxosporidia found in Fresh-water Fishes of Illinois. Arch,
f. Protist. 65: 364-375.

Nigrelli, R. F. & Smith, G. M.

1938. Tissue Responses of Cyprinodon variegatus to the Myxosporidian
Parasite, Myxobolus lintoni Gurley. Zoologica. 23: 195-202.

1939. Studies on Lymphocystis Disease in the Orange Filefish, Ceratacanthus
schoepfii (Walbaum), from Sandy Hook Bay, N. J. Zoologica. 24:
255-264.

Reed, H. D.

1924. The Morphology of the Dermal Glands in Nematognathous Fishes.
Zeitschr. f. Morph, u. Anthropo. 24: 227-264.

SCHAFERNA, KAREL & OTTO JlROVEC.

1931. tjber eine neue Myxobolus — art aus dem Zander Nebst einigen Bemer-
kungen iiber Henneguya acerinae (Schroder). Zeitschr. f. Parasit.
3: 148-159.

1940]

Nigrelli & Smith: Papillary Cystic Disease of Ameiurus

95

EXPLANATION OF THE PLATES.

Plate I.

Fig. 1. Drawing of a foi-malin-fixed specimen of Ameiurus nebulosus, showing
papillary cystic disease at the bases of the dorsal barbels. Drawing by
Clare Smith.

Fig. 2. Photograph of a front view of the catfish, demonstrating the papillary
cysts of the dorsal barbels and the barbel at the angle of the mouth on
the right side.

Fig. 3. Photomicrograph showing distribution, number, size and shape of the
myxosporidian cysts in a cross-section of one of the catfish barbels.

X 50. H-E.

Plate II.

Fig. 4. Papillary cystic tumor caused by Henneguya ameiurensis arising from
the base of a dorsal barbel. X 16. H-E.

Fig. 5. Higher magnification of section shown in Fig. 3. The organisms fill the
entire cavity of each cyst. Note the vegetative masses at the periphery
of the cysts. Note also the intercystic septa. X 125. H-E.

Plate III.

Fig. 6. Photomicrograph showing the vegetative masses of Henneguya ameiuren-
sis at the periphery of the cyst. Pansporoblasts, sporonts and young-
spores can be recognized. X 350. H-E.

Fig. 7. Photomicrograph of a slightly later stage than that shown in Fig. 6.
Here sporogenesis is almost completed. X 500. H-E.

Plate IV.

Fig. 8. Photomicrograph of young spores. X 400. H-E.

Fig. 9. Fully developed spores of Henneguya ameiurensis. Posterior extensions
of the valves not in focus. X 1400. H-E.

Plate V.

Fig. 10. Higher magnification of the spores. Note outline of the shell. X 1700.
H-E.

Fig. 11. Mature spores. Spore in upper right-hand corner shows split posterior
process and posterior basophilic granule. X 1500. H-E.

Plate VI.

Fig. 12. Photomicrograph showing cysts with stringy mucoid material, lympho-
cytes, fibroblasts and isolated spores. Perhaps an early stage in healing
giant dermal gland cells X 125. H-E.

Fig. 13. Photomicrograph showing the host tissue responses. Note the thickened
epithelium, composed of many layers of epithelial cells, mucus cells and
giant dermal gland cells. X 125. H-E.

Plate VII.

Fig. 14. Higher magnification of the thickened epithelium shown in Fig. 13.
X 350.

Fig. 15. Giant dermal gland cells found in the skin of the barbel of the catfish.

A slight hyperplasia is to be noted for these cells in the infected barbel.
X 400. H-E.

96

Zoologica: New York Zoological Society

Plate VIII.

Fig. 16. Sensory end-organ in the hyperplastic skin overlying the myxospordian
tumors. No degenerative changes are to be noted for these structures.
X 350. H-E.

Fig. 17. Melanophores in the skin of infected portion of the barbels. The frequent
occurrences of these cells in the papillary epithelium suggests a
definite hyperplasia of this type of cell. X 275. H-E.

NIGRELLI & SMITH.

PLATE I.

FIG. 1.

FIG. 3.

A PAPILLARY CYSTIC DISEASE AFFECTING THE BARBELS OF AMEIURUS NEBU-
LOSUS (LE SUEUR), CAUSED BY THE M YXOSPORI Dl AN HENNEGUYA AM EIURENSI S

SP. NOV.

NIGRELLI & SMITH.

PLATE II

FIG. 4.

FIG. 5.

A PAPILLARY CYSTIC DISEASE AFFECTING THE BARBELS OF AMEIURUS NEBU-
LOSUS (LE SUEUR), CAUSED BY THE M YXOSPORI Dl AN HENNEGUYA AM EIURENSIS

SP. NOV.

N1GRELLI a SMITH.

PLATE III.

FIG. 7.

A PAPILLARY CYSTIC DISEASE AFFECTING THE BARBELS OF AMEIURUS NEBU-
LOSUS (LE SUEUR), CAUSED BY THE M YXOSPORI Dl AN HENNEGUYA A M El UR ENSI S

SP. NOV.

NIGRELLI & SMITH.

PLATE IV.

FIG. 9.

A PAPILLARY CYSTIC DISEASE AFFECTING THE BARBELS OF AMEIURUS NEBU-

LOSUS (LE SUEUR). CAUSED BY THE M YXOSPORI DI AN HENNEGUYA AM EIURENSIS

SP. NOV.

NIGRELLI St SMITH.

PLATE V.

FIG 10.

FIG. II.

A PAPILLARY CYSTIC DISEASE AFFECTING THE BARBELS OF AMEIURUS NEBU-
LOSUS (LE SUEUR), CAUSED BY THE M YXOSPORI Dl AN HENNEGUYA AM EIURENSIS

SP. NOV.

N1GRELLI & SMITH.

PLATE VI.

FIG. 13.

A PAPILLARY CYSTIC DISEASE AFFECTING THE BARBELS OF AMEIURUS NEBU-
LOSUS (LE SUEUR), CAUSED BY THE M YXOSPORI Dl AN HENNEGUYA AM El URENSIS

SP. NOV.

NIGRELLI & SMITH.

PLATE VII.

FIG. 15.

A PAPILLARY CYSTIC DISEASE AFFECTING THE BARBELS OF AMEIURUS NEBU-
LOSUS (LE SUEUR), CAUSED BY THE M YXOS PORI Dl AN HENNEGUYA AMEIURENSIS

SP. NOV.

NIGRELL! ft SMITH.

PLATE VIII.

FIG. 16.

FIG. 17.

A PAPILLARY CYSTIC DISEASE AFFECTING THE BARBELS OF AMEIURUS NEBU-
LOSUS (LE SUEUR), CAUSED BY THE M YXOSPORI Dl AN HENNEGUYA AMEIURENSIS

SP. NOV.

Hollister: Caudal Skeleton of Bermuda Fishes

97

9.

Caudal Skeleton of Bermuda Shallow Water Fishes. IV.
Order Cyprinodontes : Cyprinodontidae, Poecilidae.1

Gloria Hollister

Department of Tropical Research.

(Text-figures 1-17).

Outline.

Page

Introduction 97

Bermuda Cyprinodontes :

Cyprinodontidae : Fundulus bermudae 98

Key 99

Poecilidae: Mollienisia sphenops 101

Gambusia holbrooki 106

Lebistes reticulatus 109

Summary 112

Introduction.

This is the fourth of a series of papers dealing with the caudal skeleton
of Bermuda fishes.2 The shallow-water Iniomi of Bermuda is represented
by two families, four genera and four species. Of these, Fundulus bermudae,
a Cyprinodontidae, is a native, whereas all the Poecilidae have been intro-
duced. Gambusia holbrooki, a common species of the Eastern States from
Delaware to Florida and now thriving in Bermuda marshes, was brought
from fresh-water ponds near Washington, D. C., and planted in Pembroke
Marsh and others in May, 1928. Gambusia thrive in our Bermuda fresh-
water pools at Nonsuch as well as in the brackish marshes where they serve
to control the breeding of mosquito larvae. Dr. Henry Wilkinson of the
Medical and Health Department of Bermuda reports that, “for the most
part they have done extremely well in brackish marshes. In some places,
however, as Shelly Bay, they do not always last, but I think birds are the
difficulty there and not salinity or other chemical substances. This fish is
most adaptable. They stay near the surface, use their upturned mouth for
mosquito larvae and give birth to a multitude of live fish. About half the
Gambusia died coming here from Washington. The survivors were put
directly into the various marshes and did well everywhere except in the
marsh at Baileys Bay (which has since been filled). For this marsh a few
of them had to be adapted by gradually stepping up the salinity in a bowl.”

1 Contribution No. 593, Department of Tropical Research, New York Zoological Society.

Contribution from the Bermuda Biological Station for Research, Inc.

2 Caudal Skeleton of Bermuda Shallow Water Fishes. I. Order Isospondyli : Elopidae,

Megalopidae, Albulidae, Clupeidae, Dussumieriidae, Engraulidae. Zoologica, New York Zoological
Society, Vol. XXI, Dec. 31, 1936.

Caudal Skeleton of Bermuda Shallow Water Fishes. II. Order Percomorphi, Suborder Percesoces :
Atherinidae, Mugilidae, Sphyraenidae. Zoologica , New York Zoological Society, Vol. XXII,
Oct. 7, 1937.

Caudal Skeleton of Bermuda Shallow Water Fishes. III. Order Iniomi: Synodontidae. Zoologica ,
New York Zoological Society, Vol. XXII, Dec. 31, 1937.

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During a series of experiments to determine “Salt and Fresh-water
Viability of Fish”3 I found that Gambusia need a few days (one day was
successful in several cases) in a 50-50% solution, before thriving in pure
sea-water. Placed directly in sea-water they survive about three hours.
Later it was found that of the minnows the guppy is the least adaptable to
salt-water, needing gradual changes over a period of about a week. Fundulus
adapts itself more readily than the others to either fresh or salt water, with-
out any intermediate steps in half-and-half solutions. Recently Lebistes and
Mollienisia have been taken to Bermuda and placed in garden lily pools,
where they thrive all the year around.

For Caudal Fin Terminology, Caudal Bibliography, and method of pre-
paring specimens for this study refer to Caudal I. The length of specimens
in this paper is standard length unless otherwise stated.

The symbols used in the figures are 1C, 1st caudal vertebra; 2C, 2nd
caudal vertebra; 3C, 3rd caudal vertebra; EP, epural; 1, 2, 3, hypurals; T,
last trunk vertebra.

I am indebted to Dr. and Mrs. Carl Hubbs for a complete series of
Mollienisia sphenops which range in age from less than 24 hours to 9 weeks,
as well as adults with an exact record of their ages when they were placed
in spirits; to Curator Lee S. Crandall, Keeper Scott and Head Gardener
George Skene for adult and young Lebistes; to Dr. Henry Wilkinson for
information concerning the introduction of Gambusia to Bermuda; to
Thatcher Adams for collecting Gambusia from the Bermuda marshes; to
Dr. William Beebe, Director of this Department, and Mr. John Tee-Van,
General Associate, for their cooperation.

The drawings are by Miss Harriet Bennett and Miss Janet Wilson.

1. Fundulus bermudae Gunther.

( Text-fig. 5 ) .

Diagnostic Characters :

Caudal fin count not less than 20, usually 8 + 12.

llT 7 + 12

Caudal raylets extend anteriorly to approximately the fifth posterior
neural and haemal spines.

No modification of ventral structure of anterior caudal vertebrae
adjacent to the anal fin in male. Anal fin in male and female
similar in this oviparous fish.

5 posterior neurals and haemals elongated into the caudal contour.

Vertebral count: 12 to 14 trunk, plus 19 to 21 caudal. Total 31,
33, 35.

1 epural.

1 large fan-shaped hypural and 1 slender ventral hypural.

Material Studied.

Length.

KOH Cat. No.

Cat. No.

81 mm.

1105

25,113

76 mm.

1106

25,113

72 mm.

2226

25,112

(4) 66-61 mm.

1109

25,113

59-56 mm.

2226

25,112

(4) 54-46 mm.

1108

25,113

44 mm.

2226

25,112

(8) 42-26 mm.

1107

25,113

3 Hollister, 1934. Salt and Fresh Water Viability of Fish. New York Zoological Society
Bulletin , Vol. XXXVII, No. 6, Nov.-Dee., 1934.

Breder, 1934. Ecology of an Oceanic Fresh-water Lake. Zoologica, New York Zoological
Society, Vol. XVIII, No. 3, August, 1934. See Bibliography.

1940]

Hollister: Caudal Skeleton of Bermuda Fishes

99

C

Y

P

R

I

N

o"

D

0

N

T

E

S

Key to Caudal Fin of Bermuda Shallow Water Cyprinodontid Fishes.

(Text-figs. 1-4).

Group I CYPRINODONTIDAE

Fundulus bermudae

Caudal fin count not less than 20, usually 8 + 12.

19 7+12

Caudal raylets extend anteriorly to approximate-
ly the fifth posterior neural and haemal spines.

No modification of ventral structure of anterior
caudal vertebrae adjacent to anal fin in male.

Sub-Group A Mollienisia sphenops

Caudal fin count usually 15-16 or 16.

16 15

Median hypurals completely divided in mid-
line forming a dorsal and ventral half.

Group II POECILIDAE

Caudal fin count less than ^0.

19

Caudal raylets extend anter-
iorly to approximately the
third posterior neural and
haemal spines.

Modification of ventral struc-
ture of anterior caudal ver-
tebrae adjacent to anal fin
in male.

Division I Gambusia

Modification of anterior three
caudal vertebrae in male.
Caudal vertebrae 18 to 20.
Total vertebrae 31 to 33;
13 + 18 to 20.

Sub-Group B

Caudal fin count less than A, usually or _13.

13 13

Median hypural not completely divided in
mid-line.

Division II Lebistes reticulatus

Modification of anterior two
caudal vertebrae in male.

Caudal vertebrae 14 to 17.

Total vertebrae 26 to 29;

12 + 14 to 17.

Text-figure 4.

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Caudal Osteology.

Urostyle : Separate segments of the urostyle cannot be found in any
specimens examined which range in size from 26 mm. to 81 mm. The urostyle
is conical in shape and the blunt posterior end can be seen in some speci-
mens at the base of the fan-shaped hypural. The urostyle appears com-
pletely consolidated with the median hypural.

Uroneurals : In several of the smaller specimens a slender paired bone
extends along the dorsal surface of the fan-shaped hypural. This may be a
uroneural. In larger specimens this cannot be seen but in all there is a
marked irregularity in the shape and size of the reduced neural on the an-
terior part of the urostyle.

Text-figure 5.

Fundulus bermudae. Length 81 mm. and largest in collection. Shows solid
dorsal hypural, raylets reaching as far forward as fifth from last neural and
haemal, and wing growths on the posterior neurals and haemals which are
irregular in the different specimens. X 15.4.

Hypurals: There are two hypurals in all specimens studied. The dorsal
fan-shaped hypural is three and a half times wider on the distal margin
than the second hypural. The larger bone is median and terminal in position
and supports about ten caudal rays. There are no lines to indicate that this
large hypural is a fusion of several bones, but very young specimens are
not available. The smaller hypural is entirely ventral in position with its
base abutting the ventral surface of the cone-shaped urostyle. In all speci-
mens the size and shape of the smaller hypural is irregular. In larger fish
there is a thin wing on the anterior surface. There is also marked variation

1940]

Hollister: Caudal Skeleton of Bermuda Fishes

101

in position of the base of the hypural. In some specimens this is close to the
urostyle but in others there is considerable space between. In one large
specimen the distal half of the smaller hypural is fused with the adjacent
and ventral side of the larger hypural.

Epural: There is one epural in all of our specimens. This bone is sim-
ilar in shape and position to the ventral hypural and the two flank respec-
tively, the anterior ventral and the anterior dorsal surfaces of the median
fan-shaped hypural. The basal end of the epural varies in exact position in
the different specimens, which is also seen in the base of the hypural which
is opposite.

Caudal Fin Ray Count :

7 or 8 + 12 = 19 or 20.

7 or 8 + 12 19 or 20

2. Mollienisia sphenops (Cuvier & Valenciennes) .
(Text-figs. 6-11).

Diagnostic Characters :

Caudal fin count usually 16 or 1 more or less.

16

Mollienisia sphenops. Adult female, length 38 mm. This is typical of the
adult male and female and shows the two large median hypurals characteristic
of this species. X 27.5.

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Diagnostic Characters (continued) :

Caudal raylets extend anteriorly to approximately the third pos-
terior neural and haemal spines.

Modification of two anterior caudal vertebrae in male. Anal fin in
male specialized in this viviparous species.

3 posterior neurals and haemals elongated into the caudal contour.
Vertebral count: 12 trunk, plus 16 to 18 caudal. Total 28 to 30.

1 epural.

2 large hypurals and 1 slender hypural.

Material Studied.

Drawings and descriptions are made from Mollienisia sphenops from
Acapulco, Mexico. The key is based on this material also.

Length.

KOH Cat. No.

Text-jig. No.

(2) 1 day or less old

7 mm.

2211

7

(2)1 week old

8 & 9 mm.

2212

8

(2) 2 weeks old

10 mm.

2213

9

(1)3 weeks old

12 mm.

2214

10

(2) About 3 weeks old

12 mm.

2215

(2) About 4 weeks old

14 mm.

2216

(2) About 5 weeks old

15 mm.

2217

(2) About 6 weeks old

16 mm.

2218

(2)7 weeks old

17 mm.

2219

(2)9 weeks old

18 mm.

2220

(2) About 9 weeks old

18 mm.

2221

(5) Adults (S AC stock),
3 females

31 mm. & (2) 38 mm.

2248

6

2 males

26 mm. & 29 mm.

2248

11

Text-figure 7.

Mollienisia sphenops. 1 day old, length 7 mm. Shows delicate ossification
of the caudal area which diminishes gradually toward the distal ends of the
neurals, haemals and hypurals. The urostyle is upturned and longer than the
older specimens. The column is unsegmented. Wing growths have not devel-
oped on the posterior neurals and haemals. As in the adults, the two large
hypurals are differentiated. X 73.5.

1940]

Hollister: Caudal Skeleton of Bermuda Fishes

103

Text-figure 8.

Mollienisia sphenops. 1 week old, length 9 mm. Shows general increase
of ossification and the urostyle elongated and upturned and the column still
unsegmented. Appearance of dorsal posterior zygapophyses and wing growths
on the last ventral haemal spine and the first hypural. X 57.2.

Mollienisia sphenops. 2 weeks old, length 10 mm. The centra show the
spool-shape of the adult. The column is unsegmented and there is a continued
increase in ossification. X 50.

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Mollienisia sphenops ssp.
From Lake Peten, Guatemala.

(2) Less than 24 hours old

2232

(2)1 week old

2233

(2)2 weeks old

2234

(2)3 weeks old

2235

(1)4 weeks old

2236

(2) About 4 Y2 weeks old

2237

(1)4 weeks old

2238

(2) 5 weeks old

2239

(2) 6 weeks old

2240

(2)7 weeks old

2241

(1) About 7 weeks old

2242

(2) About 8 weeks old

2243

(2)9 weeks old

2244

(6) Adults

2249

From Subine River, Guatemala.

(2) Less than 24 hours old
(2)1 week old
(2)2 weeks old
(2) 3 weeks old

(2) Adults
(1) Adult male

2228

2229

2230

2231

“Sail-fin” variety.

From New York City — Tank-bred.

2227

2206

Caudal Osteology.

Urostyle: In adult specimens the terminal tip of the urostyle is very
much reduced and in some hidden by the basal structure of the hypurals.

Mollienisia sphenops. 3 weeks, length 12 mm. Degree of ossification and
general shape of bones the same as stages between this age and the adult.
Column still unsegmented but in a specimen slightly older the centra are
separate. X 39.8.

1940]

Hollister: Caudal Skeleton of Bermuda Fishes

105

No separate segments can be found in any growth stages. In the young fish
from one day to nine weeks of age the terminal end of the urostyle is up-
turned and prolonged. Reduced paired neurals are present in all stages and
there is variation in size and shape.

Uroneurals: There is no evidence of uroneurals.

Hypurals: There are three hypurals in young and adult specimens of all
stages — two large median bones and one slender ventral hypural. The two
large bones are approximately the same size and together form the posterior
caudal contour and support approximately twelve rays. With high magnifica-
tion a line can be found continuing anteriorly from the open median slit in
all specimens. In one small specimen there is space between the two bases
but in others the bases appear as one. There is considerable ossification in
the caudal region of the young of this viviparous species. The third hypural
is ventral in position and the basal part of the two lateral sides of the arch
overlap the base of the adjacent hypural. The thin wing of bone on the
anterior side varies in length and shape in the adult and is not present in
the very young specimens.

Epural : There is a single epural in all specimens, which resembles the
small hypural without the arch. In young specimens there is no wing-growth
present.

Caudal Fin Ray Count :

3 + 12, 3 + 13, 3 + 12 = 15, 16, 16.

3 + 13 3 + 13 2 + 13 16 16 15

In the subspecies from Lake Peten, Guatemala, the count is;

4 + 12, 5 + 12 r= 16, 17.

5 + 12 5 + 12 17 17

In addition to the increase in the anterior raylets, in most specimens
the fourth posterior neurals and haemals are prolonged into the caudal con-
tour. In none of the sphenops is this found.

Additional Characters Worthy of Note : A paper on the caudal skeleton
of cyprinodonts would hardly be complete without mention of the specialized
caudal haemals found in the male poecilids. As in Gambusia and Lebistes,

Mollienisia sphenops. Anterior caudal haemals of adult male, length 29
mm. The last trunk, or abdominal process, is followed by the first and second
caudal which are specialized in the male. The first caudal has an opening
in the tip for the insertion of the dorsal ends of the posterior interhaemals.
The two openings in the second caudal are not constant in all of the specimens.
The ribs and neurals have been omitted for clarity. X 27.5.

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[XXV :9

the anterior caudal haemals are specialized in the male. Also the total num-
ber of caudal vertebrae is identical in the males and females. But in the
adult males the anterior two caudal haemals are specialized. Each is a long
heavy bone slanting in an anterior direction as far as the dorsal ends of the
interhaemals. As in Lebistes, paired pointed projections extend posteriorly
from the lateral sides of the closed haemal arch and can overlap the next
posterior haemal process. No caudal haemal specialization can be found in
the young up to nine weeks of age and material between this age and adults
is not at hand.

3. Gambusia holbrooki (Girard).

(Text-figs. 12-14).

Diagnostic Characters :

Caudal fin count usually 12, 12, 13.

13 14 13

Caudal raylets extend anteriorly to approximately the third pos-
terior neural and haemal spines.

Modification of anterior three caudal vertebrae in male. Anal fin in
male specialized in this viviparous species.

4 posterior neurals and haemals elongated into the caudal contour.

Vertebral count: 13 trunk, plus 18 to 20 caudal. Total 31 to 33.

1 epural.

1 large fan-shaped hypural and 1 slender hypural.

Median hypural not completely divided in mid-line.

Material Studied.

Length. KOH Cat. No

(26) Males 16-25 mm. 2225

(27) Females 15-31 mm. 2225

(12) Embryos 5 mm. 2225A

(6) Dissected male haemals 2225

Caudal Osteology.

Urostyle : The urostyle is almost a perfect half centrum and conical in
shape. The posterior tip is blunt and upturned and can be seen in most
specimens at the base of the fan-shaped hypural. No separate segments of
the urostyle can be found even in the 5 mm. embryos. In these young speci-
mens the upturned tip of the urostyle is less reduced and proportionately
larger than in the full-grown fish. In both the embryos and adults the uro-
style appears consolidated with the median hypural. In embryos and adults
either one or two reduced neurals are present on the urostyle.

Uroneurcds : There is no trace of uroneurals in any of the specimens.

Hypurals : There are two hypurals present in all specimens both embryo
and adult. The large median hypural is fan-shaped and approximately five
and a half times broader on the distal margin than the ventral hypural. The
larger hypural supports about ten caudal rays. In all but a few of the
specimens there is a median slit near the base of the large hypural. The
distance that this opening extends posteriorly is variable and cannot be cor-
related with sex or age. In one of the embryos and two adults a delicate line
is seen with the high power lens which extends through the anterior base
of the hypurals to the slit-opening. The study of younger embryos might
determine whether or not this large hypural is at one time definitely divided.

Text-fig. No.

12

13

14

1940] Hollister: Caudal Skeleton of Bermuda Fishes 107

But in none of the specimens is there any indication of a division of the
distal part of the hypural. Considerable ossification exists in the caudal
region of the young of these ovoviviparous embryos. The smaller hypural is
entirely ventral in position. The bases of the two lateral sides, comparable
to the haemal arches in the anterior processes, overlap the base of the larger
hypural. In all of our specimens there is variation in the size and details of
the shape of these haemal bases.

Text-figure 12.

Gambusia holbrooki. Adult female, length 28 mm. This caudal represents
a typical adult male and female. The single fan-shaped hypural is char-
acteristic of this species. X 20.84.

Epural: There is one epural in all specimens. This bone is similar in
shape and position to the corresponding element, the ventral hypural, and
the two flank respectively the anterior ventral and anterior dorsal surfaces
of the median fan-shaped hypural. Similar to the hypural, the basal end
of the epural varies in exact position in the different specimens.

Caudal Fin Ray Count :

2 or 3 + 10 = 12 or 13.

3 -1- 10 or 11 13 or 14

Additional Characters Worthy of Note : The total number of caudal
vertebrae is identical in the males and females of this series of Gambusia
holbrooki. But in the males the three anterior caudal haemals are highly
specialized. Each one is relatively large and dagger-shaped with the tips
of the spines directed anteriorly as far as the dorsal ends of the interhaemal
spines. Extending posteriorly from the region of the closed haemal arches
are paired projections which are capable of overlapping the next posterior

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[XXV :9

Gambusia holbrooki. Embryo, length 5 mm. There is considerable ossi-
fication in the caudal area but not in the head region. The column is unseg-
mented and the centra not spool-shaped as in the adult. The fan-shaped
hypural shows no indication of separate elements. X 86.84.

haemal process. For description of the mechanical functioning of these
specialized male haemals see Collier’s4 splendid article on Gambusia afinis
afinis (Baird & Girard). In comparing the detailed drawings of both the
total skeleton and the specialized haemals of this species with the specialized
haemals in our Gambusia holbrooki,5 it is conspicuous that the anterior, or
first, caudal of Gambusia afinis afinis is more slender and simple than that
of holbrooki. It is to be assumed that Collier’s illustrations are typical, and
so this difference may be found to be a definite caudal character in the two
species. No male caudal specialization can be determined in the twelve
embryos. In the smallest adult males, (16 mm.), the anterior caudal haemals
are differentiated but small, and the general appearance of the anal fin is like
that of the young females. Although we lack stages between the 5 mm.
embryos and the 16 mm. specimens, it would seem that the development of
the specialized caudal haemals probably commences at approximately this
latter age, and earlier than the development of the prolonged stout third
anal ray characteristic of the adult male. In all specimens but two, up to
20 mm. long, there is no marked development of this ray which is, on very

4 Collier, Albert, 1936. The Mechanism of Internal Fertilization in Gambusia. Copeia, No. 1,
May 10, 1936.

5 Langer, W. Fr., 1913. Beit rage zur Morphologie der viviparen Cyprinodontiden. Morpho-
logisches Jahrbuch, 1913, vol. 47. pp. 193-307. 150 figs., 244 bibliographical references. Illustration
of the specialized haemals in the male Gambusia holbrooki.

1940]

Hollister: Caudal Skeleton of Bermuda Fishes

109

Text-figure 14.

Gambusia holbrooki. Anterior caudal haemals of adult male, length 24
mm. Shows the last trunk or abdominal process, and first three caudal which
are specialized in the male. The ribs and neurals have been omitted for
clarity. Unlike Mollienisia and Lebistes there are no openings in the haemals
excepting those for the haemal vessel. X 18.

close examination, found to be only a little stouter than the ray of the female
of similar size. But in two specimens of 20 mm. the third ray is noticeably
longer and the fourth and fifth are also prolonged. In the other males slightly
larger the anal fin is completely developed.

4. Lebistes retieulatus (Peters).

(Text-figs. 15-17).

Diagnostic Characters :

Caudal fin count usually 12, or 13.

13' 13

Caudal raylets extend anteriorly to approximately the third neural
and haemal spines and sometimes slightly beyond.

Modification of anterior two caudal vertebrae in the male. Anal
fin specialized in this viviparous species.

4 posterior neurals and haemals elongated into the caudal contour.
Vertebral count: 12 trunk plus 14 to 17 caudal. Total 26 to 29.

1 epural.

1 large fan-shaped hypural and 1 slender hypural.

Median hypural not completely divided in mid-line.

Material Studied.

Length.

KOH Cat. No.

Text-fig. No.

(3) Males

15, 18, 20 mm.

2252

(3) Females

26, 28, 31 mm.

2252

15

(4) Males

15-18 mm.

2253

(3) Females

25-27 mm.

2253

(2) Males

13 & 18 mm.

2035

(1) Female

(6) Young, 2V2 months old.

29 mm. -

2095

2nd generation living in
sea water.

8-12 mm.

2100

(4) Young, 1 and 2 days

6, 8, 10 mm.

2311

(2) Embryos

6 mm.

2309

16

(4) Dissected male haemals

2252

17

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Zoologica: New York Zoological Society

[XXV :9

Caudal Osteology.

Urostyle : The slender upturned end of the urostyle can be seen in most
of the specimens in the base of the large hypural. No separate segments can
be found either in the 6 mm. embryo or the 6, 8 and 10 mm. young. In these
small specimens the upturned end of the urostyle is proportionally larger
and longer than in the adult fish. The urostyle and the base of the large
hypural appear consolidated in specimens of all sizes. Reduced paired neurals
are present on the urostyle in all specimens and there is variation in their
size and shape.

Text-figure 15.

Lebistes reticulatus. Adult female, length 28 mm. This caudal represents
a typical adult male and female with the exception of the irregular lines on
the second hypural. The growth lines in the large hypurals are not present
in the young or the males. X 16.9.

Uroneurals : There is no trace of uroneurals in our specimens.

Hypurals : There are two hypurals present in young and adult specimens
of all lengths. The large hypural is fan-shaped and median in position and
approximately nine times broader on the distal margin than the other ventral
hypural. The larger hypural supports approximately ten caudal rays. In
most specimens there is a median slit in the basal end of the larger hypural.
The length varies in the different specimens and cannot be associated with
sex or age. In one embryo a faint line is present extending from the anterior
end of the slit through the base of the hypural, but there is no line through
the distal end of the hypural to indicate two distinct bones. Because of
negligible evidence the larger hypural is considered a single structure. In
most of the large females lines of various lengths are present radiating from
the base of the large hypural toward the distal margin. These are not seen

1940]

Hollister: Caudal Skeleton of Bermuda Fishes

111

in the young. There is considerable ossification in the caudal region of the
embryos of this viviparous species. The second and smaller hypural is ven-
tral in position and the basal part of the two lateral sides overlap part of the
larger hypural near the urostyle. There is a thin wing of bone in the adults
extending almost the full length of the anterior side of the hypural. This
varies in length and shape in all specimens.

Lebistes reticulatus. 1 day old, length 6 mm. Delicate ossification is
present in the caudal area and considerably less in the head region. The
distal tips of the spines and hypurals are unossified and the column is unseg-
mented. X 66.3.

Lebistes reticulatus. Anterior caudal haemals of adult male, length 18 mm.
Shows the last trunk process and the two anterior caudal haemal processes
specialized in the male. The first caudal spine has an opening for the insertion
of the dorsal ends of the posterior interhaemals. The ribs and the neurals
have been omitted for clarity. X 25.

Epural : There is a single epural in all specimens. This bone resembles
its counterpart, the smaller hypural, in shape and size.

Caudal Fin Ray Count :

3 + 9, 2 + 10, 3 + 10 = 12, 13.

4 + 9 3 + 10 3 + 10 13 13

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Zoologica: New York Zoological Society

Additional Characters Worthy of Note: As in Gambusia, the anterior
caudal haemals are specialized in the male. The total number of caudal verte-
brae is identical in the males and females of this collection of Lebistes reti-
culatus. But in the adult males, the anterior two caudal haemals are special-
ized. Each one is a long heavy bone and slants in a ventral and anterior
direction as far as the dorsal ends of the interhaemals. Paired, pointed
projections extend posteriorly from the lateral sides of the closed haemal
arch and are capable of overlapping the next posterior haemal process. No
male caudal haemal specialization can be found in the embryos and young
specimens, and the anal fin shows no specialization in any of these young
fish up to 12 mm. But in the smallest of the adult males, 13 mm., both
caudal haemals and anal fins are completely developed.

No gross structural differences can be found in the six young, two and
a half months old, which were the second generation reared in sea water.6

Summary.

The following facts correlate the salient similarities found in the study
of Bermuda Cyprinodontes.

Caudal fin count: Fundulus has more raylets than the poecilids: Mol-
lienisia, Gambusia , Lebistes. The total count is increased in Fundulus by
the greater number of raylets. The total count, less raylets, is nearly the
same in the four species.

Caudal raylets extend anteriorly to approximately the fifth posterior
neural and haemal in Fundulus but in the three species of poecilids the
caudal raylets only extend to about the third posterior neural and haemal
spines. In Mollienisia subspecies there is an increase in the raylets and
along with this an additional neural and haemal is prolonged into the caudal
contour.

Fundulus is oviparous and there is no modification of the ventral hae-
mals in the male. The anal fin in the male and female is similar in Fundulus.

The poecilids are viviparous and there is modification of the ventral
caudal haemals in the male, 2 in Mollienisia and Lebistes, with an opening
in the tip of the first spine for the insertion of the posterior interhaemals,
and 3 in Gambusia with no opening in the tip of the first spine.

In Fundulus the 5 posterior neural and haemal spines are elongated to
the margin of the caudal contour. In Mollienisia sphenops 3 posterior spines
are elongated and in Mollienisia sphenops subspecies, in most specimens ex-
amined, 4 posterior spines. In Gambusia and Lebistes 4 posterior spines are
elongated.

Vertebral count : The total number of trunk vertebrae in the four species
is almost the same. Fundulus 12-14, Mollienisia 12, Gambusia 13, Lebistes
12. The total number of caudal vertebrae is more variable, Fundulus having
19-21, Mollienisia 16-18, Gambusia 18-20, Lebistes 14-17.

All four species have 1 epural.

Fundulus, Gambusia and Lebistes have 2 hypurals — 1 large fan-shaped
hypural and 1 slender hypural. Mollienisia sphenops has 3 hypurals, 2 large
and 1 slender bone. No separate elements can be found in the urostyle of
any specimens of the four species studied.

- Hollister, 1934. Salt and Fresh Water Viability of Fish.

Gresser & Breder: Histology of Anoptichthys Eye

113

10.

The Histology of the Eye of the Cave Characin, Anoptichthys.
E. B. Gresser & C. M. Breder, Jr.

Department of Ophthalmology , New York University College of Medicine ,

New York Aquarium.

(Plates I-III).

Introduction.

The blind cave characin, Anoptichthys jordani Hubbs & Innes, de-
scribed in 1936, represents the first of the order Heterognathi to show loss
of ocular structure. The Eventognathi and Nematognathi both long have
been known to have cave representatives with defective or absent vision.
Since cave fishes are not exactly numerous nor spread through many of
the orders of fishes, it is interesting to find species in each of the major
divisions of the Ostariophysi. In a list presented by Hubbs (1938) in which
he names all blind fishes and cave fishes, whether blind or not, exclusive
of deep sea forms, 20 belong to the Ostariophysi, while all the rest contribute
only 27. Of the former, 19 are derived from barbeled forms, and, as Hubbs
points out, this may well have to do with their relatively numerous presence
in caves.

Eye Structure.

The eyes of Anoptichthys jordani are somewhat translucent and de-
pressed below the surface of the ectoderm. A pigment layer is apparent
by direct examination, this layer forming, as far as is evident by the eye,
a cyst wall without an anterior opening.1 As will be shown later, this is
evidenced microscopically to be an intact cyst without a pupillary space.

In a fish 56 mm. long, the gross measurements of the optic organ are
1.5 mm. vertically, 1.5 mm. horizontally, and 1.2 mm. antero-posteriorly.
The eye is definitely below the level of the skin and is over-lapped by folds
of epidermis which in some specimens continue inwardly to form a solid
stalk of epithelial tissue and in others an invagination of epithelium form-
ing a blind sac. In both formations the epithelial column is directly in
contact with a solid mass of condensed mesodermal tissue corresponding to
the cornea. The attachment of this epithelial column to the corneal layers
is within the corneal surface area and not beyond. Intervening from all
sides around the stalk is a loose mesynchymal structure.

In general the eye has recognizable definitive layers, although of poor
development. The external musculature is present and an optic nerve has
an outline. The primitive optic vesicle without question has undergone its
first invagination and at this level apparently ceased to differentiate beyond
a pigment layer and a retinal layer. At this point it may be well to
acknowledge that evidence set forth below shows the primary vesicle after
its invagination to be definite in character by the presence of the pigment

1 The distal portion of the eye is here referred to as anterior.

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Zoologica: New York Zoological Society

[XXV :10

epithelium and several layers of retinal elements, although there has taken
place a degenerative change in which the edges of the vesicle have fused,
with poor attempts at the formation of a rod and cone layer and absence
of an outer nuclear layer. It is conjectural at this stage of development
whether the force — genetic, hormonal or intracellular metabolic control —
has failed to stimulate the overlying epithelial cells to the formation of
lenticular tissue, or perhaps the epidermal column does represent this at-
tempt and is shown by its failure to progress beyond the stage of forming
a secondary vesicle. Whatever the answer may be, whether the eye stops
in development because no lens develops and produces in turn the fusion of
the poorly differentiated primary vesicle into a cyst, or vice versa, the eye
has not progressed to any stage wherein vision can be served.

The cartilaginous-walled orbital cavity contains six extraocular muscles
that probably are functionally useless, judging from the weak striations in
the fibers.

The outer mesodermal layers are the best developed, a well marked
cartilaginous sclera, cup-shaped as is characteristic of piscine eyes, with a
posterior aperture for the entrance of an optic nerve and anteriorly replaced
by a laminated translucent fibrous tunic representing the interstitial tissue
of the cornea. It is apparent from what has been said above that the
primary vesicle, although differentiated and invaginated, still forms a com-
plete cyst, thereby precluding the presence of a pupillary opening or per-
mitting the development of iris tissue as such. Some basis, however, for
the possible formation of iris tissue is present, as is evident by the pigment
epithelium and in addition the extension of a layer of small blood vessels
derived from the choroidal network. Between this layer and the interstitial
mesoderm of the corneal analogue there is a solid mass of large, light-stain-
ing cells with a faint amount of chromatin material, mesenchymal in appear-
ance, that is present in all the eyes examined. It is probable that this is a
primitive tissue of supporting nature that would have disappeared if the
eye had progressed to maturity. In an occasional eye, not in every one
examined, there has been seen a well-defined cleft, unfilled with cells, that
stimulates the imagination to conceive it as a possible anterior chamber,
inasmuch as it possesses a definite single layer of flat lining cells.

The choroidal layer is as well defined as the sclera. A definite choroidal
gland is present, with its system of blood vessels, and warrants no further
description.

The retinal pigment epithelium is the best-defined of the retinal layers
and throughout is a layer of irregular hexagonal cells within which are
massed brown melanin pigment granules. This layer, the outer part of the
optic vesicle and without question continuous with the layer of nerve ele-
ments on its inner surface, does not contain a space between the borders of
invagination but is apposed to its opposite side, forming a complete cyst.
In several of the eyes examined there has been further growth of the wall
of the optic vesicle with the formation of subsidiary cysts within the main
body, each lined with an aborted type of retina. Of the usual stratification
of the retina that is discernible, the inner nerve fiber layer is weak and thin
throughout, even at the entrance of the optic nerve where it might be ex-
pected to have a thickness of fibers. The ganglion cell layer is in the main a
single layer of cells which is neither continuous throughout nor well devel-
oped. The inner plexiform layer is found without obvious defects in its
architecture. The inner nuclear layer is definite but lacks structural delinea-
tion in its finer distribution and in the quality of its clear arrangements.
The outer plexiform layer or network is weak and thin. Both the outer
nuclear layer and the layer of rods and cones show the poorest attempts at
development. Where the one fraction of a tissue such as the rods or cones
is not to be found, it is evident that their nuclear parts would be absent.
It is not possible either from our material or present knowledge to state upon
which deficiency is predicated the non-development of the elements. In sev-

1940] Gresser & Breder: Histology of Anoptichthys Eye 115

eral areas, islands of rods with small nuclei are found, but in which no sign
of a limiting membrane between the two parts is seen. Another frequency
are areas or islands of embryonal retinal cells that have a pigment mantle
but cannot be differentiated as to type. The abortive attempts to form rods
or cones are usually found in the more peripheral parts of the cyst wall.

Only at the center of the vesicle is there any evidence of a vitreous body.
The l’etinal tissue cyst walls in most areas are apposed and only at the mid-
part are slightly separated by a faint, loose syncitium of cells representing a
vitreous mass. This tissue, in turn, has nuclear elements in a faintly gran-
ular cytoplasmic matrix that stains faintly with hemotoxylin, a primitive
type of vitreous.

The optic nerve has not been separated into fascicles and no intraneural
bundle supporting framework has been noted. Scattered throughout the
nerve stalk are large, faintly basophilic nuclei containing fine, granular ma-
terial. They are mainly round, evidently nuclei of nerve cells. Their appear-
ance and their faint cytoplasmic prolongations, which are quickly lost, sug-
gest a nerve cell of embryonic character. Nerve fibers are few in the stalk
and poorly developed.

Discussion.

A prime step in any attempt to understand the nature of these peculiari-
ties would seem to be to determine, if possible, whether these changes are
degenerative in each fish or whether they are in the nature of an arrested
development. The foregoing descriptive matter would seem to indicate
clearly that this species lacks vision because of a stoppage in development of
the eye structures at an early stage rather than a cellular inactivity at a
more advanced state.

Although we have not had very young specimens available, reports that
- they show more evident eye structure suggest, in the light of the foregoing,
that the gross appearance of the young may be accounted for by the eye
capsule being more visible from the exterior because of the thinner nature
of the overlying tissue.

The more interesting question of whether the fish lost its visual equip-
ment after entering cave waters or whether a mutation with defective vision
simply found it possible to survive under such conditions, even today is a
matter of speculation. An experimental approach would undoubtedly be of
great value in this connection.

There is evident in the literature of blind vertebrates a marked tendency
toward speculation, as is usual in cases where the evidence does not point
very conclusively in any given direction. This centers about the significance
of phylogenetic or ontogenetic factors, the entire subject of genetics in rela-
tion to mutations and about which came first — the cave environment or the
blindness. Franz (1934) sums up the school of thought in which condition-
ing by darkness allegedly results in blindness by the acquired increments
of steadily degenerating eyes both physically and physiologically, each gen-
eration adding its own amount of increment to the transmitted factor (or
loss of factor).

Eigenmann (1900) in discussing the eye of Amblyopsis, also attributes
the lack of ocular development to a genetic cause, based on comparison with
Forbesella ( Chologaster ) agassizii, through the transmissibility of accumu-
lated degenerative changes, stressing that the phylogenetic tendency is evi-
dent in the structural formation, although an irregular individual abiotrophy
ensues almost at the start.

Since fishes living in regions of low light intensity are frequently pro-
vided with large eyes, and are found in association with a lesser number
with reduced eyes, as in deep sea fishes, it should follow that at least some
large-eyed forms should be found in cave environments as representing a
stage in the gradual accommodation to life without light at all. Actually no

116

Zoologica: New York Zoological Society

such condition has been found in those forms that are true cave dwellers.
Hubbs (1938) describes a new subspecies of Cichlasoma urophthalmus, C. u.
ericymba, as having a very slightly larger eye than the other subspecies of
this form. These fish are, moreover, apparently not true cave fishes but
occupy the more or less open cenotes of Yucatan and the difference in eye
size noted by him may have nothing to do with their habitat, especially since
the variation is not nearly as great as that frequently found among closely
related fishes living in open waters, and also as the Cichlidae have not pro-
duced a single blind cave form. The pigment of this fish is only slightly
reduced, but more striking is the unusual development of the sensory pores
about the head.

The view, on the other hand, that posits a chance mutant form with an
accidental association with a cave, as is discussed by Hubbs, would seem
more reasonable in the light of modern evolutionary thought. Genetic ex-
periments crossing Anoptichthys with Astyanax, rearing several generations
of Anoptichthys in light, an examination of the food intake of wild fish from
the caves, and an assay of their vitamin A content, should go far to elucidate
the concepts involved. Until at least some of these are undertaken, it would
seem to be pointless to attempt further speculation.

References.

Eigenmann, C. H.

1900. The Blind Fishes. Biol. Lect. Woods Hole, Boston, pp. 113-126.

Franz, V.

1934. Vergleichende Anatomie der Wirbeltiere, 2, pp. 1051-1054. Springer,
Berlin.

Hubbs, C. L.

1938. Fishes from the caves of Yucatan. Carnegie Instit. Wash. Pub. (491) :
261-295, pi. IV.

Hubbs, C. L. & Innes, W. T.

1936. The first known blind fish of the family Characidae: a new genus
from Mexico. Occ. Pap. Mus. Zool. Univ. Mich. (342) : 1-7, pi. I.

EXPLANATION OF THE PLATES.

(Photomicrographs by Dr. R. F. Nigrelli)

Plate I.

Fig. 1. Anoptichthys jordani. Mid-vertical section; ocular layers differentiated
but underdeveloped, except sclera and choroid. Note absence of iris, pupil,
anterior chamber and crystalline lens.

Plate II.

Fig. 2. Anoptichthys jordani. Mid-horizontal section. Retina lacks normal archi-
tectonics; pigment epithelium fused, forming vesicle.

Plate III.

Fig. 3. Anoptichthys jordani. Horizontal section at periphery. Retinal vesicle
subdivided into two cystic spaces. An epithelial stalk connects the sub-
surface eye with epithelial surface.

Fig. 4. Anoptichthys jordani. Retinal pigment epithelium of normal appearance;
retina lacks rods and cones, and lamination of plexiform and ganglion
cell layer.

GRESSER & BREDER.

PLATE I.

THE HISTOLOGY OF THE EYE STRUCTURE OF THE CAVE CHARAC1N, ANOPTICHTH YS.

GRESSER & BREDER.

PLATE II.

THE HISTOLOGY OF THE EYE STRUCTURE OF THE CAVE CHARACIN, ANOPTICHTHYS.

GRESSER a BREDER.

PLATE

THE HISTOLOGY OF THE EYE STRUCTURE OF THE CAVE CHARACIN, ANOPTICHTHYS.

J^ebj Horfe Ecological Society

General Office: 90 Broad Street, New York City

Officers

President, W. Redmond Cross

Vice-Presidents, Alfred Ely and Laurance S. Rockefeller
Chairman, Executive Committee, W. Redmond Cross

Treasurer, Cornelius R. Agnevv
Secretary, Fairfield Osborn

Scientific Staff

Zoological $arfe
W. Reid Blair, Director

Raymond L. Ditmars, Curator of Mammals and Reptiles
Lee S. Crandall, Curator of Birds
Leonard J. Goss, Veterinarian

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H. C. Raven, Prosector
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Charles M. Breder, Jr., Director
Christopher W. Coates, Aquarist
Ross F. Nigrelli, Pathologist

G. M. Smith, Research Associate in Pathology
Homer W. Smith, Research Associate in Physiology

JOepartment of tropical 5&egearcf)

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Aquarium

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, v v ■ ° ' J

ZOOLOGICA

SCIENTIFIC CONTRIBUTIONS
OF THE

NEW YORK ZOOLOGICAL SOCIETY

VOLUME XXV
Part 2

Numbers 11-18

PUBLISHED BY THE
THE ZOOLOGICAL PARK,

July 3, 1940

SOCIETY
NEW YORK

CONTENTS

PAGE

11. Plankton of the Bermuda Oceanographic Expeditions.

IX. The Bathypelagic Caridean Crustacea. By Fenner
A. Chace, Jr. (Text-figures 1-64). 117

12. Eastern Pacific Expeditions of the New York Zoological

Society. XIX. Actiniaria from the Gulf of California.

By Oskar Carlgren. (Text-figures 1-8). 211

13. Morphological and Embryological Studies on Two

Species of Marine Catfish, Bagre marinus and
Galeichthys felis. By Daniel Merriman. (Plates
I-V; Text-figures T9). 221

14. Propagation of the Electric Impulse Along the Organs of

the Electric Eel, Electrophones electricus (Linnaeus).

By C. W. Coates, R. T. Cox, W. A. Rosenblith &

M. Vertner Brown. (Plate I; Text-figures 1-3). 249

15. Notes on the Display Forms of Wahne’s Six-plumed Bird

of Paradise. By Lee S. Crandall. (Text-figures 1-3). 257

16. Acute Hemorrhagic Gastro-enteritis in a Giant Panda.

By Leonard J. Goss. 261

17. Two New Species of Trematodes from the Deep Sea Scor-

pion Fish, Scorpaena madurensis Cuv. & Val. By Ross
F. Nigrelli. (Plate I; Text-figures 1 & 2). 263

18. Report of the Hospital and Laboratory of the New York

Zoological Park, 1939. Mortality Statistics of the So-
ciety’s Collection. By Leonard J. Goss. 269

Chace: Bathypelagic Caridean Crustacea

117

11.

Plankton of the Bermuda Oceanographic Expeditions. IX.
The Bathypelagic Caridean Crustacea.1

Fenner A. Chace, Jr.

Assistant Curator of Marine Invertebrates, Museum of
Comparative Zoology, Harvard University

(Text-figures 1-64).

Contents.

Page

Introduction 117

Systematic Discussion 119

Key to the Bermudian Families of Bathy-
pelagic Caridea 121

Family Pasiphaeidae 121

Genus Pasiphaea 122

Pasiphaea liocerca, sp. nov 122

Pasiphaea hoplocerca, sp. nov 124

Genus Parapasiphae 126

Parapasiphae sulcatifrons 126

Parapasiphae macrodactyla 128

Parapasiphae , sp. a 129

Parapasiphae , sp. b 130

Genus Leptochela 131

Leptochela bermudensis 131

Family Oplophoridae 132

Genus Acanthephyra 133

Acanthephyra purpurea 134

Acanthephyra haeckelii 140

Acanthephyra curtirostris 143

Acanthephyra stylorostrata 144

Acanthephyra acutifrons 146

Acanthephyra eximia 147

Acanthephyra brevirostris 148

Acanthephyra gracilipes, sp. nov 149

Acanthephyra, sp. ? 161

Genus Notostomus 162

Notostomus vescus 163

Notostomus compsus, sp. nov 156

Notostomus marptocheles, sp. nov 158

Notostomus miccylus, sp. nov 161

Notostomus mollis 164

Notostomus distirus, sp. nov 166

Page

Notostomus robustus 168

Notostomus perlatus ? 170

Notostomus westergreni ? 171

Genus Ephyrina 172

Ephyrina hoskynii 173

Ephyrina bifida 174

Genus Hymenodora 175

Hymenodora gracilis 175

Genus Systellaspis 179

Systellaspis braueri 180

Systellaspis debilis 181

Genus Oplophorus 184

Opiophorus spinicauda 184

Oplophorus grimaldii 187

Family Bresiliidae 189

Genus Lucaya 189

Lucaya bigelowi 189

Family Pandalidae 190

Genus Plesionika 190

Plesionika martia 190

Genus Parapandalus 192

Parapandalus richardi 192

Family Physetocaridae, fam. nov 196

Genus Physetocaris, gen. nov 196

Physetocaris microphthalma, sp. nov.... 196

General Discussion and Summary 199

Taxonomy 199

Summary of Quantitative Results 200

Geographical Distribution 201

Color of Bathypelagic Caridea 203

Luminescence 204

Food of Bathypelagic Caridea 206

Bibliography 206

Introduction.

This report deals with the caridean decapod Crustacea taken during the
course of the Bermuda Oceanographic Expeditions from 1929 to 1935 in a
roughly circular area, eight miles in diameter, whose center is at 32° 12'
north latitude and 64° 36' west longitude or nine and one-quarter miles
south-southeast of Nonsuch Island, Bermuda. The total depth of water

l Contribution No. 697, Department of Tropical Research, New York Zoological Society.

Contribution from the Bermuda Biological Station for Research, Inc.

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in this area increases from 1,000 fathoms at the inner rim of the circle to
nearly 1,500 fathoms at the outer edge. Complete data on the 1574 nets
towed will be found in Beebe (1931, 1932 and 1936).

Inasmuch as open nets were employed throughout these investigations,
any precise distributional evidence disclosed by the collection must be ac-
cepted with some reserve. Nets usually were towed, six at a time, at 100
fathom intervals, and the towing time began when all six nets reached their
designated depth and ended when the nets started to come up from that level.
Nets towed at the deeper levels, therefore, were actually fishing a good deal
longer than the time given and the catches from these deeper hauls should be
somewhat greater proportionately than those made by the shallower tows.
Although it was convenient to use as a quantitative unit the average number
of specimens taken by a one meter net in one hour of towing, it is evident
that this figure probably never represents the actual number of specimens
per towing hour at any one level or in any one month, and it should be used
with caution when comparing the relative abundance of species of bathype-
lagic Caridea in the Bermuda area with those taken elsewhere by other
expeditions. Since this figure does not mean exactly the same at different
levels, its use in computing the relative abundance of a species at different
depths fails to give a perfectly true picture of existing conditions. However,
although it varies with the depth, the number of specimens per towing hour
can be used as a basis for comparing the relative abundance of different
species at each level and distribution graphs for different species based on
this unit can be compared directly with little fear of distortion of fact.

Most of the material which forms the basis of this report was taken in
1929, 1930 and 1931; the gi'eat majority of the tows made after 1931 were
surface hauls and contained no adult Caridea. Since all but one series of
the deep water nets were towed in the daytime — most of them in the morn-
ing— the collection fails to yield much data on the diurnal migrations of
these animals; I have seen no specimens taken in Nets 224 to 229 which
were towed at night between 500 and 1,000 fathoms. In general, specimens
from several nets from the same level were put together, but in 1929 and
1930 these combined hauls seldom represented a period of more than two or
three weeks and a rough idea of the seasonal distribution of each species
could be obtained. In 1931, however, tows as much as six to eight weeks
apart were put in the same container so that it was found impractical to
derive any seasonal distribution data from this 1931 material.

After taking all of these factors into consideration, it was felt that the
greatest value in the collection was its completeness. This, the first extensive
expedition to devote its resources to the investigation of the deep water fauna
of a relatively small column of water, accumulated such a vast amount of
material that it is unlikely that future work in the Bermuda area will find
many forms not represented in this collection unless they be extremely rare
or accidental strays from another oceanic area. In the belief that the Ber-
mudas offer an ideal base for oceanographic exploration and, therefore, that
more and more work of this nature will be carried on there in the future,
it seemed advisable to make of this report a guide to the identification of the
bathypelagic caridean prawns of the Bermudian region. The known geo-
graphical distribution of many of the species indicates that the fauna is
representative of a large part, if not all, of the Sargasso Sea area as well,
and the report may therefore serve as a supplement to the excellent guide to
the more northern forms of the North Atlantic to be found in Kemp, 1910.

Wherever practical the figures have been made from Bermuda speci-
mens. In the few cases where it has been necessary to copy previously pub-
lished figures, due acknowledgment is given. Among the references dealing
with each species an attempt has been made to include, besides the reference
to the original description, all synonyms and one of the best recent references
from which a complete list of authors may be traced. The bibliography, in

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Chace: Bathypelagic Caridean Crustacea

119

the main, is made up only of those works to which reference is made in the
text, but these include most of the important papers dealing with bathpelagic
Caridea published to date.

This opportunity is taken to express my gratitude to those who have
assisted in this report. First, I wish to thank Dr. William Beebe for entrust-
ing this unique collection to me and for supplying valuable notes from life on
certain of the prawns. Dr. Stanley Kemp has been most helpful in solving
several perplexing problems both by sharing his wealth of carcinological
knowledge and by supplying specimens for direct comparison. Dr. Robert
Gurney, as an expert on larval development, has been ever willing to lend his
aid in verifying or refuting certain conclusions. Mr. John C. Armstrong,
of the American Museum of Natural History, offered many helpful sugges-
tions on methods of attacking the quantitative aspects of the report while
engaged in postgraduate work at the Museum of Comparative Zoology. My
deepest thanks go to Miss Jocelyn Crane of the Department of Tropical Re-
search of the New York Zoological Society. Miss Crane has most generously
turned over to me copious notes made from life on many of the prawns —
color notes, notes on luminescence, activity and feeding — which add im-
measurably to the value of the report; all such remarks on color and other
aspects of the living animal, unless otherwise credited, have been taken from
Miss Crane’s notes. I wish to express my sincere gratitude to Mr. M. D.
Burkenroad of the Bingham Oceanographic Foundation at Yale University
for the great amount of valuable time he gave up to sorting out this material
and sending it on to me. Acknowledgment is also hereby made of the kindness
of the Secretary of the Smithsonian Institution in permitting mention to be
made of specimens of certain species of Oplophoridae in the collections of the
U. S. National Museum.

It is unfortunate that lack of time prevents a more complete investiga-
tion of certain aspects of this collection. When there is insufficient time to
cover all angles of a study such as this, it is only natural that only those
phases of greatest interest to the individual worker are treated. It is there-
fore with the realization that the possibilities of the present collection are
far from exhausted that this report is submitted, and it is hoped that the
few problems brought to light may lead to further investigations of a similar
nature.

Systematic Discussion.

The Caridea form one of the tribes of the most primitive suborder of
the Decapoda, the Natantia or the shrimps and prawns. The suborder is
divided into three tribes; the Penaeidea, the Caridea and the Stenopidea.
The Caridea are characterized by usually having the abdomen more or less
sharply bent at the third somite, the pleura of the second abdominal somite
overlapping those of the first, the first maxillipeds with a lobe at the base of
the exopod, the third maxillipeds composed of from four to six joints — not
seven, the third pereiopods never chelate and the gills phyllobranchiate. The
simplest and most useful character for distinguishing carideans at a glance
is the overlapping of the first abdominal pleuron by the second.

Text-fig. 1 is a diagram of an ideal caridean showing the more impor-
tant structures. An attempt has been made to employ the simplest terms in
common usage where there is no confusion as to their meaning.

The standard measurement used is the carapace length, measured from
the posterior margin to the hind margin of the orbit. Although this proce-
dure has its disadvantages in that it fails to give a clear idea of the entire
size of the animal and the ratio of carapace length to abdomen is not always
constant, it is far more accurate in general than the total length. In most
Caridea it is extremely difficult to extend the abdomen to its greatest length
after the specimen has been in preservative for any length of time, and the

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Text-figure 1.

Lateral view of a caridean prawn (in part after Schmitt, 1921).

1940]

Chace: Bathypelagic Caridean Crustacea

121

length of the rostrum varies so much from one species to another that its
inclusion in the measurement only leads to greater confusion.

As a standard size in comparing one species with another it has been
found convenient to use the carapace length of young males in which the
appendix masculina has not attained its full length. It is found that this
appendix grows so rapidly that the carapace seldom increases more than a
millimeter in length from the time the appendix first appears until it at-
tains its full growth. The stage at which the appendix first appears also is
remarkably constant in specimens from one locality; whether a different
environment affects the size of the prawn at this stage remains to be seen.
Although the size of the smallest egg-bearing female may more nearly
define the actual size of the animal at maturity, a long series of specimens is
necessary to determine this measurement, whereas a single young male will
yield a fairly accurate standard figure by the first method.

It is nearly impossible to make any definite rule for the determination
of sex in Caridea. In general, the safest procedure is to look for the ap-
pendix masculina on the inner margin of the endopod of the second pleopods.
In the Oplophoridae, the family to which the greatest number of bathy-
pelagic species belong, the endopod of the first pleopods is usually sexually
modified so that this character is frequently easier to use when one becomes
familiar with its form in both sexes. There are certain species however,
scattered among several families, in which there is no appendix masculina
in the male and the endopod of the first pleopod fails to show any clear
sexual dimorphism; in such instances the abdominal pleura are usually
modified in one sex, but familiarity with the form in each species is neces-
sary in order to distinguish the sexes in these species.

Key to the Bermudian Families of Bathypelagic Caridea.

1. Exopods present on some or all pereiopods ; first two pairs of pereiopods

obviously chelate; second pereiopods with an undivided carpus 2

No exopods on any pereiopods; first pereiopods either simple or micro-
scopically chelate; second pereiopods with a segmented carpus 4

2. First two pairs of pereiopods much longer and stouter than last three

pairs Pasiphaeidae

First two pairs of pereiopods shorter than last three pairs 3

3. Third abdominal somite not produced as a broad cap over the fourth

somite; mandibles imperfectly cleft Oplophoridae

Third abdominal somite produced as a broad cap over the fourth somite;
mandibles distinctly cleft Bresiliidae

4. Integument firm ; carapace not inflated ; mandibles deeply cleft with a

three-jointed palp Pandalidae

Integument thin and fragile ; carapace inflated ; mandibles without in-
cisor process or palp Physetocaridae

Family Pasiphaeidae.

Rostrum short or absent ; in the latter case it is represented by a post-
frontal gastric spine. First two pairs of pereiopods chelate, much longer
and stouter than last three pairs, and with the carpus undivided. Exopods
present on all pereiopods, much reduced or absent on second maxillipeds.
Terminal joint of second maxillipeds normal, not applied as a strip to the
end of the preceding joint. Mandibles without a molar process and with or
without a palp.

122 Zoologica: New York Zoological Society [XXV:11

Key to the Bermudian Genera of Pasiphaeidae.

1. Fourth pereiopod much shorter than either third or fifth; size medium

or large, adults more than two inches long 2

Last three pereiopods progressively diminishing in length from the third
to the fifth; size small, less than an inch in length; rostrum arising
from frontal margin; mandible with a one-jointed palp Leptochela

2. Rostrum in the form of a post-frontal spine; mandible without a palp

Pasiphaea

Rostrum arising from the frontal margin; mandible with a two-jointed
palp Parapasiphae

Genus Pasiphaea Savigny, 1816.

Rostrum represented by a post-frontal gastric spine. Orbits poorly
defined. Fourth pereiopod shorter than fifth. Mandible without a palp.

Key to Adults of the Bermudian Species of Pasiphaea.

Carapace dorsally carinate; abdomen neither carinate nor dentate; fingers of

chelae of second periopods shorter than palm P. liocerca

Carapace not dorsally carinate in posterior half ; abdomen carinate on second
to fifth somites and dentate on fourth to sixth ; fingers of chelae of sec-
ond pereiopods longer than palm P. hoplocerca

Pasiphaea liocerca, sp. nov.

Text-figs. 2 and 3.

Type: Holotype ovigerous female, Cat. No. 31,1229, Department of
Tropical Research, New York Zoological Society; Net 1160; August 10,
1931; 800 fathoms.

Diagnosis: Carapace dorsally carinate. Abdomen smooth, not carinate.
Telson strongly forked. Gastric spine not attaining anterior margin of
carapace. First pereiopods unarmed. Fingers of chelae of second pereio-
pods not longer than palm.

Description: Carapace dorsally carinate for nearly its entire length.
Gastric tooth short, horizontal, falling short of the anterior margin of the
carapace. Branchiostegal sinus obtuse, with a small submarginal spine
anterior to it. Abdomen without a dorsal carina or armature on any somite.
The sixth somite is strongly compressed near the dorsal margin, but still
there is no trace of a carina. Telson sulcate for the entire length of its
dorsal surface and terminating in a fork which is armed with 21 barbed
spines. Eyes pale yellow in alcohol, without trace of an inner pro-
tuberance. Stylocerite spatulate and armed with a small terminal spine
which falls far short of the end of the first antennular segment. Antennal
scale convex along its outer margin and bearing a triangular terminal
spine which projects but little beyond the end of the blade. There is a
small spine on the antennal peduncle below the base of the scale. First
pereiopods without spines on any segment. The fingers of the first chelae
are about two-thirds as long as the palm. Second pereiopods armed with
11 to 13 spines on the merus and one spine at the lower angle of the carpus.
The fingers of the second chelae are very slightly shorter than the palm.

Measurements: Total length about 46 mm.; length of carapace 14 mm.
The eggs measure about 1.3 by 2.0 mm. in shorter and longer diameter.

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123

Text-figure 2.

Pasiphaea liocerca. Holotype female. X 2.16.

Color in Life: Body and appendages scarlet with the chelae and the
tips of the pleopods deeper scarlet. Eyes bronze with golden centers.
Eggs scarlet.

Remarks: The carinate carapace, smooth abdomen and strongly forked
telson distinguish this species from all but five of the described species of
Pasiphaea. Of these five, P. longispina Lenz, 1914, and P. scotiae (Stebbing,
1914) may be separated by the extremely long gastric spine; P. alcocki
(Wood-Mason, 1891), P. rathbunae (Stebbing, 1914) and P. corteziana
Rathbun, 1902, differ in having spines on the meri of the first pereiopods
and the latter have the fingers of the second chelae distinctly longer
than the palm.

In the present collection there are two young specimens of Pasiphaea
which have been tentatively assigned to this species. Each has a carapace
length of about 6.0 mm. One was taken in Net 63; May 3, 1929; 600
fathoms, and the other in Net 626 or 637; May 23 or 28, 1930; 500

Text-figure 3.

Pasiphaea liocerca. A. Antennal
scale. B. Tip of telson.

B

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fathoms. These specimens differ from the type of P. liocerca in having a
slightly more pronounced gastric spine, no discernible dorsal carina on the
posterior two-thirds of the carapace and only five spines on the merus of
the second pereiopods. Although the absence of a complete carina on the
carapace and the small number of meral spines on the second pereiopods
would seem to ally these specimens more closely to the following species,
the fact that the fingers of the second chelae are shorter than the palm
very nearly precludes that possibility. Of these two species, P. liocerca and
P. hoplocerca, the only representatives of the genus so far known from the
Bermudian area, these young specimens are best accommodated in the first,
and the differences noted between them and the type may well be due to
their immaturity.

Types: Holotype female, Cat. No. 30,1688 Department of Tropical
Research, New York Zoological Society; Nets 713, 753, 778; June-July,
1930; 700 fathoms.

One young; Net 22; April 15, 1929; 600 fathoms.

One young; Net 171; June 15, 1929; 600 fathoms.

One female, one young; Net 195; June 20, 1929; 700 fathoms.

One young; Nets 199, 205, 217, 224, 232, 238 and 248; June 21-July 4,
1929 ; 500 fathoms.

One young; Nets 258, 264, 270, 276, 282 and 289; July 5-11, 1929;
900 fathoms.

Diagnosis : Carapace not sharply carinate dorsally behind gastric tooth.
Abdomen carinate on second to fifth somites inclusive and dentate on fourth,
fifth and sixth; telson strongly forked. Gastric spine reaching slightly
beyond anterior margin of carapace. First pereiopods unarmed except for
a blunt tooth at lower end of basis; fingers of chelae of second pereiopods
much longer than palm.

Description: Carapace not sharply carinate dorsally except on the
strong gastric tooth which extends slightly beyond the anterior margin
of the carapace. Behind this tooth a blunt carina extends backward along
the dorsal mid-line and disappears at about the posterior fifth of the cara-
pace. Branchiostegal sinus well marked, forming nearly a right angle.

Pasiphaea hoplocerca, sp. nov.
Text-figs. 4 and 5.

Text-figure 4.

Pasiphaea hoplocerca. Holotype female. X 1.59.

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125

Anterior to this sinus is a strong branchiostegal spine arising from the
margin of the carapace. On the lateral surface of the carapace, the supra-
branchial ridge is obsolete; the only apparent ridges are a pair of short
ones running backward from the orbital margin. Abdomen sharply carinate
on the second, third, fourth and fifth somites and terminating in strong
posterior teeth on the fourth, fifth and sixth. At the lower posterior angle
of the sixth somite is a minute recurved spine which can be distinguished
only with high magnification. Telson dorsally sulcate for nearly its entire
length and forked at the end where it is armed with 21 barbed spines.
Stylocerite reaches very slightly beyond the first segment of the antennular
peduncle where it tapers to a point. There is a well marked spine on the
antennal peduncle below the base of the scale. The latter bears a long
outer spine reaching far beyond the end of the blade. Third maxillipeds do
not extend quite as far as the tip of the antennal scale. First pereiopods with
a blunt tooth at the lower end of the basis and no spines on the merus or any
other segments. Second pereiopods with a similar basal tooth, merus
armed with five to eight ventral spines and a single spine at the lower
distal angle of the carpus. Fingers of the first chelae slightly shorter than
the palm, those of the second one-fourth again as long as the palm.

A

E

Text-figure 5.

C

Pasiphaea hoplocerca. Paratype. A. Tip of antennal scale.
B. Tip of telson. C. End of outer branch of uropods. D. Mandible.
E. First maxilla. F. Second maxilla. G. First maxilliped. H. Sec-
ond maxilliped.

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Measurements : Total length of holotype 52.5 mm.; length of carapace
15.5 mm.

Color in Life-. Unknown.

Remarks : The four smallest specimens (carapace length 8.3 to 9.3 mm.)
differ from the type in having much less distinct abdominal carinae and
the posterior spines of the fourth and fifth somites minute, although still
visible.

Of the 34 known species of Pasiphaea, 24 apparently have no teeth on
any of the abdominal somites, seven have a tooth on the sixth somite and
one, P. orientalis Schmitt, 1931, has teeth on the third and sixth somites but
none on the fourth and fifth. P. hoplocerca can therefore be readily distin-
guished from all other described forms with the exception of the two men-
tioned below by the posterior spines on the fourth, fifth and sixth somites of
the abdomen. Two species, P. amplidens Bate, 1888, and P. longispina Lenz,
1914, have been described from specimens in which part or all of the abdomen
was missing. From the first, P. hoplocerca differs in the smaller gastric
spine and in having the fingers of the second chelae longer rather than
shorter than the palm. P. longispina apparently differs in having a very
long horizontal gastric spine similar to that in P. scotiae (Stebbing, 1914)
with which species it may prove to be synonymous.

Genus Parapasiphae Smith, 1884.

Rostrum arising from frontal margin. Fourth pereipod shorter than
fifth. Mandible with a two-jointed palp.

Key to the Bermudian Species of Parapasiphae.

1. No spines on anterior margin of lateral portion of carapace; not more

than two spines on dorsal carina of carapace 2

Antennal and branchiostegal spines present; more than two spines on
dorsal carina of carapace; fingers of second chelae shorter than palm 3

2. Dorsal carina of carapace unarmed, but sulcate anteriorly; fingers of

second chelae not longer than palm P. sulcatifrons

One or two spines on dorsal carina of carapace; fingers of second chelae
distinctly longer than palm P. macrodactyla

3. Rostrum terminating in a short spine which does not reach beyond middle

of eyestalks P., sp. a.

Rostrum a long, stout spine which reaches nearly to tips of eyes P., sp. b.

Parapasiphae sulcatifrons Smith.

Text-fig. 6.

Parapasiphae sulcatifrons Smith, 1884, p. 384, pi. 5, fig. 4, pi. 6, figs. 1-7 ;
Stephensen, 1935, p. 34.

Diagnosis : Dorsal carina of carapace unarmed above, but the anterior
fourth is broadened and provided with a well marked groove in the adult.
Rostrum attains middle of eyestalk. No spine on anterior edge of lateral
portion of carapace. Abdomen with a small median spine at end of fourth
somite. Fingers of second chelae not longer than palm.

Measurements : The smallest recognizable male specimen, in which the
appendix masculina of the second pleopods is no more than a bud, has a
carapace length of 16.0 mm. The largest male has a carapace length of 22.5

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127

mm. and the appendix masculina is about two-thirds as long as the other
appendix. The ovigerous females range in carapace length from 22 to 26
mm., but only one specimen has a carapace length of less than 23 mm.,
whereas the females without eggs have a carapace length of 16 to 23 mm.
and only one is more than 22 mm. Most of the young are little more than
post-larvae, having a carapace length of only five or six mm.

Color in Life: General color scarlet, with the carapace and chelae
brighter than the rest. Eyes light amber to bronze. Antennae salmon-orange
and the other appendages like the body. Eggs scarlet with the developing
embryo salmon-orange.

Text-figure 6.

Parapasiphae sulcatifrons. Male taken in September, 1929,
from 700 fathoms. X 1.66.

General Remarks: The collection contains 497 specimens of this species
but only 36 are adult; 10 males and 26 females of which 14 are ovigerous.
Whether this is merely indicative of the normal mortality of the young,
whether proportionately larger numbers of adults might be expected in
greater depths or whether some other factor is involved cannot be ascertained
from the present material. P. sulcatifrons is one of the few bathypelagic
prawns in which the stomach is occasionally everted upon being brought to
the surface. Three specimens of this species were found in the stomachs of
fish ; one fi'om the stomach of Photostomias guernei and two from Gonostoma
elongatum.

Seasonal Distribution: Obviously the number of adults is too small to
supply any information on the seasonal abundance of this form. It may be
worthy of mention that ovigerous females were taken in April, June, July,
August and September and attention is called to the fact that all but one of
the female specimens having a carapace length of more than 22 mm. were
carrying eggs. The number of immature and young, likewise, is not sufficient
to warrant any definite conclusions. However, a comparison of the number
of young taken in 1929 and 1930 with the towing hours at each depth indi-
cates that the smallest number of specimens is found from April to June,

128 Zoologica : New York Zoological Society [XXV: 11

with an increase in July and August and a very slight falling off again in
September.

Vertical Distribution: Of the 36 adult specimens two were taken in 600
fathoms, four in 700, eight in 800, twelve in 900 and ten in 1,000. Compared
with the towing hours at each depth and realizing that open nets were em-
ployed, the center of abundance appears to be between 800 and 900 fathoms.
The immature and young were found between '500 and 1,000 fathoms with
two specimens being taken in 400 fathoms. Although there may be some sea-
sonal shift in the vertical range of this population, the center of abundance
seems to be between 600 and 800 fathoms, or somewhat nearer the surface
than is the case for the adults.

Geographical Distribution: The known distribution of the species in-
cludes the following areas; off the east coast of the United States from 35°
to 41° north latitude, Davis Straits, between Greenland and Iceland, south
of Iceland, southwest of Ireland, the Bay of Cadiz and, finally, off Cape
Point, Cape Colony, South Africa, according to Stebbing, 1914a. This dis-
tribution would suggest that P. sulcatifrons is not a true Sargasso Sea form,
but it is dangerous to form such conclusions until more is known of the
caridean fauna of the Sargasso area.

Parapasiphae macrodactyla Chace.

Text-fig. 7.

Parapasiphae macrodactyla Chace, 1939, p. 33.

Diagnosis : Dorsal carina of carapace armed with one or two teeth above
the orbit. Rostrum reaches as far as the cornea. No spine on anterior edge
of lateral portion of carapace. Abdomen with a median spine at end of
fourth somite. Fingers of second chelae distinctly longer than palm.

Parapasiphae macrodactyla. Holotype female from off the coast
of Cuba. X 1.88.

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129

Measurements : The type female from off Cuba has a carapace length of
24.8 mm. The male in the present collection has a carapace length of 19.0
mm. and the female, 21.3 mm. Similar measurements in the young range
from 4.8 to 11.0 mm. In the male the appendix masculina is less than half as
long as the other appendix.

Color in Life : Unknown.

General Remarks : There are 30 specimens of this species in the col-
lection, one male, one female and 28 young. P. macrodactyla is easily dis-
tinguished from the preceding species, even in the smallest specimens, by
the presence of one or two spines at the base of the rostrum and, particularly,
by the much longer fingers of the second chelae.

Seasonal Distribution: In 1929, fifteen specimens were captured, of
which nine were taken in September and the remaining six in May, June,
July and August. Of the four specimens taken in 1930, three were found in
September and the other in June. In 1931, eleven specimens were captured
but were separated in such a way that monthly totals could not be obtained.
The adult male was taken in June, 1930, and the female in September, 1929.

Vertical Distribution : Both adult specimens were found in 1,000 fath-
oms. Of the young, three were taken in 600 fathoms, ten in 700, four in 800,
five in 900 and six in 1,000.

Geographical Distribution: Aside from the present specimens, P. mac-
rodactyla is known only from the type taken off Bahia de Guantanamo, Cuba.

Parapasiphae, sp. a.

Text-fig. 8.

Diagnosis: Dorsal carina of carapace armed with about six small, un-
equal spines in its anterior half. Rostrum short, terminating in a small
spine which falls far short of the tips of the eyes. Antennal and branchio-
stegal spines present. Abdomen carinate on third and fourth somites, the
latter armed with a large, sharply carinate spine overhanging the fifth
somite. Fingers of second chelae distinctly shorter than palm.

Text-figure 8.

Parapasiphae, sp. a.
A. Lateral view of
carapace. B. Chela of
second pereiopod.

B

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Remarks : This form is represented by a single young specimen having
a carapace length of about 8 mm., taken in Net 819, 844, 862 or 867, August
29 to September 10, 1930, from 800 fathoms. It is apparently related to
P. serrata Rathbun, 1902, (p. 904) and 1904 (p. 25, fig. 7), from off the
coast of southern California, as indicated by the form of the rostrum, the
tooth on the fourth abdominal somite and the proportionate lengths of the
palm and fingers of the second chelae.

Parapasiphae, sp. b.

Text-fig. 9.

Diagnosis: Carapace denticulate on anterior half of dorsal carina and
drawn out to a slender, pointed rostrum which reaches nearly to the tips of
the eyes. Antennal and branchiostegal spines present. Fourth abdominal
somite armed with a small tooth or spine. Fingers of second chelae some-
what shorter than palm.

Text-figure 9.

Parapasiphae, sp. b. A.

Carapace of specimen
from net 1502. X 11.

B. Chela of second per- A

eiopod of specimen
from net 657.

B

Remarks: Two specimens, one having a carapace length of about 6.5
mm. from Net 657, June 2, 1930, 700 fathoms, the other with a carapace
length of about 5.0 mm. from Net 1502, July 25, 1934, 500 fathoms, are
assigned to this indeterminable species. The larger specimen is moulting
but seven distinct spines, exclusive of the long, slender rostrum, can be seen
on the anterior half of the dorsal margin of the carapace. This specimen also
has a distinct spine on the fourth abdominal somite. The smaller figured
specimen has a similar rostrum but the spines behind it are much less distinct
and the spine on the fourth abdominal somite is scarcely visible. The form
of the rostrum and the denticulate carina of the carapace recalls similar
structures in P. gilesii Wood-Mason, 1893 (p. 166), and 111. Investigator
(pi. 3, fig. 8) from the Indian Ocean.

1940] Chace: Bathypelagic Caridean Crustacea 131

Genus Leptochela Stimpson, 1860.

Rostrum arising from frontal margin. Last three pereiopods similar,
diminishing in length from third to fifth. Mandible with a palp which
consists of a single joint. Adults usually less than an inch in length.

There is but one species of the genus known from the Bermuda area.

Leptochela bermudensis Gurney.

Text-fig. 10.

Leptochela bermudensis Gurney, 1939, pp. 426-433, figs. 1-10.

Diagnosis'. Orbital margin smooth, not denticulate. Abdomen not car-
inate on fourth or fifth somites; telson with one pair of dorsal and one pair
of lateral spines.

Measurements'. The two males in the collection have a carapace length
of 3.2 and 4.5 mm. The females have corresponding measurements of 2.9 to
3.2 mm., both ovigerous females having the carapace 3.2 mm. long. In
the young the carapace is from 1.3 to 1.8 mm. long.

Color in Life : Integument transparent, showing the pinkish or brownish
heart and other whitish internal organs. Bases of all legs and middle of
second segments of pleopods bright spectrum red; in one ovigerous female
these markings were pure scarlet instead of spectrum red. All appendages
otherwise transparent with a faint violet tinge on the distal segments. In
one lot, faint scarlet markings were noticed on the maxillipeds. Eyes deep,
dull violet-blue. Eggs orange-pink.

A number of these prawns were kept alive in a refrigerator for about
eighteen hours. At about the tenth hour, lateral spectrum red spots ap-
peared on the abdomen, the violet tinge became more pronounced and in the
larger specimens the internal organs appeared greenish-yellow.

General Remarks : Since there are only sixteen specimens in the collec-
tion, two males, six females, of which two are ovigerous, and eight young,
it is probable that the species is found only sporadically in the Bermuda area.
There also is a good indication that swarming occurs in this species, for
Gurney obtained no less than 45 adult individuals from a single net. As in
other species of the genus, ovigerous females have three dorsal carinae on
the carapace, whereas males and females without eggs have only the median

Text-figure 10.

Leptochela bermudensis. Male from net 989. X 6.

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carina developed for the entire length. Of the specimens kept alive in the
refrigerator, a few of the smaller ones became very lively after their periodic
removal from the ice, but the larger specimens were at all times sluggish
and feeble.

Luminescence : The following notes were made by Miss Crane on speci-
mens taken in Net 1028; June 20, 1931; surface at night.

“These abundant small shrimps seemed to be the source of most of the
luminescence in this net (save for the minute sparks furnished by very
small copepods). They emitted, both when isolated and when in the company
of their kind, short, tiny flashes of bright white light. The part of the body
from which these flashes came could not be determined. Dissection was not
particularly helpful, as no structures were found which resembled typical
light-organs of any kind I have yet observed : the best possibility seems to be
that the pigment spots, seemingly superficial, at the bases of the pleopods,
are in reality photophores.

“Although a couple of specimens were kept alive until the following
evening by frequently changing the water and by alternately placing them in
and removing them from the refrigerator, no luminescence whatever was
produced even by tapping the glass and directly prodding them. They were
dead the following morning.’’

Seasonal Distribution : The type specimens, of which twelve were oviger-
ous, were taken on June 14, 1938. In the present collection one male and
three females were captured on September 25, 1930; one female on September
26, 1930; one male, one ovigerous female and five young on June 3, 1931;
three young on June 20, 1931; and one ovigerous female on November 2, 1931.

Vertical Distribution : There is evidence of a marked diurnal migration
in this species. Between one and two P.M., specimens were found at 700
fathoms; between nine-thirty A.M. and one P.M. they were taken in 400
fathoms; and three lots were taken in surface nets between seven-thirty and
eight P.M., between eight-thirty and nine P.M. and between nine-thirty and
ten P.M. Apparently L. bermudensis is normally found at or near the surface
at night and sinks to considerable depths during the hours of sunlight.

Geographical Distribution : According to our present knowledge, L.
bermudensis is known only from the Bermuda region.

Family Oplophoridae.

Rostrum always present but extremely variable in form. First two pairs
of pereiopods chelate, usually shorter than the last three pairs and with the
carpus undivided. Last three pairs not abnormally long. Exopods present
on the maxillipeds and on all of the pereiopods, but there is no lash on the
exopod of the first maxilliped. Terminal joint of second maxilliped applied
as a strip to the end of the preceding joint. Mandibles imperfectly cleft and
provided with a palp.

Key to the Genera of Oplophoridae.

1. Exopods of at least the third maxillipeds and first pair of pereiopods
foliaceous and often rigid; outer margin of antennal scale usually
armed with a series of spines ; telson not truncate at tip, but ending in

a sharp point; eyes large and well pigmented Oplophorus

None of the exopods of the pereiopods foliaceous or rigid 2

2. Last four abdominal somites, at least, dorsally carinate 3

Sixth abdominal somite not dorsally carinate 4

1940]

Chace: Bathypelagic Caridean Crustacea

133

3. Usually no straight ridge or carina running entire length of lateral sur-

face of carapace along median lateral line ; hind margin of hepatic
furrow not usually cut off abruptly by an oblique ridge or carina.
Incisor process of mandible toothed for entire length of the cutting

edge Acanthephyra

Carapace provided with at least one straight carina traversing the lateral
surface from orbit to hind margin of carapace ; hind margin of hepatic
furrow abruptly cut off from branchial region by an oblique carina.
Anterior half of cutting edge of, incisor process of mandible unarmed
Notostomus

4. Ischial and meral joints of pereiopods abnormally broad and flat Ephyrina

Pereiopods normal 5

5. Eyes very small and poorly pigmented ; anterior margin of first abdominal

somite entire, not toothed; telson terminating in a truncate, spinose

tip Hymenodora

Eyes very large and well pigmented ; anterior margin of first abdominal
somite armed with a distinct lobe or tooth overlapping hind margin of
carapace; telson terminating in a sharp-pointed end-piece laterally
armed with spines Systellaspis

Genus Acanthephyra A. Milne Edwards, 1881a.

Carapace usually without a straight ridge or carina running along the
median lateral line from orbit to hind margin, and without an oblique ridge
or carina delimiting the hind margin of the hepatic furrow. Abdomen car-
inate on the last four somites, at least. Telson more or less truncate at the
tip, not terminating in a sharply pointed end-piece laterally armed with
spines. Pereiopods not abnormally broad and flattened. Exopods of third
maxillipeds and pereiopods neither foliaceous nor rigid. Eyes well pigmented
even when small. Outer margin of antennal scale not armed with a series of
spines. Incisor process of mandible toothed for the entire length of its
cutting edge.

Key to the Bermudian Bathypelagic Species of Acanthephyra.

1. Second somite of abdomen dorsally carinate; integument firm 2

Second somite of abdomen not dorsally carinate; integument membran-
ous V

2. Carapace not dorsally carinate on posterior third 3

Carapace dorsally carinate throughout its length 6

3. Branchiostegal spine supported by a very short carina 4

Branchiostegal spine supported by a long carina which extends back to

posterior half of branchial region 5

4. No posterior spine on fourth abdominal somite; telson armed with about

four pairs of lateral spines A. purpurea

A posterior spine on fourth abdominal somite; telson armed with eight
to eleven pairs of lateral spines A. haeckelii

5. Rostrum triangular and drawn out to a sharp point; telson armed with

six to fifteen pairs of lateral spines A. curtirostris

Rostrum very convex dorsally so that it appears somewhat semicircular
in outline; telson armed with about three pairs of lateral spines
A. stylorostrata

134 Zoologica: New York Zoological Society [XXV: 11

6. First abdominal somite dorsally carinate; telson obscurely sulcate dor-

sally A. acutifrons

First abdominal somite not dorsally carinate; telson obscurely ridged
dorsally A. eximia

7. Third abdominal somite armed with a large, fleshy spine which usually

overreaches fourth somite A. brevirostris

Spine of third abdominal somite, if present, not abnormally large 8

8. A spine on third abdominal somite A. gracilipes

No spine on 1 bird abdominal somite A., sp. ?

Acanthephyra purpurea A. Milne Edwards.

Text-figs. 11-17.

Acanthephyra purpurea A. Milne Edwards, 1881, p. 933; Murray &
Hjort, 1912, pi. 3, fig. 2, (col. fig.) ; Stephensen, 1923, p. 44; Kemp, 1939,
p. 576.

Diagnosis: Integument firm. Rostrum long and slender. Carapace not
dorsally carinate posteriorly. Branchiostegal spine supported by a short
carina. Abdomen carinate on all but the first somite and with a median
posterior spine on the third, fifth and sixth somites, but without any on the
fourth; telson dorsally sulcate on proximal half and armed usually with
four pairs of lateral spines.

Measurements: The smallest recognizable males in which the appendix
masculina is little more than a bud have a carapace length of 7.5 mm. The
largest male in the collection has a carapace length of 18.8 mm. and all of
the larger specimens, in which the carapace is more than 17 mm. long, are
males. The smallest ovigerous female examined has the carapace measuring
9.5 mm. Incidentally, the eyes of the embryos were easily seen in the eggs
of this specimen, indicating that they were nearly ready to hatch. The
youngest specimens have a carapace length of about 3.0 mm.

Acanthephyra purpurea. Male taken in August, 1931, from 600
fathoms. X 1.58.

1940J

Chace: Bathypelagic Caridean Crustacea

135

Color in Life: Body and appendages brilliant scarlet-red with setae of
flame-scarlet. Eye black. Eggs scarlet-red, with the abdominal portion of
the developing embryo peach-red.

General Remarks: The collection contains 4,572 specimens of this
species; 909 males, 796 females, of which 278 are ovigerous, and 2,867
young. Of the ovigerous specimens, 79 are carrying eggs in which the eyes
are visible.

A. purpurea is distinguished from the following species by the absence
of a tooth or spine on the fourth abdominal somite, by having fewer lateral
spines on the telson and by its somewhat smaller size. There is but one speci-
men in the collection, a female with a carapace length of 9.0 mm., which has
the combination of four pairs of telson spines and a tooth on the fourth
abdominal somite. Of the 4,572 specimens, only 158 do not have four spines
on either side of the telson. This does not take into consideration about 3.7
per cent of the remaining 4,414 specimens in which the telson is damaged
so that the determination of the number of telson spines is impossible. If
allowance is made for this, however, probably all but three or four per cent
of the specimens have the normal armature of four spines on either side of
the telson. Text-fig. 12 shows graphically the numerical distribution of the
spines in those specimens which do not have four pairs of lateral telson
spines. Of these 158 specimens, 122 have the normal number of four spines
on either the right or the left side of the telson. If the number of specimens
is doubled to include the spines on either side, it is found that in only 26
out of 316 cases are there more or less than three to five spines, and in prac-
tically all of these few instances the telson is so malformed that it might be
classed as a freak or as the result of injury.

The following notes on the eggs and embryos of this species were made
by Miss Crane from a specimen taken in Net 1107; July 27, 1931; 400
fathoms.

“Silky threads bind the eggs to one another, the substance seemingly

/Vvwtef of /ct/'o/'O’/ fto/s-o'? *r/

Text-figure 12.

Distribution of lateral telson spines in individuals of Acanthe-
phyra purpurea which do not have four spines on either side of
the telson. Solid bars represent spines on the left side of the telson;
hatched bars those on the right.

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continuous with the egg membrane itself. There are usually half a dozen
points of attachment on each egg — one at each end and four in the vicinity
of the equator. These threads are very fine, though tough and elastic, and
in at least several I can quite clearly see that they are tubular in structure —
as if each egg were enclosed in the common atrium into which the six tubes
open.

“The well-formed, salmon-orange embryos are nearly ready to hatch,
the tip of the abdomen in some cases already projecting from the ruptured,
transparent egg membrane. The scarlet-red yolk sac remains only along the
dorsal side of the head and thorax of each embryo, which is curled up in the
longitudinal plane of the egg, the abdomen flexed beneath the thorax. The
uropods and telson, perfectly formed except for the four or five pairs of
spines and the feathery setae of the adult, fit exactly between the two pairs
of antennae, which are already the longest of the appendages. I am reason-
ably certain that the enormous structure almost covering the side of the
head is a compound eye, comparable to that of a Hyperiid amphipod, but
further observation is required. The anlage of the rostrum is visible as a
small, blunt projection. The thoracic appendages are outlined, but fixed to
one another, and there are, of course, no pleopods.’’

In three instances, A. purpurea was found in the stomach of the fish,
Gonostoma elongatum.

Text-figure 13.

Seasonal distribution of adults (solid bars) and young (hatched
bars) of Acanthephyra purpurea computed from specimens taken
in 1929 and 1930 between 500 and 1,000 fathoms.

1940]

Chace: Bathy pelagic Caridean Crustacea

137

For remarks on the food of this species, see the general discussion at
the end of this paper.

Seasonal Distribution-. Text-fig. 13 shows the fluctuation in the popula-
tion of A. purpurea from April to September for the years 1929 and 1930.
Although the population, particularly in regard to the proportion of young
or immature specimens, is reasonably uniform throughout this period of the
year, it will be noted that there is apparently a perceptible increase in the
total population in July, August and September. It will also be noticed that
there is a much greater proportion of young in September than in June.
Although a slightly larger proportion of the September tows were made in
shallow water where young are somewhat more abundant, this difference is
hardly great enough to produce such a change in the total population. It
must be kept in mind that no attempt was made to segregate various sizes
of young and the presence of more than one age class might very well serve
to confuse the general aspect of the results. However, there is certainly an
indication of an increase in the numbers of young individuals early in the
fall.

Text-figure 14.

Seasonal distribution of ovigerous females of Acanthephyra
purpurea computed from specimens taken in 1929 and 1930 between
500 and 1,000 fathoms. Solid bars represent the percentage of all
females which were ovigerous; hatched bars the percentage of all
females which carried eggs in which the eyes were visible.

The distribution of ovigerous females over the same six months period,
as shown in Text-fig. 14, indicates little more than that there is no sharply
defined “breeding period” during this part of the year. Whether the eggs
are carried throughout the year in this area can be ascertained only by addi-
tional towing in the winter months. The suggestion has been advanced that
the eggs are carried by the female for a considerable length of time but
hatch only during a short period when conditions are most favorable, but if
it is assumed that eggs in which the eyes of the larvae are visible are nearly
ready to hatch the present material would seem to refute this hypothesis.

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Vertical Distribution : The vertical distribution of the total catch of
A. purpurea from 1929 through 1935 is given in the following table:

Fathoms

Males

Females

Young

300

2

2

19

400

2

1 (ovig.)

31

500

57

80 (31 ovig.) . . . .

584

600

161

220 (79 ovig.)

782

700

227

193 (80 ovig.) . . . .

662

800

203

356

900

138

70 (16 ovig.) . . . .

260

1,000

115

171

1,100

4

1

2

1,200

1

Text-fig. 15 shows the distribution of all adult and young specimens
captured between 300 and 1,000 fathoms. All of these specimens were taken
in the daytime; in fact, I have seen no specimens from nets 224 to 229
which were towed at night between 500 and 1,000 fathoms. This may or
may not indicate that the center of distribution of the species is above 500
fathoms at night (see Welsh, Chace & Nunnemacher, 1937, p. 190). It is
obvious that in the daytime most adults are to be found between 600 and 800
fathoms and, inasmuch as open nets were employed, it is not unlikely that
there are fewer individuals at 900 and 1,000 fathoms than indicated here.
The center of abundance of the young population, on the other hand, is
slightly nearer the surface, between 500 and 700 fathoms, and the greatest
number of young in proportion to the number of adults is found at 500
fathoms.

This collection has disclosed the interesting fact that the male portion

Text-figure 15.

Vertical distribution of adults (solid bars) and young (hatched
bars) of Acanthephyra purpurea.

1940]

Chace: Bathypelagic Caridean Crustacea

139

5

|

*

\

*

6

//7 o'O’i/// /4?s?

Text-figure 16.

Relative abundance of males and females of Acanthephyra pur-
purea from 500 to 1,000 fathoms. Solid bars represent the male
fraction of the adult population; hatched bars the female.

of the adult population is found in somewhat deeper water than the female
portion, as shown in Text-fig. 16. If these findings accurately represent the
actual conditions, one would expect to find equal numbers of males and
females at about 675 fathoms, whereas above that level the females are in
the majority and below it, the males. Whether the female population in-
creases slightly again in proportion at 1,000 fathoms as shown by these data
is problematical since it must be kept in mind that catches from the deeper
tows must have been somewhat contaminated with specimens from lesser
depths through the use of open nets. It might be supposed that this curious
distribution of the two sexes in A. purpurea could be accounted for by the
possibility that ovigerous females came up into lesser depths to hatch their
eggs, but this is hardly borne out by the present material as shown in Text-
fig. 17, which shows the percentage of all females which are ovigerous at
each depth. Although there is a slightly larger proportion of ovigerous
females from 500 to 800 fathoms, the largest fraction is found at 700 fathoms
where more than half of the population is made up of males. These data also
tend to refute the theory that the eggs are hatched near the surface, for the
largest proportion of specimens carrying eyed eggs was taken in 1,000
fathoms.

Geographical Distribution: Inasmuch as many workers, including my-

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S00 700 <200 900 /000

JP&yo/'A <S7 Ta<7Aarr7<r

Text-figure 17.

Vertical distribution of ovigerous females of Acanthephyra pur-
purea. Solid bars represent the percentage of all females which
were ovigerous; hatched bars the percentage of all females which
carried eggs in which the eyes were visible.

self, have previously confused several closely related species with A. pur-
purea, the distribution of this form is somewhat in doubt. However, as
Kemp (1939) has shown, it is not a world-wide form as generally consid-
ered, but is probably restricted to the north Atlantic between about 20° and
53° north latitude.

Acanthephyra haeckelii (von Martens).

Text-figs. 18-20.

Ephyra Haeckelii von Martens, 1868, p. 54, pi. 1, figs. 7a-b.

Acanthephyra a gassizii Smith, 1884 (not 1882), p. 372 (part), pi. 8,
fig. 1.

Acanthephyra sica Bate, 1888, p. 739, pi. 125, fig. 1.

Acanthephyra rectirostris Riggio, 1901, p. 20.

Acanthephyra purpurea var. multispina Coutiere, 1905, p. 10.

Acanthephyra multispina Murray & Hjort, 1912, pi. 3, fig. 1 (col. fig. ) ;
Stephensen, 1923, p. 44.

Acanthephyra haeckeli Kemp, 1939, p. 575.

Diagnosis: Integument firm. Rostrum long and slender. Carapace not
dorsally carinate posteriorly. Branchiostegal spine supported by a short
carina. Abdomen carinate on all but the first somite and with a medium
posterior spine on the third, fourth, fifth and sixth somites; telson dorsally

1940]

Chace: Bathypelagic Caridean Crustacea

141

sulcate on proximal half and armed usually with eight to eleven pairs of
lateral spines.

Measurements : The smallest recognizable male has a carapace length
of 10.0 mm. The largest male in the collection has a carapace length of 24.0
mm., and the largest female, 24.5 mm. Of thirteen specimens in which the
carapace measures 20.0 mm. or over, ten are males and three females. In
the smallest specimen seen, the carapace measures 2.6 mm.

Color in Life : Body and appendages brilliant scarlet-red, with the setae
somewhat lighter and with the antennae, antennal scales and proximal seg-
ments of the pleopods tinged with salmon-orange. Eyes black.

General Remarks : This species is much less common in the Bermuda
area than the last. The collection contains only 217 specimens, of which 35
are males, 36 females and 146 young.

A. haeckelii can be readily distinguished from A. purpurea by the tooth
on the fourth abdominal somite and the greater number of dorso-lateral
spines on the telson. It will be seen from Text-fig. 19 that the most common
number of lateral telson spines on either side is nine, followed in order by
ten, eight, and eleven. Six, seven, twelve and thirteen spines occur so rarely
that such a condition might well be regarded as abnormal. This species also
attains a larger size than the preceding, and the rostrum is usually some-
what shorter proportionately in the Bermuda specimens. In studying the
present collection, it was noticed that in the preserved specimens of A.
haeckelii the abdomen was generally fully flexed as in the accompanying
figure, whereas in those of A. purpurea it is usually merely bent sharply at
the third somite, and the dorsal margins of the fourth and fifth somites most
frequently form a straight line.

It is noteworthy that none of the females carried eggs. On the basis of
the percentage of ovigerous specimens in A. purpurea, one would expect
about twelve of the 36 females of A. haeckelii to be ovigerous also. The most
plausible explanation would seem to be that this region is not included in the
normal range of the species and that breeding does not occur due to the
unfavorable conditions. There is the possibility, of course, that ovigerous
females might be found during the winter months, but the apparent absence
of any well defined “breeding season” in A. purpurea would tend to preclude

Text-figure 18.

Acanthephyra haeckelii. Male taken in September, 1929, from
900 fathoms. X 1.45.

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Text-figure 19.

Distribution of the lateral telson spines in Acanthephyra
haeckelii. Solid bars represent the spines on the left side of the tel-
son; hatched bars those on the right.

this possibility. Finally, one must not entirely disregard the chance that a
greater number of females might disclose some with eggs, especially since
Stephensen, 1923, reports finding only seven out of 72 females of this species
carrying eggs in contrast to 17 out of 58 in the case of A. purpurea.

Seasonal Distribution: The number of specimens is obviously too small
to give any indication of the seasonal distribution of the species. Both
adults and young were taken in every month of 1929 and 1930 in which
towing was done, i.e. from April to September.

Vertical Distribution: The following table shows the vertical distribu-
tion of the total catch of A. haeckelii from 1929 through 1935:

Fathoms

Males

Females

Y oung

400

2

500

1

600.

1

16

700

3

7

36

800

8

6

32

900

15

8

33

1,000

8

15

26

When correlated with the number of hours of towing at each depth,
these data are shown graphically in Text-fig. 20. All of these specimens
were taken in the daytime. A comparison with the vertical distribution of
A. purpurea leaves little doubt that A. haeckelii is usually found at greater
depths in the area in question. In A. purpurea the center of the adult pop-
ulation was found to be between 600 and 800 fathoms, whereas in A. haeckelii
it seems to be at about 900 fathoms. The greatest number of young, al-
though higher in the water than the adults, were not as near the surface as
those of A. purpurea. In the latter, the young were most numerous at from

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143

Text-figure 20.

Vertical distribution of adults (solid bars) and young (hatched
bars) of Acanthephyra haeckelii.

500 to 700 fathoms, while in A. haeckelii the center of abundance seems to be
between 700 and 900 fathoms.

Geographical Distribution: This species is known from the North
Atlantic as far north as Davis Strait, the Mediterranean and the South
Atlantic, South Pacific and Indian Oceans. In the western North Atlantic
A. haeckelii seems to be much commoner than A. purpurea west of the Gulf
Stream, whereas the present collection certainly indicates that the reverse
is true in the Sargasso area. Our pi’esent knowledge of the distribution of
the two species in the North Atlantic would lead to the assumption that
A. haeckelii is normally a colder water form than A. purpurea.

Acanthephyra curtirostris Wood-Mason.

Text-fig. 21.

Acanthephyra curtirostris Wood-Mason, 1891, p. 195; Balss, 1925, p.
261, fig. 30 (mandible) ; Chace, 1937, p. 111.

Diagnosis: Integument firm. Rostrum not reaching beyond antennular
peduncle and with a single spine on lower margin. Carapace not dorsally
carinate posteriorly. Branchiostegal spine supported by a carina which ex-
tends back to posterior part of branchial region. Abdomen carinate on all
but the first somite and with a median posterior spine on the third, fourth,
fifth and sixth somites; telson dorsally sulcate on proximal half and armed
with six to fifteen dorso-lateral spinules.

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Color in Life : Crimson (Alcock, 1901). From recently preserved speci-
mens, both from the Atlantic and the Pacific, I would infer that the general
color of this species is very similar to that in A. purpurea and A. haeckelii.

Remarks: This species is represented by a single young specimen (cara-
pace length, 4.0 mm.) from Net 1273, 1279, 1285, 1293, 1316 or 1328, Sep-
tember 7 to 19, 1932, from 800 fathoms. Had I not recently seen several
adult specimens of this species from the West Indian region, I should have
been very hesitant about the determination of this young specimen, but
there is little doubt that it belongs to this species.

Text-figure 21.

Acanthephyra curtirostris. Male from Tongue of the Ocean,
Bahamas. X 1.44.

Geographical Distribution : A. curtirostris ranges from the east coast
of Africa through the Indian Ocean, Malay, Philippine and Japanese regions
to the Hawaiian Islands and the west coast of America from San Diego to
Peru. In the Atlantic it has been known only from a specimen in the U. S.
National Museum taken off British Guiana. Recently I have seen several
specimens taken by the Pawnee in the Bahaman region and by the Atlantis
off the north and south coasts of Cuba. Although apparently less common
in the Atlantic than off the Pacific coast of America, the limits of range in
both areas appear to be similar.

Acanthephyra stylorostrata (Bate).

Text-fig. 22.

? Bentheocaris exuens Bate, 1888, p. 724, pi. 123, figs. 3-3o.

Bentheocaris stylorostratis Bate, 1888, p. 726, pi. 123, figs. 4-4o.

Acanthephyra stylorostratis Caiman, 1925, p. 14; Chace, 1936, p. 30.

Diagnosis: Integument rather firm but by no means hard. Rostrum
short and high and somewhat semicircular in outline. Carapace not dorsally
carinate posteriorly. Branchiostegal spine supported by a long carina which
passes back onto the posterior part of the branchial region. Abdomen car-
inate on all but the first somite and with a median posterior spine on the

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145

third, fourth, fifth and sixth somites; telson dorsally sulcate and armed
normally with three pairs of lateral spines.

Measurements : The smallest recognizable males have a carapace length
of 8.5 mm. The largest male has a carapace length of 16.2 mm., and the
largest female, 13.5 mm. In the smallest of the five ovigerous females taken,
the carapace is 10.0 mm. long. The carapace of the smallest young individual
measures 2.9 mm. In this and the preceding species the carapace length is
shorter in relation to the abdomen than in most other species of the genus,
so the total body length, exclusive of the rostrum, will be somewhat greater
in these two forms than in specimens of other species of similar carapace
measurements.

Color in Life: Body and appendages brilliant scarlet, with salmon-orange
setae on antennae, pereiopods and pleopods. Eyes golden-bronze, speckled
with black. Eggs scarlet.

Text-figure 22.

Acanthephyra sty loro strata. Male taken in June, 1930, from
900 fathoms. X 1.74.

General Remarks : This species is represented by 91 specimens; 17
males, 15 females, of which 5 are ovigerous, and 54 young.

The form of the rostrum distinguishes this species at a glance. It is
very probable that Bate’s Bentheocaris exuens is the same species but, in-
asmuch as Caiman states that the type is in such condition that comparison
is almost impossible and as Bate’s figure is inadequate, it seems best to
retain the much more suitable name, styloro strata, for this form.

Practically all of the specimens in the collection have three pairs of
dorso-lateral spines on the telson ; I have seen but two which vary from this
pattern and they have four on the left side and three on the right.

Vertical Distribution : The following table shows the depths at which
this species were found:

Fathoms

600

Males

Females

Young
2

700

6

800

2

11

900

21

1,000

1 100

10

5 (1 ovig.) . . . .

13

1,200

1

Seasonal Distribution : There is no obvious seasonal or yearly variation
in the numbers of specimens found in the Bermuda area. Of the five oviger-

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ous females, two were taken in July, 1930, one either in June, July or August,
1931, one in August, 1931, and one in October, 1931. In one of the specimens
taken in July, 1930, the eyes of the larvae can be seen through the egg
membranes.

Obviously, A. stylorostrata is a deep water species usually found from
800 to 1,000 fathoms. Although the number of adults taken was small, it
will be noticed that, as in A. purpurea, the males seem to be found at some-
what deeper levels than the females. The single female which carried eyed
eggs was found at 1,000 fathoms. Even the young of this species seem to be
found in considerable depths, since the greatest number came from 900
fathoms.

Geographical Distribution : A. stylorostrata has been recorded previously
in the eastern North Atlantic in the region of Madeira, the Canaries and
the Cape Verde Islands as far west as 45° west longitude. A few specimens
have been seen from the region of the Gulf Stream east of New Jersey, and
Caiman records specimens from off Natal, South Africa. Bentheocaris
exuens, which is probably the same species, came from the South Pacific
south of the Tuamotu Archipelago.

Ac anthephyra acutifrons Bate.

Text-fig. 23.

Acanthephyra acutifrons Bate, 1888, p. 749 (part), pi. 126, fig. 3; Kemp,
1906, p. 20; Balss, 1925, p. 261.

Diagnosis : Integument firm but not hard. Rostrum rather short and
deep with a single ventral tooth. Carapace dorsally carinate for its entire
length. Branchiostegal spine supported by a short, rounded ridge. Abdomen
carinate on every somite and with a median posterior spine on the third,
fourth, fifth and sixth somites; telson dorsally sulcate and armed with five
or six pairs of lateral spinules.

Color in Life : Unknown.

Text-figure 23.

Acanthephyra acutifrons. Ovigerous female from Tongue of the
Ocean, Bahamas. X 0.80.

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147

Remarks : Two young specimens (carapace lengths, 5.1 and 7.1 mm.)
from Net 813, 823, 830, 839, 843, 849, 855, 861 or 866, August 28 to Septem-
ber 10, 1930, 700 fathoms, and Net 1272, 1278, 1284, 1292, 1297, 1315 or 1327,
September 7 to 19, 1931, 700 fathoms, are assigned to this species. Although
they bear a striking resemblance to A. curtirostris , the completely carinated
dorsal margin of the carapace and the absence of a long carina supporting
the branchiostegal spine eliminates that species.

Despite the fact that the Valdivia obtained a specimen of A. acutifrons
in a vertical net, one would guess that adults of this heavy-bodied species
are largely benthonic and it is possible that only young individuals are
usually encountered in midwater.

Geographical Distribution: Only two specimens of this species are re-
corded in the literature: the type from the Indian Ocean (Kemp believes the
other specimens included under this species by Bate to be A. curtirostris )
and the Valdivia specimen collected south of Sumatra. In addition, I have
examined seven specimens taken by the Albatross, Pawnee and Atlantis in
the Gulf of Mexico and the West Indian region.

Acanthephyra eximia Smith.

Text-fig. 24.

Acanthephyra eximea Smith, 1884, p. 376; Smith, 1886, pi. 14, fig. 1.

Acanthephyra angusta Bate, 1888, p. 737, pi. 124, fig. 6.

Acanthephyra edwardsii Bate, 1888, p. 747, pi. 126, fig. 1.

Acanthephyra brachytelsonis Bate, 1888, p. 753, pi. 126, fig. 7.

Acanthephyra eximia Wood-Mason & Alcock, 1892, p. 361, fig. 3.

Acanthephyra brachytelsonis Wood-Mason & Alcock, 1892, p. 362, fig. 4.

Acanthephyra eximia Kemp, 1906, p. 20.

Acanthephyra eximea Chace, 1936, p. 27.

Acanthephyra eximia Caiman, 1939, p. 191.

Diagnosis: Integument firm but not hard. Rostrum rather long and
slender. Carapace dorsally carinate throughout its length. Branchiostegal
spine not supported by a ridge. Abdomen carinate on all but the first somite
and with a median posterior tooth or spine on the third, fourth, fifth and
sixth somites, that of the third much the largest and those of the fourth and
fifth often very small; telson faintly ridged dorsally and armed with from
three to five pairs of lateral spinules.

Color in Life: Crimson (Alcock, 1901).

Remarks : Three specimens, all young, are referred to this species. One
specimen with a carapace length of 8.9 mm. was taken in Net 257, 263, 269,
275, 281 or 288, July 5 to 11, 1929, from 800 fathoms; one specimen with a
carapace length of 3.9 mm., from Net 427, 448, 462, 466, 467 or 476, Sep-
tember 5 to 13, 1929, from 900 fathoms; and one specimen with a carapace
length of 5.2 mm. from Net 931, 935, 949, 962 or 969, September 22 to 30,
1930, from 800 fathoms.

This is another of that group of species which are apparently usually
found on or near the bottom as adults.

There seems to be no excuse at the present time for separating A.
brachytelsonis from this species, as the distinction between the two rests
largely on the number of teeth on the lower margin of the rostrum. Appar-
ently specimens having three teeth on the lower margin are commoner in
the Pacific and those with four in the Atlantic, but the number varies from
one to five, and I have seen one individual which was entirely devoid of ven-
tral rostral teeth.

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Acanthephyra eximia. Holotype male (after Smith, 1886).

X 1.03.

Caiman, 1939, has pointed out that Smith, 1884, in one place in the
original description used the correct form of the specific name, eximia, so it
is probably advisable to regard the spelling, eximea, as a lapsus calami.

Geographical Distribution : This is a widely distributed species, having
been found in the western Atlantic off the coasts of North and South Amer-
ica and in the Indian and Pacific Oceans from off Cape Natal, Africa, to the
Malay Archipelago, Japan and the Hawaiian and Society Islands. A few
specimens were also taken by the Challenger near the Kermadec Islands.
A. eximia seems to be most prevalent in the vicinity of land and has been
found off most temperate and tropical coasts with the exception of the west
coasts of the Americas and the west coasts of Europe and Africa. Acanthe-
phyra pulchra A. Milne Edwards from the Mediterranean is probably a
closely related form.

Acanthephyra brevirostris Smith.

Text-fig. 25.

Acanthephyra brevirostris Smith, 1885, p. 504; Smith, 1886, p. 670
(66), pi. 14, fig. 2, pi. 15, figs. 2 and 8, pi. 16, figs. 1 and 6; Lenz & Strunck,
1914, p. 327.

Hymenodora duplex Bate, 1888, p. 843, pi. 136, fig. 3.

Diagnosis : Integument soft and almost membranous. Rostrum short.
Carapace not dorsally carinate posteriorly. Branchiostegal spine not sup-
ported by a ridge. Abdomen carinate on all but the first two somites and
with a median posterior tooth or spine on the third, fourth, fifth and sixth
somites, that of the third extremely large and fleshy and usually over-
reaching the fourth somite; telson dorsally sulcate and armed with about
five pairs of lateral spinules.

Color in Life : Unknown.

Remarks-. This species, which is easily recognized by the large, fleshy
spine on the third abdominal somite, is represented in the Bermuda collec-
tion by six specimens: a young male having a carapace length of 14.0 mm.,
and five young in which the carapace length varies from 5.5 to 9.0 mm. The
young male was taken in June, July or August, 1931, from a depth of 1,000

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149

Acanthephyra brevirostris. Male from off Chesapeake Bay.

X 1.55.

fathoms. The young were taken on the following dates: June 19, 1929,
June 7, 1930, June 28 to July 4, 1930, September 13 to 19, 1930, and June 2
to August 5, 1931, in depths of 1,000, 1,000, 1,000, 700 and 900 fathoms
respectively.

Examination of the mandible discloses that the incisor process is dentate
throughout so that there is little doubt that this species is correctly assigned
to the genus Acanthephyra and does not belong in the genus N otostomus as
might be assumed from its soft integument and the fact that it has no close
relatives among the species of the former genus.

It is not improbable that adults of A. brevirostris are usually found on,
or close to, the bottom. At least it is evidently a deep-water form since no
adults have been recorded from less than 1,000 fathoms.

Geographical Distribution-. The majority of the recorded specimens of
this species were taken off the east coast of the United States, but it has
also been found in the Equatorial Atlantic, off Marion Island, southeast of
the Cape of Good Hope and in the eastern Pacific off the coast of Ecuador.

Acanthephyra gracilipes, sp. nov.

Text-figs. 26 and 27.

Type : Holotype ovigerous female, Cat. No. 301689 Department of Trop-
ical Research, New York Zoological Society; Net 689 or 734; June 9 or 27,
1930; 1,000 fathoms.

Diagnosis : Integument very thin and fragile. Rostrum short. Carapace
not dorsally carinate posteriorly. Branchiostegal spine not supported by a
ridge or carina. Abdomen carinate on all but the first two somites and with
a median posterior tooth on the third, fourth, fifth and sixth somites; telson
dorsally sulcate.

Description : Carapace with a depression in the dorsal margin in the
region of the cervical groove. Rostrum triangular, reaching about as far as
the third segment of the antennular peduncle and armed dorsally with six

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small teeth, two of which are behind the line of the orbit; there are no teeth
or spines on the lower edge. The carina supporting the rostrum in the dorsal
midline disappears before reaching the cervical groove. Antennal spine so
minute as to be scarcely discernible. Branchiostegal spine much more prom-
inent but without any supporting ridge or carina. The integument is so
membranous that it is difficult to determine the distinctive grooves and
ridges, but they apparently consist of an hepatic groove, a faint cervical
groove passing over the dorsal midline and a distinct carina-like ridge above
the branchial region passing backward from the hepatic region. Abdomen
dorsally smoothly rounded on the first two somites, sharply carinate on the
four following. Each of the latter four is provided with a prominent tooth,
those of the third, fourth and fifth somites being offset to the left side of
the animal as in Notostomus compsus. The end of the telson is missing; the
remaining portion is dorsally sulcate and armed with two pairs of small
spines. Eyes dark brown in alcohol and provided with a papilla on the inner
surface just behind the cornea. Antennal scales broken. Third maxillipeds
and all of the pereiopods very long and slender; the third pereiopods, which
are the longest, exceed the antennal scale by the dactyl and probably nearly
half of the propodus.

In the single specimen, the eggs were apparently nearly ready to
hatch as evidenced by the presence of eye spots.

Text-figure 26.

Acanthephyra gracilipes. Holotype ovigerous female. X 3.49.

Measurements : Total length about 42 mm. ; length of carapace, 12.4
mm.; length of rostrum, 2.6 mm. The eggs measure about 0.8 mm. in
longer diameter.

Color in Life: Unknown.

Remarks: A. gracilipes belongs to that group of the genus comprising
A. sibogae de Man, 1916, A. cucullata Faxon, 1893, A. tenuipes (Bate, 1888)
and A. indica Balss, 1925. All of the species of this group have a more
or less membranous integument and closely resemble the less specialized

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151

Acanthephyra gracilipes. A. Mandible. B. First maxilliped.

members of the genus Notostomus, so much so that their generic status
cannot easily be determined except by examination of the mandibles. A.
sibogae differs from A. gracilipes by having the carapace dorsally carinate
throughout its length, by having no antennal spine, the branchiostegal spine
supported by a carina and no spine or tooth on the third abdominal somite.
A. cucullata has a much higher and shorter rostrum, the branchiostegal spine
supported by a long carina and the second abdominal somite dorsally
carinate. A. tenuipes more closely resembles A. gracilipes and it is diffi-
cult from Bate’s description and figure to make a very definite comparison
with the present species; apparently, however, in A. tenuipes the rostrum
is higher and somewhat more compressed laterally and (Bate’s figure to
the contrary) the fourth, fifth and sixth abdominal spines are less promi-
nent than those of A. gracilipes. Finally, A. indica, which closely resembles
the Bermuda species in general form, can be distinguished by the presence
of a branchiostegal carina, a carina on the dorsal margin of the second
abdominal somite and probably by a much shorter telson. A. gracilipes
probably has longer and more slender pereiopods than any of the other
species of the group.

Acanthephyra, sp. ?

Text-fig. 28.

Diagnosis : Integument thin and soft. Rostrum short. Carapace not
dorsally carinate posteriorly. Branchiostegal spine supported by a very
short carina. Abdomen carinate on all but the first two somites and with,
or without, a small posterior spine on the fifth and sixth somites, telson
dorsally sulcate and armed with three pairs of lateral spinules.

Color in Life : Unknown.

Remarks : Eight specimens, ranging in size from a carapace length of
3.8 to 5.8 mm., belong to this undetermined species. They were taken in

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A

B

Text-figure 28.

Acanthephyra, sp. ?. A. Carapace of specimen taken in June,

July or August, 1931, from 1,000 fathoms. B. Telson of same speci-
men.

June, 1929, June to September, 1930, and June to September, 1931, in 900
and 1,000 fathoms. Since none of the specimens are ovigerous or have an
appendix masculina on the second pleopods, it is very likely that they are
immature. They most closely resemble A. rostrata (Bate, 1888) and may
prove to be the young of that species, but they differ from Bate’s description
and figure in that they have fewer rostral teeth and no long carina-like
ridge supporting the branchiostegal spine. The fact that the larger speci-
mens have a small tooth at the end of the sixth abdominal somite and the
largest also has a small tooth on the fifth somite indicates that possibly
the dentition of the abdominal somites may be further developed in adult
individuals if, indeed, the present specimens are actually young.

Genus Notostomus A. Milne Edwards, 1881a.

Carapace with at least one straight carina running along the median
lateral line from the orbital region to the hind margin, and with an
oblique carina delimiting the hind margin of the hepatic furrow. Abdomen
carinate on the last four somites, at least. Telson more or less truncate
at the tip, not terminating in a sharply pointed end-piece laterally armed
with spines. Pereiopods not abnormally broad and flattened. Exopods of
third maxillipeds and pereiopods neither foliaceous nor rigid. Eyes well
pigmented even when small. Outer margin of antennal scale not armed
with a series of spines. Incisor process of mandible toothed for only half
the length of the cutting edge.

153

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Key to the Bermudian Species of N otostomus.

1. A single longitudinal carina on lateral surface of carapace; dorsal mar-

gin of carapace not denticulate on posterior three-fourths of its length ;
abdomen not dorsally carinate on first somite 2.

More than one longitudinal carina on lateral surface of carapace; dorsal
margin of carapace denticulate for nearly its entire length; abdomen
dorsally carinate on every somite 6.

2. No posterior tooth or spine on third abdominal somite 3.

A posterior tooth or spine on third abdominal somite 5.

3. Integument firm; eyes at least as broad as eyestalks in lateral view 4.
Integument extremely soft and fragile; eyes much narrower than eye-

stalks N. mollis

4. Sixth abdominal somite about twice as long as fifth; second abdominal

somite not dorsally carinate N. vescus

Sixth abdominal somite little longer than fifth; second abdominal somite
dorsally carinate N. compsus

5. Posterior tooth on third abdominal somite broad and truncate or con-

cave distally; dactyls of chelipeds terminating in two small, blunt

spines; size very small N. miccylus

Posterior tooth on third abdominal somite triangular like those on fol-
lowing somites; dactyls of chelipeds terminating in two long, slender
spines; size moderately large N. marptocheles

6. Not more than two prominent longitudinal carinae on lateral surface

of carapace N. di stilus

Four or five longitudinal carinae on posterior half of lateral surface of
carapace 7.

7. Four longitudinal carinae on posterior half of lateral surface of carapace.

8.

Five longitudinal carinae on posterior half of lateral surface of carapace
N. westergreni ?

8. Post-orbital not continuous with rostral carina N. robustus

Post-orbital continuous with rostral carina N. perlatus ?

Notostomus vescus Smith.

Text-figs. 29 and 30.

Notostomus vescus Smith, 1886, p. 677 (72).

Acanthephyra brevirostris Bate, 1888, p. 751, pi. 126, figs. 5-6, (Not
Acanthephyra brevirostris Smith, 1885).

Acanthephyra batei Faxon, 1895, p. 167.

Acanthephyra batei Hansen, 1908, p. 77, pi. 4, fig. 2a.

Notostomus batei Balss, 1925, p. 267.

Notostomus vescus Chace, 1936, p. 28.

Diagnosis : Integument soft, but not so fragile that the body easily loses
its shape. Carapace not inflated, dorsal margin in line with rostrum and
not dentate on posterior three-fourths of its length. A single lateral carina
on carapace passing from orbit to hind margin. Branchiostegal spine with-
out a carina. Rostrum usually with one or two ventral teeth in addition

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Notostomus vescus. Male taken in June, July or August, 1931,
from 700 fathoms. X 2.91.

to the dorsal series. Abdomen carinate only on the third, fourth, fifth and
sixth somites and with a posterior tooth on the fourth, fifth and sixth;
sixth somite about twice as long as fifth. Eyes at least as broad as eye-
stalks. Dactyls of first two pairs of pereipods ending in two short spines,
one of which falls on either side of the fixed finger when fingers are closed.

Measurements: The smallest recognizable males have a carapace length
of about 8.0 mm. The largest male has a carapace measuring 18.0 mm.
The largest female gives a corresponding measurement of 16.7 mm. and the
smallest ovigerous female, 13.0 mm. The youngest specimen has a carapace
4.3 mm. long.

Color in Life: Of this species, Hansen, 1908, writes, "The carapace
is greenish, lighter or darker chiefly according to the colour of the tissues
underneath. The dorsal aspect of the first three abdominal segments is
gray-green, the lateral surfaces much lighter.” Although no color notes
were made upon specimens of this species in the Bermuda material, I
cannot help but feel that Hansen’s specimen may have attained this colora-
tion artificially, possibly from being kept in a copper container which will
often affect specimens in this way. I have seen living specimens of this
species and, although no color notes were taken, my impression is that the
animal was a very dark, dull crimson, the carapace being somewhat darker
than the abdomen and very much darker than other bathypelagic forms
commonly met with.

General Remarks: This species is represented by 152 specimens; 28
males, 50 females, of which 15 are ovigerous, and 74 young. Seven of the
ovigerous specimens are carrying eggs in which the eyes are visible.

An examination of Smith’s type of Notostomus vescus in the U. S.
National Museum leaves little doubt that Acanthephyra brevirostris Bate
( = A. batei Faxon) is a synonym of Smith’s species.

Out of 123 of the Bermuda specimens in which the rostrum was exam-
ined, 5 have no ventral rostral teeth, 84 have one such tooth, 32 have two
and 2 have three. In several of the very young specimens, the posterior
spine on the fourth abdominal somite is minute and sometimes entirely
absent.

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155

Seasonal Distribution : Any conclusions as to seasonal distribution based
on such a small number of specimens must be accepted with caution. Both
adults and young were taken in every month from April to September in
1929 and 1930. Ovigerous females were found in April, May, June, July
and August, and those in which the eggs were eyed occurred in May, June,
July and August. It is of interest that of the thirteen females taken in
September none were carrying eggs, whereas out of a like number taken in
June six were ovigerous. If one examines the proportion of specimens
carrying eyed eggs out of the number of ovigerous females taken in each
month, it is found that the single ovigerous specimen taken in April did not
carry eyed eggs, of four taken in May the eggs of one were eyed, of six in
June three were eyed and the single ovigerous specimen taken in August
carried eyed eggs. One might theorize from this that the eggs usually
hatched in June or July, but much more material would have to be accumu-
lated before such a hypothesis could be verified. It will be noted that the
total catch contained nearly twice as many females as males. This dissimi-
larity is found to hold true for every month in which towing was done with
the possible exception of September, in which month eleven of the twenty-
four adults taken were males.

Vertical Distribution : The following table shows the vertical distribu-
tion of the total catch of N. vescus :

Fathoms

M ales

Females

Young

500

1

1

11

600

4

22

700

10

14(3 ovig.) . . . .

13

800

6

12

900

4

1 (ovig.)

7

1,000

3

9

Text-figure 30.

Vertical distribution of adults (solid bars) and young (hatched
bars) of Notostomus vescus.

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These data, correlated with the towing time at each depth, are shown
in Text-fig. 30. The optimum level for adults appears to be at about 700
fathoms, and for the young at about 600 fathoms. All of these specimens
were taken in the daytime. It is difficult to explain the increase in numbers
of adults at 1,000 fathoms, but a smooth distribution curve is hardly to be
expected with so few individuals and with the use of open nets. Ovigerous
females carrying eyed eggs were taken at 600, 700, 800, 900 and 1,000
fathoms, one at each of the first four levels and three at 1,000 fathoms.

Geographical Distribution : N. vescus has been recorded from the North
Atlantic south of Iceland, off the easf coast of the United States, in mid-
Atlantic near the equator and in the Bay of Bengal. I have also seen speci-
mens from the Philippine region.

Notostomus compsus, sp. nov.

Text-figs. 31 and 32.

Types : Holotype male, Cat. No. 30,322, Department of Tropical Re-
search, New York Zoological Society; Net 748; June 30, 1930; 900 fathoms.

One ovigerous female; Net 421; September 4, 1929; 1,000 fathoms.

One ovigerous female; Net 552; May 9, 1930; 600 fathoms.

Diagnosis : Integument firm but not hard. Carapace not inflated ; dorsal
margin in line with rostrum and not dentate on posterior three-fourths of its
length. A single lateral carina on carapace passing from orbit to hind
margin. Branchiostegal spine supported by a short ridge. Rostrum without
teeth or spines on lower margin. Abdomen carinate on all but the first
somite, but that of the second is indistinct; a posterior spine on the fourth,
fifth and sixth somites; sixth somite little longer than fifth. Eyes as broad
as eyestalks, dactyl of first two pairs of pereiopods with a minute spine far
back from the tip.

Description : Carapace dorsally sharply carinate for nearly its entire
length; this carina is somewhat concave in the central portion and rises
slightly anteriorly to an acutely triangular rostrum which reaches beyond
the antennular peduncles but not quite as far as the tips of the antennal

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157

Text-figure 32.

Notostomus compsus. A. Left eye from above. B. Antennal
scale. C. First pereiopod. D. Second pereiopod. E. Mandible. F. First
maxilla. G. Second maxilla. H. First maxilliped. I. Second maxilliped.

scales. There is a series of about six minute teeth at the base of the ros-
trum, two of them being behind the line of the orbit, but the anterior third
of the rostrum is unarmed. There is no armature on the lower margin.
A single lateral carina passes back from the orbit to the hind margin and
the usual carina marking the posterior limit of the hepatic furrow is present.
Branchiostegal spine supported by a short ridge which disappears before
reaching the hepatic furrow. First somite of abdomen smoothly rounded
dorsally; second somite with a very faint carina, and this carina increases in
importance on each succeeding somite. Third somite unarmed; fourth,
fifth and sixth somites with a median posterior spine, those on the fourth
and fifth offset to the left side of the mid-line. Sixth somite about one-fifth
again as long as the fifth. Telson slightly shorter than the combined lengths
of the fifth and sixth somites, deeply sulcate dorsally, armed with four or
five pairs of lateral spinules and truncate at the end. Eyes brown in alcohol
and about as broad as the stalks; there is a small papilla on the inner side
of the stalk. Stylocerite reaches to the end of the first segment of the an-
tennular peduncle. Antennal scale broad with a broad, truncate tip, the
inner angle of which overreaches the outer spines. Third maxillipeds ex-
tend about to the end of the antennal scales. The fingers of the first two
pairs of pereiopods close normally, although there is a minute tooth back
of the terminal spine of the movable finger. The eggs are proportionately
very small, about 1.0 mm. in diameter, nearly spherical and very numerous.

Measurements-. Total length of holotype about 112 mm.; length of cara-
pace 28.7 mm.; length of rostrum 15.6 mm.

The ovigerous female from Net 421 has a carapace length of 27.0 mm.,
and the one from Net 552, 32.2 mm.

Color in Life : Unknown.

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Remarks: Except for its much larger size, this species might easily be
confused with N. vescus. It differs from that species by apparently never
having any ventral spines or teeth on the rostrum, by having the branchios-
tegal spine buttressed by a sharp ridge, by having the spines of the fourth
and fifth abdominal somites offset to the left and, finally, by having the
sixth abdominal somite very little longer than the fifth, rather than about
twice as long as in N. vescus. N. compsus may be readily separated from
N. mollis, the only other described species of the genus which lacks a spine
on the third abdominal somite, by its much firmer integument, more slender
carapace, longer rostrum and larger eyes.

Notostomus marptocheles, sp. nov.

Text-figs. 33 and 34.

Types: Holotype female, Cat. No. 1060, Bingham Oceanographic Collec-
tion, Yale University; Atlantis Station 1478; lat. 25° 29' N., long. 77° 18’ W. ;
N. E. Providence Channel, Bahamas; February 20, 1933; 875 fathoms of
wire.

One young; Pawnee Station 46; lat. 21° 46' N., long. 72° 50' W. ; north
of Little Inagua Island, Bahamas; April 4, 1927; 1,667 fathoms of wire.

One ovigerous female, one young; Pawnee Station 48; lat. 21° 44' N.,
long. 72° 43' W. ; north of Little Inagua Island, Bahamas; April 6, 1927;
1,167 fathoms of wire.

One young; D.T.R. Net 81; May 8, 1929; 900 fathoms.

One young; Net 138 (?), 143, 149, 154, 161, 167, 173, 179, 185 (?),
190 or 196; May 30 to June 20, 1929; 800 fathoms.

One young; Net 247; July 1, 1929; 1,000 fathoms.

One ovigerous female; Net 365; August 10, 1929; 1,000 fathoms.

Two young; Net 496 (?) or 516; September 23 or 27, 1929; 1,000
fathoms.

One young; Net 716, 740 or 761; June 17 to July 2, 1930; 900 fathoms.

One young male; Net 726 (?), 733 or 771; June 26 to July 4, 1930; 900
fathoms.

One young; Net 827 or 858; September 1 or 6, 1930; 1,000 fathoms.

One young male; Net 983 (?), 993, 1001, 1007, 1024, 1025, 1105, 1114
(?), 1116, 1124 or 1134; June 2 to August 5, 1931; 800 fathoms.

One female; Net 1165, 1166, 1171, 1178, 1185, 1190, 1203, 1204, 1207,
1208, 1213, 1219, 1230, 1239, 1245, 1251, 1258 or 1264; August 11 to Sep-
tember 4, 1931; 900 fathoms.

Diagnosis: Integument firm but not hard. Carapace not inflated, dorsal
margin in line with rostrum and not dentate on posterior three-fourths of its
length. A single lateral carina on carapace passing from orbit to hind mar-
gin. Branchiostegal spine supported by a very short carina. Rostrum with-
out teeth or spines on lower margin. Abdomen carinate on all but the first
somite, but the carina of the second is extremely indistinct; a posterior spine
on the third, fourth, fifth and sixth somites; sixth somite more than twice
as long as fifth. Eyes nearly as broad as eyestalks. Dactyls of first two
pairs of pereiopods ending in two long spines, one of which falls on either
side of the fixed finger when fingers are closed.

Description: Carapace dorsally sharply carinate for its entire length;
this carina is nearly straight throughout and in line with the dorsal margin
of the rostrum. The latter is acutely triangular and reaches just beyond the
end of the antennular peduncle; it is armed with two to five, usually three or
four, dorsal spines at the base, one or two being placed behind the line of

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Cha.ce: Bathypelagic Caridean Crustacea

159

Notostomus marptocheles. Holotype female. X 2.03.

the orbit, but the distal third, at least, is unarmed dorsally and there are no
teeth or spines on the lower margin. A single lateral carina passes from the
orbit to the hind margin, and the usual carina delimiting the hepatic furrow
posteriorly is present. Branchiostegal spine supported by a sharp carina
which is only three or four times as long as the spine, itself. First somite of
abdomen smoothly rounded dorsally. Second somite with an almost invisible
line representing a dorsal carina. Succeeding four somites with a sharp
dorsal carina ending posteriorly in a prominent tooth, those of the third,
fourth and fifth folded over to the left (Text-fig. 34A). Sixth somite
slightly more than twice as long as the fifth. Telson about as long as the
sixth somite, dorsally sulcate on the distal half, armed with four pairs of
lateral spines along the margin of the sulcus and more or less truncate at
the tip, with three spines on either side of a median one. Eyes nearly as
wide as the dorso-ventral diameter of the stalk; there is a prominent papilla
arising from the inner surface of the stalk and extending at least as far as
the tip of the cornea. Stylocerite does not reach the end of the first segment
of the antennular peduncle. Antennal scale tapering to a rounded end which
is slightly surpassed by the outer spine. Third maxillipeds extend about as
far as the antennal scales. Dactyls of chelae of first two pereiopods ter-
minate in two long, curved spines, between which the fixed finger fits when
the fingers are closed (Text-fig. 34E-F). Meri of last three pereiopods spiny.
Eggs proportionately small, about 1.1 mm. in longer diameter, broadly oval
and numerous.

Measurements : Total length of holotype about 62 mm.; length of cara-
pace 17.2 mm. ; length of rostrum 4.9 mm.

The two young males, in which the appendix masculina is shorter than
the other stylet, have the carapace 17.0 mm. long in each. The two ovigerous
females have a carapace length of 20.1 and 22.8 mm., and the latter is the
largest specimen examined. The smallest specimen has a carapace length
of 5.5 mm.

Color in Life: Deep scarlet — nearly black. Rostrum and mouth parts
bright red.

Remarks: Inasmuch as the specimens from the Bingham Oceanographic
Collection were sent for identification before the Bermuda collection, Dr.
Beebe has kindly consented to the selection of the holotype from that series.

It will be seen from the net data that N. marptocheles was found at

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Text-figure 34.

Notostomus marptocheles. A. Dorsal margins of third, fourth,
fifth and sixth abdominal somites viewed from the right side. B.
Telson and uropods. C. Left eye from above. D. Antennal scale.
E. Carpus and hand of first pereiopod. F. Carpus and hand of sec-
ond pereiopod. G. Mandible. H. First maxilla. I. Second maxilla.
J. First maxilliped. K. Second maxilliped.

Bermuda in every month of the year in which towing was done with the
exception of April. Apparently it is a relatively deep water species, since
two specimens were taken in 800 fathoms, four in 900 and five in 1,000. The
eggs on the ovigerous female from Pawnee Station 48 were eyed, whereas
those on the Bermuda specimen from Net 365 were not.

Although this species bears a close resemblance to N. compsus, it may be
readily distinguished by the presence of a spine on the third abdominal
somite, the much longer sixth somite, differently formed eyes and antennal
scales and the long terminal spines on the movable fingers of the chelipeds.

Apparently most, if not all, of the species of Oplophoridae have the
dactyls of the chelipeds more or less split at the end for the reception of
the fixed finger when the fingers are closed, but in no others that I have
seen are these terminal spines so long as to be easily visible to the naked eye.

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161

Notostomus miceyhss, sp. nov.

Text-figs. 35-37.

Types : Holotype female, Cat. No. 1061, Bingham Oceanographic Col-
lection, Yale University; Pawnee Station 48; lat. 21° 44’ N., long. 72° 43' W.;
north of Little Inagua Island, Bahamas; April 4, 1927 ; 1,167 fathoms of wire.

Two males: Pawnee Station 11; lat. 23° 58' N., long. 77° 26' W.; Tongue
of the Ocean, Bahamas; March 2, 1927; 1,167 fathoms of wire.

One female: Pawnee Station 25; lat. 24° 51' N., long. 76° 37' W.; Exuma
Sound, Bahamas; March 17, 1927; 1,333 fathoms of wire.

Two females (one ovigerous) : Pawnee Station 52; lat. 21° 30' N., long.
71° 11' W.; Turks Island Passage; Bahamas; April 11, 1927; 1,333 fathoms
of wire.

One female: Paivnee Station 58; lat. 32° 24' N., long 64° 29' W. ; off
Bermuda; April 20, 1927; 833 or 1,667 fathoms of wire.

Two females: Atlantis Station 1478; lat. 25° 29' N., long. 77° 18' W.;
N.E. Providence Channel, Bahamas; February 20, 1933; 875 fathoms of
wire.

47 males, 73 females (4 ovigerous), 25 young; D.T.R. Bermuda Expedi-
tions; 1929 to 1931; 500 to 1,000 fathoms.

Diagnosis : Integument thin but not membranous. Carapace not inflated,
dorsal margin in line with rostrum and not dentate on posterior nine-tenths
of its length. A single lateral carina on carapace passing from orbit to hind
margin. Branchiostegal spine not supported by a carina. Rostrum without
teeth or spines on lower margin. Abdomen carinate on all but first somite
but the carina of the second is hardly visible; a posterior spine on third,
fourth, fifth and sixth somites, that on the third being very broad with a
concave distal margin ; sixth somite more than two and one-half times as long
as fifth. Eyes as broad as eye-stalks. Dactyl of first two pairs of pereiopods
ending in two small, blunt spines.

Description : Carapace dorsally sharply carinate for its entire length;
the carina is nearly straight and continuous with the dorsal margin of the
rostrum. Rostrum acutely triangular and not reaching beyond the second
segment of the antennular peduncle; it is armed with from two to six,
usually three to five, dorsal spines at the base, one sometimes being placed
behind the line of the orbit; there are no teeth or spines on the lower margin.
Of 135 specimens in which the rostrum is undamaged, one specimen has two
dorsal rostral spines, 19 have three, 71 have four, 40 have five and four
have six. A single lateral carina runs from the orbit to the hind margin of
the carapace, and there is the usual carina delimiting the hepatic furrow
posteriorly. Branchiostegal spine outstanding but not supported by any car-
ina or sharp ridge. First abdominal somite smoothly rounded dorsally. Sec-
ond somite with a faint line representing a dorsal carina. The next four
somites are sharply carinate. The third somite is provided with a distinctive
broad tooth which is truncate or concave distally. Fourth, fifth and sixth
somites end in a small, sharp spine. Sixth somite more than two and one-
half times as long as the fifth. Telson slightly longer than the sixth somite,
deeply sulcate dorsally, armed with three or four pairs of dorso-lateral
spinules and with a more or less truncate tip which bears three spines on
either side of a central one. Eyes nearly black and about as wide as the
dorso-ventral width of the stalk; in dorsal view, the stalk is much wider than
the cornea and bears a small, rounded papilla on the inner margin. Stylocer-
ite narrow and drawn out to a slender point which reaches just beyond the
first segment of the antennular peduncle. Antennal scale narrow, sharply
concave along its external margin and truncate distally, the outer spine
extending beyond the blade. Third maxillipeds extend about as far as the
antennal scales. All of the pereiopods are normal. The eggs are small.

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Text-figure 35.

Notostomus miccylus. Holotype female.

X 3.38.

Measurements: Total length of holotype about 33 mm.; length of cara-
pace 9.2 mm. ; length of rostrum 2.0 mm.

The smallest recognizable male has a carapace length of 4.9 mm. The
ovigerous females have carapace lengths ranging from 7.2 to 9.5 mm. In
this species the female apparently attains a larger size than the male;
females reach a carapace length of 10.0 mm., whereas the largest male exam-
ined had a carapace measurement of only 8.1 mm. The smallest specimens
seen have the carapace 3.2 mm. long.

Color in Life: Abdomen solid scarlet; cephalothorax and telson heavily
dotted with scarlet. Eyes dark brown.

General Remarks: Inasmuch as the specimens from the Bingham Ocean-
ographic Collection were sent for identification before. the Bermuda collec-
tion, Dr. Beebe has kindly consented to the selection of the holotype from
that series.

Although this species might be confused with N. vescus because of the
similarity in size, N. miccylus is even smaller than N. vescus and can be dis-
tinguished from it and all other known species of the genus by the charac-
teristic shape of the tooth on the third abdominal somite.

Seasonal Distribution: The seasonal distribution of the population of
N. miccylus over the months in which tows were made in 1929 and 1930 is
shown in the following table:

1929

April 3 females (1 ovig.), 2 young

May 1 male, 2 females, 1 young

June 1 male, 12 females, 1 young

July 6 males, 14 females (2 ovig.), 3 young

August 1 male, 3 females, 1 young

September 8 males, 9 females, 2 young

1930

May 4 males, 1 female, 1 young

June 6 males, 5 females, 3 young

July 3 males, 4 females, 2 young

August 1 male

September 4 males, 8 females (1 ovig.), 1 young

It will be noted that the distribution over these months is reasonably
uniform and that ovigerous females were taken in April, July and Sep-
tember. Curiously, each of the four ovigerous specimens carried but one
egg and there was evidence that the remaining eggs had recently hatched;

163

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K

L

M

N

Text-figure 36.

Notostomus miccylus. A. Telson and uropods. B. Left eye from
above. C. Antennal scale. D. First pereiopod. E. Second pereiopod.
F. Third pereiopod. G. Fourth pereiopod. H. Fifth pereiopod. I. First
pleopod of male. J. Second pleopod of male. K. Mandible. L. First
maxilla. M. Second maxilla. N. First maxilliped. O. Second
maxilliped. P. Third maxilliped.

the remaining egg was eyed and obviously ready to hatch in three of the
specimens and in the other, taken in September, 1930, the egg was not eyed
and was possibly infertile. Whether the fact that only about 5 per cent, of
the adult females carried eggs and in those few most of the eggs were miss-
ing indicates that the Bermuda area is not included in the normal “breeding
range” is problematical. The table also discloses the anomaly that in 1929
only about 28 per cent, of the adult specimens were males, while in 1930 there
were like numbers of each sex.

Vertical Distribution : The vertical distribution of the total catch of
N. miccylus is shown in the following table:

Fathoms

Males

Females

Y oung

500

1

1

2

600

8

700

15

7

800

7

20

3

900

4

1,000

1

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Text-figure 37.

Vertical distribution of adults (solid bars) and young (hatched
bars) of Notostomus miccylus.

These data, correlated with the towing times, are shown in Text-fig. 37.
All of these specimens were taken in the daytime. A close parallelism be-
tween the vertical range of this species and that of the other small species
of the genus, N. vescus, will be noted.

Notostomus mollis (Smith).

Text-fig. 38.

Meningodora mollis Smith, 1882, p. 74, pi. 11, figs. 8-9, pi. 12, figs. 5-9.

Hymenodora mollis Bate, 1888, p. 841, pi. 136, fig. 5.

Notostomus fragilis Faxon, 1893, p. 207 ; 1895, p. 170, pi. 44, figs. 2-2b.

Notostomus mollis Balss, 1925, p. 266, fig. 37 (mandible).

Diagnosis : Integument extremely soft and fragile. Carapace somewhat
inflated, dorsal margin evenly convex from rostrum to hind margin and not
dentate on posterior nine-tenths of its length. A single lateral carina on
carapace passing from orbit nearly to hind margin. Branchiostegal spine
supported by a short sharp ridge or carina. Rostrum without spines on
lower margin. Abdomen carinate on third, fourth, fifth and sixth somites
only, and with a posterior tooth on fourth, fifth and sixth ; sixth somite about
one and two-thirds times as long as fifth. Eyes narrower than eyestalks.
Dactyls of first two pairs of pereiopods ending in two small, blunt, unequal
teeth between which the fixed finger lies when the fingers are closed.

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165

Measurements: A young male, in which the appendix masculina of the
second pleopods is rudimentary, has a carapace length of 17.0 mm. The
largest male has the carapace 23.0 mm. long, the largest female 24.2 mm.
and the single ovigerous female 23.4 mm. The smallest specimen has a
carapace length of 4.2 mm. The eggs are 0.9 mm. in greater diameter.

Color in Life : Cephalothorax maroon-tinged black shading to nopal red
on the abdomen. Antennae, thoracic epipods and pleopods all nopal red.
Thoracic legs dark maroon shading to lighter toward their tips. Eyes
blackish.

General Remarks: The collection contains 56 specimens of this species;
2 males, 10 females, of which 1 is ovigerous, and 44 young. The eyes are
visible in the eggs of the single ovigerous specimen.

N. mollis may be distinguished from the other members of the genus in
which the carapace is not markedly inflated and the first abdominal somite
uncarinated by its much more membranous integument. It differs from all
members of the genus in having the eyes noticeably smaller in diameter than
the somewhat swollen eyestalks.

An examination of the types of Meningodora mollis Smith and Noto-
stomus fragilis Faxon discloses no points of difference between the Atlantic
and Pacific specimens.

N. mollis forms a connecting link between the primitive members of the
genus belonging to the N. vescus group and the typical forms of the N.
robustus group.

Seasonal Distribution: Specimens were found in every month of 1929
and 1930 in which tows were made, and the numbers taken are more or less
evenly distributed over these periods. The single ovigerous female was taken
in May, 1930.

Text-figure 38.

Notostomus mollis. Female taken in June, 1931, from 900 fath-
oms. X 1.64.

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Vertical Distribution : The vertical distribution is shown in the follow-
ing table:

Fathoms

Males

Females

Young

500

2

600

700

1

13

800

2

3 (1 ovig.)

7

900

3

8

1,000

3

11

Geographical Distribution : N. mollis has been found off the east coast
of the United States, off the coast of Brazil, in the Bay of Biscay, the
Indian Ocean, north of the Philippine Islands (specimen in the U. S. National
Museum) and off the west coast of Central America between Panama and
the Galapagos Islands.

Notostomus distirus, sp. nov.

Text-figs. 39 and 40.

Types : Holotype female, Cat. No. 311915, Department of Tropical Re-
search, New York Zoological Society; Net 1281; August 9, 1931; 1,000
fathoms.

One female; Net 684; June 7, 1930; 1,000 fathoms.

Diagnosis : Integument soft. Carpace inflated, dorsal margin elevated
somewhat above level of rostrum but nearly straight and denticulate for the
greater part of its length. Only two prominent lateral carinae on carapace,
the semi-carina below post-orbital represented by a deep sulcus and the sub-
marginal carina not prominent; post-orbital and post-antennal carinae rela-
tively close to each other and not markedly divergent behind hapatic region ;
post-orbital not continuous with rostral carina; no median lateral carina at
base of rostrum. Rostrum with three spines on lower margin. Abdomen car-
inate on every somite and with a posterior tooth on third, fourth, fifth and
sixth; sixth somite about one and one-third times as long as fifth. Eyes
very large and black. Dactyls of first two pairs of pereiopods ending in
two small teeth or spines between which the fixed finger lies when the fingers
are closed.

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167

Text-figure 40.

Notostomus distirus. A. Mandible. B. First maxilla. C. Sec-
ond maxilla. D. First maxilliped. E. Second maxilliped.

Description : Carapace moderately inflated, although not as much so as
in some of the more typical members of this group. Dorsal margin sharply
carinate, sparsely denticulate for its entire length, nearly straight for the
greater part of its extent and somewhat, but not greatly, elevated above the
level of the rostrum. Rostrum slender, reaching about as far as the tips of
the antennal scales and armed on the slender portion with two dorsal and
three ventral teeth. There is no median lateral carina on the rostrum. The
ventral rostral carina is not continuous with the post-orbital. The latter
forms an obtuse angle at the junction with the post-hepatic carina and
passes backward and somewhat upward to the posterior margin. Immediately
below the posterior half of this carina is a narrow longitudinal sulcus but
no prominent semi-carina. The post-antennal carina rather closely approxi-
mates the post-orbital on the hepatic region and then passes backward and
disappears at about the middle of the posterior branchial region. These two
main carinae diverge slightly behind the hepatic region but not as much so
as in N. robustus. The submarginal carina passing from the dorsal surface
around to the hepatic region is no more prominent than in the species of
other genera. Abdomen carinate on every somite, although the carina on the

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first is very low. The third, fourth, fifth and sixth somites are armed with
large posterior teeth. Sixth somite about one and one-third times as long
as the fifth. Telson about as long as the outer branches of the uropods,
deeply sulcate dorsally and armed with three pairs of minute dorso-lateral
spinules and three terminal spines on either side of a median one. Eyes very
large with a minute papilla on the inner side of the stalk just back of the
cornea. Stylocerite reaches nearly to the end of the first segment of the
antennular peduncle. Antennal scale tapering to a very bluntly angular end
which reaches about as far as the outer spine. Third maxillipeds extend
about as far as the antennal scales. Movable fingers of both pairs of cheli-
peds terminating in two inconspicuous spines which lie on either side of the
fixed finger when the fingers are closed.

Color in Life : Unknown.

Measurements: Total length of holotype about 87 mm.; length of cara-
pace 26 mm. ; length of rostrum 14 mm.

The paratype is somewhat smaller, having a carapace length of about
20.5 mm.

Since there are no males or ovigerous females, there is naturally no way
of determining whether these specimens are females or immature specimens.

Remarks: N. distirus is apparently one of the more primitive members
of the robustus group of the genus as evidenced by the relatively slightly
inflated carapace and few lateral carapacic carinae. It most closely resembles
N. murrayi Bate, 1888, and N. japonicus Bate, 1888. The former was very
possibly described from an immature specimen, but it apparently differs
from N. distirus in the much greater number of teeth on the lower margin
of the rostrum and the prolongation of the post-antennal carina nearly to the
posterior margin of the carapace. N. japonicus agrees very well with the
present species except that the post-orbital and post-antennal carinae are
farther apart and the telson is shorter than the inner branch of the uropods.

For the possible presence of young of this species in the collection see
the remarks under N. robustus.

Notostomus robustus Smith.

Text-fig. 41

Notostomus robustus Smith, 1884, p. 377, pi. 7, fig. 2.

Notostomus beebei Boone, 1930, p. 39, fig. 6.

Notostomus robustus Chace, 1936, p. 28.

Diagnosis: Integument soft. Carapace greatly inflated, dorsal margin
highly arched above level of rostrum and minutely dentate for its entire
length in the adult. Four lateral carinae on posterior half of carapace in-
cluding the semi-carina just below post-orbital and the submarginal carina
below post-antennal; post-orbital and post-antennal carinae distinctly diver-
gent behind hepatic region; post-orbital not continuous with rostral carina;
no median lateral carina at base of rostrum. Rostrum usually with two or
three spines on lower margin. Abdomen carinate on every somite and with a
posterior tooth on third, fourth, fifth and sixth ; sixth somite about one and
one-half times as long as fifth in adult. Eyes very large and black. Dactyls
of first two pairs of pereiopods ending in two small, blunt, unequal teeth
between which the fixed finger lies when the fingers are closed.

Measurements: There is no young male in the collection to determine
the minimum size of adults. The only male has the carapace 49.0 mm. long.
The largest female is the type of Notostomus beebei in which the carapace
measures 45.0 mm. There are seven specimens in which the carapace is

1940]

Chace: Bathy pelagic Caridean Crustacea

169

from 19.0 to 34.0 mm. long, and the remaining 31 specimens are obviously
young, the carapace measuring 3.7 to 14.8 mm.

Color in Life : Scarlet.

General Remarks: The collection contains 20 specimens which undoubt-
edly belong to this species; one male, one female, seven specimens which may
be either females or young and eleven young. There are 20 other specimens
in which the carapace is less than 7 mm. long which may belong either to
N. robustus or to N. distirus. These young individuals closely resemble the
adults of the N. vescus group in that the carapace is not greatly inflated, the
dorsal carina of the carapace is not denticulate posteriorly, the lateral car-
inae are inconspicuous, the teeth on the third, fourth and fifth abdominal
somites may be absent and the sixth abdominal somite is more than twice
as long as the fifth. They can always be distinguished from the species of
the vescus group, however, by the very large eyes.

There is little doubt in my mind that N. beebei Boone is a synonym of
N. robustus. An examination of the type specimens of both species fails to
disclose any differences between the two except for the presence or absence
of the curious excavation in the dorsal carina at the base of the rostrum.
If this character was found in but a single specimen one would readily as-
sume that it was caused by an injury but, curiously enough, the female
cotype of N. robustus has almost exactly the same kind of excavation armed
anteriorly with a “quadrispine.” Smith failed to mention this abnormality
in his description and apparently assumed that it was the result of an
injury. Since only four positively adult specimens of this species are known,
two males and two females, and since both females have this curious depres-
sion in the dorsal carina, the possibility that this is a sexual character can-
not be entirely disregarded.

Text-figure 41.

Notostomus robustus. Young female taken in August, 1930,
from 800 fathoms. X 1.32.

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[XXV :11

Seasonal Distribution: The adult male was taken June 19, 1929, and the
type female of N. beebei on May 25, 1929. Other specimens with a carapace
length exceeding 19 mm. were taken in June and July, 1929, and in August
and September, 1930. Specimens with a carapace length of 7 to 15 mm. were
found in June, July and September, 1929, and in May and September, 1930.

Vertical Distribution: The adult male was taken in 1,000 fathoms and
the adult female in 900. Of the intermediate group in which the carapace
measures from 19 to 34 mm., three were found at 800, one at 900 and two at
1,000 fathoms. Of the smallest specimens with a carapace length of 7 to 15
mm., one was taken in 600 fathoms, three in 700, one in 800, four in 900 and
one in 1,000.

Geographical Distribution: N. robustus is known only from the western
North Atlantic between 42° north latitude and Bermuda.

Notostomus perlatus Bate ?

Text-fig. 42.

? N otostomus perlatus Bate, 1888, p. 831, pi. 134, fig. 2.

? Notostomus brevirostris Bate, 1888, p. 832, pi. 134, fig. 3.

? N otostomus perlatus Balss, 1925, p. 268.

Diagnosis: Integument soft. Carapace greatly inflated, dorsal margin
highly arched above level of rostrum and minutely dentate for its entire
length. Four lateral carinae on posterior half of carapace, including the
semi-carina just below the post-orbital and the submarginal carina below
the post-antennal; post-orbital and post-antennal carinae not markedly
divergent behind hepatic region; post-orbital continuous with rostral carina;
no median lateral carina at base of rostrum. Rostrum with six teeth on
lower margin. Abdomen carinate on every somite and with a posterior tooth
on third, fourth, fifth and sixth ; sixth somite about and one three-fourths
times as long as fifth. Eyes very large and black. Dactyls of first two pairs
of pereiopods ending in two small unequal teeth between which the fixed
finger lies when the fingers are closed.

Color in Life : Unknown.

Remarks: A single young specimen (carapace length 14.6 mm.) from
Net 999; June 5, 1931; 700 fathoms, is tentatively assigned to this species.

The only apparent difference between this specimen and N. perlatus lies
in the length of the rostrum. There are two adult male specimens (carapace
lengths 51 and 52 mm.) in the U. S. National Museum which were taken by
the Albatross in the Gulf of Mexico southeast of the Mississippi delta which
almost certainly belong to the same species as this young Bermuda specimen.
These two specimens differ from all other known specimens of N. perlatus
in having the rostrum reaching as far as the ends of the antennal scales
rather than no more than half that distance. Since the rostrum is usually
longer in immature specimens in this genus, its greater length in the Ber-
muda example is by no means unexpected. Since these three specimens
agree with N. perlatus in all other particulars, it does not seem desirable
at present to separate them merely on the basis of the rostral length.

Another possibility is that this form is really N. gibbosus A. Milne
Edwards, 1881, which was taken by the Blake off Grenada. It corresponds
reasonably well with the figure of that species (A. Milne Edwards, 1883)
except for the absence of a semi-carina below the posterior portion of the
post-orbital carina. Another discrepancy is the fact that Milne Edwards
describes the telson as being distinctly shorter than the outer branch of the
uropods. The type of N. gibbosus cannot be found in the Museum of Com-

1940]

Chace: Bathypelagic Caridean Crustacea

171

Text-figure 42.

Notostomus perlatus ?. Young taken in June, 1931, from 700
fathoms. X 2.56.

parative Zoology. It is to be hoped that it is in the Paris Museum, for it is
only by examination of this type specimen that the correct status of the
species can be determined.

Geographical Distribution: All of the recorded specimens of N. perlatus
have been taken within ten degrees of the equator; off the coast of Brazil,
off the west coast of Africa, off the east coast of Africa, in the Indian Ocean
and south of the Philippines. I have also seen a typical specimen of N. per-
latus in the U. S. National Museum which was taken east of the Marquesas
Islands, also within ten degrees of the equator. The only records from
outside of this equatorial belt in addition to the present specimen are the two
males, mentioned above, from the Gulf of Mexico.

Notostomus westergreni Faxon ?

Text-fig. 43.

? Notostomus westergreni Faxon, 1893, p. 208; 1895, p. 171, pi. F.

? N otostomus westergreni Balss, 1925, p. 269.

Diagnosis: Integument soft. Carapace moderately inflated, dorsal mar-
gin arched but slightly above level of rostrum and denticulate for its entire
length. Five lateral carinae on posterior half of carapace, including the
semi-carina just below post-orbital and the submarginal carina along lower
edge ; post-orbital and post-antennal carinae nearly parallel ; post-orbital not
continuous with rostral carina; a median lateral carina at base of rostrum.
Rostrum with eight or nine teeth on lower margin. Abdomen carinate on
every somite and with a posterior tooth on third, fourth, fifth and sixth;
sixth somite nearly twice as long as fifth in the young. Eyes very large and
black. Dactyls of first two pairs of pereiopods ending in two small spines
between which the fixed finger lies when the fingers are closed.

Color in Life: The type specimen of N. westergreni was bright red. No
color notes were taken on the Bermuda specimens.

Remarks: Two immature specimens are referred to N. westergreni

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[XXV :11

N^tostomus westergreni ?. Young taken in July or August,

1929, from 900 fathoms. X 3.70.

with some hesitation. One specimen, with a carapace length of 9.0 mm.,
was taken in Net 325, 330, 362 or 382; July 24 to August 16, 1929; 900
fathoms. The other, with a carapace length of 7.8 mm., is from Net 613 or
614; May 21, 1930; 600 fathoms.

These specimens differ from the male type of N. westergreni as follows;
the dorsal margin of the carapace is less elevated, there are fewer teeth on
the lower margin of the rostrum and the sixth abdominal somite is pro-
portionately longer. All of these characters might well be attributed to the
obvious immaturity of the specimens. Although Faxon described this species
as having a posterior tooth on the “third, fourth and fifth” somites of the
abdomen, the type has a well developed tooth on the sixth somite as well, as
shown in Faxon’s figure. I have also seen a specimen in the U. S. National
Museum in which the carapace is 12.5 mm. long, which corresponds in every
particular with these Bermuda specimens except that the semi-carina fuses
anteriorly with the post-orbital. This specimen was taken by the Albatross
off the coast of Peru not far from the type locality of N. westergreni.

It is obviously impossible at present to determine with certainty young
specimens of N otostomus due to the comparative scarcity of material, but
of the three known species of the genus which have five lateral carinae on
the posterior half of the carapace, N. westergreni, N. patentissimus Bate,
1888, and N. longirostris Bate, 1888, the present specimens are much nearer
the first than to the other two.

Geographical Distribution : There are but three recorded specimens of
N. westergreni; the type from off the coast of Ecuador, a specimen from
near the Keeling Islands in the Indian Ocean and the third from off Cape
Point, South Africa.

Genus Ephyrina Smith, 1885.

Carapace with a blunt ridge running along the median lateral line from
the orbital region to the hind margin, and with an oblique ridge delimiting
the hind margin of the hepatic furrow. Abdomen smoothly rounded, not

1940]

Chace: Bathypelagic Caridean Crustacea

173

carinate, throughout its length. Telson more or less truncate, not terminat-
ing in a sharply pointed end-piece laterally armed with spines. Ischial and
meral joints of the pereiopods abnormally broad and flattened. Exopods of
third maxillipeds and pereiopods neither foliaceous nor rigid. Eyes well pig-
mented. Outer margin of antennal scale not armed with a series of spines.
Incisor process of mandible toothed for only half the length of the cutting
edge.

Key to the Known Species of Ephyrina.

1. A posterior tooth or spine on third abdominal somite 2.

No posterior tooth or spine on third abdominal somite; ten to thirteen

pairs of lateral spines on telson E. hoskynii.

2. Posterior tooth on third abdominal somite triangular; twenty to twenty-

five pairs of lateral spines on telson E. benedicti

Posterior tooth on third abdominal somite broad and bifid at the end;
five to eight pairs of lateral spines on telson E. bifida.

Ephyrina hoskynii Wood-Mason.

Text-fig. 44.

Ephyrina hoskynii Wood-Mason & Alcock, 1891, p. 194.

Ephyrina hoskyni Kemp, 1910, p. 68, pi. 7, figs. 1-6.

Ephyrina benedicti Chace, 1936, p. 29 (part).

Diagnosis: Abdomen without dorsal teeth or spines on any somite; tel-
son armed with ten to thirteen pairs of lateral spinules. Eyes not broader
than eyestalks.

Color in Life : Dark red, according to Alcock, 1901. No color notes were
taken on the Bermuda specimen.

Ephyrina hoskynii. Female (after Kemp, 1910). X 1.19.

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Remarks : This species is represented by a single young specimen having
a carapace length of 6.8 mm. taken in Net 896; September 16, 1930; 700
fathoms.

The large number of specimens of various sizes with bifid spines on the
third abdominal somite in the present collection have forced me to the
conclusion that there are in reality three known species of Ephyrina rather
than one variable one. There appears to be no difficulty in separating these
forms since they differ, not only as regards the teeth on the third and fourth
abdominal somites, but also in the number of lateral telson spines and the
size of the cornea of the eye. There is probably little in the form of the
rostrum that is of taxonomic importance since all of the described forms
may be found among the various age groups of a single species.

Geographical Distribution : E. hoslcynii is known from off the southwest
coast of Ireland, the Bay of Biscay, off the coast of Spain or Portugal, the
Arabian Sea in the vicinity of the Laccadive Islands and the Bay off Bengal
off Ceylon.

Ephyrina bifida Stephensen, 1923, p. 58, fig. 18.

Ephyrina benedicti Balss, 1925, p. 269, figs. 38 and 39.

Ephyrina benedicti Chace, 1936, p. 29 (part).

Diagnosis: Abdomen with a broad, bifid tooth on third somite and a
small triangular tooth on fourth somite in the adult; telson armed with five
to eight pairs of lateral spinules. Eyes distinctly broader than eyestalks.

Measurements : The collection contains no adult male or ovigerous fe-
male specimens by which the size of adults might be gauged. The largest
specimen, apparently a female, has a carapace length of 21.3 mm. In the
smallest specimen the carapace measures 3.9 mm., but in this specimen, as in
several others slightly larger, the telson still retains its larval form.

Color in Life : Unknown.

Ephyrina bifida Stephensen.

Text-fig. 45.

Text-figure 45.

Ephyrina bifida. Female taken in September, 1931, from 1,000
fathoms. X 1.19. On the right is shown the development of the
tooth on the third abdominal somite in this species with the cara-
pace lengths in mm.

1940]

Chace: Bathypelagic Caridean Crustacea

175

General Remarks: The collection contains 70 specimens, five of which
have a carapace length exceeding 15 mm. and are tentatively designated as
females.

In specimens of E. bifida having a carapace length of 5 mm. or less
there are no teeth on any of the abdominal somites. At about 6 mm. the
tooth on the third somite is represented by a broad, shallow lobe which be-
comes faintly notched at about 7 mm. From this stage on, the tooth grad-
ually increases in length and becomes more distinctly bifid, the mature con-
dition being reached in specimens having a carapace length of about 9 mm.
The small tooth on the fourth somite first appears in specimens having a
carapace length of 8 or 9 mm. Young specimens lacking either of these
spines may be distinguished from those of E. hoskynii by the smaller num-
ber of lateral spines on the telson.

One young specimen of this species was found in the stomach of the fish,
Gonostoma elongatum.

Seasonal Distribution: Specimens were taken in every month of 1929
and 1930 in which towing was done, with the largest numbers appearing in
September of each year. Specimens of various sizes were scattered at ran-
dom over these months and individuals having carapace lengths between 4
and 5 mm. were taken in every month.

Vertical Distribution: This species was taken in depths ranging from
400 to 1,000 fathoms, but only one specimen was found above 600 fathoms
and no specimens having a carapace length greater than 5.5 mm. were
taken in less than 800 fathoms. The largest number of specimens were
found at 800 fathoms, but in none of these did the carapace exceed 10 mm.
in length; only at 900 and 1,000 fathoms were possibly adult specimens
collected.

Geographical Distribution: Only four specimens of this species have
been recorded previously. These came from the Bay of Biscay, the Gulf of
Guinea off the west coast of Africa, the Indian Ocean east of the Maidive
Islands and the southern Indian Ocean west of Australia.

Genus Hymenodora G.O. Sars, 1877.

Carapace with no ridge or carina running the entire length of the median
lateral line from the orbital region to the hind margin, and without an
oblique ridge or carina delimiting the hind margin of the hepatic furrow.
Abdomen smoothly rounded throughout, without a dorsal carina on any
somite. Telson truncate at the tip, not terminating in a sharply pointed
end-piece laterally armed with spines. Pereiopods slender. Exopods of
third maxillipeds and pereiopods neither foliaceous nor rigid. Eyes small
and very slightly pigmented. Outer margin of antennal scale not armed with
a series of spines. Incisor process of mandible toothed for the entire length
of its cutting edge.

Only one species is known from the Bermuda area.

Hymenodora gracilis Smith.

Text-figs. 46-49.

Hymenodora gracilis Smith, 1886, p. 681, pi. 12, fig. 6.

Hymenodora gracilis Stephensen, 1923, p. 60.

Hymenodora glacialis Chace, 1936, p. 29 (part) .

Diagnosis: Integument extremely soft and fragile, rostrum reaching
about as far as tips of eyes, a groove passing upward and backward from

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[XXV :11

Hymenodora gracilis. Male taken in August or September,

1931, from 900 fathoms. X 2.66.

the middle of the hepatic furrow but no similar groove converging toward
the first from the supra-branchial groove, end of squamous portion of an-
tennal scale truncate and not reaching beyond base of outer spine.

Measurements : The smallest recognizable males have a carapace length
of 9.0 mm. The largest males have the carapace about 13.0 mm. long. The
smallest ovigerous females are about the same size as the smallest males or
even slightly smaller. The male appears to attain a somewhat larger size
than the female in this species; if, as seems likely, the females mature at a
smaller size than the males there is no way of distinguishing these small, non-
ovigerous females from immature specimens. The smallest specimens seen
have a carapace length of 3.0 mm. Since specimens of this species are usually
in poor condition due to the extremely fragile integument, it is often difficult
to obtain accurate measurements. The eggs are very large in comparison
with the size of the animal, measuring 2.3 by 2.0 mm.

Color in Life : Two slightly different shades of color were noted in Ber-
muda specimens of this species. In two ovigerous females taken in Net
1099 the body and appendages were bright scarlet with a faintly yellowish
tinge to the tips of the appendages and to the setae. The eyes were buffish,
with the facets outlined in brownish-black. The eggs were pansy-purple
with the developing embryo pale peach-colored. In another ovigerous speci-
men taken in Net 1106 the body and appendages were grenadine red with
the scarlet red heart showing through the dorsal part of the carapace. The
eyes were as in the above specimens. The eggs, which were not well-developed
in this specimen, were spectrum red.

General Remarks: The collection contains 1,829 specimens; 73 males,
206 females, of which 78 are ovigerous, and 1,550 young.

Dr. Stanley Kemp has convinced me that H. gracilis is distinct from
H. glacialis with which it has often been confused. I believe that Dr. Kemp
will clarify the differences in a forthcoming paper; the diagnosis given
above is sufficient to recognize the species, the only one I have seen from the
Bermuda area. The only additional point of taxonomic interest is the fact

1940]

Chace: Bathy pelagic Caridean Crustacea

177

Text-figure 47.

Seasonal distribution of ovigerous females of Hymenodora
gracilis computed from specimens taken in 1929 and 1930.

that a few of the specimens have five or six terminal spines on the telson
instead of the usual four.

Seasonal Distribution: There is no apparent monthly fluctuation in the
numbers of either adults or young taken during that part of the year in
which towing was carried on. Naturally, there were proportionately more
young taken in those months in which a large part of the tows were made
above 900 or 1,000 fathoms and conversely, a greater percentage of adults
were taken during the months when more of the deeper tows were made,
but when all of these factors are taken into consideration there are no strik-
ing disclosures regarding the seasonal distribution of the population of
H. gracilis. In contrast to the results obtained with Acanthephyra purpurea,
however, there appears to be an increase in the numbers of ovigerous females
during the latter part of the summer as shown in Text-fig. 47. Although
these results may be entirely due to chance, it is difficult to suppose that the
percentage error would be great enough to account for the fact that out of
forty females taken in June only nine were ovigerous, whereas out of forty-
two females in September twenty-one were carrying eggs.

Vertical Distribution : The vertical distribution of the entire catch of
H. gracilis is shown in the following table:

Fathoms

Males

Females

Y oung

400

1

1

500

1

17

600

1

1

83

700

193

800

10

334

900

15

417

1,000

45

503

1,100

1

2

1,200

..........1 .............

The distribution of the adult and young when reduced to the number
of specimens taken in an average hour of towing at each depth is shown in
Text-fig. 48. All of these specimens were taken in the daytime. This distribu-
tion naturally suggests that a large part of the adult portion of the popula-

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[XXV : 11

■700

J~00 700 £00

900

/000

Text-figure 48.

Vertical distribution of adults (solid bars) and young (hatched)

bars) of Hymenodora gracilis.

tion is usually found at depths exceeding 1,000 fathoms. That they were
not taken in greater numbers at 1,100 and 1,200 fathoms may be due to the
fact that these depths are too near the bottom in the area investigated.
It is not unlikely that additional towing at greater depths in this general
region might reveal a much greater proportion of adults. An increase in the
proportionate number of males as the depth becomes greater, similar to the
vertical distribution of the sexes in Acanthephyra purpurea, is apparent
in this species as shown in Text-fig. 49. Since the females are more numer-
ous at every haul, it is probable that this sex is normally more abundant
than the males in this species, but one cannot entirely ignore the possibility
that if the increase in the male component of the population continues with
increase in depth to 2,000 fathoms or over this sex ratio might be more nearly
equalized; if, in the case of Acanthephyra pur-purea, only specimens from
600 fathoms and above had been seen, one would assume that the females
were distinctly in the majority, but when the deeper hauls are considered
just the reverse proves to be true. It will be noticed from the table that the
ovigerous females were fairly well scattered from 700 to 1,000 fathoms,
varying from 35 to 42 per cent, of the adult females at those levels.

Geographical Distribution : Since H. gracilis has so frequently been con-
fused with H. glacialis, its distribution is very imperfectly known. All of
the specimens which 1 have seen from the temperate North Atlantic, with
the exception of a single specimen taken southeast of Georges Bank, appear

1940]

Chace: B at hy pelagic Caridean Crustacea

179

'****/'& j- /n ae/is//" jeo^r t//trfSO*r

7S SO

1

I

700

X

\ 300

—

X

* 900

—

\

/ooo

bboi:"

I I

2S SO 7S

Text-figure 49.

Relative abundance of males and females of Hymenodora
gracilis from 700 to 1,000 fathoms. Solid bars represent the male
fraction of the adult population; hatched bars the female.

to belong to H. gracilis; possibly H. glacialis is usually confined to the arctic
and subarctic regions in the North Atlantic. However, the specimens from
the Gulf of Panama referred to this species by Faxon (1895, p. 168) prove
to be typical specimens of H. glacialis.

Genus Systellaspis Bate, 1888.

Carapace without a straight ridge or carina running along the median
lateral line from the orbit to the hind margin, and without an oblique ridge
or carina delimiting the hind margin of the hepatic furrow. Abdomen never
dorsally carinate on the sixth somite. Telson not truncate at the tip but
terminating in a sharply pointed end-piece latei'ally armed with spines.
Pereiopods not abnormally broad and flattened. Exopods of third maxilli-
peds and pereiopods neither foliaceous nor rigid. Eyes well pigmented and
usually large. Outer margin of antennal scale not armed with a series of
spines. Incisor process of mandible toothed for the entire length of its cut-
ting edge.

Key to the Bermudian Species of Systellaspis.

Rostrum triangular and less than half as long as carapace; hind margins of
fourth and fifth abdominal somites not denticulate; sixth somite at
least twice as long as fifth S. braueri.

Rostrum slender and longer than carapace; hind margins of fourth and fifth
abdominal somites denticulate; sixth somite less than twice as long
as fifth S. debilis.

180

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Zoologica: New York Zoological Society

Systellaspis braueri (Balss) .

Text-fig. 50.

? Systellaspis echinurus Coutiere, 1911, p. 158.

Acanthephyra braueri Balss, 1914, p. 594.

Systellaspis densispina Stephensen, 1923, p. 57, fig. 17.

Systellaspis braueri Balss, 1925, p. 245, text-figs. 17-20, pi. 21.

Diagnosis'. Rostrum triangular and less than half as long as carapace.
No sharp dorsal carina on any abdominal somite; hind margins of fourth
and fifth somites not denticulate; a blunt lobe on anterior margin of first
abdominal somite overlapping carapace; a deep notch in hind margin of fifth
pleuron ; sixth somite at least twice as long as fifth.

Color in Life : Unknown.

Remarks'. The collection contains thirteen specimens of this species,
ranging in size from 4.2 to 14.3 mm. in carapace length. All but two of
these specimens are obviously young. The two largest specimens, possibly
mature females, were taken in June and September, 1929, and the others
were scattered from May to September of 1929, 1930 and 1931. The depths
in which they were found varied from 700 to 1,000 fathoms, only the smaller
specimens being taken in the shallower depths and the two largest in 1,000
fathoms.

As Stephensen has pointed out, it is very likely that S. echinurus Cou-
tiere belongs to this species, but the description is much too inadequate to
permit recognition of the earlier name. Balss is probably correct in com-
bining S. densispina Stephensen with this species.

Text-figure 50.

Systellaspis braueri. Female taken in September, 1929, from
1,000 fathoms. X 2.13.

Geographical Distribution : Only five or six specimens of S. braueri have
been recorded heretofore. They were found in the Bay of Biscay, in the
Gulf of Guinea off the west coast of Africa and in the Bay of Bengal off
Ceylon. There is one additional specimen in the U. S. National Museum
which was taken by the Albatross off Los Angeles, California, and a large
female has recently been acquired by the Museum of Comparative Zoology
from southeast of Georges Bank.

1940] Chace: Bathypelagic Caridean Crustacea 181

Systellaspis debilis (A. Milne Edwards).

Text-figs. 51-53.

Acanthephyra debilis A. Milne Edwards, 1881a, p. 13.

Miersia gracilis Smith, 1882, p. 70, pi. 11, figs. 4-4d.

Acanthephyra debilis var. Europoea A. Milne Edwards, 1883.

Acanthephyra gracilis Smith, 1886, p. 672.

Systellaspis debilis Coutiere, 1905, p. 5, fig. 2.

Systellaspis bouvieri Coutiere, 1905, p. 8, fig. 3.

Systellaspis debilis var. indica de Man, 1920, p. 51, pi. 6, figs. 11-llf.

Systellaspis debilis Balss, 1925, p. 242.

Diagnosis : Rostrum slender and usually longer than carapace. Abdomen
dorsally carinate on most of third and posterior portion of fourth somites;
hind margins of fourth and fifth somites denticulate either side of median
spine; an acute lobe on anterior margin of first abdominal somite overlapping
carapace; a notch in hind margin of fifth pleuron, sixth somite less than
twice as long as fifth.

Measurements : The smallest recognizable males show some slight varia-
tion in size, having a carapace length of from 7.2 to 7.8 mm. The largest
males have a carapace length of about 14.0 mm. and the largest females about
14.2 mm. The smallest ovigerous females examined had a carapace length
of 10.9 mm. The youngest specimens have the carapace 2.8 mm. long. The
eggs vary somewhat in size depending on their degree of development and
state of preservation, but their average measurements are about 3.7 by
1.9 mm.

Color in Life : The largest adult specimens have the entire body and
appendages, except for the eyes and light organs, scarlet-red, with the
antennal scales, antennae, pleopods and uropods tinged with salmon-orange.
The eyes are dark brown; the light organs have deep bluish-purple centers
and dark red reflectors.. The eggs are spectrum red. In somewhat smaller
individuals the body color varies from scarlet-red to white splotched with

Text-figure 51.

Systellaspis debilis. Male taken in June, 1929, from 600 fath-
oms. X 2.12.

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Text-figure 52.

Seasonal distribution of adults (solid bars) and young (hatched
bars) of Systellaspis debilis computed from specimens taken in 1929
and 1930.

scarlet, the abdomen frequently showing considerable white in young speci-
mens; in those specimens which are not of maximum size the tips of the
antennae, rostrum, pleopods, uropods and telson are nearly white.

General Remarks : The collection contains 1,949 specimens of this
species; 470 males, 567 females, of which 327 are ovigerous, and 912 young.
Of the ovigerous specimens, 66 carry eggs in which the eyes are visible.

S. debilis can be distinguished from all other members of the genus by
the denticulate hind margins of the terga of the fourth and fifth abdominal
somites.

Luminescence : The position of the photophores in S. debilis is given by
Kemp, 1919 (p. 64). Harvey, 1931, discusses the chemistry of the lumin-
escence in this form and also reports that specimens can be stimulated to emit
“a cloud of bluish luminescent secretion . . . from glands near the mouth.”
Observations on the luminescence of living specimens will be found on page
204 of the present paper.

Seasonal Distribution : The monthly distribution of the adults and young
of this species in 1929 and 1930 is shown in Text-fig. 52. Obviously, the
fluctuation in the population does not appear to follow any definite plan.
Curiously enough, the young population seems to remain more stable than
the adult. The seasonal distribution of ovigerous females similarly discloses
little of significance; in each month of 1929 and 1930 in which towing was
done, with the exception of April, more than half of the females were carry-
ing eggs and more than six per cent, of the female population carried eggs
in which the eyes are visible. When one examines the yearly hauls for 1929,
1930 and 1931, it is found that in 1929 the yield of this species per hour
of towing was 0.249 adults and 0.224 young, whereas in 1930 and 1931 this
figure was 0.190 for the adults in each year and 0.200 and 0.114 respectively
for the young. However, it is found that the numbers of ovigerous females
per hour of towing in 1929 and 1930 were nearly the same, 0.075 and 0.073

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183

Text-figure 53.

Vertical distribution of adults (solid bars) and young (hatched
bars) of Systellaspis debilis.

respectively. Of course, this also reveals a dissimilarity in the proportion of
adult females which were ovigerous in the two years; in 1929 only 58.2 per
cent, were ovigerous, while in 1930 75.5 per cent, were carrying eggs, and
these proportions remain fairly uniform when these years are analyzed
month by month. It is evident from an examination of the collection that
practically all of the females in which the carapace is more than about 11.5
mm. long were ovigerous, so the explanation of the smaller proportion of
ovigerous specimens in 1929 must be that young females with a carapace
length of from 7.5 to 11.5 mm. were more abundant in that year than in
1930. On the other hand, although adult specimens seemed to be just as
abundant in 1931 as in 1930, only 0.048 ovigerous females were taken per
hour of towing and only 43.6 per cent, of the adult females were carrying
eggs. Since towing in 1931 was carried on from July to October, while in
1930 it was done from May to September, it is somewhat hazardous to com-
pare these two years in this manner. However, it would appear that young
females were even more abundant in 1931 than in 1929.

Vertical Distribution : The vertical distribution of the total catch is as
follows :

Fathoms

100

200

Males

Females

Young

300

2

14

400

13

23 (3 ovig.)

90

500

153

329

600

127

125 (73 ovig.) . . . .

222

700

92

120 (80 ovig.) . . . .

103

800

34

32 (21 ovig.)

41

900

28

29 (20 ovig.)

59

1,000

1 100

21

52

T200

1

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The relation of these specimens to the number of hours of towing is
shown in Text-fig. 53. All of these specimens were taken in the daytime.
Apparently the center of distribution of this species is at 500 or 600 fathoms
for the adults and 400 fathoms or above for the young. The rapid thinning
out of the population of S. debilis above 500 fathoms and the fact that a
single young specimen was found at 100 fathoms tend to support observa-
tions made in the bathysphere by Beebe (1934, pp. 303 and 340) that speci-
mens are present in the upper levels, but that they may be able to avoid the
nets in this better lighted zone. From night hauls made by Atlantis in this
general area, it is known that S. debilis can be taken at levels at least as
shallow as 220 fathoms during the hours of darkness (Welsh, Chace &
Nunnemacher, 1937, p. 190). It will be seen from the table that at every
depth at which significant numbers of this species were taken more than half
of the adult females were ovigerous. Likewise, there was no strikingly large
proportion of those carrying eyed eggs at any particular level. There is no
evidence in this species that males are proportionately more abundant at
deeper levels as was noted in Acanthephyra purpurea; in fact, the greatest
proportion of females was found at 1,000 fathoms.

Geographical Distribution: S. debilis appears to be most abundant in
the North Atlantic, but it has been recorded as well from the eastern South
Atlantic, the Indian Ocean, the Malay Archipelago and the Hawaiian Islands.

Genus Oplophorus H. Milne Edwards, 1837.

Carapace without a straight ridge or carina running along the median
lateral line, and without an oblique ridge or carina delimiting the hind mar-
gin of the hepatic furrow. Abdomen armed with remarkably long spines on
either the second, third and fourth or the third, fourth and fifth somites;
it is usually carinate on those somites provided with spines and not on the
others. Telson terminating in a sharp point which may or may not be armed
with lateral spines. Pereiopods short and slender. Exopods of third maxil-
ilpeds and pereiopods foliaceous and usually rigid. Eyes large and well pig-
mented. Outer margin of antennal scale usually armed with a series of
spines. Incisor process of mandible toothed for the entire length of its cut-
ting edge.

Key to the Bermudian BathypElagic Species of Oplophorus.

Abdomen armed with long spines on second, third and fourth somites

O. spinicauda.

Abdomen armed with long spines on third, fourth and fifth somites

O. grimaldii.

Oplophorus spinicauda A. Milne Edwards.

Text-fig. 54.

Oplophorus spinicauda A. Milne Edwards, 1883.

Oplophorus foliaceus Rathbun, 1906, p. 922, pi. 20, fig. 8.

H oplophorus foliaceus Balss, 1925, p. 249.

Acanthephyra anomala Boone, 1927, p. 104, fig. 21 (holotype only).
Oplophorus spinicauda Chace, 1936, p. 30.

Diagnosis : Second, third and fourth abdominal somites terminating in
a long spine. No spine at postero-lateral angle of carapace. Outer margin of
antennal scale spinose. Exopods of all pereiopods foliaceous but not rigid.
Telson terminating in an end-piece armed with lateral spines.

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Chace: B at hy pelagic Caridean Crustacea

185

Color in Life: Entire integument transparent with scarlet chromato-
phores scattered over the antennal peduncles, the mouth parts, the pereio-
pods, the basal segment of the pleopods and the sides of the abdomen. The
anterior internal thoracic organs scarlet, with a more posterior mass of
spectrum red. Eyes blackish-brown.

Remarks: There are nine specimens of this species in the collection;
one with a carapace length of 6.2 mm. that is possibly a young male, and
the rest, apparently young, with a carapace length of from 4.0 to 5.2 mm.
The young male was taken September 15 to 29, 1930, in 600 fathoms, and
the young were found in August or September, 1929, July and September,
1930, June and July, 1931, and July, 1934, in depths ranging from 300 to
800 fathoms.

O. spinicauda shows a relationship to members of the genus Systellaspis
in the form of the telson, the non-rigid exopods of the pereiopods and its
relatively soft integument. In other characters as well, such as the presence
of spines on the second, third, and fourth abdominal somites rather than on
the third, fourth and fifth, the very much reduced molar process of the man-
dible and the third segment of the endopod of the second maxillipeds ves-
tigial, this species differs strikingly from all other known members of
Oplophorus.

Oplophorus spinicauda. Young male taken in September, 1930,
from 600 fathoms. X 3.39.

Examination of larger series of the species from other localities indi-
cates that there is no appendix masculina on the second pleopod in this form
and the endopod of the first pleopod fails to disclose any prominent sexual
differences. The sexes can apparently be distinguished, however, by the
presence in the male of a broad, shallow indentation in the lower margin of
the first abdominal pleuron; in the male, then, the lower margin of this
somite is more or less sinuous in outline, whereas in the female and young
it is more evenly convex.

There is little doubt in my mind, after examining the type of O. foliaceus
Rathbun, that these species are identical. Miss Rathbun has informed me
that at the time she described the species Milne Edwards’ “Recueil de Fig-
ures” was not available. The holotype of Acanthephyra anomala Boone is
a young specimen of O. spinicauda, while the partype mentioned in the de-
scription is apparently a young specimen of O. grimaldii Coutiere.

The following remarks were made by Miss Crane on a specimen of this
species :

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“The shrimp showed a great deal of spasmodic activity both on the ice
and off until placed in formalin twenty-eight hours after it was brought in ;
at all times it kept up a continuous and vigorous kicking of its pleopods.”

Luminescence : The following description of the distribution of photo-
phores in an immature specimen of this species is taken from notes taken
from fresh material by Miss Crane.

There is an elongate scarlet patch on a background of pale violet blue
situated on the anteroventral surface of the eye, while on the posterior side
is another, much larger, scarlet patch. One or both of these may be lumin-
escent. On the thorax, light organs are found on the third maxilliped and
the third, fourth and fifth pairs of walking legs. Those on the maxilliped are
in two series at the proximal and distal ends of the dactyl, and consist of
six and three minute organs respectively; they are posteriorly directed and
of a golden color set in scarlet cups. Both the series arrangement and the
general appearance bear a striking resemblance to the photophores of
Argyropelecus. At the proximal end of the carpus of the third walking leg
is situated a much larger organ, gold set in scarlet as above, but perfectly
round and directed only very slightly downward and forward. On the fourth
walking leg are two similar organs, one in a corresponding position and the
other at the proximal end of the propodus. Three such organs are present
on the fifth leg, two arranged as on the preceding, and the third (much
longer — about four times as long as broad) is located on the distal end of
the propodus. In addition to the pereiopodal organs, an elongate scarlet
patch, without any gold visible and possibly glandular in nature, is placed
at the base of the last pair of legs beneath the carapace. On the abdomen
there is a photophore directed forward and downward at the base of each
pleopod; these are of about the same size as those on the thoracic legs, but
the scarlet reflector is more prominent than the golden inner part. Finally,
there is a smaller, similarly designed light at the inferior base of the uropods.

The following remarks are taken directly from notes made on the same
specimen by Dr. Beebe.

“This small shrimp was taken alive in the big wire net at 5-7 knots on
June 12th.

“It was invisible in the dark-room until the glass was tapped or the
water stirred when a solid luminous spot of pale, bluish-white light shone
out steadily. If the shrimp was disturbed further this light vanished and a
squirt of luminosity appeared, bright at the source and rapidly fading out
as it shot into the water. The visible length seemed about three times the
length of the shrimp, and once at least the direction was at right angles
downward from the body (about 5:30 P.M.). This phenomenon was re-
peated four times and it had been noticed before once. This first time
(noticed by Miss Crane) there was the steady glow for a moment, then a
greater amount of fire shot out than at any of the later times. The last time
I saw it, it was as strong as the first. After dark (8 P.M.) the shrimp gave
out no light whatever, but the water had a considerable amount of flocculent
material on the bottom. This showed no especial structure under the lens.”

Geographical Distribution: O. spinicauda has been recorded from off the
coast of British Honduras, off the coast of Morocco, the Indian Ocean north
of Madagascar and south of India and the Hawaiian Islands. In addition,
I have seen specimens in the U. S. National Museum from off the east coast
of Florida, the West Indies and the Philippine region. In the North Atlantic
this species seems to be much more abundant in the West Indian region than
in the Bermuda area.

1940] Chace: Bathypelagic Caridean Crustacea 187

Oplophorus grimaldii Coutiere.

Text-fig. 55.

Hoplophorus grimaldii Coutiere, 1905a, p. 1114; 1905b, p. 1, fig. 1.

Hoplophorus grimaldii Coutiere, 1925, p. 249, fig. 24.

Diagnosis : Third, fourth and fifth abdominal somites terminating in a
long spine. Usually no spine at postero-lateral angle of carapace. Outer
margin of antennal scale spinose, and inner margin usually with a barb near
the tip. Exopods of all pereiopods foliaceous and more or less rigid. Telson
terminating in a long spine, but not an end-piece armed with lateral spines.

Measurements : Immature males in which the appendix masculina is
shorter than the usual appendix interna have a carapace length varying
from 7.3 to 8.0 mm. The largest male in the collection has the carapace 12.2
mm. long and the largest female, 14.2 mm. In the smallest ovigerous female
the carapace measures 11.2 mm., and all females exceeding this size are
carrying eggs. The smallest specimen has a carapace length of 2.7 mm. In
the three specimens in the collection in which the carapace measures less
than 3.0 mm. the spine on the third abdominal somite and the rostrum are
extremely long, the latter being from two and two-thirds to three times as
long as the carapace. The eggs measure 2.8 by 1.8 mm.

Color in Life : Thorax and abdomen white with a heavy wash of scarlet-
red chromatophores on the sides; since these are confined on the abdomen
to the central part of each somite, a banded appearance is produced. There
are golden reflections along the smooth, shiny sides of the carapace. Fre-
quently the abdominal pleura are tinted pale violet. The antennae are trans-
parent except for scarlet-red chromatophores scattered at their bases. Mouth
parts brilliant spectrum-red. Pereiopods somewhat lighter, shading to al-
most transparent scarlet. Pleopods salmon-orange proximally with the exo-
pods and endopods translucent white. Uropods, telson and abdominal spines
transparent. Eye dark-brown, the central portion gleaming with reddish
gold and orange. Eggs carmine to scarlet-red.

Luminescence : The following notes on the distribution of light organs
in this species were made from living material by Miss Crane.

“Behind the eye is a translucent scarlet-red projection from the eye
itself enclosing a darker oblong organ; in front a similar but lighter struc-
ture is found. One or both are possibly luminescent. On the distal seg-
ment of the last maxilliped, extending the entire length of the posterior bor-
der (ventral when flexed), is a row of about 45 minute, spectrum-red organs
above a long, unbroken line of blue violet which probably acts as a reflector.
The two organs, with their segment of the blue line, at the extreme tip are
isolated somewhat from the others. A large silvery or golden organ set in
purplish-blue occupies most of the interior of the short carpus on the third,
fourth and fifth legs. The last (fifth) leg has in addition two organs on the
long hand — an elongated oval one proximally about the size of that in the
carpal segment, and a much elongated one distally. These leg organs are
visible from both the outer and inner side through clear, window-like trans-
parencies in the cuticula. From the inside, though, they appear brilliant
scarlet-red as if seen through shiny red cellophane. They cannot be said to
be directed in any special direction, except perhaps slightly backwards. Be-
hind the last leg is an elongated transverse bar, the outer half spectrum-red,
the inner deep violet-blue. A series of five peculiar tri-colored spots is strung
in a broad arc along the carapace, cutting off the lower posterior corner.

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On close examination it appears that each spot consists of an upper, upwardly
directed bright red light with a violet-blue half-moon reflector placed below
it, the interval between light and reflector being white. The abdominal or-
gans consist of six pairs of deep bluish-purple lights set in scarlet-red
sockets and placed beneath the epimera on the anterior outer side of the
first segment of each pleopod and uropod. They are directed forwards and
downwards.”

No luminescence was produced in this species by stimulation of live
specimens. However, Miss Crane has noted the following of one specimen
placed in preservative while still alive :

“Immediately after being placed in alcohol a large quantity of white
flocculent material was exuded, apparently chiefly from the posterior part
of the ventral side of the thorax, perhaps from the elongated, transverse,
violet and red bar previously described.”

Text-figure 55.

Oplophorus grimaldii. Male taken in June, 1931, from 300 fath-
oms. X 2.71.

General Remarks : There are 48 specimens in the collection ; 8 males, 19
females, of which 8 are ovigerous, and 21 young.

O. grimaldii can be readily distinguished from the preceding species by
the different arrangement of the abdominal spines, but it is more difficult
to separate it from O. gracilirostris A. Milne Edwards (1881). In general,
O. grimaldii has the end of the antennal scale barbed on the inner margin
and no tooth or spine at the postero-lateral angle of the carapace, whereas
O. gracilirostris has no barb on the antennal scale and bears a prominent
spine at the postero-lateral angle of the carapace. However, four of the
present specimens lack the barb on the scale, possibly due to injury, and
three of the ovigerous females have a blunt tooth at the postero-lateral angle
of the carapace. Fortunately neither of these conditions occurs in the same
individual of the present series, but it is not improbable that specimens may
be found which might be assigned to either species.

Although there is a well developed appendix masculina in the adult
males of this species, there is also a sexual distinction involving the first
abdominal pleuron as in the preceding species; in the male, the lower
anterior angle of this pleuron is deeply incised so that the angle formed
approaches a right angle, while in the female this angle, if present at all,
is very obtuse.

1940]

Chace: Bathy pelagic Caridean Crustacea

189

One specimen of this species was found in the stomach of the rock fish,
Mycteroperca bowersi, caught off Challenger Bank, July 30, 1930.

Seasonal Distribution: By far the largest number of specimens of both
adults and young were taken in June and July of 1929, 1930 and 1931. Since
there was no great concentration of 300 to 500 fathoms tows during these
months, it is possible that the species really is most abundant in the Bermuda
area in mid-summer. The largest numbers of shallow hauls were made in
May, but only one young specimen was found in that month of any year.
However, at least one specimen was taken in every month from April to
September and ovigerous females occurred in June, July, August and Sep-
tember, the only specimen in which the eggs were eyed being taken in
August, 1929.

Vertical Distribution: The largest number of specimens were found at
300, 400 and 500 fathoms, although two or more specimens were taken at
every level from 300 to 1,000 fathoms. As this species has been taken at
night in the South Atlantic in some numbers only ten meters below the sur-
face, it is possible that a large part of the population occurs above 300
fathoms.

Geographical Distribution: O. grimaldii has been recorded from the
North Atlantic west of the Madeira Islands, the South Atlantic north of
Tristan da Cunha and the Indian Ocean west of Australia. I have also seen
specimens from off the east coast of the United States, the Bahamas, south
of Japan and northeast of Easter Island in the eastern Pacific.

Family Bresiliidae.

Rostrum well developed. First two pairs of pereiopods chelate, shorter
than the last three pairs and with an undivided carpus. Last three pairs
not abnormally long. Exopods present on the maxillipeds and some or all
of the pereiopods. Terminal joint of second maxilliped normal, not applied
as a strip to the end of the preceding joint. Mandible distinctly, but not
deeply cleft.

One genus only is known from the Bermuda area.

Genus Lucaya Chace, 1939.

Carapace without a branchiostegal spine. Rostrum unarmed on the
lower margin. All five pairs of pereiopods provided with exopods, and the
last three pairs decrease in length posteriorly. Eyes well pigmented.

There is but one known species.

Lucaya higelowi Chace.

Text-fig. 56.

Lucaya bigeloivi Chace, 1939, p. 34.

Diagnosis: Rostrum subquadrate in cross section. Abdomen without
carina or armature of any kind; telson with two pairs of dorso-lateral spin-
ules. Antennal scale broad with the blade extending far beyond the outer
spine.

Color in Life: Entire animal pale salmon except for the rostrum, an-
tennae and antennal scales which were scarlet.

Remarks: Two males in the present collection bring the total number
of recorded specimens of this species to three. One male, with a carapace
length of 4.2 mm., was taken in Net 18, April 12, 1929, in 700 fathoms; the
other, with a carapace length of 4.7 mm., came from Net 940, September 24,
1930, 1,000 fathoms.

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Lucaya bigelowi. Holotype male from the Bahamas. X 6.02.

More complete figures of this species are to be included in the final
report on the Crustacea of the Atlantis Expeditions to the West Indies.

Geographical Distribution : The holotype came from east of Great Abaco
Island in the Bahamas.

Family Pandalidae.

Rostrum prominent and dentate. First pair of pereiopods simple or
microscopically chelate. Second pair of pereiopods with the carpus seg-
mented. No exopods on any of the pereiopods. Terminal joint of the second
maxillipeds usually applied as a strip to the end of the preceding joint.
Mandibles deeply cleft, with a three-jointed palp.

Key to the Bermudian Bathypelagic Genera of Pandalidae.

Epipods present on all but the last pair of pereiopods Plesionika.

No epipods on any of the pereiopods Parapandalus.

Genus Plesionika Bate, 1888.

Carapace without lateral longitudinal carinae. Rostrum immovable and
armed dorsally with fixed spines and sometimes with movable spines also.
Eyes much wider than eyestalks. Stylocerite pointed. Third maxillipeds
with an exopod. Epipods present on all but the last pair of pereiopods.
Carpus of second pereiopods composed of more than three segments.

One bathypelagic species is known from the Bermuda area.

Plesionika martia (A. Milne Edwards).

Text-fig. 57.

Pandcdus martius A. Milne Edwards, 1883, pi. 21.

Plesionika semilaevis Bate, 1888, p. 644, pi. 113, fig. 3.

Plesionika martia Kemp, 1910, p. 93, pi. 12, figs. 1-4.

Plesionika martia Balss, 1925, p. 278.

1940]

Chace: Bathypelagic Caridean Crustacea

191

Plesionika martia. Ovigerous female (after Kemp, 1910). X 0.64.

Diagnosis : Rostrum longer than antennal scale, dorsally smooth beyond
antennular peduncle and regularly serrate below. Hind margin of third
abdominal somite rounded. Second pereiopods subequal in length.

Color in Life : The Bermuda specimen was pink and white. The follow-
ing more complete color notes are quoted from Kemp, 1910 (p. 95) :

“The carapace and abdomen are thickly sprinkled with bright red
chromatophores ; the former is dorsally of a dark purple tint, while in the
latter the red pigmentation is darker on the posterior portions of each
somite. The rostrum is bright red distally, less deeply coloured proximally.
The eyes are black, with golden reflections; the antennules are red, and the
antenna and antennal scale are more sparsely pigmented with the same
colour. The outer maxillipeds and pereiopods are more or less thickly
spotted with red; the pleopods, telson and uropods are light red. All the
fringes of the setae are golden in colour.”

Remarks: A single young specimen, having a carapace length of 7.0
mm., was taken in Net 860, September 8, 1930, in 600 fathoms.

This is the first record for this species in the western Atlantic and one
of the very few specimens ever taken with a mid-water net. Apparently P.
martia generally remains on or near the bottom as indicated by the long,
slender pereiopods, since most of the known specimens have been taken
with a dredge or trawl. However, Balss records an ovigerous female cap-
tured by the Valdivia in the Gulf of Guinea with a vertical net between
437 fathoms and the surface.

Geographical Distribution: P. martia, including its varieties, has been
found off Ireland, in the Bay of Biscay, the Mediterranean Sea, the Gulf of
Guinea off the west coast of Africa, off the Cape of Good Hope, off the east
coast of Africa from Tanganyika to Italian Somaliland, in the Arabian Sea,
the Andaman Sea, the Malay Archipelago, Sagami Bay, Japan, the Fiji
Islands, off Sydney, Australia, the Kermadec Islands and the Hawaiian
Islands.

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Zoologica: New York Zoological Society

Genus Parapandalus Borradaile, 1900.

Carapace without lateral longitudinal carinae. Rostrum immovable and
armed dorsally with fixed spines and sometimes with movable spines as well.
Eyes much wider than eyestalks. Stylocerite pointed. Third maxillipeds
' with an exopod. No epipods on any of the pereiopods. Carpus of second
pereiopods composed of more than three segments.

Only one bathypelagic species is known from the Bermuda area.

Parapandalus richardi (Coutiere).

Text-figs. 58-61.

Pandalus ( Stylopandalus ) Richardi Coutiere, 1905b, p. 18, fig. 6.

Parapandalus Richardi de Man, 1920, pp. 108 and 140.

Diagnosis: Rostrum about three times as long as carapace and armed
with more or less widely spaced teeth along the entire length of its dorsal
margin, including two larger teeth above the eye. Third abdominal somite
at middle of its posterior margin with a slender, movable spine. Sixth ab-
dominal somite at least three times as long as high and about two and two-
thirds times as long as fifth somite.

Measurements : The smallest recognizable males in which the appendix
masculina is no more than a bud have a carapace length of 4.2 mm., and the
smallest ovigerous females have the carapace 5.8 mm. long. In ovigerous
females in which the eggs are not eyed, the average carapace length is 6.7
mm., whereas in those in which the eggs are eyed this figure is 6.5 mm.;
this may be due to a slight shrinking of the integument prior to the moult
following the hatching of the eggs. The carapace of the largest male meas-
ures 7.2 mm., and of the largest female 7.5 mm. In the smallest specimen
seen the carapace is 2.9 mm. long, but in most of the young specimens this
measurement exceeds 3.5 mm.

Color in Life : In mature individuals, the entire prawn is bright scarlet,
though paler on the abdomen where the color is resolvable into individual

Parapandalus richardi. Male taken in June, July or August,
1931, from 500 fathoms. X 3.05.

1940]

Chace: Bathy pelagic Caridean Crustacea

193

chromatophores on a white background. In young specimens, the anterior
part of the body as far back as the third abdominal somite dorsally and the
fourth pereiopod ventrally is bright scarlet including the appendages ; be-
hind this the prawn is white, with the line of demarcation between the two
regions yellowish sprinkled with scarlet chromatophores; there is a faint
sprinkling of chromatophores along the sides of the abdomen also. Eyes
brownish-black. Eggs pomegranate purple.

General Remarks : There are 404 specimens of this species in the collec-
tion; 166 males, 195 females, of which 105 are ovigerous, and 43 young.
Of the ovigerous females, 30 carry eggs in which the eyes are developed.

Table 1.

Seasonal Distribution of P. richardi in 1929.

Carapace

Length

(mm.)

3.0

.3.5

April

May

June

July

Aug.

Sept.

1

1

4

1

4.0

4

—

4

4

6

4.5

7

1

2

—

1

4

5.0

12

3

5

1

1

2

5.5

6

4

—

1

1

6.0

4

7

20

6

3

8

6.5

1

—

21

20

4

5

7.0

—

—

3

17

5

7

7.5

Table 2.

1

1

Seasonal Distribution of P. richardi in 1930.

Length

Carapace

April

May

June

July

Aug.

Sept.

(mm.)

3.0

£

—

—

—

—

1

3.5

1

—

—

—

2

4.0

o»

—

1

—

—

7

4.5

ci

4

1

1

—

2

5.0

10

3

1

1

4

5.5

6.0

in &

o

8

7

2

8

11

—

4

3

6.5

2

3

12

3

4

7.0

o

1

—

7

—

10

7.5

—

—

—

1

1

Table 3.

Vertical Distribution of

P. richardi.

Carapace

Length

(mm.)

Fathoms

300

400

500

600

700

800

900

1,000

3.0

3.5

—

1

1

3

—

—

1

3

1

2

4.0

3

2

12

2

1

1

1

2

4.5

—

4

10

1

—

—

1

5

5.0

1

8

16

10

3

4

1

3

5.5

1

1

11

4

7

1

2

1

6.0

—

3

25

19

11

6

9

9

6.5

3

4

31

25

14

5

6

9

7.0

3

6

34

10

14

5

1

4

7.5

—

—

3

3

1

—

—

—

194

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Text-figure 59.

Average carapace lengths of specimens of Parapandalus
richardi taken in 1929 (solid bars) and 1930 (hatched bars).

This species is very similar to P. zurstrasseni Balss (1914) from the
Indo-Pacific region. Whether or not they are identical can be determined
only when more specimens of the latter species are discovered. P. zurstras-
seni apparently lacks the slender spine on the third abdominal somite, but
this is frequently broken off in specimens of P. richardi.

Seasonal Distribution'. Analysis of the tows made in 1929 and 1930
reveals a very definite increase in the proportion of ovigerous females from
about 10 per cent, of all females in April and May to 70 or 80 per cent, in
July, August and September. However, as shown in Tables 1 and 2 and in
Text-fig. 59, this is largely due to the fact that most of the females present
in the spring are too young to bear eggs. If only those females in which the
carapace exceeds 6.0 mm. are considered, it is found that from 60 to 100 per
cent, are ovigerous and 15 to 30 per cent, are carrying eggs in which the eyes
are visible in every month in which towing was done. The seasonal distribu-
tion of the size groups shown in Text-fig. 59 indicates that one group matures
in July or August and that another group of younger individuals appears at
this time. Study of the tables, however, shows that this lowering of the aver-
age carapace length is due fully as much to the decrease in the numbers of
large specimens as to the increase of the small. The graph might lead to
the assumption that the season was retarded somewhat in 1930 over that of
1929, but this is due largely to the increase in average size in August, 1930,
and in this month only five specimens were taken, certainly not enough to
yield significant results. It is difficult to account for this seasonal distribu-
tion and, except for the fact that a similar trend was found in both 1929
and 1930, little importance would be laid on results gained from such a com-
paratively small number of specimens. Whether this trend is caused simply
by the migration of fully mature individuals through the towing area in
greatest numbers in June and July or whether it is due to the normal growth
of more than one age class can hardly be determined from these data. There
is evidence that swarming occurs to some extent in this species since 65 of

1940]

Chace: Bathypelagic Caridean Crustacea

195

a/s _

300 000 S00 600 700 000 900 /000

/>? faff/fasn'T

Text-figure 60.

Vertical distribution of adults (solid bars) and young (hatched
bars) of Parapandalus richardi.

the specimens were taken in only seven nets; one in May, two in June and
four in July. This, of course, may be due merely to a general increase in the
number of individuals for a short period rather than actual swarming, but
it has a definite effect on the results.

Vertical Distribution: The vertical distribution of the total catch is
given in the following table:

Fathoms

Males

Females

Young

300

4

4 (3 ovig.)

3

400

12

14 (3 ovig.)

4

500

59

23

600.

34

40 (20 ovig. ) . . . .

700

22

28 (17 ovig.) . . . .

1

800

11

5

900

8

11 (5 ovig.) . . . .

1

1,000

14

18 (11 ovig.) . . . ,

5

Reduced to the number of specimens per towing hour, these data pro-
duce the graph shown in Text-fig. 60, and the average carapace length of the
specimens at each depth is shown in Table 3 and Text-fig. 61. The optimum
level for the species is at about 500 fathoms and, since open nets were em-
ployed, it might be assumed that specimens taken in over 800 fathoms were
captured actually at higher levels as the nets were taken in ; the unexpectedly
large proportion of young specimens at these deep levels would point to the
some conclusion. However, this question can be settled only by the use of clos-
ing nets. As will be seen from the table, ovigerous females were dispersed
fairly uniformly from 300 to 1,000 fathoms, and the same applies to those
carrying eyed eggs. As in Systellaspis debilis, there is no indication that
males are proportionately more abundant at the deeper levels.

Geographical Distribution: The species has been previously recorded
only from the Canary Islands and west of the Madeira Islands. Since it has
not been taken by any of the expeditions in the northern part of the North
Atlantic, P. richardi is probably a warm water form.

196

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[XXV :11

300 #00 300 £00 700 S00 900 /000

Text-figure 61.

Variation in the average carapace length of catches of Para-
pandalus richardi made at 100 fathom intervals.

Family Physetocaridae, fam. nov.

Rostrum present as a broad prolongation of the carapace. First pereio-
pods simple. Second pereiopods chelate, with the carpus segmented. No
exopods on the third maxillipeds or any of the pereiopods. Terminal joint
of the second maxillipeds not applied as a strip to the end of the preceding
joint. Mandible without an incisor process or palp. •

Genus Physetocaris, gen. nov.

Carapace greatly inflated. Carpus of second pereiopods consisting of
four segments; chela flattened with a very short, broad dactyl. Branchial
formula as follows:

VII

VIII

IX

X

XI

XII

XIII

XIV

Podobranchiae

ep.

ep.

ep.

ep.

ep.

ep.

Arthrobranchiae

Pleurobranchiae

1

1

1

1

1

Physetocaris microphthalma, sp. nov.

Text-figs. 62 and 63.

Types : Holotype female (?) Cat. No. 30,523, Department of Tropical
Research, New York Zoological Society; Net 798; July 15, 1930; 600 fathoms.

1940]

Chace: Bathypelagic Caridean Crustacea

197

Text-figure 62.

Physetocaris microphthalma. Holotype. X 6.00.

One female (?) ; Net 983, 990, 997, 1003, 1014, 1015, 1016, 1102, 1108,
1115, 1121, 1131, 1137 (?), 1138 (?), 1144, 1149 or 1150; June 2 to August
8, 1931 ; 500 fathoms.

Diagnosis : Carapace with two lateral carinae. Abdomen without any
dorsal carinae or spines. Telson deeply sulcate dorsally and broadly truncate
at the tip. Eyes very small and set on outside of stalks.

Description : Integument extremely thin and fragile. Carapace markedly
inflated dorsally and anteriorly to form a very broad, inflated rostrum. The
dorsal margin is carinate for its entire length and minutely denticulate on
the anterior third of its length. In addition to the dorsal carinae there is a
lateral carina originating behind the eye, which passes backward and slightly
downward for about three-fifths of its length and then turns upward and
terminates just before reaching the hind margin of the carapace. A second
lateral carina starts at the branchiostegal spine, runs backward nearly
parallel with the first for about half the length of the carapace, then turns
upward and joins the first carina at the point where the latter bends upward.
Orbit not defined, the raised line bordering the lower lateral margin of the
rostrum forming a smooth curve and teminating in the prominent branchi-
ostegal spine. Abdomen without any dorsal carina or spine on any soinite,
the only ornamentation being a raised ridge at the junction of the terga and
pleura on the fourth and fifth somites; this ridge also extends onto the
anterior part of the sixth somite. The sixth somite is about three and one-
half times as long as the fifth. Telson longer than the sixth somite, deeply
sulcate dorsally and broadly truncate at the tip where it is armed with a row
of ten small spines, the outermost pair being about twice as long as the eight
central ones. In the holotype, the tip of the telson is slightly concave rather

198 Zoologica: New York Zoological Society [XXV:11

Text-figure 63.

Physetocaris microphthalma. A. Telson and uropods. B. Tip
of telson. C. First pereiopod. D. Second pereiopod. E. Chela of sec-
ond pereiopod. F. Fifth pereiopod. G. Mandible. H. First maxilla.

I. Second maxilla. J. First maxilliped. K. Second maxilliped. L.

Third maxilliped.

than nearly straight as in the paratype (Text-fig. 63 B). Eyes small and
nearly hidden beneath the rostrum; the minute cornea is set at the antero-
external angle of the stalk so that the latter extends beyond the cornea.
Stylocerite is a broad, fleshy lobe of the first segment of the antennular
peduncle and its outer spine reaches about to the distal third of that segment.
Antennal scales damaged in both specimens; there is a small spine on the
peduncle at the base of the scale. First pereiopod with a sickle-shaped dactyl.
Second pereiopod with the carpus divided into four joints, the first of which
is nearly as long as the combined length of the other three; chela flattened
and provided with a curious, broad dactyl (Text-fig. 63 E). Third and fourth
pereiopods missing. Fifth pereiopod reduced and simple (Text-fig. 63 F).
The mouth parts are shown in Text-fig. 63 G-L. The mandible lacks both
an incisor process and a palp and the remaining mouth parts are corre-
spondingly reduced. The first three segments of the second maxilliped are
more or less fused and the terminal segment bears three prominent spines
and is applied normally to the end of the preceding joint.

Measurements: Carapace of holotype, measured from the base of the
eyestalk, measures 9.2 mm. The paratype is somewhat smaller, having a
carapace length of only 6.2 mm.

Color in Life : Carapace translucent white over a deep red organ (pos-
sibly ovaries). Abdomen scarlet with red at the base of the pleopods.

Remarks: In the absence of adult males or ovigerous females it is, of
course, possible that this prawn may be a larval form. The fully developed
appendages and lack of exopods do not bear out this possibility, however.
Whether it is a larval stage or not, its systematic position is extremely ob-
scure. It has been impossible to accommodate it in any known caridean
family and even its relative position among the established families is un-

1940]

Chace: Bathypelagic Caridean Crustacea

199

certain. There is little doubt that it is one of the most specialized bathy-
pelagic carideans known, and it apparently shows affinities with the Process-
idae and the Cragonidae as evidenced by the reduced gill structure, the form
of the mandibles and other mouth parts, the simple first pereiopods (as in
one of the first legs in Processa) and the multiarticulate carpus of the second
pereiopods. The chela of the second pereiopod is unlike that found in any
other caridean except possibly that on the first pereiopod in the genus
Discias; obviously, however, its relation to Discias is remote.

General Discussion and Summary.

Taxonomy. The name of the family to which the largest number of
bathypelagic carideans belong has been the cause of considerable confusion ;
at various times it has been known as the Oplophoridae, Ephyridae, Miersii-
dae and Acanthephyridae. Although the latter name, which I have employed
previously, seems the more suitable, it now appears advisable to use the
name Oplophoridae since Kingsley’s use of the subfamily name Oplophorinae ,
in 1878 and 1879, antedates all others and logically proposes Oplophorus as
the type genus, even though Kingsley originally erected the subfamily to
include Oplophorus elongata (= Xiphocaris elongata, an atyid). In 1879
(p. 416) he proposed Ephyrinae as a subfamily of the Atyidae, whereas his
Oplophorinae (p. 426) was included in the Palaemonidae. Ephyra, the type
genus of the Ephyrinae, was proposed by Roux, 1831, for the type species,
Alpheus pelagicus Risso, 1816, and later replaced with Miersia by Kingsley,
1879, since Roux’s name was preoccupied. Miersia pelagica is apparently
unknown to modern carcinologists, but it seems not unlikely that Risso’s
species may be the same as the form now known as Acanthephyra pulchra
A. Milne Edwards, 1890. Risso’s crude figure is obviously incorrect as re-
gards the form of the rostrum, and there is nothing in the description which
would not apply equally well to A. pulchra except that all of the abdominal
somites are said to be sharply carinate and no mention is made of the pres-
ence of exopods on any of the pereiopods. Even if these discrepancies are
overlooked, however, Risso’s description can never be used with any cer-
tainty and, since the type is probably not extant, the best procedure seems
to be to discard Ephyra and Miersia entirely and use the name Oplophoridae
for the family as it is known today.

There has also been some difference of opinion as to the proper spelling
of the name of the type genus. The correct transliteration is, of course,
Hoplophorus, but even though the aspirate is seldom if ever used in modern
French and so might well have been overlooked by Milne Edwards, there is
no choice under the present rules of nomenclature but to preserve the origi-
nal orthography, Oplophorus, as proposed by the original author.

It is likely that future investigations will reveal that the systematics of
the Oplophoridae as known at present are very inadequate. Systellaspis and
Oplophorus are more or less well defined genera and there is little likeli-
hood of confusing the species of those genera with any other. Likewise, the
more typical members of Acanthephyra cause little trouble, but the line of
division between Acanthephyra and the three other genera, Hymenodora,
N otostomus and Ephyrina, is not sharply defined. Although there is little
chance of confusing the species of Hymenodora with those of Acanthephyra,
it is difficult to find any character of systematic importance for distinguish-
ing the two groups. The reverse is true of the division between Acanthe-
phyra and Notostomus; it is almost impossible to assign species like
Acanthephyra gracilipes and N otostomus compsus to the proper genera at a
glance, but the form of the mandible reveals a good diagnostic character.
The genus Notostomus, as here defined, is a very heterogeneous group. It
may be necessary at some future date to split off the species of the N.

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[XXV :11

vescus group under the inappropriate name, Meningodora. Possibly this
group, even as known at present, deserved at least subgeneric recognition,
but I have not made this distinction because of the way in which the gap
between the two groups is filled by such species as N. mollis and N. distirus.
Ephyrina is another easily recognizable genus which exhibits no character
of great systematic importance to separate it from N otostomus ; the form
of the mandibles and the presence of a lateral ridge on the carapace ally
it so closely to N otostomus that the broadened form of the pereiopods is the
only distinctive character separating it from that genus. Notostomus, then,
is the genus which causes the greatest trouble; it is made up of about fifteen
species at present and more are being found by nearly every expedition.
Until our knowledge of this genus is more nearly complete, it is probably
advisable to follow the systematic scheme already established for the family.

Since so many of the bathypelagic Caridea belong to relatively primitive
groups, it is surprising to find such a highly evolved new form as Physe-
tocaris in such a habitat. Although this animal appears to be one of the
most specialized carideans known, as evidenced by the absence of exopods,
the reduced gill series, the simplified mouth parts and the jointed carpus
and complex chela of the second pereiopods, it has apparently been able
to assume a pelagic mode of life by adding to its buoyancy by means of a
membranous integument and an inflated carapace.

Summary of Quantitative Results : The following table lists the species
taken during the expeditions, arranged according to the number of adult
specimens procured.

Species

Acanthephyra purpurea

Systellaspis debilis

Parapandalus richardi

Hymenodora gracilis

N otostomus miccylus

N otostomus vescus

Acanthephyra haeckelii . . . ,
Parapasiphae sidcatifrons . .
Acanthephyra stylorostrata
Oplophorus grimaldii

Notostomus mollis

N otostomus robustus

Leptochela bermudensis
Notostomus marptocheles . ,

Ephyrina bifida

Notostomus compsus

Parapasiphae macrodactyla

Systellaspis braueri

Pasiphaea hoplocerca

Notostomus distirus

Lucaya bigelowi

Physetocaris microphthalma
Oplophorus spinicauda . . . . ,
Acanthephyra brevirostris .

Pasiphaea liocerca

Acanthephyra gracilipes . . ,

Acanthephyra, sp. ?

Acanthephyra eximia

Parapasiphae, sp. b

Acanthephyra acutifrons . .
Notostomus westergreni ? . . .

Parapaiiphae, sp. a

Acanthephyra curtirostris . .

Notostomus perlatus ?

Ephyrina hoskynii

Plesionika martia

Adults

Young

1705

2867

1037

912

361

43

279

1550

120

25

78

74

71

146

36

461

32

54

27

21

12

44

9

11

8

8

6

9

5

65

3

—

2

28

2

11

2

5

2

—

2

—

2

—

1

8

1

5

1

—

1

—

—

8

—

3

—

2

—

2

—

2

—

1

—

1

—

1

1

—

1

1940]

Chace: Bathypelagic Caridean Crustacea

201

Acanthephyra purpurea and Systellaspis debilis are by far the com-
monest species in this area but even these species, at the levels at which
each was most abundant, were captured at the average rate of only one
adult for every 1.7 and 2.1 hours of towing respectively. In the case of
the tenth species in the list, Oplophorus grimaldii, 39 hours of towing were
required for every adult specimen taken at 300 fathoms, the optimum level
for that species in the daytime. A one meter net is far from being perfectly
efficient in straining a one meter column of water, but if the assumption of
100 per cent, efficiency is allowed and if the nets are drawn through the
water at the rate of one knot, which is probably conservative, these results
would indicate an average concentration of the commonest caridean, Acan-
thephyra purpurea, of one adult in every 88,269 cubic feet of water at 700
fathoms which is the optimum level for that species. This gives some
indication of the rarity of many of the species toward the end of the above
list. Several are probably only accidental strays into this region from some
other oceanic area, but the total results of all past expeditions seem to indi-
cate that certain bathypelagic species are everywhere very rare. It is un-
wise, however, to make such general statements when one realizes how little
of the volume of the sea has been investigated. Many species, now con-
sidered rare, may prove to be relatively common when their center of con-
centration is discovered. This point is clearly emphasized by the fact that
the fifth species in order of abundance in the Bermuda area, N otostomus
miccylus, was previously unknown.

Text-fig. 64 may be useful in picturing the relative abundance of adults
of the ten commonest species in the Bermuda area at each level from 400 to
1,000 fathoms. The fact cannot be stressed too strongly that this graph
is applicable only to the particular region in which this work was done, or
at most to the Sargasso Sea. During the course of several cruises of
Atlantis of the Woods Hole Oceanographic Institution to the Sargasso Sea
and to waters off the continental shelf to the west and north of the Gulf
Stream, species common to both areas were found in much greater depths
in the Sargasso area than on the coastal side of the Stream.

One of the interesting but unexplainable points brought to light by the
collection is the change in the ratio between the sexes of certain species
with an increase in depth. In both Acanthephyra purpurea and Hymenodora
gracilis, males were proportionately more numerous near the lower limits of
their range than they were nearer the surface. On the other hand, in two
other species which were represented by an equally large number of speci-
mens, Systellaspis debilis and Parapandalus richardi, no such change in the
sex ratio was found. Whether the females of the first two species are for
some reason more buoyant than the males or whether the phenomenon has
some more involved explanation can hardly be decided at present. The
logical conclusion that females with eggs are found nearer the surface is
not confirmed by this material.

As practically all of the tows were made in the daytime, the collection
fails to increase our knowledge of the diurnal vertical migration of these
forms. Only in the case of Leptochela bermudensis was there evidence of
diurnal migration. Although the number of specimens involved was small,
this species was found at the surface in the evening but only at 400 and
700 fathoms during the daytime.

No definite evidence of any seasonal distribution could be discovered,
with the exception of Parapandalus richardi (see remarks on seasonal dis-
tribution under that species). In all other cases where significant numbers
of specimens were available, no seasonal fluctuation of any sort could be
detected over the period in which towing was done. Ovigerous females of
most species were taken in every month from April to September.

Geographical Distribution : As noted above, that portion of the sea
through which nets have been drawn is extremely small and our knowledge

202

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[XXV : 11

Text-figure 64.

The vertical distribution of adults of the ten commonest species
of bathypelagic Caridea in the Bermuda area during the daytime.
1. Acanthephyra purpurea. 2. Systellaspis debilis. 3. Parapandalus
richardi. 4. Hymenodora gracilis. 5. Notostomus miccylus. 6. Notos-
tomus vescus. 7. Acanthephyra haeckelii. 8. Parapasiphae sulcati-
frons. 9. Acanthephyra stylorostrata. 10. Oplophorus grimaldii.

1940]

Chace: Bathypelagic Caridean Crustacea

203

of oceanic barriers to the dispersal of bathypelagic organisms is correspond-
ingly inadequate. From such distributional data as are available at present,
it is obvious that the temperature and chemical composition of the sea have
a much greater effect on the dispersal of such species than have any land
barriers. Probably, then, oceanic currents will be found to delimit faunistic
zones in the ocean depths much as do mountain ranges, deserts and bodies
of water on land. In most cases, it is certainly misleading to define the
limits of pelagic organisms on the basis of parallels of latitude and longi-
tude. In reviewing the known distributions of the species of Caridea found
off Bermuda, I have been forced to employ the more or less arbitrary
regions listed below. These areas are actually regions in which more or
less extensive investigations of the bathypelagic fauna have been carried
on and if any of them also represent definite oceanic faunistic regions it is
largely accidental. The following is a list of the zones from which species
taken by the Bermuda Expeditions have been recorded previously, with the
figures in parenthesis representing the numbers of species common to
both those areas and to Bermuda.

Sargasso Sea (4) I

North Atlantic (north and west of the Gulf Stream) (6)

Off the coasts of Ireland (5)

Eastern North Atlantic (Bay of Biscay to Cape Verde Ids.) (11)
Mediterranean Sea (2)

Bahamas and the West Indies (9)

Equatorial Atlantic (11)

South Atlantic (2)

Off the Cape of Good Hope (6)

Indian Ocean (15)

Malay Archipelago and Philippine Ids. to Japan (9)

South Pacific (4)

Hawaiian Islands (5)

Eastern Pacific (California to Peru) (6)

In many instances this list is no more than a key to the amount of
exploration carried on in each of these zones; there is little doubt that addi-
tional towing in the West Indian region and in the equatorial Atlantic will
reveal more forms found also at Bermuda. The small number of species
previously known from the Sargasso Sea is due entirely to the lack of reports
on that area, but there is an indication that the Sargasso fauna extends well
to the east in the region of the Bay of Biscay and off the coast of Africa;
several forms found both in that area and off Bermuda have not been
recorded from north and west of the Gulf Stream or off the coasts of
Ireland. The one striking fact illustrated by the list is the unimportance
of distance or land barriers when dealing with the distribution of pelagic
animals; even though the number of species may have been proportionately
increased by the amount of deep-sea work carried on there, the fact that
no less than fifteen of the twenty-five species from Bermuda, which had been
described heretofore, are also found in the Indian Ocean should be of par-
ticular interest to the student of zoogeography.

Color of Bathypelagic Caridea'. So much has been written on the color
of deep-sea Crustacea that there is little necessity for reviewing the subject
here. The excellent color notes on the Bermuda material supplied by Miss
Crane reaffirms the well established fact that most bathypelagic prawns are
of some shade of red in life. All of the species of Acanthephyra in which
the color was noted, as well as Pasiphaea liocerca, Parapasiphae sulcati-
frons, Notostomus robustus, Hymenodora gracilis and Systellaspis debilis,
have the body entirely scarlet or scarlet-red. Certain of the species of
Notostomus of the N. vescus group have the abdomen scarlet but the cara-
pace much deeper in color, at times nearly black. Lucaya bigelowi is unique
in being pale salmon in life. There is another group of species, many of
which are found relatively near the surface, in which the scarlet coloring

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is reduced to patches or even to isolated chromatophores. This tendency
is noted in Notostomus miccylus and Parapandalus richardi which have the
carapace evenly scarlet but the abdomen, especially in immature specimens
which frequent shallower depths, heavily splotched with scarlet. The same
condition is noted in young specimens of Systellaspis debilis, and even in
mature specimens of this species which have not attained their maximum
size the uropods are often transparent. In Leptochela bermudensis , Oplo-
phorus spinicauda, Oplophorus grimaldii and Plesionika martia, all of which
may be found relatively near the surface, the entire animal is more or less
transparent with scattered scarlet markings or dots. Specimens of Lepto-
chela bermudensis which were kept alive for some time after their capture
showed a slight deepening of color after several hours; it is not incon-
ceivable that some species undergo a certain amount of color change depend-
ing upon the amount of light to which they are exposed.

Luminescence: Four of the bathypelagic Caridea of the Bermuda area
are shown to be luminescent from notes made on living material; these
species are Leptochela bermudensis , Systellaspis debilis, Oplophoims spini-
cauda and 0. grimaldi. All of these forms apparently possess photophores,
or dermal light organs of a definite structure, although this conclusion could
not be substantiated in Leptochela bermudensis. Only from this latter
species, however, could luminescence of the photophore type be produced.
To my knowledge, the photophores of captured specimens of Systellaspis
debilis have never been seen to emit light, but that such is their function can
hardly be doubted, and Beebe (1934, p. 304) records the following observa-
tion made from the bathysphere, “Again, one good-sized shrimp at 1300
feet had six or eight scattered lights, and one long, slit-like light near the
center of its body, which identified it with almost complete certainty as
Systellaspis debilis.” Although the presence of a transverse, slit-like organ
does not necessarily prove that the species observed was S. debilis since
several oplophorids have similar photophores behind the fifth pereiopods
and those in Oplophorus grimaldii are very like the comparable organs in
S. debilis, these observations lend further assurance that these types of
photophores are functional light organs. During one of the cruises of
Atlantis to the Sargasso Sea, S. debilis was obtained in considerable num-
bers and it was possible to keep them alive for at least a week by refrigera-
tion, but all attempts to shock them into producing light were futile.
Similar experiments tried on living material of Oplophorus spinicauda and
Oplophorus grimaldii from the present collection were likewise negative.
One of the theories advanced to explain the function of these light organs
suggests that they are used as specific recognition signs to enable indivi-
duals to find specimens of the opposite sex of the same species. Is it not
possible, if such is the explanation, that the ability to produce light may be
controlled by the ripening of the eggs and sperm? This would greatly
reduce the likelihood of seeing the photophores light up in any given speci-
men. Since these animals can be obtained in considerable numbers in the
Bermuda area and since they can be kept alive and apparently healthy
for some length of time by regulating the temperature of the water, it is to
be hoped that this theory may be checked in the near future.

Another type of luminescence noted in bathypelagic Caridea is the pro-
duction of the so-called “luminous cloud.” This appears to be a purely
defensive mechanism whereby a prawn, when startled, can emit a cloud of
luminous material from some internal gland. This spectacle was seen in
Systellaspis debilis and Oplophorus spinicauda (see notes under that species)
among the Bermuda material. Dr. Beebe assures me that this phenomenon
was repeatedly observed in specimens of the former species and has very
kindly provided me with a photograph of a specimen which had been seen
to emit such a cloud; there is no doubt that it belongs to this species. Miss
Crane has also observed that on many occasions when specimens of this
species were placed in alcohol, a mass of flocculent material was immediately

1940]

Chace: Bathypelagic Caridean Crustacea

205

ejected. On several occasions this was tested in the dark-room and found
to be fluorescent. Curiously enough, Miss Crane has also noted the presence
of this flocculent material when specimens of Acanthephyra wei'e put in
preservative.

Dr. Beebe (1934, p. 304) records as follows the results of his observa-
tions on this subject made during descents of the bathysphere: “Whereas
the photophore-like organs of euphausids may serve chiefly as recognition
marks, or other non-defensive capacities, the luminous matter [of large,
Acanthephyra- like shrimps] was obviously discharged only when a shrimp
was startled, as when it bumped against the bathysphere window. When
this happened, a rocket-like burst of fluid was emitted with such violence
that the psychological effect was that of a sudden explosion. This occurred
time and again at the lower levels, and I learned to distinguish two separate
types of discharge, one uniformly luminous, the other dimmer but inter-
spersed with dozens of brilliant stars and pinheads. For an instant the
shrimp would be outlined in its own light-vivid scarlet body, black eyes,
long rostrum — and then would vanish, leaving behind it the confusing glow
of fluid. The light died out gradually, but the discharge disappeared even
more slowly. It was not until one of my last dives that I learned that certain
grayish bodies which I had been reporting as unlighted fishes were in
reality these burnt-out masses of fluid.”

Dr. Harvey (1931) has discussed the chemistry of luminescence in
Systellaspis.

Among the Oplophoridae, luminescence apparently is found in but two
genera, Systellaspis and Oplophorus, where both photophores and luminous
glands may be found in the same species, but the possibility can not be
entirely ruled out that luminous glands may be found in other genera as
well. More observations made directly from living material should.be en-
couraged to increase our knowledge of the number of species which are
luminescent. It is also helpful to preserve some of each species in formalin,
for photophores which disappear in a few days in alcohol, will be distinct
several years later in material put up in formalin. This recommendation
is made with some reservation, however, for alcohol is much to be pre-
ferred to formalin as a general preservative.

Food of Bathypelagic Caridea: The following remarks on the food of
Acanthephyra purpurea and Systellaspis debilis are based on such incomplete
data that they are included here merely in the hope that other workers may
be induced to investigate the problem more thoroughly. Miss Jocelyn Crane
dissected out the stomachs of a dozen specimens of Acanthephyra purpurea
and found them usually crammed with well digested food containing a few
identifiable animals. Very much the same type of stomach contents were
found in a few specimens of Systellaspis debilis which I dissected. In all
but one or two cases, surprisingly large fragments of at least one blackish
fish were found. Also present were various crustaceans ranging from
copepods to comparatively large shrimp-like forms several species of ptero-
pods; worms, some of which may have been parasitic; and radiolarians. The
most striking point about these stomach contents was the size of the frag-
ments. In some cases one fish apparently took up the entire space within the
stomach and it was often so well preserved as to permit its generic deter-
mination. Several of the crustaceans could likewise be determined from the
available pieces. Apparently, then, the mandibles do not necessarily crush
the food into an amorphous mass before it is passed into the stomach.

One can only guess whether these prawns are able to capture their prey
alive or whether they are purely scavengers which feed upon dead or dying
organisms. It is very possible that much of the material found in the
stomachs of these specimens was consumed after the specimens were in the
net. When the contents of a net have been dumped into a tank of water
immediately after being taken aboard ship, I have observed specimens of

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Systellaspis debilis feeding with apparent enthusiasm upon dead fish as well
as dead specimens of their own species. The fact that the stomachs of these
prawns were usually distended with food when examined may be entirely due
to the plentiful meal provided by the contents of the net and one cannot be
certain that this food is representative of their normal diet.

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Carlgren: Actiniaria from the Gulf of California

211

12.

Eastern Pacific Expeditions of the New York
Zoological Society. XIX.

Actiniaria from the Gulf of California.1

Oskar Carlgren

University of Lund, Sweden.

(Text-figures 1-8).

[This is the nineteenth of a series of papers dealing with the collections
of the Eastern Pacific Expeditions of the New York Zoological Society made
under the direction of William Beebe. The present paper is concerned with
specimens taken on the Templeton Crocker Expedition (1936). For data on
localities, dates, dredges, etc., refer to Zoologica, Vol. XXII, No. 2, pp.
33-46.]

Contents.

Page

Introduction 211

Family Aliciidae

Alicia beebei sp. nov 212

Family Nemanthidae fam. nov 212

Nemanthus gen. nov 212

Nemanthus nitidus (Wassilieff) 214

Nemanthus calif ornicus sp. nov 215

Family Sagartiidae

Actinothoe calif ornica sp. nov 217

Species not determined 219

Introduction.

The Actiniaria from the Templeton Crocker Expedition to Lower Cali-
fornian waters contained four species, of which three apparently are new
to science. Of especial interest is Nemanthus calif ornicus, nearly related to
Nemanthus ( Sagartia ) nitidus (Wassil.) from the waters of Japan, both
belonging to a new genus and a new family characterized by having curious,
acontia-like formations without any nematocysts at the termination of the
filaments.

Family Aliciidae.

Alicia beebei, sp. nov.

Diagnosis : Pedal disc, wide, thin, with fairly distinct insertions of the
mesenteries. Column in its uppermost part smooth, forming a capitulum,

1 Contribution No. 598, Department of Tropical Research, New York Zoological Society.

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[XXV :12

lower parts provided with sessile vesicles crowded at the base, upwards
more scattered ; at the border to the capitulum the vesicles are pedunculate,
each peduncle bearing 2-7 vesicles. Tentacles long and slender, about 96.
Mesenteries 96 pairs of perfect mesenteries. Nematocysts of the lower part
of the column 12-14 x 2 g; those of the vesicles partly 59-72 x about 12 g,
common, macrobasic amastigophors, partly 60-77 x 6-8.5 g, very numerous,
microbasic amastigophors; those of the capitulum 12-17 x (almost) 2-2 g,
common ; those of the tentacles partly 43-51 x 5-5, 5 g common, microbasic
amastigophors, partly 13-18 x (about) 2 g, common; those of the actino-
pharynx (26)36-41(46) x 4, 5-5 g, common, microbasic amastigophors, those
of the filaments partly 24-29 x over 4 g, microbasic amastigophors, partly
7.5-10 x about 2, 5 g, microbasic p - (penicilli-like) mastigophors ; the small
nematocysts of the column and tentacles probably basitrichs. Spirocysts of
the capitulum up to 36 x 6 g] those of the tentacles 17 x 2, 5- (about) 41
x 6 g.

Color in Alcohol : Basal parts of the tentacles yellowish-brown.

Size: Height 2.5 cm., breadth of pedal disc 2 cm., of the oral disc 1.5
cm. ; length of the tentacles 3.5 cm., or longer.

Locality: St. 136 D-30, Arena Bank, 23° 27' N. Lat., 109° 24' W. Long.,
35 fms., May 1, 1936, 1 specimen.

Family Nemanthidae fam. nov.

Acontiaria with mesogloeal sphincter. No acontia proper but acontia-
like organs situated at the termination of the filaments and not forming any
batteries of nematocysts.

I refer the family for the present to Acontiaria, because it seems to
me that it is more closely related to this subtribe than to the Mesomyaria.
The diagnosis of the subtribe Acontiaria may be altered in such a way that
it encloses also the family proposed above.

Nemanthus gen. nov.

Nemanthidae with wide pedal disc, smooth column and long tentacles,
considerably fewer than the mesenteries at the base. Sphincter mesogloeal,
well developed. Longitudinal muscles of tentacles and radial muscles of oral
disc ectodermal. Actinopharynx with distinct, fairly broad siphonoglyphs,
as a rule 2, but possibly varying in connection with asexual reproduction.
As a rule 2 pairs of directives; perfect mesenteries at least 6, up to 12
pairs, frequently irregularly arranged, so that one side has more mesen-
teries than the other, sterile ; stronger imperfect mesenteries fertile. Longi-
tudinal muscles of mesenteries not forming distinct pennons; parietobasilar
muscles moderately weak or weak. Acontia-like organs, not forming any
batteries of nematocysts, present at least on some of the mesenteries. Amas-
tigophors absent in the tissues.

The type of the genus proposed above is Sagartia nitida Wassilieff ;
described from Sagami Bay on the coast of Japan. As the anatomical de-
scription given by Wassilieff (1908, p. 31) of the species in several points
is imperfect, it was necessary to re-examine the species. It was so much
more important as, owing to the imperfect preservation of N. calif ornicus
described below, I could not describe with certainty some details of its
anatomy.

Before discussing the species it seems suitable to describe the curious
acontia-like organs so characteristic of the genus. They occurred in all six
specimens examined, (4 of nitidus and 2 of calif ornicus) but always in

1940]

Carlgren: Actiniaria from the Gulf of California

213

small number and only on some of the mesenteries. Like the acontia they
are attached to the mesenteries at the termination of the filaments. They
are either short and thick, or longer and sometimes more slender, but
always many times thicker than the filaments; in the latter case they were
twice found to be squeezed out through the actinopharynx through the
contractions of the animals.

N emanthus nitidus. Lowest part of mesentery with filaments and acon-
tium-like thread.

Their appearance (Text-fig. 1) is in itself somewhat different from that
of an acontium, so it calls for an anatomical examination. I have made
slides of two threads, one from nitidus and one from calif ornicus , and ex-
amined several in maceration preparations. Text-figure 1 shows the lowest
part of a mesentery with filaments and the acontium-like thread of nitidus.
Text-figure 2 a, b, c, shows three transverse sections of the same thread,
the section on Text-fig. 2 a taken at the insertion of the thread on the
mesentary, and Text-fig. 3 a, b, two similar sections of a thread of cali-
f ornicus. Thus the cross-sections are seen to recall somewhat that of an
acontium. There is an axis of mesogloea, as it seems always thick and
surrounded for the most part by a very thick epithelium (ectoderm) and in
the part where the mesogloea runs out in a more or less fin-like outgrowth,
by flattened endoderm cells. The fin varies very much, from small (Text-
fig. 3 a, b) to elongated (Text-fig. 2 c). No muscles are attached to the
fin in contradistinction to the conditions of the acontium. The epithelia are
not provided with any nematocysts. (True, I have found a few nematocysts

Text-figure 2.

Nemanthus nitidus. Transverse sections of acontium-like thread shown
in Text-figure 1.

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Text-figure 3.

Nemanthus calif omicus. Transverse sections of an acontium-like thread.

in some of the maceration preparations, but they belonged probably to the
filaments as they agreed in size as well as in types with those. If they oc-
casionally are really present they are at least extraordinarily rare). The
question is how to interpret these acontia-like organs which Wassilieff
(1908, p. 34) has anticipated to be acontia. They may be degenerated
acontia which have lost their nematocysts, or formations sui generis, or
acontia which have not yet developed their nematocysts. The last supposi-
tion is not likely, as the nematocysts appear in the acontia in an early stage
of their development (Carlgren, 1934). At present the question must be
left unanswered. In certain respects the acontia-like organs recall the acon-
tioids of the Ceriantharia inasmuch as the ectoderm here also consists only
of supporting and gland cells.

Nemanthus nitidus (Wassilieff).

Sagartia nitida sp. nov. Wassilieff, 1908, p. 31. pi. I, figs. 18, 19; pi.
VIII, figs. 84-87; text-figs. 14-17.

Diagnosis: Pedal disc wide. Column smooth with longitudinal furrows
indicating the insertions of the mesenteries. Margin distinct. Sphincter
strong, mesogloeal, alveolar but here and there more reticular, separated
from the muscles of the endoderm and from the ectoderm by a broad
mesogloeal lamella. Tentacles fairly numerous, up to about 130, not half
the number of the mesenteries at the base, conical, the inner ones long, the
outer ones considerably shorter. Longitudinal muscles of the tentacles and
radial muscles of the oral disc weak. Actinopharynx smooth with indis-
tinct longitudinal furrows and 2 (always?) fairly broad siphonoglyphs.
Mesenteries variable in number, up to about 280 at the base, often asymme-
trically situated. Pairs of perfect mesenteries 6-12, sterile, the stronger
imperfect ones fertile. Two pairs (always?) of directives. Longitudinal
muscles of mesenteries weak, forming no distinct retractors, parietobasilar
muscles very weak. Acontia-like organs forming no nematocyst batteries,

1940]

Carlgren: Actiniaria from the Gulf of California

215

present on a few mesenteries. Dioecious. According to Wassilieff reproduc-
tion by any kind of longitudinal fission. Nematocysts of the column 14-17
x 2.5 g, basitrichs; those of the tentacles 22-31 x (about) 2.6-3 g, numerous,
probably basitrichs; those of the actinopharynx partly 15.5-24 x (about)
2.5 g, probably basitrichs, partly 17-18 x 2.6-3 g, microbasic p-(penicilli-
like) mastigophors ; those of the filaments partly about 10 x 2.5 /a, partly
13-18 x (about) 2.7-3 g, microbasic p-mastigophors. Spirocysts of the
tentacles 19 x 2.5 - 41 x 5 /a.

Color: According to Wassilieff and Gislen, rose-colored.

Localities: Japan. Sagami. Koaziro, 6 m. T. Gislen, 1930, 2 specimens;
Bonin Isl. N.W. of Port Lloyd, 70 fms. S. Block, 1914, 1 specimen; 33° 15' N.,
129° 15' E., 45 fms., temp. 73° F. Svenson, 1890, 1 specimen; 33° 5' N.,
129°16' E., 36 fms., temp. 51°F. Svenson, 1890, 3 specimens.

As to the perfect mesenteries, I have examinated three specimens, one
of which was fertile and had well developed testes in the stronger, imperfect
mesenteries. The fertile individual and the single specimen from Bonin
Island had 12 pairs of perfect mesenteries, two of which were directives,
symmetrically arranged. Both showed the same arrangement of the per-
fect pairs, 4 pairs on the one side of the directive plane, 6 on the other; the
third specimen had only 6 pairs of perfect mesenteries, symmetrically sit-
uated. The last specimen had 90 tentacles but about 275 mesenteries at the
base, another specimen 136 tentacles and about 280 mesenteries at the pedal
disc.

As Wassilieff’s figure of the sphincter is very schematic, I give here a
more detailed figure of it (Text-fig. 4).

Nemanthus ealifornieus sp. nov.

Diagnosis: Pedal disc wide. Column smooth with more or less distinct,
longitudinal furrows. Sphincter strong, in its uppermost part occupying al-
most the whole breadth of the mesogloea and here reticular, with tendency to
transverse stratification on its external side, gradually diminishing in
breadth downwards, in its lower part alveolar. Tentacles in number up to
about 124, long, conical, the inner ones about twice as long as the outer
ones. Longitudinal muscles of tentacles ectodermal, weak; radial muscles
of oral disc ectodermal, weak at the insertions of the mesenteries, thickened
between them, the folds delicate but not much branched. Oral disc with
deep radial furrows at the insertions of mesenteries. Actinopharynx with 2
rather deep siphonoglyphs. Mesenteries at the base up to about twice as
numerous as the tentacles. Pairs of perfect mesenteries 8 or more. Two
pairs (always?) of directives. Longitudinal muscles of mesenteries form-
ing low folds and uniformly distributed over almost the whole surface of the
mesenteries. Parietobasilar muscles not situated on an outgrowth of the
mesogloea, reaching upward for some distance, their folds sometimes tall
but delicate. Acontia-like organs and probably also reproduction as in N.
nitidus. Dioecious. Nematocysts of the column 14-19 x 2.5 g, basitrichs;
those of the. tentacles 23-29 x 2.5 (about) 3 /a, rod-like, probably basitrichs;
those of the actinopharynx partly 17-24 x 2-2.5 g rod-like, fairly common,
partly 18 x 3 g microbasic p-mastigophors, rare; those of the filaments
partly 14-18 x 2.5-3. 5 g microbasic p-mastigophores, partly 8.5-12 x 1-1.5 g
probably basitrichs. Spirocysts of the tentacles 22 x (about) 2.5-43 x 5 g.

Color in Alcohol: Column with numerous, close, greenish-gray spots
distributed over the whole surface.

Size: Largest specimen, average height about 1 cm., breath about 1.5
cm., (pedal disc broader) ; length of the inner tentacles up to 1.5 cm., of
the outer ones about 0.7 cm.

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Zoologica: New York Zoological Society

Text-figure 4.

Nemanthus nitidus. Section of
sphincter.

Text-figure 5.

Nemanthus calif omicus. Sec-
tion of sphincter.

Locality. St. 136 D-l, Arena Bank, 23° 29' N. Lat., 109° 25' W. Long.,
45 fms., April 3, 1936, several specimens. (No. 36171). Some individuals
were attached to the stalk of an octocoral.

1940]

Carlgren: Actiniaria from the Gulf of California

217

Text-figure 5 shows a section of the sphincter. Two individuals ex-
amined had 8 pairs of perfect mesenteries, a third sectioned had at least 8,
probably a few more. It was, however, wrinkled and not so well preserved
that I was able to count their exact number. The largest example had about
124 tentacles and 196 mesenteries at the base, another had 112 tentacles
and about twice as many mesenteries at the base. The perfect mesenteries
were apparently sterile and the development of the generative organs is
probably limited to the stronger imperfect mesenteries. A sectioned speci-
men was a female with numerous, small ova. As with N. nitidus the indi-
viduals of the present species live together in colonies, sometimes so closely
packed that the borders of the pedal disc touch each other intimately.
Wassilieff has described longitudinal fission in nitidus — perhaps it would
be more adequate to speak of budding. Probably asexual reproduction
occurs also in calif ornicus, since I have found a small bud at the base of
one specimen.

Remarks : The species is closely related to nitidus but the color is
different, the muscles of the oral disc and the mesenteries are stronger, the
sphincter stronger and more reticular in calif ornicus, etc.

Family Sagartiidae.

Aetinothoe californica sp. nov.

Diagnosis : Pedal disc broad, column smooth, cinclides?, margin dis-
tinct. Sphincter strong, alveolar, with tendency to be reticulated, occupy-
ing in its upper part the whole breadth of the mesogloea, diminishing down-
wards, and separated from the circular muscles of the column by a fairly
thin lamella. Tentacles about 192. Longitudinal muscles of tentacles and
radial muscles of oral disc ectodermal. Two distinct siphonoglyphs. Prob-
ably not more mesenteries than tentacles. Two pairs of directives. At least
12 pairs of perfect mesenteries. Retractors of the older mesenteries strong,
fairly concentrated with tall folds. Parietobasilar muscles apparently weak.
Nematocysts of the column partly 17-34 x 3. 5-5.5 g, partly 12-17 x 1.5-2 g,
basitrichs; those of the tentacles partly 21-34 x 2.5-3 g, rod-like, partly
21-22 x (about) 4 g, those of the actinopharynx 24-29 x (about) 2.5 g;
those of the acontia partly 48-53 x (about) 6 g ; microbasic amastigophors,
partly 19-28 x 2-2.5 g, basitrichs; spirocysts of the tentacles 15-29 x 2. 5-4.5
g.

Color in Alcohol : Column with irregular red-brown spots arranged at
least in its proximal part in distinct longitudinal rows about as in A. un-
data var. anguicoma and lacerata. (see Stephenson, 1935).

Size: Largest, strongly contracted example; height 1.5 cm., pedal disc
2.5 x 2 cm.

Locality : Sta. 142 D-3, Santa Inez Bay, 27° 04' N. Lat., 111° 54' W.
Long., 40 fms., April 11, 1936. (No. 36,253). 2 specimens on Murex re-
curvirostris, one shell inhabited by a pagurid.

Text-figure 6.

Aetinothoe californica. Lower
part of column, showing arrange-
ment of spots.

Text-figure 6 shows the arrangement of the spots in the lower part of
the column, Text-fig. 7 a section of the upper half of the sphincter and Text-

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[XXV :12

Text-figure 7.
Actinothoe calif or-
nica. Section of upper
half of sphincter.

1940]

Carlgren: Actiniaria from the Gulf of California

219

fig. 8 a retractor of a perfect mesentery. Because the pedal disc was ex-
tended over the spines of the shells and the body so strongly contracted
that the summits of the contracted tentacles almost reached the pedal disc,
and the preservation of the individual furthermore was not too perfect, the
examination of the specimen was difficult and some observations uncertain.
The examined example was a male, but the distribution of the gonads was
impossible to decide. As to the broader nematocysts of the column, the
shorter ones (about 17-18 /a) were certainly microbasic amastigophors —
the capsules were often exploded and the “hampe” somewhat longer than
the capsule; the longer ones were probably similar. (They were usually ex-
ploded but with the “hampe” torn to pieces, but as I have seen the apex
of the “hampe” and it was not provided with a thread, I think that they all
were amastigophors though somewhat different in their structures).

The rod-like nematocysts of the tentacles were not examined, the broader
being exploded but with the “hampe” damaged, wherefore I cannot de-
cide their category. Probably they were microbasic amastigophors.

Besides the species described above, a fourth species has been taken
by the expedition on Gorda Bank, 23° 06' N. Lat. and 109° 25' W. Long.,
at a depth of 50 fathoms. Two small individuals, both attached to the stem
of an octocoral, were present in the collection. One specimen had 72 tentacles
and about 136 mesenteries at the base disc, the other about 80 tentacles
and about 170 mesenteries at the base. In one individual 10 mesenteries
were perfect. The sphincter was strong and mesogloeal-alveolar ; the longi-
tudinal muscles of the tentacles were ectodermal. I have made maceration
preparations of the tissues in order to examine the nematocysts, but, in
spite of careful examination I have not found any nematocysts which could
be supposed to belong to acontia. The general appearance of the animals
and the characters enumerated here, apart from the absence (?) of acontia,
recall those of the genus Amphianthus, but as the individuals also were
sterile, I have preferred not to name the species.

LITERATURE CITED.

Carlgren, O.

1934. Ceriantharia, Actiniaria and Zoantharia. Rept. Scient. Results
“ Michael Sars” North Atlantic Exped., 1910. Vol. V. Bergen.

Stephenson, T. A.

1935. The British Sea Anemones. Vol. II, Roy. Soc., 121.

Wassilieff, A.

1908. Japanische Actinien. Abhandl. mat-phys. Klasse. R. Bayer. Akad. d.
Wissensch., 1908, 1 Suppl. Band, 2te Abh. Munchen.

Merriman: Studies on Marine Catfish

221

13.

Morphological and Embryological Studies on Two Species of
Marine Catfish, Bagre marinus and Galeichthys felis.

Daniel Merriman

Osborn Zoological Laboratory, Yale University.

(Plates I-V; Text-figures 1-9).

Introduction.

The material presented in this paper is based on a large number of
preserved eggs, young, and adults, of the gaff-topsail catfish, Bagre marinus,
and the silver or sea catfish, Galeichthys 1 felis. The collection of these speci-
mens was made by Mr. M. B. Bishop of the Peabody Museum, Yale Uni-
versity, during 1938 and 1939, while working on a cooperative study of the
tarpon at the field laboratory of the New York Aquarium at Palmetto Key,
Florida. The entire collection was then put at the author’s disposal through
the generosity of Dr. C. M. Breder, Jr., Acting Director of the New York
Aquarium, and Prof. A. E. Parr, Director of the Peabody Museum.

These two ariids occur commonly along the coasts of the southern
Atlantic and Gulf states, but are comparatively rare north of Chesapeake
Bay, although the northern limit of their range is Cape Cod (Breder, 1929).
Galeichthys felis has been taken as far south as the Yucatan Peninsula
(Hubbs, 1936), and Bagre marinus in Panama (Breder, loc. cit.).

The nomenclature of these two species has been discussed recently in
papers by Hubbs (loc. cit.) and Lee (1937). Former names for Galeichthys
felis (Linnaeus) include Galeichthys milberti, Hexanematichthys felis, Arius
milberti, etc.; among those for Bagre marinus (Mitchell) are Felichthys
felis and marinus, and Ailurichthys (emended to YElurichttiys) felis and
marinus. Hubbs (loc. cit.) has also pointed out that Regan’s (1907)
Galeichthys guentheri is not separable from G. felis.

Both of these species are particularly interesting from a number of
points of view. Thus it is well known that their eggs are among the largest
found in the teleosts, averaging 14-19 mm. in diameter before fertilization
and during the early stages of development. Furthermore, these catfish ex-
hibit the phenomenon of oral gestation (Smith, 1907; Gudger, 1916 and
1918; and Lee, 1931 and 1937), the males picking up the eggs shortly after
fertilization and carrying them in their highly modified mouths through
the entire period of development up to the nearly complete absorption of
the yolk-sac. Another interesting point about these species is the fact that
both of them exhibit sexually dimorphic characters in the pelvic fins; the
extraordinary hook-like protuberance on the adaxial surface of the pelvic
fin in Galeichthys felis females has been described by Lee (loc. cit.) and
Gowanloch (1933). The osteology of these forms is also of considerable

1 The word Galeichthys comes from yaW weasel, and ixOi’s, fish (Jordan & Evermann, 1896).

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[XXV :13

importance because it illustrates the high degree of specialization so char-
acteristic of the Nematognathi. Finally, the inner ears of G. felis and
B. marinus are interesting because they show important structural modifi-
cations not found in other groups of fishes. Although most of these points
have received some attention from a number of authors, many interesting
details have been neglected. This paper takes up various morphological and
embryological aspects of these two species, and treats them under the fol-
lowing headings: External Anatomy, Morphology of the Alimentary Tracts
and the Feeding Habits, Osteology, Anatomy of the Inner Ear, and Em-
bryology.

External Anatomy.

The external characters of Galeichthys felis and Bagre marinus which
may be used in distinguishing these two species have been adequately de-
scribed by many authors, among them Jordan & Evermann (1896), Smith
(1907), and Breder (1929). A character which seems, however, to have
been generally overlooked is the posterior margin of the anal fin, which in
B. marinus has a prominent V-shaped indentation and in G. felis a compara-
tively straight border (Text-fig. 1, C and D). This does not show in the
illustrations in Jordan & Evermann (1896) and Smith (1907), but Meek
& Hildebrand (1923) describe the anal fin in members of the genus
Felichthys as being “more or less emarginate.” Another character which
should be used with care is the description of the maxillary barbels as being
flat and ribbon-like in B. marinus, as opposed to the condition in G. felis
where the maxillary barbels are said to be “circular” in section. While it is
true that these barbels are flatter and more ribbon-like in B. marinus,
those of G. felis are distinctly flat over the greater part of their length, and
only approach an oval condition in section at their extreme bases.

The modifications of the pelvic fins in adult female Galeichthys felis
have been described and pictured in some detail by Lee (1937). The fleshy,

A. The adaxial surface of the pelvic fins of a mature female Galeichthys
felis, to show the fleshy protuberances which develop in the breeding season.
B. The same view of the pelvic fins of a mature male G. felis. C and D. Lateral
views of the posterior parts of the bodies of G. felis and Bagre marinus respec-
tively.

1940]

Merriman: Studies on Marine Catfish

223

hook-like protuberances on the adaxial surface of the pelvic fins of a
female nearing full maturity are shown in Text-fig. 1, A. As noted by
Beeson (Lee, loc. cit.) this striking pelvic modification of the females be-
comes larger and more pronounced as the spawning season approaches, and
then degenerates after spawning to a relatively inconspicuous affair. At the
height of maturity these fleshy protuberances develop to such an extent that
the greater part of each pelvic fin is forced into a more or less vertical
plane. What is normally the adaxial (dorsal) surface is now turned out-
wards and is lateral in position, and the original abaxial (ventral) surface
is turned inwards and is medial in position (Text-fig. 1, C). The pelvic
fins of the females are thus forced into a trough-like structure at the time
the eggs are extruded. There is some question as to the exact significance
of these changes during the breeding season (Breder, 1935, and Harden-
berg, 1935), and as to the uses which this modification might serve. In
Arius maculatus, an East Indian form which also practices oral gestation,
Hardenberg has described a similar development on the female’s pelvic
fins, and says, “This is a sexual character, which has something to do with
spawning and mating. It is clear that the male is attached by these hooks
and the fertilization of the eggs takes place perhaps inside the body of the
female or more probably outside the body just at the moment when they
leave the genital opening.” Lee (loc. cit.) points out that, “In all proba-
bility this modification is widespread through the family, though not of
universal occurrence, for Gudger failed to note the structure in Bagre
marinus and Hubbs found it undeveloped in Arius aqua-dulce (as reported
by Breder).” This author also has found no similar modification in Bagre
marinus in his examination of numerous specimens, thus substantiating
Gudger.

Lee (loc. cit.) says, “No such modification of the pelvic fin has been
found in the male, although in egg-carrying males the fin shows a white,
almost ridge-like area . . . , corresponding in position to the modification in
gravid females. The significance of the ridge, if indeed it has any, is not
understood; it is perhaps only a rudiment of a structure proper to the other
sex.” The figure accompanying this statement shows a vague area on the
pelvic fin of a gestating male.

In the majority of male Galeichthys felis examined in the present work
no modification was apparent, but in more than one-third of these male
specimens there was a definite structure on the adaxial surface of the pelvic
fins. Sometimes this modification was indistinct as indicated in Lee’s figure,
but at other times it was more conspicuous as shown in Text-fig. 1, B,
where it is clearly seen to be a hook-like development not widely different
from the condition found in immature and non-spawning females. Further-
more, this modification was found not to be confined to the gestating males.2

Lee (loc. cit.) has noted the fact that the pelvic fins of mature
female Galeichthys felis are considerably larger, in proportion to the length
of the fish, than in mature males of the same species. The author finds that
this also holds true for the pelvic fins of Bagre marinus. This marked
sexual dimorphism is shown in Text-fig. 2, where the pelvic fins of two
gaff-topsail catfish of the same size are drawn from the abaxial surface.
It is apparent that the fins of the male are not only much smaller than those
of the female, but that they are also quite different in shape. Thus the
greatest length of the male pelvics is on the outside of the fin — near its
lateral margin. Also the posterior edge of the fin in males is relatively
straight and slants obliquely inward (toward the medial margin, which has
the shortest fin rays). By contrast, the female pelvic fins have their greatest
length near the region of the middle fin rays. Furthermore, the posterior
margins of the pelvics in females of this species are rounded in contour —

2 The gonads of some of these male Galeichthys felis were sectioned to avoid any possibility
of an error in sex determination — in other words, of their having been immature females.

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Text-figure 2.

Abaxial views of the pelvic fins of male and female specimens of Bagre
marinus of the same size. Note the typical sexually dimorphic characters.

not straight as in the males. This is obviously because the lateral and medial
fin rays are shorter than those in the middle of the fin, thus making the
posterior margin semi-circular in shape. Table 1 shows measurements on
the pelvic fins of ten males and ten females of about the same size. The
lengths of the fins were taken from the point marked by the junction of
the lateral margin with the base, to the tip of the longest ray. The widths
of the pelvics were measured from a point marked by the junction of the
medial margin with the base, to the lateral margin in a line at right angles
to the plane of the fin. The results of these measurements are shown at the
bottom of Table 1, where it will be seen that the average lengths of the
pelvic fins of males and females of the same standard lengths are 4.4 and
6.4 cm. respectively; the average widths are 1.2 and 1.4 cm. respectively.

Table 1.

Comparative Lengths and Widths of Pelvic Fins of
Male and Female Bagre marinus.

Males

Females

Standard

Standard

length

length

of fish

Length of

Breadth of

of fish

Length of

Breadth of

(cm.)

fin (cm.)

fin (cm.)

(cm.)

fin (cm.)

fin (cm.)

26.0

3.6

1.0

26.5

5.2

1.2

26.5

3.7

1.1

27.0

5.4

1.2

28.5

3.8

1.1

28.5

5.6

1.2

32.0

4.5

1.1

28.5

5.8

1.2

33.0

4.6

1.1

32.0

6.6

1.6

33.0

4.4

1.2

34.5

7.1

1.3

33.5

4.7

1.3

35.0

7.0

1.8

33.5

5.0

1.4

35.5

6.2

1.3

35.0

4.8

1.2

36.0

7.8

1.6

39.0

5.3

1.7

37.0

7.2

1.6

Average 32.00 cm. 4.4 cm.

1.2 cm.

32.05 cm. 6.4 cm.

1.4 cm.

1940] Merriman: Studies on Marine Catfish 225

Morphology of the Alimentary Tracts and the Feeding Habits.

The general anatomy of the coelom and its organs is essentially the
same in Galeichthys felis and Bagre marinus, and only minor differences
exist. One of these differences is in the parietal peritoneum of the two
species. In Galeichthys felis the peritoneal lining appears almost grayish,
due to minute black pigment spots which are abundant over its otherwise
white surface. These pigment spots are more numerous on the dorsal sur-
face of the coelom than they are on the lateral and ventral walls of the
body cavity. In Bagre marinus, however, there are no corresponding pig-
ment spots, and the peritoneum is a clean, silvery white.

Among the more interesting of the differences between these two
species are those found in the morphology of the alimentary tracts. It will
be seen in Text-fig. 3 that the fundamental pattern of the gastro-intestinal
tracts is the same in both species. Thus, in each type the intestine describes
one and a quarter full loops. Yet there are distinct differences within that
basic pattern, and these are constant for the species. Thus the stomach
in Galeichthys felis (Text-fig. 3, A) is typically J-shaped. At its posterior
end it makes a sharp bend anteriorly, decreasing in diameter as it does so,
and ending in a characteristic constriction marking the pylorus. But the
stomach in Bagre marinus (Text-fig. 3, B) turns only at right angles to
the longitudinal axis at its posterior end and terminates abruptly with the
constriction for the pylorus, which is somewhat less distinct in this species.
Leaving the pylorus, in Galeichthys felis (Text-fig. 3, A), the intestine pro-
ceeds anteriorly, slightly lateral to the left side of the stomach. At the
anterior end of the stomach it curves to the right and passes ventral to the
junction of the oesophagus and the stomach. It then continues to curve
sharply and passes posteriorly as a straight tube just lateral to the right
side of the stomach. Only by the time that it reaches the posterior end of

Text-figure 3.

Diagrammatic sketches of the gastro-intestinal tracts of Galeichthys felis
(A) and Bagre marinus IB) — ventral aspect.

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the stomach does it suddenly become much smaller in diameter. Almost imme-
diately it also takes on a highly convoluted form. These convolutions con-
tinue posteriorly nearly to the end of the coelom, where the intestine again
turns anteriorly, simultaneously losing most of its convolutions. The in-
testine then runs to a point close to the pylorus, makes another sharp turn
posteriorly and continues back to the anus without any section of it being
highly convoluted. The intestinal pattern of Bagre marinus (Text-fig. 3, B),
although fundamentally the same, differs strikingly in the point at which
the convolutions first begin. Here they start just after the intestine curves
under the junction of the oesophagus and stomach; in other words, the
intestine does not continue as a straight tube so far posteriorly. Also there
is no sudden decrease in the diameter of the intestine at the point where
the convolutions first occur; instead the diameter decreases gradually. The
intestine maintains its highly convoluted form to the posterior end of the
coelom. It then curves abruptly and runs anteriorly as far as the level of the
posterior end of the stomach, bends sharply on itself again and continues
posteriorly to the anus; this latter part of the intestine has only minor
convolutions.

o

x in
Z
o o
z

ID z

< u.
O t-
H <

< o

I-
i n

LENGTH OF INTESTINE IN CMS.

Text-figure 4.

Graph of the leng-ths of the intestines of Galeichthys felis and Bagre marinus
of different sizes, to show the proportionately longer intestine of B. marinus.

Since the intestine of Bagre marinus is convoluted over a greater part
of its length than that of Galeichthys felis, it is a reasonable expectation
that it would be proportionately longer. That this is true is shown by Text-
fig. 4, where the lengths of the intestines of both species are plotted against
the standard lengths of the fish. It is clearly evident that the length of
the intestine in Bagre marinus is longer in proportion to the size of the fish
than in Galeichthys felis. Thus in two individuals 30 cm. in standard
length, the intestine would average 75 cm. in G. felis and 88 cm. in B.
marinus. This difference is fairly constant, although it is apparent from
the scatter of points on the graph that there is considerable variation and
not infrequently an overlap between the two species. However, using Tip-
pett’s (1931) method for analysis of small samples, y is less than .001 — in
other words, the chances are less than 1 in 1,000 that the difference is not
significant.

-

1 1 1 1 1 T

lengths of intestines of

GALEICHTHYS FELIS (o)

AND BAGRE MARINUS (•)

OF DIFFERENT SIZES

•

✓O

° ' o o °o

stf o

t

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Merriman: Studies on Marine Catfish

227

Stomach content analyses show that there is no essential difference in
the diets of these two species. Both forms are completely omnivorous.
Algae and various kinds of sea grasses are not infrequently eaten by them.
Among the invertebrates, coelenterates (sea anemones), holothurians, gas-
tropods, polychaets, and Crustacea (isopods, and various decapods such as
shrimps, spider crabs — Libinia sp., swimming crabs — Callinectes sapidus
and Ovalipes ocellatus, etc.) are all eaten by these species. Teleost fishes
also form a common item of their diet; among those which could be iden-
tified were: thread herring ( Opisthonema oglinum), the slender sea robin
( Prionotus scitulus), various poeciliids, haemulids, sparids and lutianids.
There is no assurance, however, that these fishes were taken alive, and there
is good reason to believe that many of them were eaten after being thrown
back into the water by net fishermen. Both of these catfishes are scavengers
and will apparently gather quickly in any place where refuse is thrown
overboard. This is shown by the fact that large cycloid and ctenoid scales,
which came from fishes that were much too large to be eaten by either
of these species, were commonly found among the stomach contents, thus
indicating that the catfishes had probably gathered near a boat or dock
where fishes were being cleaned for market. The best example of their
catholic tastes was found in the examination of the stomach of a single
individual (G. felis) ; the contents included coffee-grounds, peas, and pieces
of carrots, potatoes and meat.

In view of the fact that oral gestation is characteristic of the males
of both species, it is of some interest to consider the feeding habits in rela-
tion to reproduction. Observations on preserved egg-carrying male Galeich-
thys felis and Bagre marinus are in agreement with those of Beeson (Lee,
1937). Aside from catfish eggs (see below), nothing was ever found in
the stomachs of individuals practising oral gestation except an amorphous
greenish or yellowish material; most of the time the stomachs were com-
pletely empty. Quite often, however, the egg-carrying males were found
to have one or more eggs in their stomachs. As Lee has pointed out, these
were probably swallowed “because of the confusion resulting when the
fish are brought up in the trawl.” This conclusion is supported by the
observations of Lee and the present author — namely, that in most instances,
the eggs showed little or no signs of digestion and were (in each individual’s
stomach) in the same stage of development as those in the mouth. However,
in several specimens in the collection that forms the basis for this paper,
eggs were discovered in the stomachs in an advanced state of digestion. A
number of alternative explanations can be advanced to account for this.
First, it is possible that the fish was frightened into swallowing the eggs at
some time considerably previous to its eventual capture. Secondly, it is con-
ceivable that the gastric juices had a chance to act on the eggs for some
time, since the fish were not always preserved immediately after capture.
Both of these possibilities seem unlikely, however; the second alternative is
particularly so, since (as noted by Breder, 1935) the whole eggs should be
rather resistant and the effects of digestion should be slight in such a short
time, and since in the great majority of occasions when eggs were found in
the stomach, they showed no effects of the digestive processes. Thirdly, it is
possible that the egg-carrying males sometimes eat one or more of their
eggs for nourishment during the time occupied by oral gestation. Since the
period of incubation is apparently fairly long (see below), and any other
means of obtaining food is impossible because the mouth is so full of eggs,
this would not seem to be an unreasonable expectation. The possibility of
gastric incubation has been fully discussed by Breder ( loc . cit.) , who cites
Devincenzi’s (1933) article in which “incubation gastrica” is described in
the South American ariid, Tachysurus barbus (Lacepede). Breder places a
different interpretation on Devincenzi’s description most convincingly, de-
spite the latter’s interesting account of the high degree of vascularization of
the stomach, and is inclined to believe that the eggs were swallowed from

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fright at the time of capture. There is certainly no evidence for gastric in-
cubation in Galeichthys felis and Bagre marinus. The fact that Lee {loc.
cit.) and the present author have both found large catfish eggs in the
stomachs of female Galeichthys felis would indicate that such eggs are not
an entirely unacceptable item of diet. These eggs are not necessarily con-
fined to this species, for in one instance the author found eggs that were
almost certainly from Bagre marinus 3 in the stomach of a female Galeichthys
felis.

Osteology.

The skulls of Galeichthys felis and Bagre marinus have received little
attention other than that given them by Gudger (1916 and 1925), who pic-
tured them both, but discussed their structure only briefly in relation to the
size of the buccal cavity and the likeness of the ventral surface of each to a
crucifix and halo3 4 (Text-fig. 5, B and D), and by Regan (1911) in his paper
on the classification of the order Ostariophysi.

The present studies show that the high degree of specialization in the
skull structure so characteristic of the Nematognathi (Gregory, 1933) is
well exemplified by these two species. The bones of the roof of the skull are
solidly fused together so as to form the typical cephalic shield (Text-fig. 5,
A and C), to which the frontals, sphenotics, pterotics, post-temporals, and
supraoccipitals contribute in both forms. Viewed from the dorsal surface,
these bones are so well fused that it is extremely difficult to detect the sutures
in adults. Here again, the fundamental patterns of the skull structure are
much the same, and the main elements contributing to the skull are essen-
tially similar in Galeichthys felis and Bagre marinus. Yet it is apparent that
there are many distinct differences, not only in the shape of the individual
bones and the whole cephalic shields, but also in the fontanelles and fora-
mina. For instance, the frontals and supraoccipitals, especially the posterior
processes of the latter, are strikingly different in their proportions. And
it is clearly evident that the relations of the dermethmoids and premaxil-
laries are not precisely the same in both species (Text-fig. 5, A and C).
The prefrontals are also somewhat different in the two forms, although they
have been removed in Text-fig. 5, C and D. In Bagre marinus each one is
characterized by the presence of a long, relatively thin spine which is di-
rected posteriorly and lies ventral to the lateral edge of the frontal, eventu-
ally meeting a corresponding process of the frontal (see Text-fig. 5, D, for
this process). In Galeichthys felis the spine is much more stout and unites
with the lateral anterior portion of the frontal bone; it does not extend
under the ventral surface of the lateral edge of the frontal.

In Galeichthys felis there is a single median fontanelle which is only
partially divided about one-third of the way from its anterior end, and there
are two large oval foramina just lateral to the anterior half of the fontanelle.
The lateral edge of each foramen is formed by the fusion of the stout pos-
terior spine of the prefrontal with the anterolateral part of the frontal.

3 The eggs of these two species are not easily distinguishable (see below), but Bagre marinus
spawns considerably in advance of Galeichthys felis, and in this instance the time of year this
fish was captured and the condition of its own gonads, make it extremely unlikely that the eggs
were not those of B. marinus.

4 Norman (1931) discusses this likeness to a crucifix and halo as follows, “Travellers in
South America and the West Indies often return with tales of the so-called ‘Crucifix-fish,’ which
is said to be held in great esteem and even veneration by the natives of these parts, who look
upon it as a kind of fetish or charm against danger or sickness. These are nothing more
than the prepared skeletons of certain Cat-fishes. . . . The skulls of many of these fishes exhibit
on their lower surfaces a rough but readily recognizable resemblance to a crucifix, while the small
bones known as the Weberian ossicles form a halo. The upper surface of the skull, with its
rugose bones, has been described as resembling ‘a hooded monk with outstretched arms,’ or ‘the
breastplate of a Roman Soldier’ ; the dorsal spine is said to represent the spear ; and the otoliths,
which rattle when the skull is shaken, are the ‘dice with which the soldiers cast lots for the
garments of our Lord’!” Gregory (1933) in speaking of this resemblance of the under side of the
skull to a crucifix, says, “No better example perhaps could be found of a class of fortuitous
resemblances between wholly unrelated objects, which the late Professor Bashford Dean called
‘Unnatural History Resemblances’.”

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Merriman: Studies on Marine Catfish

229

A

C

Dorsal and ventral views of the skulls of Galeichthys felis (A and BJ and
Bagre marinus (C and D).

These fontanelles are probably for the passage of sensory nerves to the snout
and barbels. In Bagre marinus the median fontanelle is quite different. A
small posterior section of it is separated from the main anterior part by a
complete fusion of the bony elements in the mid-line (Text-fig. 5, C). The
main fontanelle narrows in the middle of its passage forward, but widens
into a V-shaped anterior end. Ahead of this terminal portion of the main
fontanelle is a single, median, fairly large foramen. The anterior, posterior,
and basal walls of this single median foramen in Bagre marinus have no
perforations, and the only means of entrance into it are through distinct well-
rounded apertures on its lateral walls. These apertures lead through canals
in the prefrontals and vomer directly to the brain. It may be, therefore, that

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2 3 4

Text-figure 6.

Different views of the nuchal shields of Galeichthys felis (A) and Bagre
marinus (B). 1. Dorsal view of the shield and spine of the dorsal fin. 2. Dorsal
view of the articulation of the spine with the shield, the small bone shown
separately in 3 having been removed. 3. Dorsal and ventral views of the wedge-
shaped bone lying just ahead of the spine of the dorsal fin. See text for descrip-
tion. 4. Lateral view of the shield and its various elements.

some of the nerves to the snout enter through the lateral walls and reach
the surface of the skull via this anterior median foramen. The large, lateral,
oval foramina which are so prominent in Galeichthys felis are covered over
by the forward and lateral expansion of the frontals in Bagre marinus, and
are only apparent when the skull is viewed from the anterior end. The skull
of Bagre marinus is also characterized by a prominent foramen on each side

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231

at the junction of the post-temporal and pterotic bones, while the corres-
ponding foramina in Galeichthys felis are small and easily overlooked.

Gregory (1933) points out that “the jaws are peculiarly specialized in
most siluroids in the reduction of the maxillae to small bones that support
the barbels. Diplomystes, however, the most primitive catfish, has a well de-
veloped maxillary, expanded distally and toothed (Tate Regan, 1929, p.
316).” Both the species under consideration show the reduction of the
maxillae clearly. And, as is further characteristic of the catfishes (Starks,
1926), the palatine is rod-like and separate from the pterygoid; however, it
retains its attachment to the maxilla and is modified to help control the
maxillary barbel. The bones supporting teeth in the roof of the mouth in
both species are the premaxillaries and vomers, although as Hubbs (1936)
has noted, there may be a wide variation in the vomerine dentition in
Galeichthys felis. The vomerine and premaxillary teeth were all present in
the specimen from which Text-fig. 5, B, was made, but the left half of each
series was removed before the skull was drawn. The left premaxillary was
removed before the drawing of the ventral surface of the skull of Bagre
marinus was made (Text-fig. 5, D). It is apparent that the vomerine teeth
in Bagre marinus form a bar of uniform width across the roof of the mouth.
In Galeichthys felis, however, the vomerine teeth patches on each side tend
to be triangular in shape, the apex of each triangle pointing posteriorly.

There are a number of other interesting points in the study of the
ventral aspects of the skulls of these two species (Text-fig. 5, B and D).
Among them are the distinct crescentic cavities in the sphenotic bones for
the articulation of the curved dorsal edges of the hyomandibulars (Text-fig.
7, A and B). Also, the paired bullae, containing the unusually large utricular
otoliths or lapilli (see below), are particularly prominent. Among the bones
which contribute to these bullae are the exoccipitals, pterotics, and prootics.
The post-temporal bones are of interest for several reasons. First, they are
definitely annexed to the skull and form a part of the cephalic shield. Sec-
ondly, they apparently (see below) contribute to the formation of a pair of
deep sockets for articulation of the sharp prong-like processes of the cleithra
of the pectoral girdle (Text-fig. 8). These sockets differ somewhat in the
two species, but have the same fundamental construction. Finally, the modi-
fications at the base of the skull are interesting because they are so extreme,
although they are essentially comparable to the condition in related forms.
The Weberian ossicles (believed to be derived from the ribs and neural arches
of the four anterior vertebrae) form a chain of bones on each side,
connecting the air-bladder with the perilymph-filled spaces surrounding the
inner ear. The tripus, in both species, as in some other Ostariophysi, con-
sists of two parts, a crescentic posterior part, and an anterior section. The
anterior vertebrae are highly specialized, possibly, as Gegory (1933) sug-
gests, “. . . in order to support the massive skull.” Regan (1929), describ-
ing these vertebrae in this group of fishes, says that the first vertebra
forms a disc “rigidly united to the basioccipital and to the second, third,
and fourth vertebrae, which are ankylosed to form a complex to which the
fifth is rigidly attached and with the parapophyses ankylosed to the centra.”
The transverse processes in this region where the vertebrae are so highly
modified are clearly different in Galeichthys felis and Bagre marinus, as is
shown in Text-fig. 5, but here again the basic patterns are similar.

Just behind the posterior process of the supraoccipital, and in contact
with its rear margin, in both species, lies a bony shield. This, according to
Gregory (1933), is “. . . formed by the expansion and coalescence of the
bony supports of the first three rays of the dorsal fin.” The shields of both
species are shown in Text-fig. 6. Together with the posterior process of
the supraoccipital, each is known as a nuchal shield. These structures are
much alike in the two forms under consideration, and differ only in the
proportion and relative sizes of the elements making up the units. One of

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A

DN.

ART.

Q U . 1 0 P.

Text-figure 7.

Lateral views of the dentary and opercular series in Galeichthys felis (A)
and Bagre marinus (B).

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Merriman: Studies on Marine Catfish

233

the more interesting of these elements is the V-shaped, wedge-like bone
just anterior to the true spine of the dorsal fin. This is shown in Text-fig. 6,
A1 and Bl, in its natural position from the dorsal aspect; in Text-fig. 6, A3
and B3, removed from the rest of the elements forming the shield from its
dorsal and ventral surfaces; and in Text-fig. 6, A4 and B4, in its normal
position from the lateral aspect. This bone appears to be of significance
in the articulation of the large spine of the dorsal fin only as a fundamental
part of the locking mechanism which keeps the spine erect when it has
been advanced to a vertical position. It possibly represents a rudimentary
fin ray. Regan’s (1911) account of the nuchal shield in siluroids describes
this condition and the relations of the spines to the “interneurals” (basalia
and radalia).

The lower jaw and opercular series in Galeichthys felis and Bagre
marinus are shown in Text-fig. 7, A and B. Two points are of interest in this
part of the skeleton. One is the complete absence of the subopercular
bones in both of these forms. This disappearance of the subopercular
(coincident with that of the parietals, opisthotic, and symplectic of the
skull) is characteristic of the nematognath fishes (Gergory, 1933, and Re-
gan, 1911). The opercle in these forms is in direct contact with the inter-
opercle, and although Text-fig. 7, B, shows a gap between the two, this is
simply due to a separation in the process of drying the skeleton, and the
natural relationship is as shown in Text-fig. 7, A. The other point to be
considered concerns the metapterygoid and the pterygoid bones. As de-
scribed by Regan (1911) for the Ostariophysi, the metapterygoid has moved
forward over the top of the quadrate, and it is relatively large and is con-
nected suturally with the quadrate and hyomandibular. This condition is
evident in Text-fig. 7, A and B. In the course of this movement it has taken
over the normal position of the pterygoid. In many of the Ostariophysi the
pterygoid has disappeared. Gregory (1933) says it has disappeared in all
of this group except the Bagridae. This, however, is not actually so, for
Regan (1911) has found it in some Ariidae. It is present in both Galeich-
thys felis and Bagre marinus (although not shown in Text-fig. 7), and lies
dorsal to the metapterygoid and hyomandibular, and somewhat out of the
normal relationship of this series. In these two forms the anterior part of
the metapterygoid comes exceedingly close to the posterior edge of the pal-
atine. The relationship of the metapterygoid with the surrounding bones
is not unlike that shown for “Amiurus” by Kindred (1919, Fig. 15).

The pectoral girdles of both Galeichthys felis and Bagre marinus are
shown from different aspects in Text-fig. 8. In both species the main element
of the pectoral girdle is the cleithrum, the postcleithrum and the scapula
apparently being completely lacking. What Regan (1911) calls the “hypo-
coracoids” (coracoids), “form an interlocking symphysis behind that of the
cleithra.” In these two species they are so solidly fused to the cleithra that
it is extremely difficult to separate them or to see the sutures. In Bagre
marinus they are slightly more easy to recognize as distinct entities than in
Galeichthys felis where the fusion is particularly complete. In both species
(Text-fig. 8), the serrated interlocking part of the pectoral symphysis is
composed of the “hypocoracoids”, and only the extreme anterior portion of
the symphysis is formed by the cleithra. The problem of the supracleithrum
is somewhat confusing. Regan (1911) describes it as follows for the silu-
roids : “. . . the supra-cleithrum ... is typically forked, the upper limb
usually rigidly attached to pterotic and epiotic, the lower to the basioccipital ;
. . . the distal part, ‘stem’, of the supra-cleithrum, beyond the fork, is deeply
cleft to form a socket for the head of the cleithrum.” And for the Ariidae
he says, “. . . supra-cleithrum with stout lower limb united by suture with
basioccipital.” Text-fig. 5, B and D (SCL), show this relationship much as
Regan has described it. Gregory (1933), however, labels the corresponding
bone in an Arius sp. (Fig. 79) as the post-temporal, in which case the supra-

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Text-figure 8.

Various views of the pectoral girdle, its elements, and the serrated spine of
the pectoral fin of Galeichthys felis (A, B, C and D) and of Bagre marinus

(E. F and G).

cleithrum must be lacking. It seems probable that Regan’s interpretation is
the more correct, but it also appears that the post-temporal contributes to
the socket for the head of the cleithrum. However, because of the high
degree of fusion in the bones of the skull in these species, it is difficult to
tell in skeletons of adults where the original sutures lay. The dorsal parts
of the pectoral girdles in both forms show three, prominent, prong-like proc-
esses. The most anterior of these cleithral prongs fit into the deep sockets
shown on the base of the skulls in Text-fig. 5, B and D (see above). In Text-
fig. 8, A and B, these articulating processes are the ones most medial in
position, and in E and G they are the longest of the three prongs.

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Merriman: Studies on Marine Catfish

235

The articulation of the large serrated pectoral spine and its role as a
sound-producing mechanism in both catfishes is of considerable interest.
This subject has been discussed by Burkenroad (1931), who worked on
Galeichthys felis (“ milberti ”) and found that this fish has two methods of
sound production. One is by means of a vibration of muscles in the dorsal
body wall above the air-bladder, which “would appear to be an ‘elastic spring’
mechanism essentially similar to that described for other Siluroids by S0ren-
sen (1884).” The other is by means of the highly complicated articulation
of the pectoral spine in its socket (Text-fig. 8). There are two aspects of
this articulation that deserve attention. One is a defensive locking mechan-
ism when the spine is brought forward to a position at right angles to the
longitudinal axis of the body. By means of this mechanism the spine is
fixed in place and cannot be pushed back into its normal position unless the
anterior (non-ser rated) edge of the spine is slightly elevated and the base is
simultanously rotated in a counter-clockwise manner (“counter-clockwise”
when the right half of the pectoral girdle is viewed from the lateral aspect) .
This is due to a small nubbin of bone (Text-fig. 8, D and G, 1) which slips
into a socket when the spine is advanced to the defensive position. When
the proper muscles exert their action (note the depressions and foramina
in the pectoral girdle for the passage of muscles in Text-fig. 8) the tongue-
like ridge (Text-fig. 8, D, 2) of the pectoral spine fits into and rotates in
a corresponding groove in the socket of the cleithrum. The other aspect of
this articulation which deserves attention is the problem of just how and
by what part of the base of the spine and its socket the stridulation is
effected. This subject is intimately associated with the locking mechanism
discussed above, and is reviewed by Burkenroad (1931) as follows:

“S0rensen, in connection with this phenomenon as observed by him in other
siluroid species, states that the pectoral spine is fixed in position as a defensive
weapon by the friction of an arched crest of its base against the adjacent
‘scouring faces’ of the articulation. He believes that the ‘brake-like’ action as the
spine is moved to various defensive positions incidentally causes the sounds,
which, however, he thinks may have a secondary function in frightening the
assailant. Thilo (1896), with whom Sprensen (1896) disagrees, while not men-
tioning sound production, believes that the arched crest (designated by Sprensen
as S) is a portion of the diarthrosis of the joint. Thilo believes that the essential
portion of the defensive locking mechanism is a prop-like projection ( Hemm -
fortsatz) of the base of the spine; Sprensen, on the contrary, believes this peg,
which he designates as /3, to be a part of the diarthrosis. Dufosse (1874), has
described the base of the pectoral spine of a siluroid both as a sound-producing
and a locking mechanism. The structure and action of t^e pectoral smne of
Galeichthys milberti seem to be similar to that of other siluroids described by
previous workers.”

Burkenroad does not think that (in Galeichthys ) S0rensen’s belief that
sound is produced incidentally to the defensive fixation of the spine is
correct. He believes, “. . . that the friction of the crest against the articulat-
ing surface ... is adapted ... to sound production.” Thilo ( loc . cit .) is
undoubtedly correct in thinking that the arched crest (Text-fig. 8, D, 2) is
part of the diarthrosis of the joint. Whether or not the arched crest is the
only element that produces the sound by friction against the articulating
surface is questionable in these two species. Burkenroad has pointed out
that, “Along the lateral contact surface of the crest are a number of fine
vertical striations which are thus probably to be considered as stridulatory
ridges.” These can be seen in Text-fig. 8, D, 2; there is no doubt that they
possess the function ascribed to them and are of primary importance in
this respect. Yet there is a third process (Text-fig. 8, D, 3) involved in the
articulation of the pectoral spine which may possibly play a part in sound
production. This process projects posteriorly, and when the spine is par-
tially advanced its medial surface articulates with a small area on the
coracoid. In prepared skeletons it is easy to manipulate the spine with
only a slight pressure and produce a grating noise from the articulation

[XXV :13

236

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Text-figure 9.

A and B. Dorsal views of the elements of the pelvic girdles of Bagre
marinus (A) and of Galeichthys felis (B). C and D. Lateral views of the caudal
fin skeleton of B. marinus (C) and of G. felis (D).

described above. It seems possible, therefore, that this process is at times
a sound-producing mechanism, but it certainly does not possess this func-
tion to the exclusion of that by the crest. It cannot function in this
capacity simultaneously with the arched crest, since when the latter has
the proper contact the former does not. It is at best no more than an inci-
dental method of sound production.

The pelvic girdles of Bagre marinus and Galeichthys felis are shown
from the dorsal aspect in Text-fig. 9, A and B. Sheldon (1937) has studied
the osteology and myology5 of the pelvic girdle of Hexanematichthys ( Gal-
eichthys) felis, and Text-fig. 9, B, is shown only for comparison with the
pelvic girdle of Bagre marinus. The girdles of both species have much
the same form. In each the basipterygia unite in the mid-line to form a
symphysis, the whole structure being a bilaterally symmetrical unit, the
“basal plate” (Sheldon). That of Galeichthys felis is slightly thicker and
more substantial. Both species have two pairs of elongate anterior pro-
cesses— an external and an internal pair. Each basipterygium has a single
neural foramen in Galeichthys felis, while in Bagre marinus each has a

5 Sheldon (1937) supports the view of Breder (1935) that the hook-like process on the
adaxial surface of the pelvic fin in Galeichthys (see before) is used as a clasper in the spawning
act, on the basis of his findings in the pelvic myology. He notes that various muscles (adductor
superficialis, adductor profundus, and abductor profundus) have slips which are clearly differen-
tiated and form almost independent muscles inserting on the most medial rays, which bear the
clasper-like processes. This appears to him to be a clear indication of a powerful grasping func-
tion for these modified rays.

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pair of somewhat less conspicuous foramina. The most prominent difference
between the two pectoral girdles is the presence of a pair of broad flat
posterior processes on the basipterygia of Bagre marinus. These are com-
pletely lacking in Galeichthys felis. They are situated on the medial part
of the posterior margin of the basal plate, and Sheldon (loc. cit.) notes
that in other nematognath forms they serve as a place of insertion for part
of the retractor ischii muscles.

The skeletons of the caudal fins of both species are shown in Text-fig.
9, C and D, from which the large soft caudal fin rays as well as the small
hypaxial and epaxial procurrent rays have been omitted. Here again the
structures are fundamentally the same. In each the elements composing the
hypural plate are similar. There are apparently six hypural bones, some
of which, however, are well fused. These, of course, vary much in shape,
but are not unusual in their distribution or size relationships. The urostyle
is a prominent element of both caudal fins, extending almost to the posterior
margin of the dorsal angle of the hypural plate. Above the urostyle there
is but one dorsal caudal radial in each species. The proximal ends of the
first three hypural bones (those most ventrally situated) contribute to the
formation of a prominent ridge-like process in each species. This process
is undoubtedly for muscle attachment, and is quite comparable to the
“hypural spine” described by Merriman (1940) in Roccus saxatilis, and
figured by Whitehouse (1910) for Serranus cabrilla and by Hollister (1937)
in Mugil curema. The caudal skeletons of these catfish are, in general,
characteristic of the less specialized homocercal tail; the long urostyle, and
the number and disposition of the hypural bones are indicative of this low
form of homocercy, as indicated by Whitehouse ( loc . cit.) .

Anatomy of the Inner Ear.

The anatomy of the inner ear of siluroid fishes has received little
attention other than that given it by Neumayer (1908) and de Burlet (1929).
De Burlet’s paper contains a full account of the auditory organ of
“Amiurus” ( Ameiurus ) nebulosus and other siluroids, and is by far the
most extensive discussion of this subject. It does not, however, contain
information on any of the Ariidae.

The otoliths of siluroids have been studied by Frost (1925), who has
also recorded a utricular otolith resembling those of siluroids from the
Upper Jurassic in England (1926). As Gregory (1933) notes, this “is
another suggestion of the l’elative antiquity of the ostariophysial fishes.”
In North America the fossil siluroids are poorly represented. Among the
Ariidae, Cope (1884 and 1891) described seven species belonging to the
genus, Rhineastes, which he erected for them.6 Lynn and Melland (1939)
have described an interesting fossil catfish ( Felichthys stauroforus) from
the Maryland Miocene. Not only the skull, but also the structure of the
otolith, clearly indicate that this individual is closely related to the two
forms that are the subject of this paper.

The inner ears of Galeichthys felis and Bagre marinus are almost
identical in structure. Each has a large rounded utriculus, with its utricular
otolith or lapillus, above, and a smaller elongate sacculus, with its saccular
otolith or sagitta, below. Each also has the small sac-like outgrowth from
the sacculus, the lagena, the forerunner of the spirally twisted cochlea of
higher vertebrates; these lagenae each have a characteristic otolith, the

6 Lynn & Melland (1939) point out that, "The remains upon which these species are based
are extremely fragmentary, consisting of small portions of skull plates, isolated fin-spines and
otoliths. Nevertheless, they are sufficiently complete to indicate clearly that the genus belongs
among the sea-catfishes so it has been assigned to the family Ariidae by Jordan (1923).” Hay
(1929) lists five other species (besides Cope’s seven) from North American deposits; these, how-
ever, are freshwater forms belonging to the family Ameiuridae.

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asteriscus. These chambers with their otoliths and the semi-circular canals
are shown in Plate I, A, B; the lapilli and asterisci removed from their
respective chambers appear in Plate II, A, B, C, D.

The semi-circular canals are of course connected with the utriculus
and are quite normal in relationships and distribution. There are the
typical anterior and posterior vertical canals and the horizontal canal in
each. The prominent ampullae are located in their usual positions and are
clearly visible in Plate I, A, B. Due to the much enlarged utriculus, how-
ever, the semi-circular canals lie close to this upper chamber, and do not
extend well above it as in other forms. Instead, the utriculus fills the
whole space between the canals, and all three of the canals run close to
and often touching the surface of the utriculus. The semi-circular canals
are not solidly embedded in the bones of the skull, and only small portions
of them pass through bone. It is also worth mentioning in passing that
the utriculus is covered with a layer of highly pigmented tissue. This
melanistic pigmentation is clearly visible in Plate I, A, B, although in A
part of this tissue was removed before the photograph was made. The pig-
mentation is not confined to the utriculus and can be seen both on the
ampullae and the semi-circular canals. The sacculus and lagena, by con-
trast, are unpigmented.

The sacculus and lagena are not unusual except in relation to the
large size of the utriculus (see below). They are, however, solidly em-
bedded in bone. The saccular otolith (sagitta) in both species is an elongate
and highly brittle affair. Unlike the asteriscus or lapillus, the sagitta
crumbles easily. It is irregularly circular in cross-section, narrowing at
both ends and rounded at either extremity — in other words, it is roughly
cigar-shaped. In adults it is 4-6 mm. in length. The otolith of the sac-like
lagena, the asteriscus, from Bagre marinus is shown in Plate II, C, D, from
the dorsal and ventral surfaces. In both forms it is a thin, disc-like affair,
averaging 3-5 mm. in diameter in adults. The spiral appearance of its flat
surface shown in Plate II, D, is characteristic of the two catfishes that
form the subject of this paper. As can be seen in Plate I, B, the asteriscus
lies directly above the sagitta, which runs in an antero-posterior direction,
and its flat surfaces form dorsal and ventral sides.

In almost all fishes the sagitta is the largest otolith and the lapillus
tends to be small and insignificant, although in some forms the sagitta is
small and the asteriscus is relatively large. In Galeichthys felis and Bagre
marinus, however, the utricular otolith or lapillus is much enlarged and
dwarfs the sagitta and asteriscus by comparison. The lapilli of these adult
catfishes may be as much as 15-16 mm. in diameter. As described by Frost
(1925) their “shape is conchoidal and biconvex.” Each has a prominent
posterior process, and each has characteristic markings that are nearly
identical in both forms (Plate II, A, B). Frost ( loc . cit.) notes that in the
Ariidae in general “. . . the lapillus is much larger, in comparison with the
other otoliths, than in other Siluroids except the Plotosidse.” In conclusion
he states, “In the Ostariophysi the saccular otolith, the sagitta, which is
generally the principal otolith in other fishes, is attenuated and diminutive.
In the Cyprinoids and Diplomystes the asteriscus is the largest otolith, in
the Siluroids (except Diplomystes and a few South American species) the
lapillus. It seems possible that the reduction of the sagitta may be related
to the development of the Weberian mechanism, and that the great develop-
ment of the lapillus or utricular otolith, in the Siluroids, may in muddy
waters compensate for the decreased use of the eyes for maintaining
equilibrium.” This last statement appears to be somewhat debatable, but
may well be a partial explanation of the relatively large size of the lapillus
by comparison with that of the sagitta. Frost also points out that the
interrelationships of the Siluroid families with each other and with the
primitive characins are clarified considerably by studies on the otoliths.

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Embryology.

Despite the fact that the eggs of Galeichthys felis and Bagre marinus
are among the largest known in the teleost fishes, only a few authors have
made any study of the embryology. This is partially because of the difficulty
of getting a complete series of developmental stages, owing to the habit of
the males in both forms of practicing oral gestation. Both Gudger (1916)
and Lee (1937) experienced the greatest difficulty in keeping the eggs alive
after removing them from the parent male’s mouth. It therefore appears
probable that the most feasible way to obtain a complete series is to collect
the egg-carrying males in sufficient numbers until all the stages are filled.
This necessitates a large collection since the eggs in a single male’s mouth
are usually all in the same stage of development.

The present studies are confined mainly to the eggs and larvae of
Galeichthys felis, and are based entirely on preserved specimens. It is
apparent from Gudger’s (1916 and 1918) figures, however, that the eggs
and larvae of G. felis are much like those of B. marinus. Comparing the
small number of eggs of B. marinus in the present collection with those of
G. felis, it is not easy to distinguish between the two forms in early stages.
Gudger’s (1918) measurements of the eggs of B. marinus would indicate
that they average a little larger than the eggs of G. felis, but there is con-
siderable overlap between the two forms since there is a wide variation in
size in each type (see below). Many of the characteristics which serve as
a means of distinguishing these catfishes when they become adults, however,
are useful in separating the later egg stages of the two forms. Thus the
barbels are sufficiently well developed some time before hatching to enable
immediate identification, although the removal of the egg shell usually
facilitates the necessary observations. The two barbels on the lower jaw
of Bagre marinus and the four on that of Galeichthys felis are differentiated
early in development. Also the much longer maxillary barbels and the fila-
ments on the pectoral and doi’sal spines in B. marinus are apparent well
before hatching.

The study of the gonads of a large series of adult and immature speci-
mens of Galeichthys felis and Bagre marinus varying in size from 17-44
cm. (standard length) has given some interesting information as to the
time of spawning, the maturation of the ova, the age of maturity, etc. The
individuals on which this study is based were collected in February, March
and April, and June, July and August. It is evident from this collection
that the main time of spawning for G. felis in 1938 on the west coast of
Florida was June and July, while B. marinus spawned about a month earlier
in this locality in that year. This information checks well with the con-
clusions of Lee (1937) on Galeichthys felis, which were based on observa-
tions over a number of years in the vicinity of Grand Isle, Louisiana.

The size of the eggs of G. felis just before spawning ranges from 12-19
mm. in diameter, the average being from 14-17 mm. Gudger’s (1918 and
1919) measurements on the eggs of B. marinus show that the average size
is 19-20 mm., and that the lower limits of size in this form are comparable
to the average size of the eggs of G. felis. It should be mentioned, however,
that Gudger’s measurements were on unpreserved material, by contrast to
those in the present study. But the limited number of measurements of the
eggs of B. marinus which the present collection allowed, indicate that the
eggs are usually somewhat larger than those of G. felis. The eggs of both
forms vary in shape from a somewhat oval or elliptical to a perfectly spher-
ical condition, the latter state, however, being much less common. The
measurements given in this paper refer to the longest diameters in every
instance. Just after spawning the largest eggs in the ovaries of these two
forms average 2-4 mm. in diameter, although there are occasionally eggs
up to 6 mm. in diameter. There are, of course, countless eggs of various

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sizes below 2 mm. The larger eggs at this time are undoubtedly those which
will be spawned the following season. It is of course impossible to obtain
much information on the progressive increase in size of the developing eggs
without material collected throughout the year. However, studies on a lim-
ited number of apparently maturing females of both forms taken in Feb-
ruary, March and April, indicate that the increase in size of the eggs at
this season is extraordinarily rapid, and that there is relatively little
change in egg size from immediately after the time of spawning in the
spring and early summer to January and February. In other words, the
tremendous increase in bulk of the maturing eggs in both forms is mainly
confined to the four or five months immediately preceding spawning; this
increase in bulk is therefore not a progressive affair over an eleven or
twelve months period, but is mainly concentrated in a much shorter space
of time. Thus the largest maturing eggs from the gonads of Bagre marinus
averaged from 5-8 mm. in diameter in February, about 14 mm. in diameter
in March, and 18-19 mm. in diameter in samples taken in April. Similar
samples of maturing eggs from the gonads of Galeichthys felis indicated
that the increase in bulk of the eggs was equally rapid but did not start
until a month or more later. In March the eggs were only 4 mm. or a
little more in average diameter, but by April they had increased to 7 mm.
in most instances, and in one individual the largest eggs were 12-14 mm.
in diameter. The somewhat later period of rapid increase in bulk of the
maturing eggs of Galeichthys felis is undoubtedly correlated with the
correspondingly later time of spawning in this form.

The present collection was unfortunately rather limited in the numbers
of immature individuals, so that it is impossible to draw many conclusions
as to the size at which these two catfishes first become mature. Lee (1937)
found one female Galeichthys felis only 12.6 cm. in standard length which
was gravid. The present author has found a considerable number of female
sea catfish up to and occasionally above 20 cm. (standard length) which
were either immature or not spawning in that particular season, for their
gonads had no eggs larger than 2-4 mm. in diameter immediately before the
spawning season, at a time when mature fish possessed many eggs 14 cm.
or more in diameter. These were not fish which had spawned in advance
of the main lot, since their gonads were firm and full and did not have
the typical flaccid, empty appearance characteristic of gonads that have just
lost the mature eggs and show empty follicles. It therefore seems probable
that there is a considerable variation in the size at which female Galeichthys
felis first mature; this range is roughly from 12-20 cm., with the majority
of individuals maturing for the first time near the upper end of the scale.
No information was obtained as to the size at maturity of male Galeichthys
felis, save from several individuals that were either immature or not spawn-
ing that season; these fish were 19 and 20 cm. in standard length respec-
tively. It may therefore be that the males mature when slightly larger
than the females. Males above '25 cm. were found to be mature. The col-
lection did not contain any immature Bagre marinus, so it is impossible
to draw any conclusions as to the length at maturity of this form. The
smallest specimen was 26.5 cm. in standard length and was a mature female.

The problem of how many mature eggs are produced each season by the
females of these two catfishes is of considerable interest in view of the large
size of the eggs. In Galeichthys felis the number of mature eggs per gonad
usually varies from 10-20, and the total number per fish 20-40. Occasionally
a larger number of eggs were found, however; one individual collected
just before the spawning season contained 53 mature eggs in its gonads,
while another had 64 (39 in one gonad and 25 in the other). The average
number of eggs produced each year by Bagre marinus is of the same order
of magnitude. Evidence that all the mature eggs in one season are not
extruded at one time in Galeichthys felis is forthcoming from the examina-

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tion of gonads from individuals in the present collection. A number of
females were found which possessed one gonad in which all the mature eggs
had been released and in which empty follicles were plainly evident, but in
which the other gonad had a few mature eggs and some empty follicles. Such
an individual had certainly spawned once or more already, and in all proba-
bility would have spawned again that season and so have rid herself of the
few remaining mature eggs, thus indicating that polygamy in this form
is apparently not uncommon.

The number of eggs carried in the mouths of the orally gestating males
has been treated by Gudger (1918) for Bagre marinus and Lee (1937) for
Galeichthys felis. Gudger found this number to vary from 2 to 55, the
average being from 15-30. The largest number of eggs in the mouth of a
male Galeichthys felis recorded by Lee is 48. The average number is slightly
less than that for Bagre marinus. In the same manner as indicated by
Gudger for Bagre marinus, a single female Galeichthys felis can apparently
produce more eggs than a single male can carry in his mouth, which is
added evidence that polygamy is not uncommon among these forms.

The eggs in the mouth of any individual male are usually all in the same
stage of development, but, as noted by Lee (1937), there are exceptions.
Sometimes there may be several infertile eggs among the developing ones,
and rarely one or two eggs that are in a much earlier stage of development
than the main lot. Undoubtedly the infertile eggs are picked up with the
fertile ones by the male at times. It may also be that if a male does not
acquire his full complement of eggs the first time, he will attempt to pick
up additional eggs later on. On the other hand, several eggs in an earlier
stage than the majority in a male’s mouth may simply be cases of retarded
development, as suggested by Lee.

As mentioned before, the young remain in the parent male’s mouth long
after hatching, in many instances up to the time when the yolk sac is com-
pletely absorbed. Gudger (1918) believes the entire incubatory period to be
60 to 70 days in Bagre marinus near Beaufort, N. C. Evidence from the
present collection indicates that the time occupied by oral gestation is some-
what shorter for both Galeichthys felis and Bagre marinus on the west coast
of Florida, and that 6 to 8 weeks is the period there; this estimate, how-
ever, is based on an inadequate amount of material (see below). Gudger is
convinced that the young Bagre marinus actually feed while still in the mouth
of the parent male, “. . . filtering out of the respired sea-water, by means of
their closely set gill rakers, minute Crustacea to satisfy their hunger.” Proof
that this supposition is correct not only for Bagre marinus but also for
Galeichthys felis is provided by the dissection of individuals in a late stage
of development but still in the parent males’ mouths. Stomach content an-
alyses showed that in almost every instance these small fish had been feeding
heavily. There is some possibility, of course, that the small fish feed outside
the parent male’s mouth and subsequently return to it or are picked up again
by the male.

The developmental stages of Galeichthys felis are shown in Plate II,
1-6, and Plate III. Plate II, 1, shows the whole egg in which the developing
embryo is at the earliest stage represented in the entire collection. The egg
shell has not been removed (as it has in the succeeding photographs in this
figure), and it is possible to see the distinct space between it and the
yolk and developing embryo. This perivitelline space is apparently formed
at or shortly after the time of fertilization, when the chorion moves out
from the yolk which it so closely surrounds before fertilization. Plate II,
2, shows the same stage with the egg shell removed, and the developing
embryo is clearly visible on the surface of the yolk. A photograph of a
whole mount of this embryo dissected away from the yolk is shown in
Plate III, 1. It is obvious that at this stage the eye and lens are clearly
differentiated, mesodermal segmentation has progressed a long way, the

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somites being evident over the greater part of the entire length of the
fish, neuromeres are distinctly visible, the pectoral limb buds are apparent,
and the otoliths have formed. The frontal and sagittal sections of embryos
of this stage shown in Plate IV, 1, 2, 3, also illustrate these points. It is
characteristic of the yolk at this stage to show an irregular trough-shaped
indentation of considerable size in the vicinity of the embryo. The embryo
usually lies at one end of this indentation, the posterior part of its body
being in it, while the anterior end extends out onto the smooth surface of
the yolk. This is probably the groove into which the embryo fits so closely
at subsequent stages.

It is of some interest to consider why no earlier stages were present
in the entire collection despite the large number of eggs which were
examined. Gudger (1916) failed to obtain any of the segmentation stages
in his studies of Bagre marinus, but got a complete series “from invagina-
tion to the free swimming young in which the walls of the belly have
closed over the diminished yolk sac and have coalesced into a raphe on
the median line.” In Galeichthys felis the absence of any early stages may
possibly be accounted for by the following statement by Smith (1907) :
“Spawning occurs in the summer, the large eggs being first deposited
in a sandy depression and subsequently taken into the mouth of one of the
parents. . . Others have made similar observations (Gudger, 1918),
which, if reliable, would account for the lack of early stages in the mouth
of any orally gestating male.

The next stage of development in Galeichthys felis is shown in its
normal relation to the yolk in Plate II, 3, as a whole mount of the embryo
dissected away from the yolk in Plate III, 2, and in sagittal section in
Plate IV, 4, 5. By this time there has been a considerable advance over the
previous stage. The eyes have become particularly prominent, the pectoral
limb buds have enlarged considerably, and the “coiling” of the body so
characteristic of the later stages has started with the twisting of the
caudal end of the embryo to a position where it is at right angles to the
longitudinal axis of the body. This coiling in a more advanced state is
apparent in Plate II, 4, 5, and Plate III, 3. Here it will be seen that the
posterior half of the body has twisted to such an extent that the caudal
fin at least touches the head and more usually overlaps it completely. This
coiling is neither universally sinistral or dextral; however, it tends to be
counter-clockwise in the majority of the specimens examined. It will be
noticed that the pigmentation of the dorsal surface of the body becomes
increasingly prominent as this coiling stage is more fully developed. The
barbels, which serve as a means of distinguishing these two catfishes (see
before), are also clearly evident by this time. Furthermore, the prominent
utricular otoliths have attained a large size at this stage and the hypural
plate has undergone considerable development (see Plate III, 3). Sagittal
sections through the anterior part of the body at the coiling stage are
shown in Plate V. The parasagittal section in Plate V, 1, shows the well-
developed eye with its retina and lens, the characteristic and perfectly
normal gill arches, and the large space in the posterior part of the neuro-
cranium that is the site of the more prominent part of the inner ear —
namely, the semicircular canals, utriculus, and lapillus. The nearly median
sagittal section at the same stage (Plate V, 2) indicates, by comparison
with sagittal sections of the previous stage, that the main steps in organo-
genesis have been accomplished during this phase of development. Thus
the swim-bladder, with its distinct partitions, the oesophagus and gut with
its rugose lining, the beginnings of the convolutions of the gastro-intestinal
tract, the kidney, etc., are now clearly evident.

The developing embryo apparently remains in the coiled stage up to
the time of hatching. Shortly after hatching the yolk sac is still large,
but the larvae are well-developed and most of the fundamental differences

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between Bagre marinus and Galeichthys felis are readily apparent — e.g.,
see the photograph of the newly hatched G. felis in Plate II, 6. The yolk
sac is now gradually absorbed, and, as mentioned before, the small fish are
regularly found in the parent males’ mouths up to and even some time after
the complete disappearance of the yolk sac.

Finally some information on the rate of development of Galeichthys felis
is provided by the following data. The eggs in the earliest stage taken
(Plate II, 2, and Plate III, 1) were found in the mouths of males commonly
in late July, and in one instance in mid-August in 1938. Those in the suc-
ceeding stages of development (Plate II, 3, and Plate III, 2) were taken
most frequently in late July and early August, although these too have
been collected in mid-August or slightly later. Embryos in the late coiling
stage were taken in greatest numbers in mid-August, although it was not
uncommon to find individuals which had hatched and even those in which
the yolk sac had been completely absorbed by this time. Unfortunately
the collection of material stopped shortly after the middle of August, so
that it was impossible to draw any conclusions as to how much later oi’al
gestation is carried on. The largest individual taken from the mouth of a
parent male at this time was 49.5 mm. in standard length, and many larvae
averaging about 40 mm. were collected at that date. The above information
indicates that in 1938 on the west coast of Florida, hatching occurred in
about a month, and that the larvae were retained in the parents’ mouths
from 2-4 weeks thereafter.

Summary.

1. The material presented in this paper is based on a large number of
preserved eggs, young, and adults, of the gaff-topsail catfish, Bagre marinus,
and the silver or sea catfish, Galeichthys felis, which were collected in the
vicinity of Palmetto Key, Florida. Various morphological and embryo-
logical aspects of these two catfishes are discussed under the following
headings :

External Anatomy.

2. A distinguishing character which seems to have been generally
overlooked, is the posterior margin of the anal fin, which in B. marinus
has a prominent V-shaped indentation and in G. felis a comparatively
straight border. The cross-sectional shape of the maxillary barbels of the
two species is not as clearly diagnostic as a distinguishing character as it
has been described to be.

3. The pelvic fins of mature (or nearly mature) female G. felis have
extraordinary, fleshy, hook-like protuberances on the adaxial surfaces; the
males not infrequently exhibit a similar, though far less well developed,
structure. Neither male nor female B. marinus possess corresponding
modifications of the pelvics.

4. The pelvic fins of mature females of both forms are considerably
larger, in proportion to the length of the fish, than in mature males of the
same species. The sexually dimorphic character of the pelvic fins of
B. marinus is apparent both as to size and shape.

Morphology of the Alimentary Tracts and the Feeding Habits.

5. The parietal peritoneum in G. felis appears almost grayish, due to
minute black pigment spots which are abundant over its otherwise white
surface, and which are more numerous on the dorsal surface of the coelom
than on the lateral and ventral walls of the body cavity. The peritoneum
of B. marinus lacks any melanistic pigmentation, and is a clean, silvery
white.

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6. The fundamental pattern of the gastro-intestinal tracts is the same
in both species. Yet there are minor differences between the two which
are constant for each form. Among these is the fact that the length of
the intestine in B. marinus is longer in proportion to the size of the fish
than in G. felis.

7. Stomach content analyses show that there is no significant difference
in the diets of these two species. Both forms are completely omnivorous.
There is certainly no evidence for gastric incubation in G. felis and
B. marinus, although the eggs of these catfish are apparently not an entirely
unacceptable item of diet at times.

Osteology.

8. The present studies show that the high degree of specialization in
the skull structure so characteristic of the Nematognathi is well exempli-
fied by these two species. Thus the bones of the skull are solidly fused
together in both forms so as to form the typical cephalic shield. The
fundamental patterns of the skull structure are essentially similar, yet
there are many distinct differences, not only in the shape of the individual
bones and the whole cephalic shields, but also in the fontanelles and fora-
mina; these are described in some detail.

9. A nuchal shield lies just behind the posterior process of the supra-
occipital, and in contact with its rear margin, in both species. These
structures are much alike in the two forms, but show characteristic dif-
ferences.

10. Studies of the lower jaw and opercular series show several in-
teresting specializations. Among these is the complete disappearance of the
subopercular (coincident with that of the parietals, opisthotic, and sym-
plectic of the skull), and the relation of the metapterygoid and pterygoid
bones.

11. The main elements of the pectoral girdles are the cleithra and
coracoids, and the articulation of these girdles in both forms is accom-
plished by prominent cleithral prongs which fit into deep sockets on the
base of the skull. The articulations of the serrated pectoral spines with the
girdles are highly complex, involving defensive locking-mechanisms and
sound-production.

12. The pelvic girdles in these catfish are much alike, although that
of B. marinus is characterized by a pair of broad flat posterior processes
on the basipterygia.

13. The caudal skeletons are characteristic of the less specialized
homocercal tail, as indicated by the long urostyle and the number and
disposition of the hypural bones.

Anatomy of the Inner Ear.

14. The inner ears of G. felis and B. marinus are almost identical in
structure. Each has a large rounded utriculus, with its utricular otolith
or lapillus, above, and a smaller elongate sacculus, with its saccular otolith
or sagitta, below. Each also has a small sac-like outgrowth from the
sacculus, the lagena, the forerunner of the spirally twisted cochlea of
higher vertebrates; these lagenae each have a characteristic otolith, the
asteriscus.

15. In almost all fishes the sagitta is the largest otolith and the
lapillus tends to be small and insignificant, although in some forms the
sagitta is small and the asteriscus is relatively large. In G. felis and B.
marinus, however, the utricular otolith or lapillus is much enlarged and
dwarfs the sagitta and asteriscus by comparison. The lapilli of these
adult catfishes may be as much as 15-16 mm. in diameter.

1940]

Merriman: Studies on Marine Catfish

245

Embryology.

16. The eggs of B. marinus and G. fells are much alike, although
those of B. marinus are, on the average, slightly larger. Many of the
characters which serve as a means of distinguishing these catfish as adults
are useful in separating the later egg stages of the two forms — e.g., the
barbels.

17. Studies of the gonads of these catfishes indicate that the main
time of spawning for G. felis in 1938 on the west coast of Florida was
June and July, while B. marinus spawned about a month earlier in this
locality in that year.

18. Observations on the gonads of a limited number of apparently
maturing females of both forms taken in February, March and April, indi-
cate that the increase in size of the eggs at this season is extraordinarily
rapid, and that there is relatively little change in egg size from immediately
after the time of spawning in the spring and early summer to January and
February. In other words, the tremendous increase in bulk of the matur-
ing eggs in both forms is mainly confined to the four or five months imme-
diately preceding spawning.

19. There is probably considerable variation in the size at which
female G. felis first mature; this range is roughly from 12-20 cm., with the
majority of individuals maturing for the first time near the upper end of
the scale.

20. The total number of mature eggs produced by the females of these
two catfish is usually from 20-40 per individual, although there are some
instances where the numbers are smaller or larger than these limits.

21. The numbers of eggs carried in the mouths of the orally gestating
males vary greatly in both forms, the average being 10-30. A single
female can apparently produce more eggs than a single male can carry
in his mouth, which, in the light of other evidence, indicates that polygamy
is not uncommon among these forms.

22. The eggs in the mouth of any individual male are usually all in
the same stage of development, although there are exceptions. The young
remain in the parent male’s mouth long after hatching, in many instances
up to the time when the yolk sac is completely absorbed. The incubatory
period on the west coast of Florida is probably about 6-8 weeks long. The
young taken from the mouths of parent males were often found to have
been feeding independently, as evidenced by the presence of small Crustacea
in their stomachs.

23. The earliest stage of development in this collection is represented
by a number of specimens in which the eye and lens are clearly differenti-
ated, mesodermal segmentation has progressed a long way, the somites
being evident over the greater part of the entire length of the fish,
neuromeres are distinctly visible, the pectoral limb buds are apparent, and
the otoliths have formed. A possible explanation of the lack of earlier
stages in the mouths of orally gestating individuals may be that the eggs
are first deposited in a sandy depression and subsequently taken into the
mouth of the parent male.

24. The later stages show the characteristic “coiling” of the body,
the time at which the main steps in organogenesis occur, and various ex-
ternal and internal developmental features of interest. All available stages
have been studied as whole mounts, and in sagittal and frontal sections.

25. Studies on the rate of development indicate that in 1938 on the
west coast of Florida, hatching occurred in about a month, and that the
larvae were retained in the parents’ mouths from 2-4 weeks thereafter.

246

Zoologica: New York Zoological Society

[XXV :13

Bibliography.7

Breder, C. M., Jr.

1929. Field book of marine fishes of the Atlantic Coast from Labrador to
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1935. The reproductive habits of the common catfish, Ameiurus nebulosus
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Burkenroad, M. D.

1931. Notes on the sound-producing marine fishes of Louisiana. Copeia, 1:
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Cope, E. D.

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de Burlet, H. M.

1929. Anatomisches zur Horfahigkeit der Siluroiden. Zeitschr. f. Anat. u.
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*Devincenzi, G. J.

1933. La perpetuacion de la especie en los Peces Sudamericanos. An. Mus.
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♦Dufosse, M.

1874. Recherches sur les sons expresifs que font entendre les poissona
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Frost, G. A.

1925. A comparative study of the otoliths of the Neopterygian fishes (con-
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433-446.

1926. Otoliths of fishes from the Jurassic of Buckinghamshire and Dorset.
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Gowanloch, J. W.

1933. Fishes and fishing in Louisiana. Bull. 23, State of La. Dept, of Con-
servation, New Orleans: 1-638.

Gregory, W. K.

1933. Fish skulls, a study of the evolution of natural mechanisms. Trans.
Amer. Philos. Soc., N. S., 23: 75-481.

Gudger, E. W.

1916. The gaff-topsail ( Felichthys felis ), a sea catfish that carries its eggs
in its mouth. Zoologica, 2 (5) : 123-156.

1918. Oral gestation in the gaff-topsail catfish, Felichthys felis. Papers
from the Dept, of Mar. Biol, of the Carnegie Instit. of Wash., 12 (Pub.
No. 252) : 25-52.

1919. The ovary of Felichthys felis, the gaff-topsail catfish; its structure
and function. Papers from the Dept, of Mar. Biol, of the Carnegie
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1925. The crucifix in the catfish skull. Nat. Hist., 25 (4) : 371-380.

♦Hardenberg, J. D. F.

1935. Miscellaneous notes on Indian fishes. IV. On the propagation of Arius
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7 A * indicates that the original paper has not been read by this author, and that the
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1940]

Merriman: Studies on Marine Catfish

247

Hay, O. P.

1929. Second bibliography and catalogue of the fossil vertebrates of North
America. Carnegie Instit. of Wash., Pub. No. 390, 1 : 1-916.

Hollister, G.

1937. Caudal skeleton of Bermuda shallow water fishes. II. Order Perco-
morphi, Suborder Percesoces: Atherinidae, Mugilidae, Sphyraenidae.
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Hubbs, C. L.

1936. Fishes of the Yucatan Peninsula. Publ. Cam. Instit. Wash., Pub. No.
457: 157-287.

Jordan, D. S.

1923. A classification of fishes, including families and genera as far as
known. Stan. Univ. Publ., Biol. Sci., 3: 79-243.

Jordan, D. S. & Evermann, B. W.

1896. The fishes of North and Middle America. Bull. U. S. Nat. Mus.,
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Kindred, J. E.

1919. The skull of Amiurus. Illinois Biol. Monogr., 5 (1): 1-120.

Lee, G.

1931. Oral gestation in the marine six-whiskered catfish, Galeichthys felis.
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1937. Oral gestation in the marine catfish, Galeichthys felis. Copeia, 1 : 49-56.
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1939. A fossil catfish ( Felichthys stauroforus) from the Maryland Miocene.
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Meek, S. E. & Hildebrand, S. F.

1923. The marine fishes of Panama. Field Museum of Natural Hist., Zool.
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Merriman, D.

1940. The osteology of the striped bass ( Roccus saxatUis) . Ann. and Mag.
of Nat. Hist. (11), 5 (25): 55-64.

Neumayer, L.

1908. Zur vergleichenden Anatomie des Gehirns und des Cavum Cranii der
Siluroiden. Anat. Anz., 32: 273-277.

Norman, J. R.

1931. A history of fishes. Ernest Benn, Ltd., London: 1-463.

Regan, C. T.

1906-1908. Pisces. Biologia Centrali- Americana: 1-203.

1911. The classification of the Teleostean fishes of the order Ostariophysi. —
2. Siluroidea. Ann. and Mag. of Nat. Hist. (8), 8: 553-577.

1929. Fishes. Encycl. Brit. (14), 9: 305-328.

Sheldon. F. F.

1937. Osteology, myology, and probable evolution of the Nematognath pelvic
girdle. Ann. N. Y. Acad. Sci., 37 (1) : 1-96.

Smith, H. M.

1907. The fishes of North Carolina. N. Car. Geol. and Econ. Surv., 2: 1-449.
S0RENSEN, W.

*1884. Om Lydorganer hos Fiske. Copenhagen. Dissertation.

1896. Some remarks on Dr. Thilo’s memoir on “Die Umbildungen an den
Gliedmassen der Fische.” Morph. Jahrb., 25: 170-189.

248

Zoologica: New York Zoological Society

[XXV :13

Starks, E. C.

1926. Bones of the ethmoid region of the fish skull. Stanford Univ. Publ.,
Biol. Sci, 4 (3) : 139-388.

Thilo, O.

1896. Die Umbildungen an den Gliedmassen der Fische. Morph. Jahrb., 24:
287-355.

Tippett, L. H. C.

1931. The methods of statistics. Williams and Norgate, Ltd., London: 1-222.
Whitehouse, R. H.

1910. The caudal fin of the Teleostomi. Proc. Zool. Soc. of London : 590-629.

EXPLANATION OF THE PLATES.

Plate I.

A. Dorso-lateral view of the utriculus and semi-circular canals of Bagre
marinus. The sacculus has been removed.

B. Side view of the inner ear of Galeichthys felis. Note the large size of the
utriculus.

Plate II.

Otoliths from Galeichthys felis and Bagre marinus.

A. Views of the two sides of the lapillus from B. marinus.

B. The same from G. felis.

C and D. Dorsal and ventral aspects of the asteriscus of B. marinus.

1-6. Various developmental stages of the eggs and larvae of Galeichthys felis.

Plate III.

1-3. Whole mounts of the embryos of Galeichthys felis in different stages
of development.

Plate IV.

1-5. Sagittal and frontal sections of Galeichthys felis embryos in various
stages of development. See text for description.

Plate V.

1 and 2. Sagittal sections through the anterior part of the body of Galeichthys
felis in the “coiling” stage of development.

MERRIMAN.

PLATE I.

UTR. OTOL.

ANT.

VERT. —
CAN.

ANT.

AMP.—

A

POST.

VERT.

CAN.

POST.

AMP.

ANT.

AMP.—

HOR. „
CAN.

ANT. VERT. CAN.

UTR. OTOL.

POST.

VERT.

CAN.

LAG.

SAC.

B.

MORPHOLOGICAL AND EM BRYOLOGICAL STUDIES ON TWO SPECIES
OF MARINE CATFISH. BAGRE MARINUS AND GALEICHTHVS FELIS.

MERRIMAN.

PLATE II

MORPHOLOGICAL AND EM BRYOLOGICAL STUDIES ON TWO SPECIES
OF MARINE CATFISH, BAGRE MARINUS AND GALEICHTHYS FELIS.

MERR1MAN.

PLATE III.

MORPHOLOGICAL AND EM BRYOLOGICAL STUDIES ON TWO SPECIES
OF MARINE CATFISH, BAGRE MARINUS AND GALEICHTHYS FELIS.

MERRIMAN.

PLATE IV.

1.

5.

MORPHOLOGICAL AND EM BRYOLOGICAL STUDIES ON TWO SPECIES
OF MARINE CATFISH, BAGRE MARINUS AND GALEICHTHYS FELIS.

MERRIMAN.

PLATE V.

1.

MORPHOLOGICAL AND EM BRYOLOGICAL STUDIES ON TWO SPECIES
OF MARINE CATFISH, BAGRE MARINUS AND GALEICHTHYS FELIS.

Coates, Cox, Rosenblith & Blown: Electric Eel

249

14.

Propagation of the Electric Impulse along the Organs of the
Electric Eel, Electrophorus electricus (Linnaeus).

C. W. Coates,

New York Aquarium,

R. T. Cox, W. A. Rosenblith,

Department of Physics, New York University

&

M. Vertner Brown,

Department of Physics, College of the City of Neiv York.

(Plate I; Text-figures 1-3).

Observations already reported1 have shown that in the discharge of the
electric organs of the electric eel a pulse of potential gradient runs along the
organ from anterior to posterior. The speed of the pulse was roughly
estimated by several methods and was found much higher than the highest
recorded speeds of impulses along nerves. The cathode-ray oscillograph used
in these observations did not, however, make possible anything better than
rough determinations of the speed. A more suitable cathode-ray oscillograph2
having lately become available, it seemed worth while to attempt more
accurate measurements.

The method employed is almost the same as one of those used before.
The fish is taken out of water and laid in an insulating trough. The trough
is grooved across one side every 10 cm. and aluminum strips inserted in
any of these grooves make good electrical connection through the skin to the
large electric organs and serve as electrodes for connecting the organ to the
oscillograph. Text-fig. 1 shows the positions of these electrodes on the fish
in one of the observations. Electrode 1 was at the anterior end of the large
organ and electrode 2 was 10 cm. behind it. With these positions a con-
veniently measurable voltage was developed in the segment of the organ
included between the two electrodes. The first two electrodes were conse-
quently kept in these positions during all the observations. Electrode 3
was placed at a distance behind electrode 2 differently chosen in the various
observations. Electrode 4 was placed far enough behind electrode 3 that
the peak voltage between 3 and 4 was comparable to that between 1 and 2.

Oscillographic traces were photographed with three different connec-
tions of the electrodes to the oscillograph. First electrodes 1 and 2 were
connected to the vertically deflecting plates of the oscillograph tube. When
the discharge of the eel developes a voltage between these two electrodes,

1 Coates, C. W., Cox, R. T., and Granath, L. P. The Electric Discharge of the Electric Eel,
Electrophorus electricus (Linnaeus). Zoologica, New York, 1937. Vol. 22, Part 1, pp. 1-32.

2 Allen B. Dumont. Type 175A Oscillograph.

250

Zoologica: New York Zoological Society

[XXV :14

this voltage applied to the plates deflects the beam of electrons in the
oscillograph tube. The luminous spot which the beam makes where it strikes
the fluorescent screen at the end of the tube rises and falls with the voltage
developed in this segment of the organ, the displacement of the luminous
spot at any instant being proportional to the voltage existing at the same
instant between the electrodes. While the discharge of the fish is causing
this vertical motion, voltage from a “sweep circuit,” applied to the hori-
zontally deflecting plates of the oscillograph tube, carries the luminous spot
horizontally across the fluorescent screen at a constant rate. The luminous
spot thus traces a graph, which can be photographed, in which the voltage
between the electrodes is plotted vertically against the time plotted horizon-
tally.

12 3 4

Position of electrodes for determining voltage at anterior and posterior ends
of electric organs.

Next the vertically deflecting plates were connected to electrodes 3 and
4 on the fish and the graph of voltage against time was obtained for the
discharge in this posterior segment. Plate I, Figs. 1 and 2, show traces
obtained for the discharge of the two segments in an eel 108 cm. long. The
sweep circuit could not be synchronized with the discharge of the organs.
Consequently traces of a number of discharges occurring during successive
horizontal sweeps of the electron beam are shown overlapping in each figure.

These two figures show the voltage-time relation for each segment, but
they give no indication of the time-lag between the pulses in the two seg-
ments. To determine the time-lag the fish was connected to the oscillograph
in a third way. The sweep circuit was disconnected from the oscillograph
tube, the anterior segment 1-2 was connected to cause a vertical deflection
of the electron beam, and the posterior segment 3-4 was connected to cause
a horizontal deflection. The trace obtained with this connection is shown in
Plate I, Fig. 3. Because the voltage rises in the anterior segment before it
begins to rise in the posterior segment, the initial deflection is vertical.
This shows as the vertical side at the right of the loop. The right-angled bend
of the trace at its upper left corner indicates that the voltage in the anterior
segment has ceased to rise but has not started to fall at the instant at which
the voltage begins to rise in the posterior segment. After this, the voltage
starts to fall in the anterior segment while remaining near its peak in the
posterior segment. Finally it falls in both together and the loop is closed.

If the time-lag between the discharges in the two segments were known,
it would be possible to construct the trace shown in Fig. 3 from measure-
ments on the two traces shown in Figs. 1 and 2. For if the time-lag were
known, corresponding instants on the time-axes of the two traces could be
paired, and voltages simultaneously existing in the two segments could be
determined. A graph showing the voltage in the anterior segment plotted
vertically against the simultaneous voltage in the posterior segment plotted
horizontally would be similar to the loop of Fig. 3, which was obtained with
the oscillograph when the two segments were connected respectively to the
vertically and horizontally deflecting plates. Conversely, by assuming various

1940]

Coates, Cox, Rosenblith & Brown: Electric Eel

251

values for the time-lag and plotting the corresponding values of the two
voltages one against the other, there will be obtained a number of graphs
of which the one most like the recorded loop will have its assumed time-lag
nearest the actual time-lag.

This procedure is illustrated by Text-fig. 2. The oscillographic traces
copied in this figure were obtained with an eel 124 cm. long. The traces of
the discharge in the anterior and posterior segments are superimposed with
an assumed time-lag of 0.6 millisec. so as to give a loop analogous to the
one which was recorded. Points A, B, and C, corresponding to three instants
during the discharge, serve to illustrate the construction. A shows the
voltage in the anterior segment (plotted vertically) at the start of the
discharge in the posterior segment. B shows the voltage in the anterior
segment at the instant at which it is equal to that in the posterior segment
(plotted horizontally). C shows the voltage in the posterior segment at the
end of the discharge in the anterior segment.

Text-figure 2.

Construction of graph of anterior voltage vs. posterior voltage by super-
position of graphs of anterior voltage vs. time and posterior voltage vs. time.
Assumed time-lag, 0.6 millisecs. (The scale of voltage is only I'oughly approxi-
mate) .

Text-fig. 3 shows a more extended example of the same construction.
The traces copied here were obtained with an eel 167 cm. long. The graphs
(a) — (i) have been constructed as in Text-fig. 2 with various assumed
time-lags between the starts of the discharge in the anterior and posterior
segments. Graph (e) in the second row is seen to be very similar to the
loop drawn next to it, which is a copy of the photographically recorded loop.

The time-lag between the starts of the rising voltage in the two seg-
ments is thus found to be 0.6 millisec. Now the instant at which the voltage
between the electrodes 1 and 2 starts to rise is the instant at which the front
of the pulse of potential gradient starts down the organ from electrode 1
at the anterior end. Similarly the instant at which the voltage starts to
rise between electrodes 3 and 4 is the instant at which the front of the pulse
reaches electrode 3, for as soon as the front of the pulse passes a given point
that point will be at a different potential from points farther along the
organ. Hence the time-lag found, 0.6 millisec., is the time required for the
front of the pulse to traverse the distance from electrode 1 to electrode 3.
This distance was 60 cm. Dividing the distance by the time-lag, we find the
average speed of tfie pulse along the anterior 60 cm. of the large organ to be
100 cm. per millisec. or 1,000 meters per sec.

With the same fish oscillographic traces were photographed with elec-

252

Zoologica: New York Zoological Society

[XXV :14

Text-figure 3.

Construction of graph of anterior voltage vs. posterior voltage with various
assumed time-lags.

Graph a b cd efghi

Assumed time-lag,

(milliseconds.) 06 0.1 0.3 0.5 0.6 0.7 0.9 1.2 1.8

Computed speed of pulse,

(meters per second.) 10,000 6,000 2,000 1,200 1,000 860 670 500 330

The photographically-recorded trace (P) is shown for comparison at (e).

1940]

Coates, Cox, Rosenblith & Brown: Electric Eel

253

trodes 1 and 2 in the same positions as in Text-fig. 1 but with electrodes 3
and 4 at 40 cm. and 60 cm. respectively from the anterior end. The time-
lag between the starts of the rising voltage in the anterior and posterior
segments was again determined by the procedure just described and was
found to be only 0.15 millisec., approximately. From this result the average
speed of the pulse along the anterior 40 cm. of the organ is found to be
about 2,500 meters per sec.

The pulse takes 0.6 millisec. to travel along the organ 60 cm. from the
anterior end and 0.15 millisec. to travel 40 cm. from the anterior end. The
difference of these two time-lags, 0.45 millisec., is the time required for the
pulse to traverse the segment of the organ between 40 cm. and 60 cm. from
the anterior end. The average speed of the pulse along this 20 cm. stretch is
thus only 450 meters per sec.

None of these determinations can be taken as at all precise. Neverthe-
less it is clear that the speed of propagation of the pulse along the organ
diminishes from anterior to posterior. The speed at the anterior end of the
organ must then be greater than the average speed along the anterior 40
cm., which was reckoned as 2,500 meters per sec. Even with allowance made
for the inaccuracy of this measurement, it seems safe to take 2,500 meters
per sec. as a minimum estimate for the speed at the anterior end. The
large electric organ of this fish was at least 120 cm. long, and the average
speed of the pulse over the segment between 40 cm. and 60 cm. from the
anterior end was found to be 450 meters per sec. Considering the fall in
speed toward the posterior end it seems safe, after allowing for the
inaccuracy of the measurement, to take 450 meters per sec. as a maximum
estimate of the speed at the posterior end. The ratio of the speed at the
anterior end to that at the posterior end must then be greater than 5 to 1.

Measurements were made in all on three fish of different lengths. The
results are summarized in the following table.

Table 1.

Propagation of pulse along large organs of three eels of different sizes.

Eel

Length

(cm.)

Circumfer-
ence at
anterior end
of large
organ (cm.)

Distance
from anterior
end to elec-
trode 3*
(cm.)

Time-lag

(millisec)

Speed
(meters
per sec.)

I

108

20.5

40

Between
0.25 and 0.3

About

1500

II

167

34

40

0.15

2500+

60

0.6

1000

III

124

25

20

0.1—

2000 +

40

0.45

900

* Electrode 1 was always at the anterior end of the large organ. Hence this distance is the
distance traversed by the pulse during the measured time-lag. The speed is computed from the
distance and the time-lag.

In the electric eel the electric organs lie parallel to the spinal cord. From
special large cells, the electric nerve cells, lying in the cord, nerves branch
out to the adjacent parts of the organs. The discharge in every part of the
organ is excited by the nerves running to that part. For if the cord is cut at
any point, parts of the organ posterior to the cut do not discharge. On the
other hand, if the organ is transsected at any point without injury to the
cord, the segment of the organ posterior to the transsection still discharges.

254 Zoologica: New York Zoological Society [XXV:14

The path traversed by the nervous impulse which initiates the discharge
at any point is certainly no shorter than the distance to that point along the
body of the eel. It seems necessary then to conclude either that the nervous
impulse travels along the cord as fast as we have found the electric pulse
traveling along the organ or else that there is some special mechanism which
delays the discharge in the anterior part of the organ while the nervous
impulse runs down the cord to initiate the discharge in posterior parts.

If we take the first of these hypotheses and assume a speed of the
nervous impulse along the fibers in the cord as high as 2,500 meters per
sec., and if we apply the rule given by Blair and Erlanger3 that the speed
is proportional to the square of the diameter of the fiber, then we find that
the fiber must have a diameter of about 140 microns or .014 cm. We have not
found a record of the size of the nerve fibers in the spinal cord of the electric
eel, but a diameter of 140 microns is several times greater than that of the
largest ordinary nerve fibers. If, instead of following the rule of Erlanger
and Gasser, we adopt the conclusion of other investigators that the speed is
proportional to a power of the diameter between the first and second, then
still larger nerve fibers must be assumed for the electric eel.

The evidence which we have on the second hypothesis, that of a special
process delaying the discharge in the anterior part of the organ, is as yet
slight and inconclusive.

It is at any rate easy to see how the effective use of the organ requires
the pulse to travel along it at a much higher speed than that at which nervous
impulses are commonly transmitted. The point is most readily discussed in
terms of the description of the organ as a series of galvanic elements effec-
tively insulated from each other except when, during the discharge, a
transient drop in the electrical resistance between adjacent elements joins
them briefly in series4. This series connection persists only for about 1.5
millisec. at any point in the organ. The front of the pulse at any instant is
the point on the organ at which the connection is just beginning to be made.
If the front travelled down the organ at a speed of 100 meters per second,
or 10 cm. per millisec. (which is near the highest speed recorded even in
mammalian nerve) the length of the segment connected at any one time
would be only 15 cm. As soon as the front had travelled that far along the
organ, the connection would be broken behind it as fast at it was made in
front. The highest voltage developed would thus be only a fraction of what
is developed with the actual speed. The discharge would, it is true, be pro-
longed by the slower propagation, but this advantage could hardly offset the
disadvantage of the lower voltage in repelling the enemies of the fish and
stunning its prey, especially since the fish can now repeat its discharge with
such rapidity as to have much the same effect as prolonging it, perhaps with
a greater effect physiologically.

It thus appears that the evolution of the electric eel has developed a
mechanism for the propagation of the pulse along the organ which circum-
vents whatever factor limits in other animals the speed of the impulse along
nerve fibers, and that without this mechanism — whatever it is — the electric
power of the eel would be much less effective. It also appears, however, that
this mechanism is itself subject to some limitation, for it does not make the
discharge exactly simultaneous at every point of the organ. Because of the
time lag, the peak voltage between the extremities of the organ is less than
the sum of the peak voltages developed in all its segments. Measurements on
an electric eel about 170 cm. long showed that the voltage developed in the
anterior four-fifths of the large organ is practically as great as that
developed between its extremities. This indicates that the posterior fifth of

3 Blair, E. A., and Erlanger, J., Am. J. Physiol., 106, 524 (1933).

4 Cox, R. T., and Coates, C. W. Electrical Characteristics of the Electric Tissue of the Electric
Eel, Electrophorus electricus (Linnaeus). Zoologica , New York, 1938. Vol. 23, Part 2, pp. 203-212.

1940]

Coates, Cox, Rosenblith & Brown: Electric Eel

255

the organ is not fully joined in series until the connection at the anterior end
has begun to be broken. Consequently in this eel the posterior fifth of the
organ has little effect other than to prolong the discharge.

Our observations afford only a narrow basis for the comparison of the
speeds of the pulse in eels of different lengths. However it seems that there
is probably an upper limit to the speed attainable and that there is conse-
quently an increase, as the eel grows, in the time lag between the discharges
at the extremities of the organ. It seems therefore a quite reasonable
inference that, because of this factor, growth of the eel beyond a certain
length would increase the need of prey faster than the additional effective-
ness of the discharge would help in its capture. What this length would be
we can not say from our observations. The longest electric eel that the New
York Aquarium has had was about 2.7 meters long, and this is about the
greatest length recorded. It would seem unlikely that growth beyond this
point would be helpful to the eel.

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Zoologica: New York Zoological Society

EXPLANATION OF THE PLATE.

Plate I.

Oscillographic traces of the discharge of an electric eel 108 cm. long.

Fig. 1. Voltage-time graph of anterior segment. Electrodes at anterior end of
Large Organ and 10 cm. behind. Length of horizontal base corresponds
to 4 milliseconds. The peak is about 100 volts.

Fig. 2. Voltage-time graph of posterior segment. Electrodes 40 cm. and 70 cm.
from anterior end of Large Organ. Scale same as in Fig. 1. (The
discharge of lower voltage is of the intermediate type).

Fig. 3. Graph of voltage in anterior segment (vertically) against voltage in
posterior segment (horizontally). (The short horizontal trace is made by
the discharge of the Bundles of Sachs.)

COATES, COX, ROSENBLITH & BROWN.

PLATE I.

FIG. 3.

PROPAGATION OF THE ELECTRIC IMPULSE ALONG THE ORGANS OF THE
ELECTRIC EEL, ELECTROPHORUS ELECTR1CUS (LINNAEUS).

Crandall: Bird of Paradise Display

257

15.

Notes on the Display Forms of Wahne’s Six-plumed
Bird of Paradise.

Lee S. Crandall

Curator of Birds, New York Zoological Park
(Text-figures 1-3).

The display forms of the short-tailed members of the genus Parotia,
as typified by those of the Greater Six-plumed Bird of Paradise ( P . sefilata)
and Lawes’ Six-plumed (P. 1. lawesi), are now well known. But those of
Wahne’s Six-plumed (P. wahnesi ) have caused much conjecture, because
of the remarkably long tail. It was to be supposed that some variation from
the usual forms would be found.

Wahne’s Six-plumed Bird of Paradise was unknown in captivity until
1931, when a single male, brought from northeastern New Guinea by Mr.
F. Shaw-Mayer, was secured by the Zoological Society of London. No further
importations were made until 1939, when Mr. Mayer arrived in London with
another shipment. Through the kindness of M. Jean Delacour, we obtained
an adult male from this lot. The bird arrived in New York on August 17,
1939, and about October 20 began to molt. By the end of February, 1940,
the change had been completed, the four months’ molting period being the
average for adult males of this genus.

Once the molt was complete, the bird became very active, in marked
contrast to the usually phlegmatic conduct of male Six-plumes of other
species. Displays were soon observed, and are recorded in the following notes.

When about to display on the ground, the bird stands with its body in a
horizontal position, with the wings held closed but high, in order to clear
the slightly loosened flank feathers. The tail is turned sideways, usually to
the left (Text-fig. 1). The bird feigns picking at the ground, then suddenly
throws the body forward and downward with the head turned under the
breast, so that its crown is nearly parallel with the ground (Text-fix. 2). The
tail now becomes the center of attention, for it is thrown straight upward,
behind the crouching body. While all else remains immobile, the graduated
lateral feathers are now rapidly opened and closed, the central pair remain-
ing fixed. After this fan-like effect has been obtained perhaps five or six
times, the bird suddenly drops the tail and throws its body into an upright
position. Simultaneously, the long feathers of the back and flanks are raised
to form the “umbrella” well known in other forms of Parotia, and the head
plumes are thrown far forward, three on each side of the crown (Text-fig. 3).
The brilliant breast plate is noticeably flat and lies between a slight extension
of the feathers of the upper neck and the erected flank and breast plumes.
With the head now extended to its greatest possible height, the bird turns
it rapidly from side to side, causing the tabs on the plumes to rotate in the
typical manner. Several mincing, short steps are now taken along the

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[XXV :15

1940]

Crandall: Bird of Paradise Display

259

Text-figure 3.

Parotia wahnesi. Final position of the display, climaxed by “wobbling” of the
neck.

ground, usually to the right. After moving perhaps a foot in this manner,
the bird suddenly becomes immobile except for the neck, which is rapidly
moved from side to side, between the head and breast plate. The display
usually ends with this maneuver. During the upright form of the display,
the tail takes a negligible part and may turn to either side or even drag on the
ground.

In preparing to display off the ground, the bird moves about the perches,
its body stiffly horizontal and the tail held to the side. When the chosen
spot is reached, it throws itself directly into the upright form, the tail-
fanning being omitted. The following display is then identical with that
performed on the ground, including movement along the perch and
“wobbling” of the neck.

While in the series of papers published by the New York Zoological
Society on the display forms of Birds of Paradise, no attempt at correlation
has yet been made, one point in relation to Wahne’s Six-plumed is so striking
that it should not be overlooked. The long tail of this species is very sugges-
tive of that of Astrapia rothschildi. When it is recalled that the latter
bird also “fans” the tail in display1, and that the distribution of the two
species is generally identical and restricted to a small area in northeastern
New Guinea (mountains of the Huon Peninsula), the desirability of further
investigation in the field is suggested.

1 Zoologica, Vol. XI, No. 7, p. 81, 1932.

Goss: Hemorrhagic Gastro-enteritis in a Giant Panda

261

16.

Acute Hemorrhagic Gastro-enteritis in a Giant Panda.
Leonard J. Goss

Veterinarian, New York Zoological Park

The male giant panda, Ailuropoda melanoleuca (Daudin), in the collec-
tion of the New York Zoological Park, was found dead in its cage on the
morning of May 6. In view of the comparative rarity of the animal in
captivity and the paucity of records as to its pathology, we present here
an account of its diet and the findings on autopsy.

The specimen was reported to be 14 months old when it was received
on May 1, 1939, from China through the agency of Mr. Dean Sage, Jr. Its
weight on arrival was 72 pounds. Its age when captured could not be
learned, but it had been in captivity in China for some months, apparently
confined by a chain or rope around its neck, for the hair was worn off in this
area in a wide band, and the coat was generally in a dry, unhealthy condi-
tion. The type and amount of food supplied the animal in China and en
route to New York could not be learned.

For eleven months previous to the arrival of the male, the female giant
panda in our collection (8 months old on arrival) had been fed the following
ration devised by Dr. C. R. Schroeder: Pablum, dried milk, raw eggs, raw
green vegetables (carrots, lettuce, celery), oranges, codliver oil, brewers’
yeast, bone meal and salt. The male specimen was put on this diet and
readily consumed it.

During the twelve months the male was in the Zoological Park it made
an average monthly weight increase of 8.1 pounds. However, 23 pounds
were gained during the first 16 days and only 20 pounds were gained the
last 4 months. This is in contrast to an average monthly gain of 9.5 pounds
by our other panda, the female, for the 24 months she has been in captivity.
Her average gain for the past 6 months is exactly the same as her two-year
average, 9.5 pounds per month.

From time to time during the year that the male was in our collection,
bamboo, sugar cane and cornstalks were introduced into the diet and both
the male and the female consumed this stalky material with evident enjoy-
ment. Eventually the source of supply became less dependable and the
stalks were discontinued.

It soon became apparent that the male giant panda was not making
satisfactory progress and the diet was suspected of being faulty. At inter-
vals of about two months it ceased to feed for several days at a time and
these periods were accompanied by frequent belching-like actions. Accom-
panying these attacks, quantities of yellow, tenacious, rubbery mucus
streaked with blood were passed from the rectum. After each of these
attacks the animal’s appetite was impaired to such an extent that it would
eat only a portion of some of the usual items in its diet — food that it Rad
taken willingly before. As these digestive disturbances progressed, anal
flatus was a marked symptom.

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Zoologica: New York Zoological Society [XXV:16. 1940]

The stool was always very loose, being about the consistency and
appearance of corn meal mush, and a formed stool was never passed. While
this was to be expected, considering the elements of the diet, it was not felt
that this was a satisfactory condition. In view of the well-developed den-
ture of the panda, the large stomach, the consistency of the stool and the
attacks of intestinal disturbances, it was decided that the animal should
again be given roughage in the form of bamboo.

A small stalk of bamboo V2" in diameter and 8" long was offered. This
aroused such interest in an otherwise lethargic animal and the results were
so gratifying that the practice was continued. A small stalk of bamboo was
given each day. Invariably the bamboo was stripped of its outside covering
by the use of a paw and the teeth and the remaining stalk crushed and
chewed in the powerful teeth and jaws. Small portions of the bamboo were
fed daily and the results anxiously awaited. The appetite seemed improved,
at least for the bamboo; the stool became formed, and the anal flatus de-
sisted. Feeling that no untowai’d dangers were developing and that the
animal was now conditioned to the bamboo, we increased the amount. At
no time were more than three sticks 8" long fed the animal. Two and a
half weeks following the introduction of bamboo into the diet, the panda
developed a fatal enteritis. The morning before it died, it was found pros-
trated and breathing hard, the coat damp over the entire body. Close ex-
amination revealed the skin to be dry; it was assumed that the moisture
on the coat was a result of lying in the animal’s own fecal material. There
was a spot of vomitus in the cage, consisting of the previous evening’s meal,
and a normal stool. The eyes were closed, the conjunctiva congested, in
marked contrast to a very pale oral mucosa. A very watery, reddish-brown
(blood) stool was passed, accompanied by griping. A diagnosis of acute
enteritis was made. At 1 P.M. the animal appeared much brighter and was
not adverse to moving about. At 7 P.M. it was up and around, steady on its
feet and begging for food. Food was witheld and 1 pint of water was given
per orum. It was felt that the crisis had passed and recovery was probable,
but the animal was found dead at 7 :30 A.M. the following morning.

Autopsy findings were: Body weight 170 pounds, about two quarts of
clear watery transudate in the abdominal cavity. The mesenteric vessels
were tremendously engorged. The serosa of the intestines was purple red
in color, the mesenteric lymph glands swollen, hemorrhagic at the periphery
and juicy. The intestines contained several quarts of blood-colored material
containing small flakes of curd-like mucus. The gastric and intestinal
mucosa were acutely hemorrhagic in color. The intestinal wall was thick-
ened. The gastric and duodenal mucosa showed numerous small pits repre-
senting partially healed chronic ulcers. The spleen was pulpy and swollen.
All other organs appeared normal. Bacterial cultures of the liver, spleen,
and heart’s blood were negative. Cultures of the intestinal content in
tetrathionate broth and on brilliant green agar plates did not reveal any
paratyphoid organisms. Death was attributed to an acute hemorrhagic
gastro-enteritis. From the cultures made, bacterial infection is precluded.
The part the bamboo in the diet played in producing death is problematical.
It is thought that the alimentary tract was particularly susceptible to the
abrasive action of bamboo in view of the fact that the animal had been on a
soft diet up to 2V2 weeks before death. The ulcers found in the gastric and
duodenal mucosa undoubtedly were contributing causes.

Nigrelli: New Species of Trematodes

263

17.

Two New Species of Trematodes from the Deep Sea Scorpion
Fish, Scorpaena madurensis Cuv. & Val.

Ross F. Nigrelli

New York Aquarium
(Plate I; Text-figures 1 & 2).

Scorpaena madurensis (= Sebastes madurensis ) inhabits the deeper
waters off the Madeira Islands. Specimens were sent to the New York
Aquarium in an exchange with the London Aquarium. Two species of
trematode pai'asites belonging to the families Hemiuridae and Allocreadiidae
were found in the stomach and intestine respectively. Morphological studies
on these worms indicated that they were new and the names Tubulovesicula
madurensis and Podocotyle atzi were given them by Nigrelli (1939).

Tubulovesicula madurensis.

(Plate I, Fig. 2; Text-figure 1).

The following description is based on four specimens. The worms are
cylindrical, spindle shaped, and in all cases the tail portion was completely
everted. Total length 7.35 mm.; tail appendage proper 2.69 mm. Width of
worms, taken in the middle third of the body, 1.63 mm. Pre-oral lip and
cervical glands present, the former measuring .0435 mm. in length. Oral
sucker .251 X .287 mm., subterminal; pharynx contiguous to oral sucker,
measuring .161 X .194 mm.; esophagus short, .111 mm. The proximal part of
the ceca is lined with cuticula for a short distance, ceca terminating at the
posterior tip of the tail appendage. Acetabulum larger than oral sucker,
slightly forward in the anterior third of the body, measuring .643 X .659
mm. Testes ovoid in shape, ventral, post-acetabular, more or less obliquely
placed, separated from one another by uterine coils. Right testis measures
.444 X .498 mm.; left testis .413 X .532 mm. The seminal vesicle is short
and broad, tubular, antero-dorsal to the testes, post-acetabular, measuring
.465 X .165 mm. Pars prostatica long, surrounded by numerous prostate
cells, post-acetabular, measuring 1.28 mm. in length. Ductus hermaphroditi-
cus enclosed in a sinus sac and opening into the genital atrium. Sinus sac
large, shield-shaped, measuring .150 X .270 mm. Opening of the atrium at
the level of the pharynx. Ovary oval shaped, immediately behind the testes,
measuring .348 X .442 mm. Seminal receptacle measures .042 X .032 mm.
Laurer’s canal present. Shell gland immediately posterior to the ovary,
.233 mm. in diameter. Vitelline glands tubular, extending to the extra-cecal
area, seven lobes in number, three left, four right, measuring 1.50 X .046
mm. Receptaculum seminis iiterinum present. The uterus descends to the
posterior end of the body but does not enter into the tail appendage, or only
slightly so. It then winds anteriorly, mostly intercecal, to one side of the

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[XXV :17

Reproductive organs of Tubulovesicula madurensis. A. Part of the terminal
reproductive system. G.P., genital pore; S.S., sinus sac; H.V., hermaphroditic
vesicle; Met., metraterm; P.Pr., pars prostatica; P.C., prostate cells; S. V.,
seminal vesicle. B. Ovarian complex. Vo., ovary; Od., oviduct; Vd., vitelline duct;

S. R., Seminal receptacle; L. C., Laurer’s canal; Oot., ootype; Meh. G., Mehlis’
gland; R. S. U., Receptaculum seminis uterium; U., uterus.

ovary, and in between the testes. The metraterm portion of the uterus
measures .38 mm. in length. Eggs thick shelled, operculated and embryo-
nated, measuring .012-.015 X .018-.025 mm. The excretory stem bifurcates
just immediately behind the level of the testes; the arms of the branches
re-uniting at the level of the pharynx, on the dorsal side.

The genus Tubulovesicula was created by Yamaguti (1934) to include
the following hemiurids: T. spari Yamaguti (1934) from Sparus macro-
cephalus; T. anguillae Yamaguti (1934) from Anguilla japonica; and T.
muraenesocis Yamaguti (1934) from Muraenesox cinereus. The first of these
species was designated as type. This investigator included in this genus

T. angusticauda (Nicoll, 1915) (= Ectenurus angusticauda Nicoll, 1915)
also from Muraenesox cinereus. According to Yamaguti, however, this
species differs from the form that he described from the same host in the
size of the eggs, and in the posterior extent of the seminal vesicula and the
pars prostatica. Yamaguti further indicated in his paper that the form
described by Layman ( 1930 ) as Lecithaster lindbergi also should be referred
to the genus Tubulovesicida. Since then two other species have been in-
cluded in the genus. Park (1936) described T. calif ornica from the stomach
of Enophrys bison, a sculpin fish from the west coast of the United States,
and Yamaguti (1938) described T. pseudorliombi from Pseudorhombus
pentophthalmus.

On the basis of the characters described for the genus by Yamaguti
(1934) there can be no question that the hemiurids from the stomach of the
scorpion fish, Scorpaena madurensis should be included in this group. The
name T. madurensis Nigrelli (1939) was designated for this species and
appears to be closely related to T. pseudorliombi. T. madurensis differs from

1940]

Nigrelli: New Species of Trematodes

265

T. pseudorhombi in body size, size and extent of the seminal vesicle and
pars prostatica, size and shape of the sinus sac, size, shape and extent of
the vitellaria, amount of uterine coils and size of eggs.

The hemiurids comprise a large group of stomach parasites of marine
fishes. As was pointed out by Looss (1907), Stunkard & Nigrelli (1934)
and other investigators, these trematodes have powerful body muscles and
are capable of contracting their body tremendously. Very great changes in
the shape and spatial relations of the internal organs may be produced,
especially when the caudal appendage is protruded or retracted. Comparisons
between related worms should be made with care. Fortunately in this present
group most of the worms described by the various authors were those with
tail appendage fully extended and the following key could be employed to
determine the recognized species.

Key to the Recognized Species of Tubidovesicula Yamaguti, 1934.

A. Vitelline lobes 7 (3 + 4).

B. Vitelline gland with three right and four left lobes.

C. Uterine coils extending into tail for some distance.

D. Seminal vesicle long and slender; tail shorter than body proper

T. spari Yamaguti, 1934 (Type sp.).
DD. Seminal vesicle short and slender; tail longer than body proper

T. anguillae Yamaguti, 1934.

E. Seminal vesicle long and broad.

F. Pars prostatica originating at the middle of the acetabulum

T. muraenesocis Yamaguti, 1934.
FF. Pars prostatica originating posterior to the acetabulum

T. lindbergi (Layman, 1930).
EE. Seminal vesicle short and slender; pars prostatica originating at the

level of the anterior border of the acetabulum

T. angusticauda (Nicoll, 1915).
BB. Vitelline gland with four right and three left lobes.

G. Seminal vesicle long and slender; pars prostatica originating at the
level of the anterior border of the acetabulum

T. pseudorhombi Yamaguti, 1938.
GG. Seminal vesicle short and broad; pars prostatica originating some
distance posterior to the acetabulum T. madurensis Nigrelli, 1939.
A A. Vitelline lobes 8 (4 + 4) T. calif ornica Park, 1936.

Podocofyle atzi.

(Plate I, Fig. 2; Text-figure 2).

The genus Podocotyle, according to Odhner (1906), is characterized as
follows: “Body elongate, quite evenly wide, sometimes flattened and ribbon-
like. Esophagus at most twice as long as pharynx, forking in front of the
ventral sucker. Genital pore to the left, lying at the level of the esophagus.
Excretory vesicle reaching to the ovary. Seminal vesicle long, coiled; pars
prostatica lacking. Cirrus of marked length, straight. Ovary three-lobed.
Vitellaria normally not reaching anterior to ventral sucker. Eggs without
filament. In intestine of marine fish. Type species P. atomon (Rud.).”

Two worms were recovered from the intestine of Scorpaena madurensis,
both measuring 4.41 mm. in length; width .707-.851 mm. Oral sucker .251 X
.289 and .266 X .281 mm.; pharynx .205 X .167 and .190 X .152 mm.;

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[XXV :17

Reproductive organs of Podocotyle atzi. A. Part of the terminal reproductive
complex. C., cirrus; G.P. genital pore; C. P., cirrus pouch; Met., metraterm;
S.V., seminal vesicle; Va. D., vas deferens. B. Ovarian complex. Ov., ovary; Od.,
oviduct; S. R., Seminal receptacle; L. C., Laurer’s canal; V.R., vitelline reservoir;
S.G., shell gland; Oot., ootype; Vd., vitelline duct; U., uterus.

esophagus .294 X .061 mm. ; ceca extending to the level of the posterior group
of vitellaria; acetabulum, prominent, somewhat protruded, measuring .519
and .590 X .540 mm. Testes sharply oblique, anterior testis .255 X .281 mm.
and .296 X .273 mm. ; posterior testis .380 X .243 and .327 X .295 mm. Ovary
immediately in front of anterior testis, non-lobulated, measuring .152 X .190
and .203 X .160 mm. Seminal vesicle, slightly coiled, .380 X .167 mm. and
.319 X .152 mm. Cirrus long and narrow, .532 X .053 mm.; pars prostatica
lacking; genital pore at the level of cecal bifurcation, to the left. Ovarian
complex consisting of seminal receptacle, vitelline reservoir and shell gland.
Laurer’s canal present. Vitellaria mostly extra-testicular and extra-cecal,
extending anteriorly to the level of the posterior border of the acetabulum
and posteriorly to a short distance from the tip of the body. Uterine coils,
intercecal, few, limited to area between the anterior border of the ovary
and posterior border of the seminal vesicle. Eggs few, operculated, embryo-
nated, measuring .038 X .062. Excretory vesicle extending to level of the
posterior testis and arms extending slightly anterior to the level of the
bifurcation of the ceca, terminating on each side of the esophagus.

From the above description there can be no doubt that the allocreadids
from the intestine of the Madeira scorpion fish belong to the genus
Podocotyle. The present species designated as P. atzi Nigrelli (1939)

1940]

Nigrelli: New Species of Trematodes

267

differs from other recognized species in body size, size of the eggs, esophagus-
pharynx size relationships, size and relationships of the seminal vesicle,
extent of the excretory branches, predominance of the acetabulum, shape of
the ovary, and nature of the ovarian complex.

Summary.

1. Two species of trematodes from the stomach and intestine of the
Madeira scorpion fish, Scorpaena madurensis , are described.

2. The stomach parasites belong to the family Hemiuridae Liihe and
were named Tubulovesicida madurensis by Nigrelli in 1939. The intestinal
parasites belong to the family Allocreadiidae Stossich and were designated
Podocotyle atzi by the same author.

3. A key to the species of Tubulovesicida Yamaguti (1934) is given.

References.

Layman, E. M.

1930. Parasitic Worms from the Fishes of Peter the Great Bay. Bull. Pacific
Scientific Fish. Res. Stat. 3: 101-102.

Looss, A.

1907. Zur Kenntniss der Distomenfamilie Hemiuridae. Zool. Anz. 31 ;585.

Nicoll, W.

1915. The Trematode Parasites of North Queensland. III. Parasitology
8: 22-40.

Nigrelli, R. F.

1939. Two Species of Trematodes from a Deep Sea Scorpion Fish, Sebastes
madurensis. Anat. Rec. 75: 156 (abstract).

Odhner, T.

1906. Trematoden des Arktischen Gebietes. Fauna Arctica 4: 291-372.

Park, J. T.

1936. Two New Trematodes, Sterrhurus magnatestis and Tubulovesicula
calif oinica (Hemiuridae) from Littoral Fishes of Dillon’s Beach,
California. Trans. Amer. Micr. Soc. 55: 477-482.

Stunkard, H. W. & Nigrelli, R. F.

1934. Observations on the Genus Sterrhurus Looss, with a description of
Sterrhurus branchialis Sp. Nov. (Trematode, Hemiuridae). Biol. Bull.
67: 534-543.

Yamaguti, S.

1934. 7. Studies on the Helminth Fauna of Japan. Part 2. Trematodes of
Fishes, I. Jap. J. Zool. 4: 249-541.

1938. Studies on the Helminth Fauna of Japan. Part 21. Trematodes of

Fishes, IV. Kyoto, Japan. 1-139.

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Zoologica: New York Zoological Society

EXPLANATION OF THE PLATE.

Photographs made by direct projection of specimens on bromide
Worms stained with Delafield’s hematoxylin.

Plate I.

Fig. 1. Tubalovesicula madurensis. X 22.7.

Fig. 2. Podocotyle atzi. X 29.

paper.

NIGRELLI.

PLATE I.

FIG. I. FIG. 2.

TWO NEW SPECIES OF TREMATODES FROM THE DEEP SEA
SCORPION FISH, SCORPAENA MADURENSIS CUV. a VAL.

Goss: Mortality Statistics, 1939

269

18.

Report of the Hospital and Laboratory of the New York Zoological
Park, 1939. Mortality Statistics of the Society’s Collection.

Leonard J. Goss

Veterinarian.

During the year 1939 autopsies were performed in the Hospital and
Laboratory of the New York Zoological Park on 135 reptiles, 221 birds and
109 mammals which died in the Society’s collection.

These figures do not represent the total number of deaths in the collec-
tion; they are merely the totals of specimens in each group which were
subjected to post-mortem examination. Certain of the rarer mammals and
birds, in particular, were deposited with the Department of Comparative
Anatomy of the American Museum of Natural History without autopsy. Out
of the 800 reptiles which died during the year, only a comparatively small
number were examined. In the great majority of cases autolysis was
advanced before the specimens reached the Laboratory.

As in the report of the Hospital and Laboratory for 1938, the titles of
the International Classification of Causes of Death (human) have been
employed, with such modification as was necessary to adapt them to the
pathology of animals. Until a standard and generally accepted classification
of the diseases of animals is compiled, a modification of the human disease
list can bring at least a measure of uniformity into the reporting of animal
mortality.

Joint or contributory causes of death have been ignored in the statistical
summaries that follow, for the sake of brevity and clarity, and only the chief
pathological changes found at autopsy have been reported. More detailed
information is, of course, available on the autopsy records of the Hospital
and Laboratory. These contain a description of both gross and microscopic
tissue changes as well as the bacteriology and parasitology encountered at
autopsy.

Grouping of specimens by age has been dropped except in the case of
mammals where it was practicable to make distinctions between immature,
adult and post-adult individuals. All birds and reptiles which came to autopsy
were adult.

It should be stressed, in connection with Table V, that the specimens
considered as “in the collection’’ during 1939 were, in some instances, of
very brief tenure, and in many cases were never placed on exhibition. In
some zoological collections it is the practice not to consider as “in the collec-
tion,” for purposes of mortality statistics, specimens which failed to survive
a minimum period which may range from thirty days to six months. The
longer the probationary period, the lower the mortality percentage for the
entire collection is likely to be. Table V includes some animals which arrived
at the Zoological Park in a moribund state and failed to survive even their
first day “in the collection.”

270

Zoologica: New York Zoological Society

Table I.

[XXV :18

REPTILES.

Infectious & Parasitic Diseases

Acid-fast Infection

Igiiana i. rhinolopha — Central American Iguana

Python m. molurus — Black-tailed Python

Agkistrodon piscivorus — Moccasin

Intestinal Cestodes

Farancia a. abacura — Red-bellied Snake

Ptyas mucosus — Indian Rat Snake

Agkistrodon blomhoffi — Blomhoff’s Pit Viper

Intestinal Helminths

Eunectes mwrinus — Anaconda

Trimeresurus flavomaculatus — Indian Green Tree Viper

nj m

•

.3 "-a 15
c ft’S
» S p,

P-t H M

10

5

7.40

3.70

2.22

1.48

Nutritional Diseases

Cachexia

Iguana i. rhinolopha — Central American Iguana

Farancia a. abacura — Red-bellied Snake

Vipera russelii — Russell’s Viper

Diseases of Circulatory System

Endocarditis with Ruptured Myocardium

Lampropeltis getulus floridana — Florida King Snake . .

Pericarditis

Python m. molurus — Black-tailed Python

Filariasis with Ruptured Myocardium

Lampropeltis getulus subsp. — Texas King Snake

2

1

1

1

1

1

Diseases of Respiratory System

Pneumonia

Zonurus giganteus — Spike-tailed Lizard

Constrictor c. constrictor — South American Boa

Python curtus — Blood Python

Python reticulatus — Regal Python

Arizona e. elegans — Smooth-scaled Gopher Snake

Boiga dendrophila — Malay Mangrove Snake

Drymarchon corais couperi — Texas Gopher Snake

Elaphe q. quadrivittata — Chicken Snake

Pituophis s. sayi — Prairie Bull Snake

Ptyas mucosus — Indian Rat Snake

Dendroaspis angusticeps — Black Mamba

Dendroaspis viridis — Green Mamba

Naja naja — Indian Cobra

Naja nivea — Yellow Cobra

Trimeresurus flavomaculatus — Indian Green Tree Viper

Cerastes cerastes — Desert Asp

Agkistrodon mokasen cupreus — Copperhead

Agkistrodon piscivorus — Moccasin

Crotalus a. atrox — Texas Diamond Rattlesnake

Crotalus v. viridis — Prairie Rattlesnake

Crotalus horridus atricaudatus — Canebrake Rattlesnake
Crotalus h. horridus — Banded Rattlesnake

1

2

1

1

1

2

1

2

1

1

1

1

2

1

1

1

1

5

5

1

1

1

Diseases of Digestive System

Hepatitis

Iguana i. rhinolopha — Central American Iguana 1

Python reticulatus — Regal Python 1

Dendroaspis viridis — Green Mamba 1

4 2.96

4 2.96

I

3 2.22

1 .74

1 .74

1 .74

34 25.18

34 25.18

I

63 146.66

3 | 2.22

1940]

Goss: Mortality Statistics, 1939

Table I. (Continued)

271

REPTILES.

Necrotic Stomatitis

Heloderma horridum — Mexican Beaded Lizard

Boa canina — Green Tree Boa

Python curtus — Blood Python

Python regius — Ball Python

Naja naja — Indian Cobra

Vipera russelii — Russell’s Viper

Enteritis

Varanus sp. — Monitor Lizard

Elaphe q. quadrivittata— Chicken Snake

Lampropeltis getulus splendida — Mexican King Snake . .

Pituophis m. melanoleucus — Pine Snake

Pituophis s. sayi — Prairie Bull Snake

Cerastes cerastes — Desert Asp

Agkistrodon mokasen cupreus — Copperhead

Crotalus a. atrox — Texas Diamond Rattlesnake

Crotalus h. horridus — Banded Rattlesnake

Necrotic Enteritis

Ctenosaura acanthura — Black Iguana

Heloderma suspectum — Gila Monster

Tupinambis teguixin — Tegu Lizard

Boa canina — Green Tree Boa

Constrictor c. constrictor — South American Boa

Epicrates cenchris — Ringed Boa

Eunectes murinus — Anaconda

Python curtus — Blood Python

Python m. molurus — Black-tailed Python

Python regius — Ball Python

Python reticulatus — Regal Python

Boiga dendrophila — Malay Mangrove Snake

Drymarchon corais couperi — Texas Gopher Snake

Lampropeltis g. getulus — Common King Snake

Dendroaspis viridis — Green Mamba

Naja hannah — King Cobra

Naja naja — Indian Cobra

Bitis gabonica — Gaboon Viper

Vipera russelii — Russell’s Viper

Agkistrodon piscivorus — Moccasin

Crotalus a. atrox — Texas Diamond Rattlesnake

Crotalus cerastes — Horned Rattlesnake

Crotalus h. horridus — Banded Rattlesnake

Mil-;

XT 'O

£ .£3 <b
c a "5
• « a

|s|

4.44

13

9.62

41 30.37

1

1

1

9 I
3

2

1

1

3

1

1

1

1

1

1

1

1

1

1

1

5

1

2

Diseases of Skin & Cellular Tissue

Abscess

Drymarchon c. corais — Cribo Rat Snake

Agkistrodon piscivorus — Moccasin

Dermatitis

Python regius — Ball Python

Agkistrodon piscivorus — Moccasin ....

I 5 | 3.70

I 2 1.48

1 I

1 I

I 3 2.22

2 I

1 I

Undetermined Diagnosis

16

11.85

272

Zoologica: New York Zoological Society
Table II.

[XXV :18

BIRDS.

Infectious & Parasitic Diseases

Salmonella Infection

Coliuspasser a. albonotatus — White-shouldered Whydah

Tangara arthus — Yellow-throated Calliste

Tuberculosis

Anas crecca carolinensis — American Green-winged Teal

Meleagris gallopavo silvestris — Eastern Turkey

Porphyrio poliocephalus poliocephalus — Gray-headed

Gallinule

Streptopelia d. decaocto — Barbary Turtle Dove

Melozone leucotis — White-eared Ground Sparrow

Aspergillosis

Guttera e. edouardi — Black-breasted Crested Guineafowl

Amazona ventralis — Santo Domingo Amazon

Chlorophonia f. frontalis — Yellow-fronted Green Tana-

ger

Liver Flukes

Ardea goliath — Goliath Heron

Florida caerulea — Little Blue Heron

Coccidiosis

Astrapia s. stephaniae- — Princess Stephanie’s Bird of

Paradise

Tangara heinei — Black-capped Calliste

Intestinal Obstruction — Nematodes

Columba livia — Swallow Pigeon

Taeniasis

Ostinops viridis — Great Green Cacique

Tangara cyaneicollis hannahiae — Blue-throated Calliste

Tangara heinei — Black-capped Calliste

Turdus m. merxda — European Blackbird

Nutritional Diseases

Slipped Tendon Disease (Manganese Deficiency)

Casmerodius albus egretta — American Egret

Malnutrition .

Tyto alba pratincola — American Barn Owl

Diseases of Circulatory System

Atheromatosis

Amazona ochrocephala oratrix — Yellow-headed Amazon

Bubo v. virginianus — Horned Owl

Anthracoceros m. malabaricus — Indian Pied Hornbill..

Bycanistes cristatus — Silver-cheeked Hornbill

Rhyticeros subruficollis — Blyth’s Wreath-billed Hornbill
Ramphastos piscivorus brevicarinatus — Short-billed
Toucan

Diseases of Respiratory System

Ulcerative Laryngitis

Ajaia ajaja — Roseate Spoonbill

Sagittarius serpentarius — Secretary Bird

Pucrasia m. macrolopha — Common Koklass Pheasant. . .
Amazona b. barbadensis — Yellow-shouldered Amazon..
Amazona ochrocephala panamensis — Panama Amazon . .
Brachypternus benghalensis benghalensis — Northern
Golden-backed Woodpecker I

Specimens.

Total specimens
autopsied.

Percentage of
135 reptiles
autopsied.

24

10.85

2

.90

1

1

10

4.52

1

1

1

6

1

3

1.35

1

1

1

2

.90

1

1

2

.90

1

1

1

.45

1

4

1.80

1

1

1

1

2

.90

1

.45

1

1

.45

1

6

2.71

6

2.71

1

1

1

1

1

i

1

23 1

10.40

6 I

2.71

1

1

1

1

1

1

i

1

i

1

I

1940]

Goss: Mortality Statistics, 1939
Table II. (Continued)

273

BIRDS.

Tracheitis

Belonopterus chilensis lamprenotus — Argentine Lapwing

Pulmonary Congestion

Agelaius ruficapillus frontalis — Chestnut-fronted Black-
bird

Pneumonia

Tragopan caboti — Cabot Tragopan

Ptilinopus jambu — Jambu Fruit Pigeon

Crinifer piscator — Brown-headed Touraco

Agapornis sp. — Masked Love Bird

Anodorhynchus leari — Lear’s Macaw

Brotogeris j. jugularis — Tovi Parrakeet

Tyto alba pratincola — American Barn Owl

Anthracothorax v. violicauda — Black-throated Mango

Hummingbird

Chlorostilbon caribaeus caribaeus — Caribbean Emerald

Hummingbird

Pharomachrus m. mocinno — Quetzal

Amblyrhamphus holosericeus — Orange-headed Blackbird
Carpodacus mexicanus frontalis — Western House Finch

Tangara m. mexicana — Yellow-bellied Calliste

Parotia l. lawesi — Lawes’ Six-plumed Bird of Paradise.
Uraeginthus b. bengalus — Cordon Bleu

Diseases of Digestive System

Impacted Crop

Gallus g. gallus — Red Jungle Fowl

Gizzard Impaction

Plectropterus gambensis niger — Black Spur-winged

Goose

Buteo jamaicensis borealis — Eastern Red-tailed Hawk . .

Circus cyaneus hudsonius — Marsh Hawk

JJrobrachya a. axillaris — Orange-shouldered Whydah..

Enteritis

Casmerodius albus egretta — American Egret

Ciconia c. ciconia— European White Stork

Cygnus olor — Mute (White) Swan

Crossoptilon mantchuricum — Brown Eared Pheasant. . .

Amazona f. f estiva — Festive Amazon

Psittacus e. erithacus — Gray Parrot

Larus argentatus smithsonianus — Herring Gull

Chlorophonia f. frontalis — Yellow-fronted Green Tana-

ger

Paradisaea gulielmi — Emperor of Germany’s Bird of

Paradise

Tangara cyanoptera — Blue-winged Calliste

Tangara flav a cayana — Black-faced Calliste

Tangara larvata centralis — Mrs. Wilson’s Calliste

Tanagra v. violacea — Violet Euphonia

Cirrhosis of Liver

Lophura rufa — Malayan Crested Fireback Pheasant. . . .

Hepatitis

Rhynochetos jubatus — Kagu

Bycanistes sharpii duboisi — Congo White-tailed Hornbill

Ramphastos swainsonii — Swainson’s Toucan

Euplectes f. franciscana — Orange Weaver

274

Zoologica: New York Zoological Society
Table II. (Continued)

[XXV: 18

BIRDS.

Mania a. atricapilla — Black-headed Mannikin

Notiopsar curaeus — Chilean Blackbird.

Paroaria nigrigenis — Black-cheeked Cardinal ....
Rubigula dispar — Sumatran Ruby-throated Bulbul

Spinus t. tristis — American Gold Finch

Thraupis c. cana — Southern Blue Tanager

Tardus o. olivaceus — Olive Thrush

Icterus ;

Aidemosyne m. modesta — Cherry Finch

Specimens.

Total specimens
autopsied.

Percentage of
135 reptiles
autopsied.

i

i

i

I

1 | .45

1

Diseases of Genitourinary System

Nephritis •_ ;

Dissoura episcopus microscelis — African White-necked

Stork 1

Theristicus caudatus — Guiana Ibis 1

Psittacula eupatria nipalensis — Indian Ring Parrakeet. 1

Nymphicus hollandicus — Cockateel 1

Musophaga v. violacea — Senegal Violet Touraco 1

Gymnoderus foetidus — Bare-necked Cotinga 1

Kittocincla m. malabarica — Shama Thrush 2

Egg Bound

Phasianus colchicus versicolor — Versicolor Pheasant... 1

9 4.07

8 3.61

1 .45

Diseases of Skin & Cellular Tissue

Abscess

Lophortyx c. calif ornica — California Quail

Pavo cristatus var. — White Indian Peafowl

Kakatoe r. roseicapilla — Roseate Cockatoo

Pitta sordida cucullata — Green-breasted Pitta

Violent & Accidental Deaths

Trauma — Head

Argusianus argus argus — Great Argus Pheasant

Chrysolophus pictus — Golden Pheasant

Gennaeus leucomelanos melanotus — Melanotus Pheasant

Philortyx fasciatus — Banded Quail

Anthropoides paradisea — Paradise Crane

Laras atricilla — North American Laughing Gull

Larus n. novae-hollandiae— Silver Gull

Leptotila c. cassini — Cassin Dove

Streptopelia d. decaocto — Barbary Turtle Dove

Kakatoe g. galerita — Greater Sulphur-crested Cockatoo

Kakatoe r. roseicapilla — Roseate Cockatoo

Kakatoe sanguinea golf ini- — Goffin’s Cockatoo

Melopsittacus undulatus — Grass Parrakeet

Pionites m. melanocephala — Black-headed Caique

Trichoglossus haematod moluccanus — Blue Mountain

Lorikeet

Ramphastos discolorus — Green-billed Toucan

Megalaima virens marshallorum — Great Himalayan

Barbet

Auripasser luteus — Abyssinian Golden Sparrow

Tangara icterocephala — Silver-throated Calliste

Chlorophonia f. frontalis — Yellow-fronted Green Tana-
ger

Corvus b. brachyrhynchos — Eastern Crow

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

1

2

1

1

1

1

1

1

1

1

4 1.80

4 1.80

71 132.12

37 116.74

1940J

Goss: Mortality Statistics, 1939

Table II. (Continued)

275

BIRDS.

Cyanerpes cyaneus — Yellow-winged Sugar Bird...

Euplectes f. franciscana — Orange Weaver

Euplectes o. orix — Grenadier Weaver

Hemixus f. fiavala — White-throated Bulbul

Myadestes unicolor — Blue Solitaire

Padda oryzivora — Java Sparrow

Ramphocelus c. carbo — Silver-beaked Tanager

Taeniopygia c. castanotis — Zebra Finch

Tangara fiava cayana — Black-faced Calliste

Tangara g. guttata — Larger Spotted Calliste

Tanagra xanthogaster exsul — Venezuelan Rufous-capped

Euphonia

Fractured Cervical Vertebrae

Rhynchotus rufescens rufescens — Rufous Tinamou.

Fractured Tibia & Fibula

Grus canadensis tabida — Sandhill Crane

Destroyed — Fractured Humerus

Pavo cristatus — Peacock

Destroyed — Blind

Dromiceius n. novaehollandiae — Emu

Drowning

Ploceus v. vitellinus — Half-masked Weaver

Poisoning — Botulism

Aix sponsa — Wood Duck

Anas platyrhynchos — White Duck

Anas p. platyrhynchos — Mallard Duck

Anas rubripes tristis — Black Duck

Chen caerulescens — Blue Goose

Cygnus olor — Mute Swan

Dendrocygna a. autumnalis — Black-breasted Tree Duck

Duck

Diagnosis Undetermined

Specimens.

Total specimens
autopsied.

Percentage of
135 reptiles
autopsied.

2

1

1

1

1

1

1

1

2

3

1

i

.45

1

i

.45

1

i

.45

1

i

.45

1

i

.45

1

29

13.12

3

1

19

1

1

1

2

1

51

23.07

276

Zoologica: Neiv York Zoological Society
Table III.

[XXV :18

MAMMALS.

Infectious & Parasitic Diseases

Tuberculosis

Cebus capucina — White-faced Sapajou

Cercocebus torquatus — Red-headed Man-

gabey

Macaca bus — Java Macaque

Macaca mulatto ■ — Rhesus Monkey

Infectious Feline Enteritis

Acinonyx jubatus — Cheetah

Felis couguar — Puma

Felis pardalis — Ocelot

Aeanthocephaliasis

Solenodon paradoxus — Solenodon

Ascaridiasis

Didelphis virginiana — Opossum

Trichinosis

Taxidea taxus taxus — American Badger* . . .

Cancer And Other Tumors

Carcinoma Esophagus

Tragelaphus angasi — Nyala

Liver Cell Carcinoma

Odocoileus osceola — Florida White-tailed

Deer

Osteogenic Sarcoma

Cervus axis — Axis Deer

Nutritional Diseases

Malnutrition

Cebus capucina — White-faced Sapajou....
Cercopithecus mona cambelli — Mona Monkey
Cercopithecus sabaeus — Green Monkey....

Hapale pygmaea — Pygmy Marmoset

Cervus canadensis canadensis — Elk

Hemitragus jemlahicus — Tahr

Diseases of Blood & Blood Producing Organs.

Anemia

Nasua nasua — White-nosed Coati

Diseases of Nervous System

Encephalitis

Ateles cucullatus — Spider Monkey

Macaca nemestrina — Pigtail Monkey

Papio anubis — Olive Baboon

Destroyed — Facial Paralysis

Cebus capucina — White-faced Sapajou

Destroyed — Posterior Paralysis

Sarcophilus harrissii — Tasmanian Devil

Destroyed — Blindness

Acinonyx jubatus — Cheetah

Diseases of Circulatory System

Vegetative Endocarditis

Didelphis virginiana — Opossum

Mephitis nigra — Skunk

Immature.

Adult.

Post-adult.

Total specimens
autopsied.

Percentage of
109 mammals
autopsied.

3

6

1

10

9.17

3

1

4

3.66

1

1

1

1

3

3

2.75

1

1

1

1

1

.91

1

1

1

.91

1

I

1

1

.91

1

1

2

1

3

2.75

1

1

.91

1

1

1

1

.91

1

1

1

1

1

.91

1

4

2

6

5.50

4

2

6

5.50

1

1

1

1

1

1

1

1

.91

1

1

1

8

8

7.33

1

4

4

3.66

1

1

1

|

2

1

1

1

.91

1

1

.91

1

2

2

1.83

2

1

1

2

| 1.83

1

1

2

1.83

1

1

1940] Goss: Mortality Statistics, 1939 277

Table III. (Continued)

o3

u

3

CO

C

0)

s

a ^

<+-( r/t
O

? =

5 g^

MAMMALS.

-£>

o3

4-S

c i

-1 ft

C 03 CO

V p ft

s

g

P

-P

4->

§

$s

o p

o c o
P o "S

►H

C

H o3

d hS Cd

Diseases of Respiratory System

9

11

1

21

18.34

Suppurative Sinusitis

1

1

.91

Lutra canadensis vaga — Florida Otter

1

Pulmonary Congestion & Edema

8

2

1

11

10.09

Macaca irus — Java Macaque

1

Macaca mulatto — Rhesus Monkey

1

Erithizon d. dorsatum — Canadian Porcupine

1

Lutra canadeyisis vaga — Florida Otter

2

1

Choeropsis liberiensis — Pygmy Hippopota-

mus

1

Cervus eldi — Eld’s Deer

1

Amnotragus lervia — Aoudad

2

Saiga tatarica — Saiga Antelope

1

Bronchopneumonia

1

7

8

7.33

Callithrix penicillata — Marmoset

1

Ateles cucullatus — Spider Monkey

1

Lagothrix humboldtii — Woolly Monkey....

1

Cercopithecus mona cambelli — Mona Monkey

1

Zalophus calif ornianus — Sea Lion

1

Euphractus sexcinctus — Six-banded Arma-

dillo

1

Procavia capensis — Hyrax

1

Dama dama — Fallow Deer

1

Gangrenous Pneumonia

1

1

.91

Arctocephalus australis — South American

Fur Seal

1

Diseases of Digestive System

3

12

2

17

15.59

Choke

1

1

.91

Suricata suricatta — Meerkat

1

Gastro-enteritis

2

8

10

9.17

Lagothrix infumatus — Woolly Monkey

3

Papio anubis — Olive Baboon

1

Didelphis virginiana — Opossum

1

Phastolomis mitchelli — Wombat

1

Macropus g. giganteus — Great Gray Kan-

garoo

1

Ursus a. arctos — European Brown Bear. . . .

1

Cervus unicolor — Indian Sambar Deer

1

Cervus hortulorum — Dybowski Deer

1

Volvulus of Intestines

1

1

.91

Odocoileus osceola — Florida White-tailed

Deer

1

Impaction of Colon with Ulceration

1

1

.91

Didelphis virginiana — Opossum

1

Actinobacillosis — Oral Tissues

2

2

1.83

Sylvicapra grimmi — Gray Duiker

2

Enteritis — Squill Poisoning

1

1

.91

Koiropotamus koiropotamus kenlae — Gray

Bush Pig

1

Icterus

1

1

.91

Lutra canadensis vaga — Florida Otter

1

Disease of Bones & Organs of Locomotion. . .

2

3

2.75

Paget’s Disease

1

1

.91

Papio anubis — Olive Baboon

1

278 Zoologica: New York Zoological Society [XXV : 18

Table III. (Continued)

73

G

<y

«4H

O ~

03

*4

3

03 T3
Qi 05

S' c

sis

MAMMALS.

cj

03 "c/5

C 03 73

05 P ft

s

3

75

$3

a

M

<

O

Cu

° 3
H is

*© p
Phh ci

Muscular Distrophy

2

2

1.83

Dendrolagus ur sinus — Tree Kangaroo

2

Congenital Malformation

1

1

.91

Congenital Anomaly — Legs

1

1

.91

Cervus axis — Axis Deer

1

Diseases of New Born

1

1

.91

Internal Hemorrhage — Umbilical Artery

1

1

.91

Lama vicugna — Vicugna

1

Senility

5

5

4.58

Destroyed — Senility

5

5

4.58

Didelphis virginiana — Opossum

1

Camelus dromedarius — Dromedary

2

Cervus duvauceli — Barasingha Deer

1

Cervus canadensis canadensis — Elk

1

Violent and Accidental Deaths

8

10

1

19

17.43

Destroyed — Trauma

8

10

1

19

17.43

Cebus capucina — White-faced Sapai'ou

1

1

Erythrocebus patas — Patas Monkey

1

Macaca irus — Java Macaque

1

Didelphis virginiana — Opossum

1

Potos fiavus — Kinkajou

1

Canis latrans — Coyote

1

Felis cacomitli — Jaguarundi Cat

1

Felis leo — Lion

1

Paradoxurus hemaphroditus — Yellow Para-

doxure

8

Dasyptis novemcinctus texanus — Nine-

banded Armadillo

1

Dama damn — Fallow Deer

2

Odocoileus osceola — Florida White-tailed

Deer

2

Amnotragus lervia — Aoudad

1

i

1

Diagnosis Undetermined

1

5

7

12

11.00

* Reported in Jour, of Parasit., April, 1940, Vol. 26, No. 2, page 157.

1940]

Goss: Mortality Statistics, 1939

279

Table IV.

Summary of Mortality Percentage in Specimens Autopsied,
by Disease Groups.

Disease

% of 135

% of 221

% of 109

Group

Reptiles

Birds

Mammals

Gastro-intestinal

46.66

14.02

15.59

Respiratory

25.18

10.40

18.34

Nutritional

2.96

.90

5.50

Infectious and Parasitic

7.40

10.85

9.17

Trauma

16.74

17.43

All Other Causes

17.80

47.09

33.97

Table V.

Summary of Mortality for All Specimens in the Collection
During 1939.

Reptiles

Birds

Mammals

Specimens in Collection During 19391. . .

1,246

2,105

621

Deaths from All Causes2

800

415

120

% of Mortality in Collection3

64,20

19.71

19.32

1 These figures represent the number of specimens that came into, or remained in, the Park
during 1939, and are the totals of specimens in the collection in the course of the year. Some died
before being placed on exhibition.

2 Includes all deaths irrespective of cause or time after arrival.

3 Based on total specimens and total deaths.

J^eto iforfe Hoologtcal Society

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Chairman, Executive Committee, Laurance S. Rockefeller
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Scientific Staff

Zoological iJQarfi

H. R. Mitchell, Acting Director
Raymond L. Ditmars, Curator of Mammals and Reptiles
Lee S. Crandall, Curator of Birds
Leonard J. Goss, Veterinarian

Claude W. Leister, Ass’t to the Director and Curator, Educational Activities
II. C. Raven, Prosector
Edward R. Osterndorff, Photographer
William Bridges, Editor and Curator of Publications

Squartum

Charles M. Breder, Jr., Director
Christopher W. Coates, Aquarist
Ross F. Nigrelli, Pathologist
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Homer W. Smith, Research Associate in Physiology

department of tropical 5&£3earcl)

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Cbitoiial Committee

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Charles M. Breder, Jr.
William Bridges

H. R. Mitchell
William Beebe

ZOOLOGICA

SCIENTIFIC CONTRIBUTIONS
OF THE

NEW YORK ZOOLOGICAL SOCIETY

VOLUME XXV

Part 3

Numbers 19-24

NOV 201940

PUBLISHED BY THE
THE ZOOLOGICAL PARK,

November 14, 1940

SOCIETY
NEW YORK

CONTENTS

PAGE

19- Eastern Pacific Expeditions of the New York Zoological
Society. XX. Medusae of the Templeton Crocker and
Eastern Pacific Zaca Expeditions, 1936-1938. By Henry
B. Bigelow. (Text-figures 1-20) 281

20. Two New Species of Trematodes ( Apharyngostrigea
bilobata : Strigeidae, and Cathaemasia nycticoracis :
Echinostomidae) from Herons, with a Note on the Oc-
currence of Clinostomum companulatum (Rud.). By

O. Wilford Olsen. (Plate I) 323

21. Nesting of the Sunfish, Lepomis auritus (Linnaeus), in

Tidal Waters. By Neil D. Richmond. (Plate I) 329

22. Eastern Pacific Expeditions of the New York Zoological
Society. XXI. Notes on Echinoderms from the West

Coast of Central America. By Hubert Lyman Clark.
(Plates I & II; Text-figures 1-4) 331

23. The Nesting Behavior of Eupomotis gibbosus (Linnaeus)

In a Small Pool. By C. M. Breder, Jr. (Plates I & II;
Text-figures 1 & 2) 353

24. Reproductive Activities of a Hybrid Minnow, Notropis

cornutus X Notropis rubellus. By Edward C. Raney. 361

Bigelow: Medusae of Zaca Expeditions, 1936-1938

281

19.

Eastern Pacific Expeditions of the New York
Zoological Society. XX.

Medusae of the Templeton Crocker and Eastern Pacific Zaca
Expeditions, 1936-1938.1

Henry B. Bigelow

Museum of Comparative Zoology, Harvard University.

(Text-figures 1-20).

[This is the twentieth of a series of papers dealing with the collections
of the Eastern Pacific Expeditions of the New York Zoological Society made
under the direction of William Beebe. The present paper is concerned with
specimens taken on the Templeton Crocker (1936) and Eastern Pacific Zaca
(1937-1938) Expeditions. For data on localities, dates, nets, etc., refer to
Zoologica, Vol. XXII, No. 2, pp. 33-46, and Vol. XXIII, No. 14, pp. 287-298.]

Contents.

Page.

Introduction 282

Hydromedusae

Anthomedusae 282

Pandeidae 282

Stomotoca 282

Stomotoca pterophylla Haeckel 283

Leuckartiara 284

Leuckartiara zacae sp. nov 284

Neoturris 287

Neoturris crockeri sp. nov 287

Calycopsis 290

Calycopsis simulans Bigelow 293

Bougainvilliidae 294

Chiarel a 294

Chiarella centripetalis Maas 294

Leptomedusae 296

Thaumantadidae 296

Polyorchis 296

Polyorchis penicillata Eschscholtz 296

Laodiceidae 297

Chromatonema 297

Chromatonema erythrogonon Bigelow. 297

Aequoridae 299

Zynocanna 299

Zygocanna vagans Bigelow 299

Trachomedusae 302

Geryonidae 302

Liriope 302

Liriope tetraphylla Chamisso &

Eysenhardt 302

Trachynemidae 302

Page.

Colobonema 302

Colobonema typicum Maas 302

Halicreidae 302

Halicreas 302

Halicreas minimum Fewkes 302

Narcomedusae 303

Solmaridae 303

Pegantha 305

Pegantha clara R. P. Bigelow 305

? Pegantha martagon Haeckel 308

Solmissus 308

Solmissus marshalli Agassiz & Mayer. . 308

Aeginidae 309

Aeginura 309

Aeginura beebei sp. nov 309

Aeginura grimaldii Maas 313

Aegina 313

Aegina citrea Eschscholtz 313

Scyphomedusae 314

Coronata 314

Periphyllidae 314

Periphylla 314

Periphylla hyacinthina Steenstrup . . . . 314

Atollidae 315

Atolla 315

Atolla wyvillei Haeckel 315

Discophora 315

Pelagidae 315

Pelagia 315

Pelagia noctiluca Forskal 315

Stomolophidae 316

Stomolophus 316

Stomolophus meleagris L. Agassiz 316

1 Contribution No. 601, Department of Tropical Research, New York Zoological Society.
Contribution No. 239 of the Woods Hole Oceanographic Institution.

NOV i @ f$f§

282

Zoologica: Neiv York Zoological Society
Introduction.

[XXV :19

The medusae described in the following pages were collected by Dr.
William Beebe on Mr. Templeton Crocker’s Yacht Zaca, off Lower California
and in the Gulf of California in April-May, 1936, and along the Pacific
coast of Central America from Costa Rica to Panama in February-April,
1938. For data of the stations where medusae were taken on the first of
these cruises (Stas. 139-165), see Beebe (1937); for those of the second
cruise (Stas. 210-234), Beebe (1938).

The locations of the stations are shown on Text-figs. 1 and 2.

Text-fig. 1.

Locations of stations 139-158, April, 1936.

Hydromedusae.

Anthomedusae.

Pandeidae.

Stomotoca.

Differences of opinion still exist as to whether the pandeids with only
two large opposed tentacles, but many rudimentary tentacles, represent two
genera, one with gelatinous peduncle ( Stomotoca ), the other without
(Amyhinema) , or whether they properly form but a single genus, to which
the name Stomotoca must be applied, on the grounds of priority. The latter

1940]

Bigelow: Medusae of Zaca Expeditions, 1936-1938

283

Text-fig. 2.
Locations of sta-
tions 210-234, Feb-
ruary-April, 1938.

viewpoint has recently been adopted by Uchida (1927) and by Ling (1937),
following Mayer (1910). To Hartlaub (1914), however, among others, to
Russell (1938), and to me (1909, 1918), the presence or absence of a peduncle
has seemed a character of generic importance, and is so considered here.

Stomotoea pterophylla Haeckel.

Stomotoca pterophylla, Haeckel, 1879, p. 52, PI. 4, Fig. 10. For subse-
quent references and discussions, see Bigelow, 1918, p. 372; Ranson, 1936, p.
63.

Material: Sta. 229, surface, 1 specimen about 20 mm. in diameter: Sta.
231, surface, 1 specimen about 22 mm. in diameter, both in moderately good
condition.

Comparison of series from the Florida-Bahaman region and from the
Eastern Tropical Pacific has failed to reveal any significant difference be-
tween the Pacific divisa and the Atlantic pterophylla (Bigelow, 1918, p. 370) ;
and this conclusion the present examples corroborate, for they agree in all
significant respects with Maas’ (1897) account of “S. divisa” from the Gulf
of Panama, and with the Albatross series reported by me under that name

284 Zoologica: Netv York Zoological Society [XXV:19

from the vicinity of the Galapagos and from the West coast of Mexico
(Bigelow, 1909).

Records for the compound species in the Pacific are confined to the
vicinity of the Galapagos, and along the west coast of America from southern
Mexico to Colombia. In the Atlantic, however, its range appears to be more
extensive, for it has been taken not only near Florida and the Bahamas in the
western side (Haeckel, 1879; Mayer, 1910; Bigelow, 1918), but also in the
central basin (Ranson, 1936). 2

Leuckartiara.

Leuckartiara zacae sp. nov.

Text-figs. 3-5.

Material : Sta. 227, 910-0 meters, one specimen about 21 mm. high by 18
mm. in diameter, somewhat shrivelled in preservation, but otherwise in fair
condition.

In 1916, Browne described, from the Indian Ocean, an interesting new
pandeid ( Leuckartiara gardineri) , the generic identity of which was estab-
lished by the structure of its gonads, but which differed from all known
representatives of Leuckartiara in the presence of exumbral nematocyst-
ribs extending upward over the surface of the bell from the bases of the
tentacles. This character it shares with Pandea conica and with Eutiara
mayeri, the gonads of which are, however, very different.

The present specimen, similarly referable to Leuckartiara by its gonads,
recalls gardineri in the presence of exumbral ribs. And (so far as can be
judged in optical section) these ribs, like those of gardineri and of Eutiara
mayeri (Bigelow, 1918, p. 375), are canalar extensions of the tentacular
bases, whereas in Pandea conica, Hartlaub (1913, p. 336) describes them as
simply thickenings of the ectoderm. It differs, however, from gardineri in
the following respects :

1. Number of tentacles : In the unique specimen of gardineri, there were
only four large (perradial) tentacles and in addition about 24 rudimentary
knobs, some with filaments, but others lacking the latter. And since the
gonads contained fairly well-advanced ova, it seems probable that four is the
final number of large tentacles in that form. In the Zaca specimen, however,
there are (besides the four perradials) two or three other well-developed
tentacles of different sizes per quadrant (a total of fifteen), each with
corresponding exumbral rib, and also 1-3 rudimentary knobs (lacking fila-
ments) between every two tentacles (Text-fig. 3).

2. Number and length of exumbral ribs : In Browne’s specimen of gar-
dineri there were four of these, one opposite each perradial tentacle and
extending nearly to the apex of the bell. In the present example, in addition
to the four perradials, there are 2, 1, 1 and 1 others in the different quad-
rants, or a total of nine, each connected with a tentacle. And while four
of the smaller tentacles lack them, exumbral ribs might later have developed
in connection with these also. On the other hand, even the perradial ribs
reach up only about 2/z of the height of the bell, i.e., to about the level where
the radial canals arise from the manubrial wall, while the others reach only
to about the mid-level of the bell.

3. Gonads : These are considerably more complex than in gardineri, there
being 8-12 primary folds, which bifurcate or trifurcate in each adradial
series, as against 4-5 primary folds only — each bifid — per adradial series
in gardineri. In gardineri, furthermore, the adradial series are united in

2 Ranson (1936, p. 64) states that the Monaco specimen was collected “au sud des Azores.” But
the position of the station in question (Monaco, Station 2116) is given as Lat. 31° 42' N., long.
42° 40' W., or roughly midway between the Azores and Bermuda.

1940]

Bigelow: Medusae of Zaca Expeditions, 1936-1938

285

Leuckartiara zacae, sp. nov., type specimen, slightly restored.

each quadrant, by a transverse fold at about the mid-level of the manubrium
(Browne, 1918, PL 39, Fig. 4), but this commissure is evident in one
quadrant only of the Zaca specimen, suggesting that its presence represents
a transitory stage in development. The specimen is apparently a male; at
least no ova are visible in surface view.

Evaluation of these differences depends on whether the new-found
specimen represents merely an advanced stage in the growth of a known
species (all the differences being of a progressive sort), or whether it is
a distinct species which, in its growth, passes through a stage in which it
closely resembles the known form, but which attains to a greater degree of
morphological complexity before reaching maturity. In the present in-
stance, the available bases for decision are conflicting, for while the Zaca

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[XXV :19

specimen is about six times as large as the type of gardineri (suggesting
an advanced stage in growth), the localities of capture for the two are on
opposite sides of the globe.. And since it seems important to emphasize the
differences rather than to minimize them, so that the matter will be more
searchingly examined by whomever is lucky enough to obtain additional
specimens, a new specific name is proposed for the medusa under discussion
here.

Text-fig. 4.

Leuckartiara zacae, sp. nov.,
margin with large and small
tentacles, camera drawing, about
X 14.

Text-fig. 5.

Leuckartiara zacae, sp. nov., lateral view
of radial tentacle with nematocyst rib, about
X 15.

Further points worth notice are that the tentacular bulbs are laterally
flattened, clasping the exumbrella (as is usual in this genus), and that the
tentacular knobs, as well as such of the fully formed tentacles as still lack
exumbral prolongations, bear pigment spots on their abaxial sides. But the
spots (“ocelli”) are lacking on the tentacles in connection with which
exumbral ribs have developed — apparently they are lost when the latter
structures appear. The edges of the notably broad radial canals are slightly
wavy ; those of the ring canal smooth.

1940]

Bigeloiv: Medusae of Zaca Expeditions, 1936-1938

287

The ocelli are brick red, the manubrium and tentacular bulbs a pale
salmon-color after preservation, which suggests a more brilliant reddish or
pinkish hue in life.

Material : Sta. 234, 910-0 meters, 1 fragmentary specimen about 28 mm.
high by about 32 mm. in diameter.

The unique example is so badly battered that reconstruction of the
general form of the bell can be only tentative, further than that it is thin-
walled throughout, without apical thickening, and apparently slightly broader
than high. Fortunately, however, (though now in fragments) it still shows
the chief anatomical characters.

At first glance, the specimen might easily be mistaken for a Pandea
rubra, thanks to its dark chocolate-red pigmentation (see below). However,
the structure of the gonads is quite different, being clearly of the “Neoturris”
type as defined by Hartlaub (1914). More precisely, each perradial belt is
flanked, on either hand, by a series of transverse folds of various sizes (Text-
fig. 6). As is typical of Neoturris, the sexual folds are not connected one
with another at their interradial ends, but are entirely independent. But
the interradial zone of the manubrial wall, between each two series of folds,
is occupied by a series of small irregular ridges — primarily longitudinal —
alternating with recesses. Comparison of Text-fig. 6 with Hartlaub’s (1914,
p. 35, Fig. 273) picture of the manubrium of N. pileata will show the essen-
tial similarity. However, while the adradial folds are simple in one of the
quadrants (Text-fig. 6), in another adjacent (Text-fig. 7) they are so com-
plexly split and cross-furrowed that it is impossible either to distinguish
between primary folds and secondary, or even, in the extreme case, to dis-
tinguish sharply where adradial folds give place to interradial mesh-work.
Locally, then, the “Neoturris” state grades into the “Pandea” state. No-
where, however, does it grade into the “Leuckar tiara” state, i.e., that in

Neoturris.

Neoturris crockeri sp. nov.
(Text-figs. 6-10).

« Neoturris crockeri, sp. nov.,
interradial view of one quadrant
of manubrium to show gonads,
somewhat restored.

Text-fig. 6.

288

Zoologica: New York Zoological Society

[XXV :19

APICAL

PER

INTER.

Text-fig. 7.

Neoturris crockeri, sp. nov., adradial series
of sexual folds in quadrant adjacent to that
shown in Text-fig. 6. Camera drawing.

which the interradial extremes of the transverse adradial folds are con-
nected. The specimen is a female, the sex folds packed with large eggs ( Text-
fig. 8) — and scattered eggs are to be seen in the interradial belts, as well.
It is interesting, as an example of a fate that no doubt causes an appre-
ciable destruction of medusan eggs, that in the present case a considerable
number are to be seen within the radial canals.

The relationship of radial canals to manubrium is of the type charac-
teristic of the more complexly organized pandeids ( Neoturris , Pandea,
Leuckartiara, Catablema ) , for the so-called “mesenteries” are so well devel-
oped (Text-fig. 6) that the canals depart from the manubrium at about
the mid-level of the latter. Above this level, consequently, the bell cavity is
subdivided into four extensive pockets by the lines of attachment between
the manubrium and subumbrella.

The canals themselves are broad, flat (as usual in this family), their
edges nearly smooth — or at most but slightly irregular. And this applies
equally to the margin of the ring canal. The manubrium occupies the greater
part of the bell cavity, hanging in its present state about to the level of the
bell opening. The lips are complexly crenulated, much as in Pandea rubra.

Text-fig. 8.

Neoturris crockeri, sp. nov., portion of one
of the sexual folds packed with ova, camera
drawing, X 60.

1940]

Bigelow: Medusae of Zaca Expeditions, 1936-1938

289

There are 38 tentacles, the numbers in successive quadrants being 10, 9,
9, 10, of various sizes (Text-fig. 9). Presumably this would have been the
final number in this particular specimen, for there are no additional rudi-
ments or knobs. The tentacle bases, broadly triangular in side view (Text-
fig. 10), and laterally compressed, clasp the exumbrella, as is so commonly
the case in this family, and taper gradually to slender filaments.

There is no trace of exumbral ridges connected with the tentacles. And
since the surface of part, at least, of the bell is in fair condition, there is
no reason to suppose that such structures were present in life, but had been
destroyed. Neither are any ocelli to be seen.

The most interesting feature of this specimen is its bathypelagic colora-
tion, the manubrium as a whole being of a dense reddish-chocolate hue,
paling at the lip, and with the opaque white eggs showing through the
overlying, pigmented ectoderm, much as do the larger eggs of the Narcome-
dusa, Aeginura grimaldii. The bases of the tentacles and a narrow peripheral
belt of the subumbrella are sparsely pigmented with the same hue, the pig-
ment granules in the latter cases so large and loosely spaced as to be indi-
vidually visible under low magnification. The remainder of the subumbrella
is faintly tinged, though without visible granules.

Superficially, as noted above, this specimen most resembles Pandea
rubra among known pandeids. However, the tentacles are considerably more
numerous, the maximum yet recorded for a considerably larger specimen of
rubra being only 20 (Bigelow, 1938, p. 107). More important, the structure
of the gonads proves that the resemblance to Pandea rubra is only super-

Neoturris crockeri, sp. nov.,
segment of margin to show the
tentacles of different sizes,
about X 7.5.

Text-fig. 9.

Text-fig. 10.

Neoturris crockeri, sp. nov., lateral view of
one of the medium-sized tentacles.

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Zoologica: New York Zoological Society

[XXV: 19

ficial and that we are actually dealing with a Neoturris. However, it differs
from all known pandeids in which the gonads are definitely known to be of
the “Neoturris” type, not only in its peculiar pigmentation, but also in the
facts that the radial canals are smooth-edged, and that the tentacular organs,
large or small, are so few in number and consequently so widely spaced. A
new species seems therefore necessary for its reception.

The depth of capture of the type (and so far only known) specimen
(910-0) argues, as does its color, that it is a member of the bathyplankton.

Calycopsis.

Up to the present time, eleven supposed species have been referred to
this genus or to Sibogita, now generally considered synonymous with it.
The necessity of decision as to the status of its representative in the Zaca
collection makes this an opportune time to summarize the features by which
the several named members of the genus have been separated one from
another. Critical revision (depending on the evaluation of variability) must,
however, await the accumulation of larger study series.

One of the species, birulai Linko (1913), can be omitted from the dis-
cussion, on the ground that it does not belong to this genus, but is more
closely allied to Eumedusa similis (Bigelow, 1920). It may, in fact, prove
identical with the latter, in which case Linko’s specific name would take
precedence.

For the remaining members of Calycopsis, the following characters have
been chiefly emphasized as specific: 1, number of tentacles and canals and
their interrelationship; 2, arrangement of centripetal canals and the degree
to which these finally fuse either with one another, with the radials, or with
the base of the manubrium; 3, morphology of the gonads; 4, presence or
absence of labial nematocyst knobs ; 5, whether or not the radial canals are
dilated at their points of emergence from the manubrium to form so-called
“mesenteries” ; 6, presence or absence of gelatinous exumbral papillae on the
marginal lobes of the bell; and 7, presence or absence of a funnel-shaped
apical exumbral depression. Among these characters, the presence of an
apical depression in typa, of labial nettle knobs in nematophora, and of
prominent papillae on the exumbral surfaces of the marginal lobes in papil-
lata, in valdiviae, and in at least one of the specimens in the collection of
the German South Polar Expedition (taken near the Cape Verdes), that
were recorded by Vanhoffen (1912) as “typa,” contrasted with their ab-
sence elsewhere, is strictly alternative. The presence or absence of subum-
bral muscle bands along the radii of the tentacles, and of pigment spots at
the bases of the tentacles, also appears to be alternative at least between
certain of the named forms. Similarly, the conformation of the gonads is of
two rather sharply opposed types, for while in borchgrevinki (at least in
the male), the sex products develop in pockets of the manubrial wall, and
open to the exterior by small pores (Browne, 1910; Vanhoffen, 1911, p. 216,
Fig. 10a; Mackintosh, 1934), the gonads of all other known members of the
genus form double rows of transverse folds, occupying the interradial sectors
of the manubrium, the minor divergences being that in nematophora, small
irregular folds occur between the chief series, and that in typa, the gonads
bear egg-shaped swellings, scattered here and there (Bigelow, 1909a, Figs.
1,2).

Other characters are less sharply discontinuous. Thus, chuni with 36-27
canals, geometrica with 32-22, typa with 21-16, and nematophora with 18-16
form a continuous series. In valdiviae, however, the number of canals re-
corded (60) is some 20 greater than in any other known Calycopsis; in
bigelowi and in simplex, it may be permanently smaller (8). In simulans
and in papillata, the number (12 in each case) appears to be determinate,

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291

probably also in borchgrevinki (16), whereas in most, at least, of the other
supposed species it is variable. In simulans and papillata, furthermore, the
centripetal canals are all adradial, whereas in all other named forms in
which the metameral arrangement has been determined, there is an inter-
radial canal in each quadrant with or without adradials according to the
number of centripetals peculiar to the particular species in question. In
this respect, typa is especially unstable for there may be 2, 3, or 4 canals
in adjoining quadrants (Bigelow, 1909a), beside the interradial member.
Ordinarily the centripetals arise singly from the ring canal. In typa, how-
ever, they may exceptionally arise in pairs (Bigelow, 1909a, Fig. 7), while
in chuni they may be either single or in groups of two or three (Vanhoffen,
1911, p. 218, PI. 22, Fig. 8).

In previous discussions of the genus, much emphasis has been laid on
the endings of the centripetal canals, i.e., whether permanently blind, or
whether finally fused with the radial canals, or with the cruciform base of
the manubrium. In bigelowi, in simplex, and in such of the specimens
recorded as typa as seem properly to have belonged to that species, the
canals are described as ending blindly — in the latter case even in a specimen
37 mm. high with large ova (Bigelow, 1909a, p. 290). In simulans and
valdiviae some at least join the base of the manubrium, but others (by
present indications) may remain permanently blind, which according to
Browne (1910, p. 17) also applies to borchgrevinki. In nematophora all
come finally to join the base of the manubrium — a junction interpretable in
some cases as with a radial canal at its point of origin from the manubrium
(Bigelow, 1913, PI. 3, Fig. 24). In papillata they all end blindly in medium-
sized specimens, but in the largest (27 mm. high, with eggs), two were
still blind, “but of the remaining six, three have joined the cruciform base
of the manubrium; two join radial canals close to their bases, and one joins
a radial canal at a considerable distance from the manubrium” (Bigelow,
1918, p. 280, PI. 2, Fig. 1). This specimen thus bridges the gap between the
foregoing group and geometrica, in which all the centripetals unite with the
radials at different levels.

The numbers of tentacles in the different members of Calycopsis over-
lap to a greater degree than do the numbers of canals, the total range (60-8)
being practically covered by three species, nematophora (57-30), typa
(30-16), and borchgrevinki (8-16). Recorded numbers of tentacles for the
other supposed species are: valdiviae, about 60; bigelowi, 48; chuni, 16-17;
geometrica, 16-22; simulans, 12-18; papillata, 12; simplex, 8.

As appears from the foregoing, the canals and tentacles may be equal
in number (a tentacle opposite every canal); they may be either equal in
number or the tentacles the more numerous ( simulans ) ; the tentacles may
be characteristically the more numerous ( borchgrevinki , bigelowi, nema-
tophora, typa ) ; or the canals the more numerous ( geometrica , chuni).

In typa, it appears that the tentacles develop before the corresponding
canals appear; apparently also in nematomorpha, hence this is likely the
general rule, except in forms in which the final number of canals is gi-eater
than that of the tentacles.

Taxonomic evaluation of labial knobs and of gonad-structure presents
no difficulty, for these both appear of full specific import. Thus, nema-
tomorpha is set apart from all other members of the genus by the presence
of the former, as well as by the presence of subumbral muscle bands along
the canalar radii; borchgrevinki by the gonads. The presence of a well-
marked apical exumbral depression, coupled with an evident tendency for the
centripetal canals to continue blind, even at maturity, is a good mark of
identification for typa, for while the depression might seem a trivial feature,
its presence, not only in Fewkes’ (1882) original specimen, but also in
others collected many years later by the Grampus, and in five more in the

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[XXV :19

Arcturus collection (all from the same general region off the northeast coast
of the United States) seems to establish its constancy.

Geometrica is equally marked off by one outstanding character, i.e., by
the union (in the adult) of all the centripetal canals with the radials. The
only other Calycopsis that at all approaches it in this respect ( papillata )
has well marked exumbral sculpture which geometrica lacks.

Simulans and typa form a natural group so far as general appearance
is concerned — so closely, in fact, do they resemble each other that Vanhoffen
(1911) classed the former as a synonym of the latter. However, all the
specimens referable to simulans that have so far been seen (including the
Zaca specimen described below) have differed from typical typa not only in
lacking any trace of apical depression, or of the egg-shaped swellings on the
gonads that appear characteristic of the latter, but also in the fact that for
them the number of canals appears to be determinate (12) instead of vari-
able. Among the examples of “ simulans ” so far recorded, one from the Philip-
pines (Bigelow, 1919) is set apart by the facts that most of the centripetal
canals are fused with the manubrium instead of ending blindly, and that
there are eight small tentacles in addition to the twelve that correspond to
the canals. Whether this represents a geographic variation — i.e., that devel-
opment progresses further in this respect in one race of simulans than in
another — or whether the Philippine example actually belongs to some form
as yet unnamed, is a question for the future.

Chuni, to judge from Vanhoffen’s (1911, PI. 22, Fig. 8) picture, must
be grouped in its general organization with the typa-simulans group. How-
ever, its canals (27, 36) are somewhat more than twice as numerous as its
tentacles (16, 17), whereas in typa and in simulans, the tentacles are at
least as numerous as the canals. Valdiviae (to judge from the only adult yet
seen) is set apart by the very large number (60) of canals, just commented
upon, combined with absence of mesenteries. In bigelowi the numerical ratio
of tentacles (48, large and small) to canals (8) is much larger than in any
other member of the genus. And while the small size of the one specimen
so far seen (only 12 mm. high) might suggest that it was a juvenile, the
well-developed gonads argue against this view. Vanhoffen’s (1911, p. 218,
Fig. 12) illustration also suggests that part of the tentacles remain per-
manently small. Simplex, finally, while agreeing with bigelotvi and borch-
grevinki in the small number (8) of canals, is separated from the former
by the fact that it has only about Ve as many tentacles (8), and from the
latter by the structure of the gonads. It may be the juvenile of some other
form.

It appears, in short, that while the various named members of the genus
are closely allied, one to another, most, at least, of them differ sufficiently to
be accepted as good species until intermediates be actually found. Their
geographic distribution tends to strengthen this conclusion. Thus, speci-
mens with apical depression (typa) have been taken only off the continental
slope of North America; those agreeing with the original specimens of
simulans only off the west coast of Central America; those with labial
nematocyst knobs ( nematomorpha ) only in the Bering Sea region; those
with the borchgrevinki type of gonads only in high southern latitudes; those
with the geometrica type of canalization are known only in the Philippines
and East Indies; the simplex type only from Norway; those of the bigelowi
and chuni types from the Gulf of Aden.

Specimens with warty marginal lobes have, it is true, been recorded
widespread, i.e., from the Florida-Bahamas region (papillata, Bigelow,
1918), from the vicinity of the Cape Verde Islands (Vanhoffen, 1912), and
from the Alguhlas current by Vanhoffen (1911) as “typa,” this last being
the specimen on which Hartlaub (1914) subsequently founded the species
valdiviae. But papillata and valdiviae differ so widely in other respects (as

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293

emphasized above) that there can be no question of their identity. And the
description of the other warty specimens recorded by Vanhoffen (1912, as
“typa”3), classed tentatively by Hartlaub (1914) as juveniles of valdiviae,
are not detailed enough for positive identification.

The following artificial key to the species of Calycopsis may be helpful
for purposes of identification :

1. With labial nettle knobs and subumbral muscle bands

nematophora Bigelow

Without labial nettle knobs or subumbral muscle bands 2

2.

All centripetal canals join the radial canals at various levels

geometrica Maas

Most of the centripetal canals either end blindly, or join the cruciform
base of the manubrium 3

3. With about 60 canals at maturity

With 40 canals, or fewer

4. Gonads in pockets

Gonads in exterior folds

5. Exumbrella with well-marked apical depression

Without apical depression

6. Many more canals than tentacles

Tentacles at least as numerous as canals

valdiviae Hartlaub

4

borchgrevinki Browne

5

typa Fewkes

6

chuni Vanhoffen
7

7. Marginal lobes with well-marked exumbral papillae papillata Bigelow

Marginal lobes smooth or nearly so 8

8. Many more tentacles than canals bigelowi Vanhoffen

Number of tentacles and canals about equal 9

9. Centripetal canals all adradial simulans Bigelow

Centripetal canals interradial simplex Kramp & Damas

Calycopsis simulans Bigelow.

Sibogita simulans, Bigelow, 1909, p. 213, PI. 5, Figs. 4, 5; PI. 41, Figs. 8,
9; PI. 43, Figs. 1, 2; 1913, p. 21; Mayer, 1910, p. 187.

Calycopsis simulans, Hartlaub, 1913, p. 360; Bigelow, 1918, p. 377.

Calycopsis typa (Partim), Vanhoffen, 1911, p. 214.

? Calycopsis typa var. simulans, Bigelow, 1919, p. 292, PI. 40. Fig. 8;
PI. 41, Fig. 1.

Material: Sta. 233, 910-0 meters, one specimen, in good condition, about
26 mm. high by 25 mm. broad.

The Zaca capture corroborates the evidence afforded by the two speci-
mens from between Panama and the Galapagos, for which the species was
originally established, that the final number of canals in simulans is 12,
i.e., 2 centripetals in each quadrant. In the original specimens these all
ended blindly, though in their growth they had reached nearly to the
cruciform base of the manubrium4. And this is true also of the Zaca speci-
men, though in this case the canalar terminations come so close to the
manubrium that careful examination was required to demonstrate the dis-
continuity. It also agrees with the type specimen in the presence of as many
large tentacles (12) as canals, one of the former opposite each of the latter.
The fact that specimens apparently adult (to judge from the gonads) from

3 Among three juveniles recorded as typa, two from the tropical Atlantic and one from the
Indian Ocean, Vanhoffen (1911, p. 364) states that the wartiness was especially prominent after
alcoholic preservation, which identifies the specimens as above.

4 As pointed out elsewhere (Bigelow, 1913, p. 23), it is probable that a specimen from Bering
Sea originally referred by me to simulans, in which the canals all join the manubrium (Bigelow,
1909, p. 214, Fig. 2), actually belonged to C. nematomorpha.

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the same general region, but taken 32 years apart, agree so closely in these
respects makes it likely that this is the final state. And for this reason it
may finally prove necessary to find a new name for the Philippine specimen,
originally described by me (1919) as var. simulans of typa, but in which all
but one of the centripetals had already fused with the manubrium though
the medusa was only 16 mm. in diameter, and in which there were eight
small tentacles, besides the twelve larger ones connected with the canals
(p. 292).

I need only note further that the tentacles, like those of the original
specimens — of most other members of the genus, in fact — are tipped with
prominent nematocyst knobs; that the sexual folds correspond closely to
those of the type in regularity and general arrangement, and that there is
no trace either of an apical depression or of exumbral papillae on the mar-
ginal lobes.

In the preserved condition the manubrium is pale ochre, the tentacles
colorless.

The fact that the Albatross specimens were taken at the surface makes
it likely that the present example was picked up by the net at a small depth,
either on its way down, or its way up.

Bougainvilliidae.

Chiarella.

Chiarella centripetalis Maas.

(Text-figs. 11, 12).

Chiarella centripetalis, Maas, 1897, p. 15, PI. 2, Figs. 1-4; 1905, p. 13;
Mayer, 1910, p. 182; Foerster, 1923, p. 29.

Material : Stas. 139, 148, 159, in hauls respectively from 549-0, 549-0,
910-0 meters, 5 specimens, about 20-25 mm. high.

This large and easily recognized species had not been reported since
originally described by Maas (1897) from specimens taken in the Gulf of
California in 1891 by the U. Si Fish Commission Steamer Albatross. The

Text-fig. 11.

Chiarella centripetalis Maas, from Sta. 159, radial group of
tentacles, camera drawing, about X 13.

1940] Bigelow: Medusae of Zaca Expeditions, 1936-1938

295

Zaca captures in the same general region are, therefore, of considerable
geographic interest.

The specimens, though all more or less damaged, are in good enough
condition to show that in most respects they agree with Maas’ account so
closely that there is no doubt of their specific identity. This applies, in par-
ticular, to the general form of the bell ; to the outlines and lengths of the spur-
like interradial centripetal canals (Text-fig. 12), to the conformation of the
gonads and to the basic arrangement of the eight groups of tentacles, by
which the genus is characterized. Conspicuous, in particular, is the epaulette-
shape of the basal folds from which the tentacular filaments arise (Text-figs.
11, 12), and the adradial series of transverse sex folds in the manubrial
wall, which, in two of the Zaca specimens, are packed with large eggs.

The peduncle is considerably longer in four of the five Zaca specimens
(the condition of the fifth is not good enough to show this) than in Maas’
(1897) illustration based on a sketch from life, occupying in the extreme
case, about Vs of the length of the bell cavity. But as there is a considerable
variation in this respect within the series, and as all of them are in a flaccid
condition in weak formalin, it is probable that this apparent difference
between the collections of 1891 and of 1936 is the result either of different
states of contraction or of preservation.

It is also worth mention that the margins of the radial canals (at least
in the better-preserved examples) are slightly irregular or wavy though
never definitely lobed — a feature not mentioned by Maas. The ring canal is
smooth-edged.

Text-fig. 12.

Chiarella centripetalis Maas from Sta. 139, interradial group of
tentacles, camera drawing, about X 16.

According to Maas (1897, p. 16) there are about as many tentacles in
the interradial groups as in the radials (more than 40 in each, a total of
more than 320). However, he does not state whether an individual count
was made of any group othetf than the one pictured. And the Zaca series
proves not only to be more variable in this respect, but with the number
averaging only about half as great in specimens of the same size, the num-
bers being as follows for the groups for which counts are possible:

Approximate Radial Interradial

BELL HEIGHT GROUPS GROUPS

24 mm.
23 mm,
21 mm.
20 mm.

25 mm.

19, 19, 20, 18
18, 19

22, 22±, 25±, 33±

not more than 20 in any group

23±, 24±, 24±

16, 16, 17, 19
16, 25

24, 23, 31, 38±

28±, 27±, 24, 24±

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The spacing around the bell margin is also variable, for while in the
24 mm. and 25 mm. specimens, all the epaulettes are approximately equal in
breadth and of approximately the same breadth as the interspaces, in the
21 mm. specimen the interradial group with 38 tentacles occupies practically
the entire interspace between the two adjacent radial groups.

Maas remarked that “die einzelnen Faden sind verhaltnissmassig kurz,
was nicht auf Rechnung ihrer Contraction zu setzen ist, wie Skizzen nach
dem Leben zeigen” (Maas, 1897, p. 16). Similarly, in the Zaca series all the
more central members of each group are not only mere stumps (Text-figs. 11,
12) but such of them as are in good condition are rounded at the ends. But
in several of the better-preserved groups the one or two tentacles near the
outer margin extend as well developed though short filaments (Text-fig. 11),
suggesting that all the tentacles may have been extensible to at least this
extent in life.

According to Maas there is an ocellus at the base of each tentacle, as
also appears in his color sketch from fresh material (Maas, 1897, PI. 2, Fig.
1). These are not visible on the Zaca specimens, but they may have faded
in the preservative.

In the preserved state the manubrium varies between dull salmon-pink
and ochre yellow; the tentacular pads are of various shades of reddish-
yellow.

The original account includes no information as to the depth of cap-
ture. But the fact that all of the Zaca specimens were taken in hauls from
considerable depths suggests that the normal habitat lies considerably below
the surface.

Leptomedusae.

Thaumantadidae.

Polyorchis.

Polyorchis penicillata Eschscholtz.

Melicertum penicillatum, Eschscholtz, 1829, p. 106, PI. 8, Fig. 4.

Polyorchis penicillata, A. Agassiz, 1862, p. 349.

To the synonymy given by Mayer, 1910, p. 218, add
P. penicillata, Foerster, 1923, p. 32; and

P. minuta, Murbach & Shearer, 1902, p. 72 ; 1903 ; p. 174, PI. 19,
Fig. 3, PI. 22, Fig. 1.

Material-. Inez Bay, 1 specimen about 10-12 mm. in diameter.

In this specimen, less than half grown to judge from the size to
which the species attains in Californian waters (Mayer, 1910, p. 218;
Foerster, 1923, Table, p. 34) there are 31 well developed tentacles, as
well as three (or perhaps four) rudimentary tentacular knobs; or a number
nearly as great as given by Mayer for the adult. However, Foerster (1923,
p. 32) records an average of 64 tentacles (or 16 per quadrant) for large
specimens, suggesting that the Zaca example was about half grown as re-
gards number of tentacles as well as in size.

According to Fewkes (1889) and to Foerster (1923) the edges of the
radial canals already bear small projections even as early as the 16-tentacle
stage, foreshadowing the lateral diverticula characteristic of the adult
Polyorchis. And the latter author (1923, p. 34) records their number as
“fairly constant, i.e., 30-40, for small as. well as for large specimens.” In
the Zaca example the edges of the canals are at most slightly wavy, at a
stage in growth at which Californian specimens show well developed diver-
ticula. Since, however, this is the only feature differentiating this specimen

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297

from its confreres in more northern waters, it does not seem sufficient to
warrant even a distinct racial name.

Previous records for Polyorchis penicillata were wide-spread between
northern Vancouver Island and Santa Barbara, California. The Zaca cap-
ture is interesting chiefly for extending the known range of the species
southward to the Gulf of California.

Laodiceidae.

Chromatonema.

I have recently (1938, p. 109) given reasons for retaining this genus
among the Leptomedusae as a member of the Laodiceidae, rather than class-
ing it among the Anthomedusae as Ranson (1936) has done. The reader is
also referred to Kramp (1919, p. 14) for discussion of the differences in
gonad structure between Chromatonema, in which the gonads are broken
up into a series of discontinuous masses, and in Ptychogena, where the
gonad developed along either side of the pinnate radial canal is continuous.
Named forms referable to Chromatonema as defined by Kramp (1919) are
rubrum Fewkes, from the north Atlantic, erythrogonon Bigelow from the
eastern tropical Pacific, and hertwigi Vanhoffen from the Indian Ocean.
These are so closely allied that they may finally prove to be races or local
varieties of one wide ranging species, but the recorded differences seem
sufficient for the retention of all three names, at least for the time being.

Chromatonema erythrogonon Bigelow.

Text-fig. 13.

Ptychogena erythrogonon, Bigelow, 1909, p. 150, PI. 5, Fig. 1; PI. 38,
Figs. 8, 9; PI. 39, Figs. 1-7; Vanhoffen, 1911, p. 220.

Chromatonema erythrogonon, Kramp, 1919, p. 11; 1933, p. 553.

Chromatonema rubrum (Partim),' Ranson, 1936, p. 102.

Chromatonema rubrum var. erythrogonon, Ranson, 1936, p. 105.

Material: Stas. 139, 148, 219, 228, 244, in hauls from 910 and 914-0
meters, a total of 14 specimens ranging in diameter from about 15-16 to
about 44 mm., all of them in more or less fragmentary condition.

The Zaca series corroborates Kramp’s (1919, 1920) conclusion, based
on the original account of erythrogonon (Bigelow, 1909), that the latter
agrees in all important anatomical features (as it does in general appearance,
including color) with the Atlantic C. rubrum. Microscopic examination of
the marginal organs in the Zaca specimens, and reexamination of the
original material of erythrogonon, also shows that absence of nematocysts
in the cordyli of erythrogonon — mentioned by Kramp (1919, 1920) as a
possible distinction — is only apparent, i.e., that my earlier illustration (1909,
PI. 38, Fig. 8) was faulty in this respect. Actually (Text-fig. 13), the tips
of the cordyli are crowded with minute, fusiform nematocysts in the Pacific
form, just as in the Atlantic. And Kramp was no doubt correct in consider-
ing what I named “cirri” (1909, PI. 38, Fig. 9) for erythrogonon as identical
with the cylindrical cordyli in rubrum; especially since one Zaca specimen
bears two cordyli intermediate between the two extremes; whereas in none
of them have I been able to detect anything that could be described as a
“cirrus.”

It appears, however, that on the average the Pacific form develops
a somewhat larger number of tentacles than does the Atlantic. True, the dis-
continuity— if such exists— is not wide, for the largest (44 mm.) of the
Zaca specimens and two others of 35-40 mm. have each 38-39 tentacles,

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contrasting with a maximum of 20-24 in C. rubrum (maximum 7-8 per
quadrant, Kramp, 1933, p. 552). But occasional specimens from the Pacific
have considerably larger numbers, witness one Zaca specimen in which 57
can still be counted, also the presence of as many as 16 in a single quadrant
of one of the original Albatross specimens, suggesting a maximum of perhaps
60-64, or a number 2-3 times as large as has yet been recorded for any
specimen from the Atlantic. The variability in different sectors of the mar-
gin is further illustrated by one of the Zaca series (40 mm., Sta. 239) in
which there are 6, 8, 13, and 11 or 12 in successive quadrants. The number of
cordyli is also greater in erythrogonon than in rubrum, i.e., not far from 60
in specimens so far studied. However, there is no discontinuity in this re-
spect between the two forms, the recorded maximum being up to 40-45 in
rubrum, in which there are more often 2 than 1 between every tentacle, and
sometimes no greater than this in erythrogonon in which there is more often
1 than 2 in each intertentacular gap. In the largest of the Zaca specimens,
for example, with 38 or 39 tentacles, there is one cordylus, each, in 19 of the
gaps, and 2 each in 14 of the gaps, i.e., a total of approximately 40. In
another specimen the numbers in successive gaps were 1, 1, 1, 1, 2. However,
three were observed in one gap.

Text-fig. 13.

Chromatonema erythrogonon
Bigelow, surface view of tip of
cordylus to show nematocysts,
from a preparation stained in
Delafield’s haematoxylin, camera
drawing, about X 350.

On the other hand, it appears that the number of sexual masses may
average slightly smaller in erythrogonon than in rubrum (again, however,
there is no discontinuity!), for in the latter this varies from 10-16 on either
side of each radial canal (Kramp, 1933, p. 552), whereas among the
erythrogonon so far seen, that were in good enough condition for the
number to be determined, the maximum was 11, and most frequently 9 or
10. A minor difference that the future may or may not prove characteristic
is that in rubrum, so far as known, the free-hanging portion of the manu-
brium is quadrate, but barrel-shaped in erythrogonon. It also appears that
at maturity, erythrogonon tends to attain a larger size (up to 44 mm.) than
does its Atlantic relative (recorded maximum, 27 mm.).

Vanhoffen’s (1911) hertwigi from the Indian Ocean — if the single
specimen was representative — appears to be characterized by a much
larger number of cordyli5 (5) between every two tentacles than has ever
been recorded, either for rubrum or erythrogonon. It agrees with the
former, however, in the fact that the mouth was distinctly quadrate, and
also in the number (20) of tentacles although it was much larger in size,
i.e., 50 mm. in diameter.

The Zaca specimens are not in good enough condition to add anything
important (other than as above) to the earlier account of erythrogonon.
They do, however, provide interesting evidence of the constancy of colora-
tion in this species, the tentacular bulbs, manubrium, and distal parts of
the radial canals of the one specimen that is in good enough condition to
show these being of the same brick-red (pale along the lines of attachment
to the subumbrella and with pale cross at the base of the manubrium) and

5Vanhoffen names them “cirri,” but his illustration (1911, p. 220, Fig. 13b) shows that
actually they correspond to the cordyli of other members of Chromatonema.

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299

the sexual blocks of the same salmon-pink as in the original series. In fact,
all known representatives of Chromatonema agree in this respect.

The geographic range of C. erythrogonon appears to be decidedly
circumscribed, for it has so far been taken only off the coasts of South and
Central America between Latitude about 13° South ( Albatross Sta. 4675)
and the Gulf of California (Zaca Stas. 139, 148). And being so large,
and so conspicuous in color, it is not likely that it would have been over-
looked in the collections that have been made around Japan, around the
Hawaiian Islands, and in Philippine waters, did it occur there as frequently
as it evidently does in the eastern side of the Pacific at comparable latitudes.
In this respect it appears to differ from its Atlantic confrere, the distribu-
tion of which is general from the continental slope of America to northern
Europe as well as seemingly more northerly, for its known range extends
up to Davis Strait and the vicinity of Iceland (Kramp, 1919, Chart I, p.
13; 1920, p. 8), whereas it is not known farther south than the vicinity of
Bermuda (Bigelow, 1938).

Aequoridae.

Zygocanna.

Zygocanna vagans Bigelow.

Text-fig. 14.

Zygocanna vagans, Bigelow, 1912, p. 255; 1919, p. 315, PI. 42, Figs. 5-7 ;
PI. 43, Fig. 6.

Material: Sta. 225, 910-0 meters, 2 specimens, 40 and 33 mm. in
diameter. Sta. 233, 910-0 meters, 2 specimens, 33 and 34 mm. in diameter.

These specimens all agree so closely with the type series of Z. vagans
(Bigelow, 1912; 1919) that there is no doubt of their identity though the
locality of capture is in the opposite side of the Pacific. They add, however,
to our knowledge of the species by showing that the early branchings of
the canals as indicated by the figures formed by the courses of the canal
stripes on the aboral wall of the manubrium, not only are decidedly irregu-
lar (Text-fig. 14), but that the outermost branching of the canals is not as
closely associated with the margin of the manubrium as it was in the
original series, but may occasionally take place considerably centrifugal to
the latter. Such for example is the case in one of the specimens from Sta.
233 (Text-fig. 14A). Furthermore, the sinus-like expansions of the gastric
cavity along the lines of the groups of successively formed canals (Bigelow,
1919, p. 316), especially well developed in this specimen, show that no sharp
line of demarcation can be drawn between manubrium and canal, in this
highly variable species.

Associated, as I have pointed out, with this type of branching, the
canals are in groups; the number of groups “varies according to the
number of branchings which each main stem has undergone, and the
number of canals varies from group to group” (Bigelow, 1919, p. 316).
In the two members of the original series in which the four primary trunks
could be identified, the number of canals from each that reached the ring
canal varied between 5 and 12. In the present series again, in two speci-
mens in which these primary trunks are identifiable, the number of canals
to which each of the latter gives rise ranges between 4 -and 7.

The smallest number of groups in which the canals are associated at the
zone where they depart from the manubrium is 7 in the present series; the
largest number is 10. The number of canals in a given group may be as
small as 1 or as great as 8 even in different sectors of a given individual,
the latter being the largest number yet observed.

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The Philippine and Zaca series combined show that the numbers of
canals and of tentacles tend to increase (though irregularly) with the
growth of the medusa, as follows:

Diameter

Canals

Tentacles

Ratio tentacles

TO CANALS

*29 mm.

33

42

1.3

31 mm.

27

32

1.2

32 mm.

22

33

1.5

34 mm.

26

30

1.2

*36 mm.

29

28

1.0

38 mm.

30

30

1.0

*39 mm.

38

54

1.4

*40 mm.

35

42

1.2

*43 mm.

46

46

1.0

*50 mm.

31

45 .

1.5

*68 mm.

45

50

1.1

*76 mm.

38

70

1.8

* Philippine specimens.

As the larger of the Philippine specimens had well-developed gonads, it is
probable that the final number of canals is not likely to be much more than
45 in any specimen — perhaps not less than 35.

Zygocanna vagans Bigelow, central portions of discs of specimens
from Stas. 225 (A) and 233 (B, C) to show canal stripes, camera
drawings, about X 3.

1940] Bigelow: Medusae of Zaca Expeditions, 1936-1938 301

In two, only, of the specimens so far examined, was the number of
tentacles less than that of the canals; greater by at least 1 in all others.
But comparison between size, number of canals and number of tentacles
(ratios — , Table, p. 300) fails to suggest any general tendency for the
disparity in number between the two classes of organs either to increase or
to decrease with the growth of the medusa.

In the one member of the original (Philippine) series, on which a
complete count of the marginal organs was made, there were from 4 to 11
rudimentary tentacular knobs between every two large tentacles. The fact
that among the Zaca specimens the number was in one instance as small as
1 but frequently as large as 7, further emphasizes the wide variations in this
respect that are to be expected in different sectors of the margin, in
individual specimens.

The large size of the excretory papillae has already been commented
upon in the original accounts of the species (Bigelow, 1912; 1919, PI. 42,
Fig. 6), which the Zaca series confirms.

In all of the present series, the gonads (immature) are confined to the
outer i/4 or so of the radial canals. The condition, however, of the type
specimen of 39 mm. (Bigelow, 1919, PI. 43, Fig. 61) shows that they may
already occupy more than % the lengths of the canals in medusae no
larger — another evidence of variability of the species.

The subumbral gelatinous papillae, characteristic of the species, occupy
the same locations in the Zaca series as in the original specimens, being
confined to a row between every two adjacent canals and to the roof of the
manubrium. The intercanalar rows consist in some cases of entirely
separate conical prominences, in others of a sharp crest bearing such; the
number along each series varies from 1-9, as against 5-10, with a maximum
of 15 in the Philippine series. As in the latter the papillae are confined to
the inner 2/i or % of the bell diameter. The chief difference in sculpture
between the Zaca and Philippine specimens is that the latter had up to 16
rounded knobs within the limits of the manubrium, whereas the largest
number of such knobs among the Zaca series was only three. But in view of
variability in other characters this difference does not seem specifically
significant.

The most interesting aspect of the present captures is evidence of the
wide-ranging nature of Z. vagans. The facts not only that vagans has now
been found in both sides of the Pacific, but that all l’epresentatives of
Zygocanna recently taken are referable to it, raises the question whether
the regular bifurcation of canals well distal to the margin of the manubrium
described and figured by Haeckel (1879) for his Zygocanna costata from
New Guinea correctly represented the normal state, or whether it actually
represents Haeckel’s idealization of the alcoholic specimen on which he based
the species (and genus). Even in the latter case, however, there remains to
separate vagans from costata the presence of the subumbral papillae in the
former, contrasted with their absence (or at least with Haeckel’s failure to
mention them) in the latter; and of radial exumbral ridges in costata of
which there is no trace in vagans. The Zaca series does not help toward a
final decision as to whether Mayer (1910) was correct in uniting costata
with Peron & Lesueur’s (1809) old species pleuronota. However, the fact
that the members of two successive series of vagans (taken so far apart in
time and in space) lack any trace of peduncle, argues against Mayer’s
(1910, p. 339) suggestion that Haeckel’s Zygocannula (with well-developed
peduncle) is an advanced stage of Zygocanna.

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Trachomedusae.

Geryonidae.

Liriope.

Liriope tefraphylla Chamisso & Eysenhardt.

Geryonia tetraphylla, Chamisso & Eysenhardt, 1821, p. 357, PI. 27, Fig. 2.

For recent discussions of this (apparently) monotypic genus, see
Browne (1924) ; Thiel (1936) ; and Bigelow (1938).

Material : Sta. 227, 210-0 meters, one specimen about 12 mm. in diameter.

In this specimen the gonads are triangular, with their bases occupying
most of the periphery of the bell, but not yet actually in contact, i.e., it is
intermediate between the so-called “rosacea” and “compacta” stages already
pictured from the Eastern Tropical Pacific (Bigelow, 1909, PL 3, Figs. 7, 8).
Liriope with gonads of this type has so frequently been reported — especially
from the high seas — that the present example adds nothing significant to
our previous knowledge of this much-vexed genus.

Trachynemidae.

Colobonema.

Colobonema typicum Maas.

Homoeonema typicum, Maas, 1897, p. 22, PI. 3, Figs. 1-3.

For recent discussion of the synonymy of this species, see Bigelow, 1938,

p. 116.

Material : Sta. 219, 540-0 meters, 1 specimen, about 25 mm. high.

This specimen — in good enough condition for positive identification —
adds nothing to previous knowledge of the morphology of this well-known
bathypelagic medusa. Having been taken widespread in the Eastern Tropi-
cal Pacific, including the offing of Costa Rica and Gulf of California
(Maas, 1907; Bigelow, 1909), it is somewhat astonishing that the Zaca
collection yielded only a single example of it.

Halicreidae.

Halier eas.

Halicreas minimum Fewkes.

Halicreas minimum, Fewkes, 1882, p. 306.

Halicreas papillosum, Vanhoffen, 1902a, p. 68, PI. 9, Figs. 7-8; PI. 11,
Fig. 30.

For reasons for the final relegation of papillosum to the synonymy of
minimum, and for lists of the more important references, see Bigelow, 1938,

pp. 121, 122.

Material : Stas. 165, 210, 225, 227, 228, 230, 234, 15 specimens, all in
hauls from 910-0 meters.

The condition of the specimens is so poor that they add nothing to
previous accounts of this well-known species. All, however, show the char-
acteristic exumbral papillae, and it is chiefly on these that their identification
rests.

Having already been recorded at many stations in the eastern side of

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303

the Tropical Pacific (Bigelow, 1909), H. minimum was to be expected in the
region explored by the Zaca. It is, in fact, now known to be one of the most
generally distributed and frequently occurring of bathypelagic medusae in
the open oceans, in low and mid-latitudes, as discussed elsewhere (Thiel,
1936, p. 72; Bigelow, 1938, p. 122).

Narcomedusae.

Solmaridae.

The majority of recent authors have concurred in uniting in this one
family all Narcomedusae that lack gastric pockets. Ranson (1936), on the
other hand, has revived the old family designation Peganthidae, for such of
them as have the gonads localized in interradial diverticulae of the gastric
wall, as opposed to those in which they extend, ring-like, right around the
margin of the stomach. To use the conformation of the gonads as a basis for
family separation among the Narcomedusae that lack gastric pockets would,,
however, run counter to another set of alternative characters, namely, the
presence or absence (at least in one generation) of a canal system and of
otoporpae. Our knowledge of the existence of solmarids with canals and
otoporpae, but with a simple annular gonad, rests not only on Haeckel’s
(1879; 1881) account and figures (from life) of his Polycolpa forskalii from
the Red Sea, but equally on Browne’s (1916) discovery of a second unnamed
Narcomedusa of this type from the Indian Ocean. In fact, the Cunina
prolifera of Gegenbaur (for synonymy, see Mayer, 1910, p. 480), which
also has an annular gonad and otoporpae, appears also to belong to this
group, for none of the published illustrations of this species show any gastric
pockets. Stschelkanowzew (1906, p. 483) indeed states definitely that it has
none, but that its stomach, is perfectly round in life, though taking an
angular outline when preserved as is shown by Mayer (1910, p. 471, Fig.
319A). C. prolifera should then be removed from the Aeginidae to the
Solmaridae (and referred to Polycolpa if that genus be recognized, see
below) as Stschelkanowzew suggested, though he did not actually go so far
as to make this alteration.

The question is further complicated by the fact that the canals and
otoporpae, so conspicuous in the primary generation of Pegantha are lack-
ing in the second parasitic generation that develops within that genus, at
least up to the most advanced stage to which development has been followed
(Bigelow, 1909, p. 93), so that morphological diagnosis that applies to one
generation would not to the other. It therefore seems wiser to maintain
the family Solmaridae as widely inclusive for the time being; admitting,
however, that this may be only a temporary stand, in the slow and spasmodic
growth of our knowledge of the Narcomedusae.

Within the Solmaridae, as thus conceived, three groups of species appear
to be rather definitely definable: A, with simple annular gonad, lacking-
canals and otoporpae; B, with simple annular gonad, with canals, and with
otoporpae; and C, with interradial sexual sacs (whether or not combined with
an annular gonad), with canals, and with otoporpae. The first of these group-
ings is now generally referred to Solmaris; the second is the genus Polycolpa
of Haeckel (made by Mayer, 1910, a synonym of Solmaris ) ; it is with the
third that we are concerned here.

For the members of this last group, Haeckel (1879) revived the old
genera Pegasia of Peron & Lesueur (1809) and Poly xenia of Eschscholtz;
(1829), as well as proposing his new genus Pegantha, And it is certain
that in any final revision these two old generic names must be taken into
account, for we have as good grounds for identifying them with the medusae
in question, as in most of the cases in which final disposition of the eai’ly
medusan names has been made.

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Actually, it seems that the earliest description of a typical peganthid was
neither of Pegasia nor of Polyxenia, but of the Medusa mollicina of Forskai
(1775). This Peron & Lesueur (1809) later referred to their genus Faveola,
but incorrectly since the first mentioned member (hence type species) of the
latter — the Medusa pilearis of Linnaeus (1766) — was an Anthomedusa and
probably what is now known as Leuckartiara. The next description of what
was probably a peganthid (though no indication was given as to presence or
absence of a canal system nor of otoporpae) was by Eschscholtz (1829) as
Polyxenia cyanostylis. Meantime, however, Peron & Lesueur (1809) had
published a brief notice of Pegasia dodecagona. And while their account
and the subsequent illustration by Blainville (1834) are unrecognizable if
taken per se, Haeckel (1879, p. 331) from personal examination of one of
Peron & Lesueur’s original specimens, placed it in his family Peganthidae —
thereby at least implying a lack of gastric pockets, but presence of a
peripheral canal system — described its gonads and even counted the otocysts.
Our choice lies, then, between ignoring Haeckel’s diagnosis as Vanhoffen
(1908) has done (partly on the ground that it was not accompanied by an
illustration ) or accepting Pegasia as the oldest generic name applicable to
the group in question. And the latter alternative appears much the more
likely to conduce toward stability of nomenclature. In other words, we can
credit Haeckel with doing for Pegasia what Chun (1897) did for Abylopsis
tetragona Otto, among siphonophores, i.e., with establishing the status of an
old species from reexamination of a type specimen, the original account of
which had been wholly inadequate, or even erroneous.

Accepting the validity of the name Pegasia, we have still to decide
whether all known members of the group to which it belongs (i.e., solmarids
with otoporpae, with canals, and with genital diverticulae) should be referred
to it, or whether they represent more than one genus. Generic grouping based
on the number of tentacles, suggested by Vanhoffen (1908; 1911; 19121
having been proven untenable in this case, decision rests on what value in
classification is to be assigned to the conformation of the gonads. And while
Vanhoffen (1908), Mayer (1910), and subsequent authors generally have
been of the opinion that this is not of more than specific import in the group
in question, it seems wiser to reserve judgment, until the time (if ever),
when some student has the opportunity to examine actual specimens in which
an annular gonad is combined with interradial sexual sacs — i.e., referable
either to Polyxenia, or to Pegasia as redefined by Haeckel.

Accordingly, the generic name Pegantha is retained here (as earlier by
me) with the Haeckelian diagnosis, i.e., for Solmaridae in which the sexual
products are confined to interradial outpocketings from the gastric margin,
irrespective of the number of tentacles. Should it prove, in the future, that
the gonads must be discarded here, as a generic character, the name Pegasia
(or if not Pegasia, then Polyxenia ) will have priority.

1940] Bigelow: Medusae of Zaca Expeditions, 1936-1938 305

Pegantha.

Anatomically, Pegantha is one of the best known of narcomedusan
genera. Our knowledge, however, of specific relationships within the genus
has not significantly advanced since 1918, when I briefly outlined the existing
status in this respect (Bigelow, 1918, p. 395). Nor does the present series
contribute toward a final solution, for one of the two specimens of Pegantha
that it contains, belongs to a species that has already been fully described,
while the other is not in good enough condition for positive identification.

Pegantha clara R. P. Bigelow.

Text-figs. 15, 16.

Pegantha clara, R. P. Bigelow, 1909, p. 80.

Pegantha smaragdina, H. B. Bigelow, 1909, p. 90, PI. 14, Figs. 1, 2;
PI. 19, Figs. 1-9, Pis. 22-26.

To the synonymy and list of references given earlier by me (Bigelow,
1938, p. 134), add P. smaragdina, Hanitzsch, 1911, p. 225.

Material: Sta. 230, 910-0 meters, 1 specimen about 55 mm. in diameter
with 25 tentacles, in fair condition.

In its low disc without exumbral sculpture, and simple gonads (hemi-
spherical when relaxed but more or less constricted when contracted), the
Zaca specimen agrees very well with the earlier accounts by R. P. Bigelow of
the Atlantic P. clara and by me almost simultaneously of P. smaragdina
from the Eastern Tropical Pacific. The number of tentacles (25, 2 much
smaller than the others) also falls well within the recorded limits (19-34).
And this also applies to the number of otocysts, of which there are five (4 of
them with well developed otoporpae, but 1 lacking such an organ) in the
only sector in which they can be counted, and three, with otoporpae, close
together in the next, so localized as to suggest a total of 6-8. In short, it
corroborates the conclusion arrived at first by Vanhoffen (1912a) and subse-
quently by me (1938) that smaragdina cannot be distinguished specifically
from clara. In my original (1909) discussion of the former it was suggested

Text-fig. 15.

Pegantha clara R. P. Bigelow, aboral view of gonads and canal
system with polyp larvae, camera drawing, about X 7.

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that the ova (difficult of detection because of their small size) may develop
irregularly over the wall of the manubrium, as is the case according to
Stschelkanowzew (1906) in Cunina proboscidea, i.e., that it may be the
males alone that develop discrete gonads. In fact, ova have not been definitely
identified as yet, in any Pegantha, so far as I am aware. And surface views
of the sexual sacs of the Zaca specimen are at least compatible with this
suggestion, no ova being visible, even after staining, but only very closely
crowded masses of minute cells, suggesting stages in spermatogenesis.

Anatomically, the specimen is interesting as showing individual tendency
toward abnormality, for the otoporpae vary greatly in length, while one of
the peronii is abnormally broad distally, and bifid proximally, with a small
tentacle associated with the one branch but none with the other, the two
branches being separated by a canal, but one so much shorter than the others
that it fails to reach the margin of the bell (Text-fig. 15).

Text-fig. 16.

Pegantha clara R. P. Bige-
low, enlarged view of three of
the larvae shown in Fig. 15,
camera drawing, X 42.

The most interesting feature of the Zaca specimen is that its canal
system contains several polyp-larvae (Text-fig. 16) with much younger
(cleavage) stages in the mesoglea, agreeing vei’y closely with those described
by me (1909) for one of the two Albatross specimens from the Eastern
Pacific. These have not been found in any Atlantic example of the species
that has yet been examined. It is worth repeating, however, in this connec-
tion, that small medusae, resembling the 2nd generation of P. clara, have
often been observed in the gastrovascular cavity of the Mediterranean
Narcomedusa usually referred to in literature as Cunina prolifer a Gegenbaur,
but which is not a Cunina but a peganthid (p. 303). Unfortunately, the
histological condition of the Zaca specimen was not good enough to add
anything of moment to my earlier account of the developmental stages.
Their presence, however, in this second example shows that it is character-
istic of the earliest stages to be confined to the mesoglea of the exumbrella of
the parent, overlying — in this case — the part of the vascular system that is

1940] Bigelow: Medusae of Zaca Expeditions, 1936-1938 307

occupied by the older larvae. It is also interesting that the latter, like those
earlier described, show stages in budding at the aboral side.

Were it alone in P. clara that this type of development had been observed,
it might reasonably be questioned whether the resultant small medusae do
indeed belong to the developmental cycle of the large medusae within which
they occur, for we lack actual factual knowledge of the source of the infesting
larvae. Since, however, Stschelkanowzew (1906) was able to show that
similar larvae, found parasitic within Cunina proboscidea, develop from the
fertilized ova of that same species, we may assume that the same is true of
Pegantha.

In the case of Cunina, it appears, indeed, (from Stschelkanowzew’s
studies), that the larvae are the offspring of the same individuals within
which they develop. Since, however, the larvae have now been found in a
specimen of P. clara lacking any trace of gonads, and in a second in which
the latter are well developed, it seems that in this species any large individual
(whether or not mature) may become infested if in the vicinity of others
that are actually setting free their eggs and sperm in the water. And this
seems the more likely, from the fact that in this particular Narcomedusa (in
contrast to Cunina proboscidea) the earliest developmental stages are to be
found in the exumbral mesoglea. In other words, there is no necessity for
assuming that the larvae found within a given individual are its own off-
spring. But this whole matter necessarily remains open, until the fertiliza-
tion of the egg and its actual penetration have been traced.

According to Hanitzsch’ (1911) interpretation, based on Stschelkanow-
zew’s (1906) cytological studies of the oogenesis of Cunina, the mitotically
dividing amoeboid cells of P. clara (Bigelow, 1909, PI. 22, Figs. 8, 9) repre-
sent the product either of the last maturation or of the first cleavage division,
and those amitotically dividing (which later act as the nurse cells for the
embryo) are oocytes that have lost their character as sex cells.

A serious gap still remains in our knowledge of the developmental cycle
of P. clara, between the medusae of generation II (i.e., the small parasitic)
which are set free at what may be termed the “Solmissus” stage when some
3 mm. in diameter, and those of generation I, within which they have been
found. In the case of Cunina proboscidea Stschelkanowzew (1906) con-
cluded that an asexual polyp generation intervenes, both because the direct
development of the medusae of generation II into those of generation I seems
precluded by differences in structure, and because he believed he could
identify as the young of C. proboscidea (by the shape of the otoporpae) the
medusae set free from one of the two species of narcomedusan stolons that
are often found parasitic within the gastric cavity of Geryonia.6 Metschnikoff
(1874, p. 34) had in fact made this same suggestion, long before. These
stolons, as is well known, are the growth products of ciliated larvae, which
Woltereck (1905, Figs. 12-14) traced back to early cleavage stages in the
mesogloea of Geryonia. But the actual process by which the latter is para-
sitized by them has not yet been observed ; we have only Stschelkanowzew’s
suggestion, accepted by Hanitzsch (1921, p. 225), that the small medusae of
C. proboscidea of generation II are eaten by the Geryonia and that their
eggs and sperm come together within the cavities of the latter.7

Perhaps a stolon-generation intervenes, in Pegantlia clara also, between
the medusae of generations I and II. But it is equally possible that in this
species the large medusae of generation I develop directly from the fertilized
eggs of the small “solmarid” medusae of generation II.

6 These are the stolons usually referred to as “Cunina parasitica Metschnikoff.”

7 The development of Cunina proboscidea has been discussed not only by Metschnikoff (1886)
and Stschelkanowzew (1906), but subsequently in great detail by Hanitzsch (1911-1921).

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? Pegantha martagon Haeckel.

Pegantha martagon, Haeckel, 1879, p. 332; Bigelow, 1909, p. 83, PI. 18,
Figs. 1-8; Mayer, 1910, p. 443.

Pegantha simplex, Bigelow, 1904, p. 260, PI. 5, Figs. 19, 20.

Material: Sta. 234, 1 fragmentary specimen, about 18 mm. in diameter,
with 12 tentacles.

The condition of this specimen is not good enough for positive identifi-
cation, further than that absence of gastric pouches and presence of a well-
developed canal system prove it a Pegantha. It is tentatively referred to
P. martagon for the following reasons : 1, the bell is high-rounded, but above
the level of the tentacular insertions, shows no trace of the exumbral sculp-
ture that is so prominent in P. triloba; 2, the gonads, while showing a cer-
tain amount of irregular wrinkling and lobing are not definitely subdivided.
The margin is so much damaged as to prevent counting the otocysts; it is
unfortunate, in particular that the otoporpae have been entirely destroyed,
because the structure of these is one of the inspects in which P. martagon
differs the most sharply from P. triloba (Bigelow, 1909).

Solmissus.

Ranson (1936, p. 206 ) in his recent discussion of Solmissus, is of the
opinion that S. incisa Fewkes is a “forme geante” of S. albescens Gegenbaur,
but that the Indo-Pacific S. marshalli Agassiz & Mayer is a well defined
species rather than a variety of albescens as I (1909; 1919) formerly sug-
gested.

Solmissus marshalli Agassiz & Mayer.

Solmissus marshalli, Agassiz & Mayer, 1902, p. 151, PI. 5, Figs. 23-25;
Bigelow, 1909, p. 64, PI. 16, Figs. 5-6, PI. 21, Figs. 4, 6-8; 1919, p. 329;
1938, p. 129; Mayer, 1910, p. 484; Thiel, 1936, p. 68; Ranson, 1936, p. 208.

Solmissus punctatus, Mayer, 1906, p. 1133.

Solmissus flavescens, Vanhoffen, 1908, p. 56, PI. 2, Figs. 9-10, PI. 3,
Figs. 20-22.

( Non Solmissus flavescens Kolliker).

Material : Sta. 233, 2 specimens respectively about 34 mm. and 52 mm.
in diameter.

Tentative identification of these two specimens rests on the fact that,
in each case, traces of the outlines of gastric pockets in the radii of the
tentacular roots are still to be seen, combined with a flat-lenticular type of
bell characteristic of Solmissus of the albescens-marshalli gi'oup. Failing any
trace of the otocysts, and lacking knowledge of the precise contour of the
gastric pockets, they are referred to marshalli on geographical grounds, be-
cause albescens— -the common Solmissus of the Mediterranean — has not yet
been taken in the Pacific.

Unfortunately, the condition of the specimens is so poor that the number
of tentacles cannot be stated exactly, further than that in one case 8 large
tentacular roots were detected and in the other nine. But these numbers are
so much smaller than is usual in full-grown marshalli (usually 12-16) as to
suggest that there had been more tentacles in life, but that some had been
obliterated.

It has long been known that marshalli (or the marshalli variant of
albescens ) is widespread in the tropical and subtropical belts of the Pacific,
Indian and Atlantic oceans, while Ranson (1936) has recently reported it
from the Mediterranean.

1940] Bigelow: Medusae of Zaca Expeditions, 1936-1938 309

Aeginidae.

Aeginura.

Recent authors agree that all representatives of Aeginura that have
come to hand, subsequent to Haeckel’s (1879) original account of the genus,
are referable to the single species grimaldii Maas (1904) — of which weberi
Maas (1905) and grimaldii var. mundi, guinensis and obscura of Vanhoffen
(1908) are synonyms. The relationship, however, of grimaldii to the medusa
( myosura ) on which Haeckel founded the genus, and to Haeckel’s Cunoctona
nausithoe and C. lanzerotae (which are described as having otoporpae) is
still in doubt; may, in fact, always remain so. Having recently (1938, p. 132)
discussed this question, there is no need for me to revive it here.

Recent students of Narcomedusae have been unanimously of the opinion
that the chief generic character of Aeginura is the constant presence of 8
tentacles and 16 gastric pockets; and it has seemed at least probable that
the presence of small secondary tentacles on the margin is equally diag-
nostic, for while Haeckel (1879; 1881) neither mentioned nor pictured any
such for myosura, their apparent absence in the latter may well have been
due to the mutilated state of the unique (alcoholic) specimen. But it is now
necessary to redefine the genus, or to institute a new one, for the Zaca collec-
tion contains a specimen that agrees with Aeginura in the arrangement of
the gastric pockets in relation to the large primary tentacles, and in the
presence of small secondary tentacles on the bell-margin between the peronii,
but in which the number of tentacles, and of primary gastric pockets is
eleven, instead of eight. Decision in this case obviously depends on the rela-
tive phylogenetic weight to be attached to metameral number, as compared
with the presence of the secondary tentacles. About this opinions may differ.
Since, however, the number of primary tentacles in the Narcomedusae may
either be strictly determinate, as for instance in Solmundella and Cunoc-
tantha, or highly variable as in some species of Pegantha and Cunina,
whereas the presence of secondary tentacles is a feature peculiar (so far as
yet known) to Aeginura, it seems to me wisest to use the latter as the
primary generic character for that genus, expanding the latter to include
species with more than 8 tentacles.

Aeginura beebei sp. nov.

(Text-figs. 17-20).

Material : Sta. 227, 910-0 meters, 1 specimen, about 100 mm. in diameter
in fair condition.

This specimen (Text-figs. 17-20) is perhaps the most interesting
find among the Zaca medusae, for while in general appearance it resembles
Solmissus, critical examination shows that it actually agrees, in all signifi-
cant anatomical characters, with the bathypelagic medusa Aeginura grim-
aldii. It differs, however, from the latter in the fact that it has (as just
stated) a larger number of antimeres (11) than has ever been recorded
for Ae. grimaldii in which the normal 8 has never been exceeded — though
Ranson (1936, p. 210) records one specimen with as few as five.

Most at least of the secondary tentacles are intact, and certainly a ma-
jority of the otocysts : the numbers of marginal organs counted per antimere,
being as follows: —

Tentacles 31221131212
Otocysts 52412363322

Thus, there is no evidence that the number of secondary tentacles ever
exceeds 3 per sector, while in at least 5 of the sectors, the condition of the

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[XXV : 19

Text-fig. 17.

Aeginura beebei, sp. nov., lateral-oblique view of type specimen,
slightly more than natural size.

Text fig. 18.

Aeginura beebei, sp. nov., oral-oblique view of type specimen,
slightly more than natural size.

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311

margin was so good as to allow positive assertion that there was only one
secondary tentacle. In grimaldii (or its synonyms), the recorded number
per marginal sector ranges from 2 to 5, the total thus being about the same
as in the new species.

Structurally, the secondary tentacles of beebei differ from the primary
tentacles in the fact that they lack entodermal roots, while their bases are
much swollen (Text-fig. 19), hence there is no reason to suppose that the
former ever develop into the latter. In detail (i.e., in the core of c-hordate
entodermal cells and in the slender filament) they agree very closely with
Vanhoffen’s (1908, PI. 9, Fig. 28) illustration of the corresponding struc-
tures in his “Cunoctana obscura,” which was undoubtedly identical with Ae.
grimaldii.

A secondary tentacle is flanked, in so many cases, by an otocyst on
either hand (Text-fig. 19) that this appears to be the basic state. However,
the presence of three otocysts each, in two of the marginal sectors but of
only one secondary tentacle, proves that neither the numerical relationship
between the two, nor their relative locations on the margin, are determinate.
A similar variation in number and spacing of otocysts has already been
recorded for grimaldii (Vanhoffen, 1908). On the average, the number of
otocysts per sector (there being fewer secondary tentacles) is somewhat
smaller in the new species than in grimaldii, but the difference may not be
great enough to be of diagnostic significance. In their structure (Text-fig.
20) the otocysts are of the usual aeginid type.

Text-fig. 19.

Aeginura beebei, sp. nov., aboral view of portion of margin, with
secondary tentacle and otocyst, camera drawing, X 66.

The specimen corroborates previous observations to the effect that in
Aeginura the interradial notches that indent the margin of the gastric
cavitj^are much deeper than the perradial, such for example was the case in
the specimen from the East Indies figured by Maas (1905) as Ae. weberi;
in the Albatross specimens from the Eastern Tropical Pacific and North-
western Pacific (Bigelow, 1909, p. 80; 1913, p. 62), and likewise in
Maas’ (1904, PI. 3, Fig. 20) original figure of Ae. grimaldii, in which the
interradial notches are pictured as about twice as deep as the perradial.
Haeckel (1879; 1881), on the other hand, pictures the reverse for Ae.
myosura, this being one of the reasons for doubting whether this species
has been seen subsequently. In the Zaca specimen of Ae. beebei, the inter-
radial notches are about 16-17 mm. deep, but the perradial only 4-5 mm. The
gastric pockets themselves are rhomboid in outline (Text-figs. 18, 20),
broader distally than proximally, with the outer angles of adjacent pockets
very close together. Surface views, in fact, suggest that they may even be
confluent, which would be unique among Narcomedusae. But the condition

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of the specimen is not good enough for positive determination of this point,
though it was easy to trace the general outlines of the pockets by surface
examination, verified by the injection of air. A specific character, which,
while minor in itself, is again unique, so far as known among aeginids, is
that the interradial margins of the pockets are jagged or lobate (Text-fig.
20). Neither surface view nor injection yielded any evidence of the presence
of a peripheral canal system, in which beebei agrees with grimaldii. The
central part of the stomach has been largely torn away — the remnants, how-
ever, suggest a simple circular mouth (Text-fig. 18).

Text-fig. 20.

Aeginura beebei, sp. nov., oral view of gastric pockets, about X 3.

The gastric wall, at the interradial sides of the pockets, is thrown into
a series of folds directed obliquely distal (Text-figs. 18, 20), no doubt the
sites of the sexual products. Surface views suggest that the specimen is a
male — at least no individual eggs are to be seen. And to judge from analogy
with Ae. grimaldii, the latter might be expected to be so large as .to be
conspicuous.

A stinking feature of the specimen is that the subumbrella is deeply
indented with triangular depressions along the lines of the interradial septa
that separate the gastric pockets, and also with furrows along the perradial
septa. Thus, were the gastric wall to be stripped off, the subumbral zone
occupied by the pockets would appear as alternate triangular prominences
and depressions, the former with their bases directed distally, and shallowly
notched in the perradii, the latter with their bases directed toward the center
of the bell. The bell is low discoid in form, about Ms as high (18 mm.) as
broad, with very shallow bell cavity, i.e., hardly deeper than the breadth of
the marginal groove. There is no exumbral sculpture, other than the shallow
furrows along the lines of the peronii that are a common feature among
Narcomedusae.

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313

The gastric wall in the preserved state (formalin) is pale purplish.
Otherwise the specimen is colorless.

The features in which Ae. beebei most sharply differ from Ae. grimaldii
are 1, larger (and probably indeterminate) metameral number; 2, presence
of prominent subumbral sculpture; 3, much larger size at maturity; 4, entire
lack of the distinctive bathypelagic pigmentation.

Obviously, the one capture, having been made in an open net, yields no
information as to the normal bathymetric occurrence of the species, but the
fact that it is so faintly pigmented suggests that it is a creature of the
upper waters.

Aeginura grimaldii Maas.

Aeginura grimaldii, Maas, 1904, p. 38, PI. 3, Figs. 19-28.

See Bigelow, 1938, p. 132, for recent discussion of Aeginura with the
reasons why it seems wiser to retain Maas’ species grimaldii as distinct
rather than to unite it with Haeckel’s (1879) old species myosura, nawsithoe,
and lanzerotae as Thiel (1936) has done in his historical survey.

Material : Sta. 227, 910-0 meters, 1 fragmentary specimen about 23
mm. in diameter.

Sta. 228, 910-0 meters, fragments.

Aeginura grimaldii is represented so regularly in collections from deep
water that it must be one of the most generally and frequently distributed
of bathypelagic medusae. As a rule, however, the specimens taken have
been badly damaged amid the mass of other plankton. And the present ex-
amples are no exception, for the one (Sta. 228) is made recognizable only
by the pigmentation in combination with the fact that a few of the conspicu-
ous white ova are still intact, while the other — in a somewhat better state — -
is so battered that only one of the secondary tentacles is still to be seen, and
none of the otocysts. Hence they add nothing to previous accounts based on
better material.

Aegina.

Aegina citrea Eschscholtz.

Aegina citrea, Eschscholtz, 1829, p. 113, PI. 10, Fig. 3.

See Bigelow, 1938, p. 130, for the most recent discussion of this genus;
Mayer, 1910, pp. 451-454, and Thiel, 1936, p. 73 (“Ae. rosea”), for lists of
references to the various named representatives of Aegina, all of which
probably refer to the one species, citrea.

Material : Sta. 210, 910-0 meters, 1 specimen about 17 mm. in diameter.

Sta. 230, 910-0 meters, 1 specimen about 22 mm. in diameter.

Both of these specimens are of the so-called “rosea” type, i.e., with the
margins of the 8 adradial gastric pockets entire. A point still to be deter-
mined is the range of variation in number of otoliths that is normal for the
species. In this connection it is interesting that in one quadrant of the
specimen from Sta. 230 there are 14 of these organs — -this probably being
the complete number, since they are evenly spaced. This is somewhat fewer
than recorded by Vanhoffen (1908a), namely, 16-20 per quadrant, but a
larger number than reported by most other authors.

Beyond this, the present series calls for no special comment, for Ae.
citrea has repeatedly been described and figured from specimens in better
condition.

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SCYPHOMEDUSAE.

CORONATA.

Periphyllidae.

Periphylla.

Periphylla hyacinthina Steenstrup.

Periphylla hyacinthina, Steenstrup, 1837.

For synonymy and recent discussion see especially Mayer (1910, p. 544,
“P. hyacinthina”) , p. 546 (“P. hyacinthina forma regina”) ; Broch, 1913,
p. 4; Stiasny, 1934, p. 342; and Bigelow, 1938, p. 155.

Material : Stas. 148, 158, 210, 225, 227, 228, 230, 233, 234; a total of
22 specimens from about 15 to about 150 mm. in diameter at the level of the
coronal furrow, all in hauls from 910-0 meters.

It now seems certain that all known representatives of the genus Peri-
phylla actually belong to a single species. It also seems established that in
general the “dodecabostrycha” form (small; low-rounded bell; transparent
peripheral zpne), the “hyacinthina” form (peripheral zone pigmented, bell
usually high-arched, pointed), and the “regina” form (dense pigmentation,
low-domed bell) represent successive stages in growth. But it is still an
open question whether every Periphylla (if it survives) normally passes
from the “hyacinthina” stage into the “regina” stage, or in what degree this
basic succession may be confused by individual variation as to development
of pigment, and as to bell-shape in relation to size. The fact that the largest
specimens among the considerable series collected by Dr. Beebe near Ber-
muda during the years 1929 and 1930 (Bigelow, 1938, p. 156) were less
than 60 mm. in diameter (at level of coronal furrow) but still referable to
the “hyacinthina” stage also suggests that certain populations may fail to
develop beyond this, before they die. It is in fact likely that this may apply
to a considerable percentage of the individuals among other populations,
some of whose members do finally attain the “regina” stage. Stiasny (1934,
p. 361) has pointed out that the latter (i.e., the largest adults) have as a
rule been taken well out from any coast, whereas the distribution of the
“dodecabostrycha” stage makes it likely that “the small stages of Periphylla

have their origin on the continental slope or near the coast and that

they are driven into the open sea and sink slowly to greater depths.” But
such a tendency toward offshore dispersal can hardly explain the absence
(or at least great scarcity) of large specimens near Bermuda, for the collec-
tion was made at a station as typically oceanic as one could ask; as much so
certainly as the Indo-Pacific localities, where the “regina” stage has been
taken, namely the Galapagos-Cocos Island region, the Zaca stations, and
the Gulf of Aden (Maas, 1897; Bigelow, 1909; 1928; Vanhoffen, 1902). To
what extent such variations may be correlated with seasonal differences be-
tween generations in a given region is equally a question for the future.

The most interesting feature of the present series is that it contains
two large specimens, respectively about 85 mm. and about 110 mm. in
diameter at the level of the coronal furrow, that agree very closely in low-
rounded bell, rectangular pedalia, stout tentacular bases and dense pigmen-
tation8, with the specimens of the “regina” stage recently recorded by
Stiasny (1934). The fact that in the 85 mm. specimen (alcoholic) the pedalia
are rather sharply marked off from the coronal zone but that in the 110 mm.
example (in formalin) and also in the largest of all (about 150 mm., frag-
mentary) they merge insensibly into the latter just as in the Discovery
examples, is additional evidence (if any were needed) that characters that
have been used as specific among medusae may actually represent nothing
more than the results of different methods of preservation.

8 Except where obviously rubbed off.

1940] Bigelow: Medusae of Zaca Expeditions, 1936-1938 315

Atollidae.

Atolla.

Atolla wyvillei Haeckel.

Atolla wyvillei, Haeckel, 1880, p. 488; 1881, p. 113, PI. 29, Figs. 1-9.

See Broch (1913, p. 13), Browne (1916, p. 203), Bigelow (1928, p. 505;
1938, p. 160), and Stiasny (1934, p. 365), for early synonymy, and Bigelow
(1938, p. 162), for a recent summary of distribution.

Material : Stas. 210, 219, 225, 228, 230, 233, 234; 24 specimens, all in
hauls from 910-0 meters. The series range in diameter from about 11 mm. to
about 95 mm.

This series — all are in more or less fragmentary condition — adds little
except by way of confirmation to what is already known of this cosmopolitan
bathypelagic medusa. Neither do the records extend the known range, for
A. wyvillei had already been recorded widespread in the eastern side of the
Pacific, not only in low latitudes, but far to the north as well as far to the
south of the areas crossed by the Zaca.

Among such of the Zaca series as are in good enough condition to show
the state of the disc, it is more or less furrowed in eleven, but smooth in
four, a considerably smaller proportion of smooth-disced than reported near
Bermuda (Bigelow, 1938, p. 161), but almost precisely the same proportions
as found by Kramp (1924) in the Mediterranean. However, various collec-
tions, combined, from the Antarctic (Stiasny, 1934), Mediterranean
(Kramp, 1924), and from various localities in the North Atlantic (Broch,
1913; Bigelow, 1928; 1938) have contained about as many of one type as the
other (230 smooth, 233 notched).

Unfortunately, the smallest specimens were not in good enough condi-
tion to show whether the gonads had appeared, as they commonly do by the
time a diameter of 10 mm. is reached. The presence, however, of one speci-
men of 25 mm., in good condition but showing no trace of gonads, proves
that some Atollas may reach a considerable size before the sexual organs
appear; a retarded development for which no explanation is as yet forth-
coming.

DISCOPHORA.

Pelagidae.

Pelagia.

Pelagia noetiluca Forskal

Medusa noetiluca, Forskal, 1775, p. 109.

For summary of synonymy, see Bigelow, 1938, p. 164.

Material : Stas. 219, 227, 230, 233, 234, and Gorgona Island, 34 speci-
mens, about 20 mm. to about 75 mm. in diameter, in hauls at the surface
and from 910-0 meters.

All of the smaller specimens are in very fragmentary condition. The
two largest, however, (about 65 mm. and about 75 mm. in diameter) are
well preserved.

These specimens of this cosmopolitan inhabitant of warm seas all fall
well within the known limits of variation of P. noetiluca. In fact, it has
yet to be proven that this does not apply to every specimen of Pelagia from
whatever source, that has yet been critically examined. Hence, the general
agreement, by recent authors (see especially Kramp, 1924; Stiasny, 1934;
Bigelow, 1928; 1938) that the genus, as now known, is monotypic, and that

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some designation other than “species” must be invoked to designate the
numerous variations as to form and distribution of exumbral nematocyst
knobs, on which specific names have been based.

Stomolophidae.

Stomolophus.

Stomolophus illustrates the situation — not uncommon among medusae —
where the Linnaean system of classification cannot adequately represent the
existing facts, for while the named representatives of the genus differ
widely, they are connected by so continuous a series of intermediates in all
characters supposedly diagnostic that it appears impossible to define distinct
species in the ordinary sense. And, correspondingly, the history of the genus
has been one of alternate multiplication and reduction. Thus, Mayer (1910)
reduced the five named forms to one species meleagris L. Agassiz and one
variety of the latter ( fritillaria Haeckel), while I (Bigelow, 1914) concluded
that there was no reason to retain fritillaria even as a varietal name. Stiasny
(1921; 1922), however, again separated fritillaria from meleagris, basing
the distinction on form of the bell, number of marginal lappets and color,
and especially on the form and size of the scapulettes which he considered a
reliable anatomical character. More recently, however, he (Stiasny, 1931)
has found from examination of excellent preserved specimens from British
Guinea and Trinidad, that while extremes of the “fritillaria” and “melea-
gris” forms are far apart, all the characters formerly considered specific are
so variable that he “der Bigelowischen Ansicht dass nur ein einziger ampla
Species von Stomolophus besteht, nicht mehr so ablehnend gegenliber stehe
als friiher” (Stiasny, 1931, p. 175). The union of meleagris and fritillaria
may therefore be allowed to stand, at least provisionally. Unfortunately, the
present series does not afford any basis for the critical study that Stiasny
(1922) thinks needful for eventual revision.

Stomolophus meleagris L. Agassiz.

Cephea rhizostoma, Gibbes, 1847, vol. 3, p. 183 ( non Lamark, 1817).
Stomolophus meleagris, L. Agassiz, 1860, PI. 14, Figs. 1-8 ; 1862, p. 138,
151; A. Agassiz, 1865, p. 40; Haeckel, 1880, p. 599; Mayer, 1910, p. 710, PI.
75, PI. 76, Figs. 1-3; Bigelow, 1914, p. 240; Stiasny, 1921, p. 170; 1922,
p. 57; 1922a, p. 499; Boone, 1933, p. 48, PI. 10.

Stomolophus agaricus, Haeckel, 1880, p. 599.

Stomolophus fritillaria, Haeckel, 1879, PI. 35, Figs. 1-9; 1880, p. 598;
Von Lendenfeld, 1884, p. 292; Vanhoffen, 1888, p. 31, 42; Stiasny, 1921, p.
170; 1922, p. 55; 1931, p. 175.

Stomolophus meleagris var. fritillaria, Mayer, 1910, p. 711.

Stomolophus chunii, Vanhoffen, 1888, p. 31, PI. 3, Figs. 4, 5, PI. 4, Fig.
1; Trinci, 1906, p. 1.

Brachiolophus collaris, Haeckel, 1880, p. 597 (juv.).

Material : Sta. 220, surface, 1 specimen about 23 mm. in diameter.

Sta. 219, 540-0 meters, 1 specimen about 40 mm. in diameter.

The larger of these specimens is at about the stage pictured by Mayer
(1910, PI. 75, Fig. 2). They do not add anything of importance to the ac-
counts of the young stages given by him, and by Stiasny (1922a).

Pacific localities of record for Stomolophus meleagris had already in-
cluded the Bay of Panama (Vanhoffen, 1888), the coast of Costa Rica
(Haeckel, 1880, as S. agaricus), and southern California (Bigelow, 1914).

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317

It was therefore to be expected in the waters traversed by the Zaca. The
genus is so far known only from the two coasts of America, North and
South, in low and mid-latitudes.

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Ranson, G.

1936. Meduses provenant des campagnes du Prince Albert ler de Monaco.
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Russell, F. S.

1938. The Plymouth offshore Medusa fauna. Jour. Mar. Biol. Assoc., Vol. 22,
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Steenstrup, J. J. S.

1837. Acta et Catalogus Musei Hafniensis. Hauniae. (not accessible).
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1921. Studien fiber Rhizostomeen Capita Zoologica, Vol. 1,

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1922. Ergebnisse der Nachuntersuchung einiger Rhizostomeen — - Typen
Haeckel’s und Chun’s aus dem Zoologischen Museum in Hamburg.
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Zur Kenntniss der Entwicklung von Stomolophus meleagris L. Agassiz.
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textfigs.

Stschelkanowzew, J.

1906. Die Entwickelung von Cunina proboscidea Metschn. Mitth. Zool. Stat.,
Neapel, Vol. 17, Pt. 4, pp. 433-486, Pis. 28, 30.

Thiel, M. E.

1936. Systematische Studien zu den Trachylinae der Meteorexpedition. Zool.
Jahrb. Abt. Syst., Vol. 69, pp. 1-92, 18 textfigs.

Trinci, G.

1906. Sopra una Discomedusa del Golfo di Paria (America del Sud). Ann.
Mnseo Zool. R. Univ. Napoli (2), Vol. 2, No. 9 pp. 1-4.

Uchida, Tohru.

1927. Studies on Japanese Hydromedusae I. Anthomedusae. Jour. Fac. Sci.
Imp. Univ., Tokyo, Sect. IV, Zool., Vol. 1, Pt. 3, pp. 145-241, Pis.
10-11, 47 textfigs.

Vanhoffen, Ernst.

1888. Untersuchungen fiber Semaeostome und Rhizostome Medu: en. Biblio-
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Bigeloiv: Medusae of Zaca Expeditions, 1936-1938

321

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textfigs.

Olsen: New Species of Trematodes

323

20.

Two New Species of Trematodes ( Apharyngostrigea bilobata :
Strigeidae, and Cathaemasia ny die oracis : Echinostomidae) from
Herons, with a Note on the Occurrence of Clinostomum
cam panulatum (Rud. ) h 2

O. Wilford Olsen3

(Plate I).

During the course of parasite examinations on black-crowned night
herons ( Nycticorax nycticorax hoactli (Gmelin)) and great blue herons
(. Ardea herodias herodias Linn.), originating from three widely separated
rookeries in Minnesota, two new species of trematodes belonging to the
families Strigeidae and Echinostomidae were collected.

Family Strigeidae.

A number of strigeid flukes collected from both the black-crowned night
heron and great blue heron constitute a new species for which the name
Apharyngostrigea bilobata n. sp. is proposed, the name being suggested by
the bilobed character of the ovary.

Apharyngostrigea bilobata n. sp.

Description: Body length 2.8-4 mm., forebody 0.766-1.33 mm. long by
0.633-0.8 mm. wide, hindbody 1.76-2.95 mm. long by 0.266-0.65 mm. wide.
Forebody approximately cubic, hindbody cylindrical, narrowing somewhat
posteriorly, caudal extremity truncated. Holdfast organ well developed, fill-
ing forebody, composed posteriorly of a cup-like structure with anterior
half forming a smaller dorsal and a larger ventral lamella. Adhesive gland in
posterior extremity of adhesive organ which is at junction of two body
regions, strongly lobulated, posterior third composed of a single large piece,
anterior two-thirds composed of numerous small lobes, shape longitudinally
oval, size 160-266 g wide by 240-333 g long. Oral sucker 76-164 g wide by
124-208 g long; acetabulum 120-280 g wide by 132-300 g long; pharynx ab-
sent, oesophagus extends to level of anterior margin of acetabulum, intestinal
crura extend along ventral surface of body to its caudal extremity. Testes
tandem, multilobated, large, fill third quarter of body length, vasa efferentia
unite at anterior margin of first testis forming the voluminous, convoluted
proximal portion of vas deferens which narrows greatly in diameter, then
passes caudad along median ventral surface of testes and forms large coiled

1 Paper No. 1734, Scientific Journal Series, Minnesota Agricultural Experiment Station,
St. Paul.

2 In cooperation with the Division of Game and Fish, Minnesota Conservation Department.

3 Resigned August 15, 1939, to enter the U. S. Bureau of Animal Industry, Zoological Division.

324

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[XXV :20

seminal vesicle which fills area between hind testis and genital cone, entering
genital pore from dorsal side. Ovary smooth, transversely elongated, bilo-
bated, with deep cleft in posterior margin, located near middle of body; width
180-483 g, length 124-400 g\ oviduct originates on posterior-ventral surface
of ovary passing dorso-caudad through cleft to dorsal surface of latter where
Laurer’s canal is given off, then swings to left side of front testis and
caudad along its lateral margin to Mehlis’ gland. Ascending limb of uterus
extends cephalad from Mehlis’ gland along ventral side of anterior testis
and ovary, rising to dorsal side of body at anterior margin of ovary and
continuing cephalad to near holdfast gland where it bends caudad to form
descending limb which passes along ventral side of body to genital cone,
entering common genital duct from ventral side. Mehlis’ gland lies between
testes, dextrad from median line. Yolk reservoir median, at caudal margin
of Mehlis’ gland. Vitelline follicles large, spread over entire surface of
body and lips of holdfast except dorsal to holdfast gland and reproductive
organs and in ventral surface of forebody. Follicles fewer laterally in region
of ovary and testes, more abundant ventrally and laterally between holdfast
gland and ovary; they encircle body caudad from hind testis. Ova 56-64 g
wide by 83-88 g long.

Host: Nycticorax nycticorax hoactli (Gmelin).

Ardea herodias herodias Linn.

Habitat: Small intestine.

Locality : Owatonna, Minnesota, U. S. A.

Type Specimen: U. S. Nat. Mus. Helm. Coll. No. 9279; paratype: 9280;
also Helm. Coll. Div. Ent. and Eco. Zool., University of Minnesota.

Discussion.

The genus Apharyngostrigea was erected by Ciurea (1927) to receive
Distoma cornu Goeze (Zeder) 1800. Szidat (1929) added A. brasiliana Szidat,
1929, and Distomum simplex Johnston, 1904. In his description of D. simplex,
Johnston (1904: 112) states “the oral and ventral suckers are large and
equal in size . . . ; the pharynx is smaller” but since his figures do not show a
pharynx Szidat concluded that “Trotz dieser Bemerkung scheint auch bei
dieser Art der Pharynx ganz zu fehlen, da Johnston 1904 ihn in seinen
Figuren nicht abbildet.” The correctness of Szidat’s conclusion in regard to
the absence of the pharynx on the grounds that it is not figured might justly
be questioned and until an examination of Johnston’s material has been made,
Distomum simplex should be considered as a provisional member of the
genus. Dubois (1938) included Holostomum repens Chase, 1921, from
herons.

Subsequent to the appearance of Szidat’s paper, the descriptions of four
additional species of Apharyngostrigea have appeared. These, together with
Apharyngostrigea bilobata n. sp., are included in the following key, which
will serve to differentiate them. A. bilobata may be readily distinguished
from all the known species by the strongly bilobated ovary with its deep
posterior cleft.

Key to the Known Species of Apharyngostrigea.

1. Testes with margins smooth or nearly smooth, rounded or ovoid 2

Testes with margins deeply lobated, variously shaped 3

2. Testes very large, being 800 g in diameter; oral sucker large, being 740 g
in diameter; from Cancroma cochlearia, Brazil A. brasiliana Szidat, 1929
Testes small, being 336-400 g and 330-338 g in diameter; oral sucker 180
g in diameter; from cattle egret, India A. egretti Verma, 1936

1940]

Olsen: New Species of Trematodes

325

3. Testes bilobated, horse-shoe shaped ; anterior limit of vitellaria at level of
acetabulum, ovary and testes in posterior half of hindbody; from Circus

macrurus, Syria A. flexilis Dubois, 1934

Testes multilobated, not horse-shoe shaped; ovary at middle of hindbody
or cephalad 4

4. Oral sucker and acetabulum equal or about equal in size, ratio being as

1:1 - 1:1.1 5

Acetabulum distinctly larger than oral sucker 6

5. Ovary near middle of hindbody, walls of hindbody nearly parallel, not

bulged by testes; oral sucker and acetabulum equal in size; from Noto-
phoyx novae hollandiae, Australia A (?). simplex (Johnston, 1904)

Ovary in posterior portion of first quarter of hindbody ; testes enormous
in size with numerous lobes, causing hindbody to bulge greatly in two
middle quarters; oral sucker slightly smaller than acetabulum, ratio as
1:1.1; from Herodias timorensis, Philippine Islands

A. garciai Tubangui, 1933

6. Ovary bilobated, being deeply cleft on posterior margin ; from Nycticorax

nycticorax hoactli and Ardea herodias herodias, North America

A. bilobata n. sp.

Ovary not bilobated 7

7. Vitellaria do not reach to acetabulum; body large, being 6 mm. long;

from Notophoyx novae hollandiae A. repens (Chase, 1921)

Vitellaria extend cephalad from acetabulum 8

8. Acetabulum about twice size of oral sucker; body large, being 6 mm. or
more in length; from Ardea cinerea and A. purpurea, Europe

A. cornu Goeze (Zeder), 1800
Acetabulum not twice as large as oral sucker, ratio as 1:1.5; body small,
up to 3.5 mm. long; from Ardeola ibis ibis, Egypt A. ibis Azim, 1935

Family Echinostomidae.

Two specimens of flukes collected from one black-crowned night heron
belong to the genus Cathaemasia Looss, 1899. The name Cathaemasia
nycticoracis n. sp. is proposed.

Cathaemasia nycticoracis n. sp.

Description : Body elongated with ends rounded, anterior end somewhat
antenuated cephalad from acetabulum; length of body 5.3-5. 4 mm., width at
level of ovary 1.05-1.06 mm. Cuticle without spines. Oral sucker subterminal,
416 p wide by 416-466 p long. Acetabulum 700 p wide by 633-650 p long,
located in anterior part of second quarter of body length. Prepharynx absent,
pharynx 208-216 p wide by 266-292 p long; oesophagus 466 p long, of uneven
diameter, being strongly constricted in some parts while others are greatly
distended, lined with intestinal epithelium; intestinal caeca extend to caudal
end of body, with small evaginations of wall. Testes and ovary in third
quarter of body length. Testes small, tandem, about equal in size, compact
and with 6-9 short dendritic processes each, anterior testis 450 p wide by
366-433 p long, posterior testis 416-450 p wide by 416-483 p long. Cirrus
retracted, seminal vesicle about 400 p long by 128 p in diameter. Genital
pore median, at anterior margin of acetabulum. Ovary transversely elong-
ated, separated from anterior testis by Mehlis’ gland and seminal receptacle,
300-433 p wide by 166-200 p long; seminal receptacle oval, 216 p wide by
150 p long, located at posterior margin of ovary; Mehlis’ gland at level of

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Zoologica: New York Zoological Society

[XXV :20

and sinistrad from seminal receptacle; uterus short, its coils lie between
ovary and acetabulum. Laurer’s canal present. Vitellaria consisting of
rather large follicles, extending from between caudal margin of acetabulum
and ovary to posterior end of body. Caudad from hind testis, the follicles
meet medially. Excretory vesicle with shallow, lateral evaginations. Ova
91-95 g long by 42-53 g wide.

Host: Nycticorax nycticorax hoact.li (Gmelin), Ardea herodias herodias

Linn.

Habitat: Small intestine.

Locality: Owatonna, Steele Co., Minnesota; Cheboygan County, Michi-
gan (Michigan Biological Station).

Type Specimen: No. 9281 U. S. Nat. Mus. Helm. Coll.; paratype: Helm.
Coll. Ent. and Eco. Zool., University of Minnesota, No. 4:14.

Discussion.

In his discussion of the genus, Harwood (1936) pointed out the similari-
ties existing between Cathaemasia Looss, 1899, and members of the family
Echinostomidae in the nature of the excretory system and the presence of
an epithelial rather than a cuticular lining of the oesophagus. For this rea-
son, he transferred it from the family Cathaemasiidae Furhmann, 1928,
back to the family Echinostomidae where Odhner (1926) had placed it
originally. Cathaemasia together with Mehlisia Johnston, 1913, and Ste-
phanopr oroides Price, 1934, lack the spined collar of the other members of
the family Echinostomidae.

Cathaemasia nycticoracis differs from all the known species of the genus
in (1) the absence of cuticular scales, (2) in the very compact nature of the
body of the testes, both of which are confined to the third quarter of the body
length, and from which radiate simple, stubby branches, (3) the equatorial
position of the ovary, and (4) the vitellaria fuse medially caudad from the
hind testis. It resembles C. reticulata (Wright, 1879) in that the uterus is
relatively short.

The following key, adopted from Harwood (1936), will further differen-
tiate the species.

Key to the Species of Cathaemasia.

1. Body without scales; ovary equatorial; testes with very short, simple
branches radiating from large compact body, confined to third quarter of
body length; vitelline follicles fuse medially caudad from hind testis;
from Nycticorax nycticorax hoactli and Ardea herodias herodias, North

America. C. nycticoracis n. sp.

Body scaled; ovary definitely pre- or postequatorial ; testes consist pri-
marily of long slender branches; extend into extreme posterior part of
body 2

2. Ovary preequatorial ; uterus relatively short; cuticular spines present on

both ventral and dorsal surfaces, from Megaceryle alcyon, North America.

C. reticulata (Wright, 1879)
Ovary postequatorial; uterus relatively long; cuticular spines confined
to ventral surface 3

3. Testicular branches relatively short; vas efferens from caudal testis

passing to right; from Ardea nigra, Europe, and Ciconia alba, Africa ....

C. Mans (Rud., 1809)

Testicular branches relatively long; vas efferens from caudal testis pass-
ing to left 4

327

1940] Olsen: New Species of Trematodes

4. Egg 160-170 p long; from Leptoptilus crumenifer, Africa

C. spectabilis Odhner, 1926
Egg 57-62 p long; from Tantalus ibis, Africa C. famelica Odhner, 1926

Note on the Occurrence of Clinostomum campanulatum
in Herons and Cormorants.

The black-crowned night herons, totalling 49 altogether, consisted of
both adult and juvenal birds taken during the course of several summers
from rookeries in Ramsey and Steele Counties, Minnesota. The great blue
herons, on the other hand, were much fewer, there being only 11 birds,
originating from a rookery in Rice County. From the total of 60 herons
examined, Clinostomum campanulatum occurred only four times, all of
which were in the black-crowned night herons.

In addition to the herons, Clinostomum campanulatum was found in 6
of 18 double-crested cormorants, Phalacrocorax auritus auritus (Lesson)
killed in Minnesota. This appears to be a new host record for C. campanu-
latum.

References.

Azim, M. Abdel.

1935. Entwicklungsgeschichte von Apharyngostrigea ibis n. sp. und seine
Entwicklung aus Cercaria Apharyngostrigea ibis n. sp. in dem Reiher
Ardeola ibis ibis. Zeitschr. Parasitenk. 7 : 608-614.

Chase, E. E.

1921. A new avian trematode. Proc. Linn. Soc. N. S. Wales. 45: 500-504.

Dubois, Georges.

1934. Etude des Strigeides de la collection du Department de Parasitologie
de l’universite Hebralque de Jerusalem. Rev. suisse Zool. 42: 571-585.

Dubois, Georges.

1938. Monographie des Strigeida (Trematoda). Mem. Soc. Neuchateloise
Sc. Nat. 6: 1-535.

Harwood, P. D.

1936. Notes on Tennessee helminths. III. Two trematodes from a kingfisher.
Tennessee Acad. Sci. 11: 251-256.

Johnston, B. S.

1904. Contributions to a knowledge of Australian entozoa. Proc. Linn. Soc.
N. S. Wales. 29: 108-114.

Looss, A.

1899. Weitere Beitrage zur Kenntnis der Trematoden-Fauna Aegyptens,
zugleich Versuch einer naturlichen Gliederung des genus Distomum
Retzius. Zool. Jahrb. Abt. Syst. 12: 521-784.

Odhner, Th.

1926. Protofasicola n.g., ein Protypus des grossen Leberegels. Arch. f. Zool.
18(a) : 1-7.

SZIDAT, L.

1929. Beitrage zur Kenntnis der Gattung Strigea (Abildg.). II. Spezieller
Teil: Revision der Gattung Strigea nebst Beschreibung einer Anzahl
neuer Gattungen und Arten. Zeitschr. Parasitenk. 1: 688-764.

Tubangui, M. A.

1933. Trematode parasites of Philippine vertebrates. VI. Description of new
species and classification. Philip. Jr. Sc. 52: 167-197.

328

Zoologica: Neiv York Zoological Society

Verma, S. C.

1936. Notes on trematode parasites of Indian birds. Part I. Allahabad Univ.
Stud. 12: 147-188.

Wright, R. R.

1879. Contributions to American helminthology. No. I. Proc. Canad. Inst.,
n. ser. 1: 54-75.

All drawings made with the aid of a camera lucida except Figs. 6 and 7,
which were made free-hand.

Fig. 1. Ventral view of Cathaemasia nycticoracis.

Fig. 2. Dorsal view of Apharyngostrigea bilobata from Ardea herodias herodias.
Fig. 3. Dorsal view of A. bilobata from Nycticorax nycticorax hoactli.

Fig. 4. Longitudinal section of A. bilobata.

Fig. 5. Cross-section of A. bilobata taken a short distance cephalad from adhesive
gland.

Fig. 6. Sinistral view of female reproductive system. Reconstructed.

Fig. 7. Sinistral view of male reproductive system. Reconstructed.

EXPLANATION OF THE PLATE.

Plate I.

Key to Abbreviations.

A. acetabulum
AG. adhesive gland
DL. dorsal lip
GA. genital atrium
GC. genital cone
GP. genital pore

OV. ovary

S. seminal vesicle
SR. seminal receptacle

T. testes

U. uterus

V. vitellaria

I. intestinal caeca
LC. Laurer’s canal
MG. Mehlis’ gland

VE. vas efferentia
VD. vas deferens

O. oral sucker
OE. oesophagus

VL. ventral lip
YD. yolk duct

YR. yolk reservoir

OLSEN.

PLATE I.

TWO NEW SPECIES OF TREMATODES (APHARYNGOSTRIGEA BILOBATA : STRIGEIDAE,

AND CATHAEM ASI A NYCTICORACIS : ECH I NOSTOM I DAE ) FROM HERONS, WITH A

NOTE ON THE OCCURRENCE OF CLI NOSTOM UM CAMPANULATUM (RUD.).

Richmond : Nesting of Lepomis auritus

329

21.

Nesting of the Sunfish, Lepomis auritus (Linnaeus), in
Tidal Waters.

Neil D. Richmond

(Plate I).

During the summer of 1939, while the writer was making herpetological
investigations in New Kent County, near Lanexa, Virginia, large numbers of
sunfish nests were observed exposed at low tide along the shore of the
Chickahominy River. These were all nests of Lepomis auritus (Linnaeus).
On July 13, 1939, there being a low tide in the afternoon, the nests were
photographed (Plate I) and examined more closely. At this time they con-
tained eggs, in different stages of development, and hatchlings. Several
nests were being constructed, and several were no longer in use, as could be
seen by the accumulated silt. During low tide adult sunfish were not seen,
although when the nests were submerged they could be seen swimming about
in them.

The sites chosen for nesting were located where the bottom was of sand,
not too deeply covered by mud and usually near some sheltering object such
as a log (PI. I, Fig. 1), stump, or as shown in PI. I, Fig. 2, along the base of
a retaining wall. In suitable locations the nests were grouped in large
numbers, in some cases so closely that only the width of the rim separated
them. This crowding was probably due to the areas suitable for nesting
being very limited in this locality. The river is always turbid and all of the
shallow water sheltered from wave and tide action has a mud bottom be-
cause of the rapid rate of silting, and is thus rendered unsuitable for
nesting. Since all of the river has a mud bottom below the level of the
normal low tide, there is left as the only possible nesting area a narrow strip
of sand between the levels of the high and low tides on the more exposed
portions of the shore. The proximity of the nests to the shore is apparently
limited by the level of the lowest high tide; presumably the length of
exposure above that level discouraged nest-building.

The normal rise and fall of the tides here is approximately two to three
feet, but varies from day to day with a range of six feet between the extreme
high tide and the extreme low. As the nests are all below the level of the
lowest high tide, they are always submerged at least twice a day and on
days when the tide is running high they may not be exposed at all. At other
times only those nests nearest the shore are exposed. Thus the amount of the
time that the nests are not submerged is extremely variable, the maximum
exposure observed being about four hours with each low tide. This results
in the nests being subjected to continually and rapidly changing environ-
mental factors. When the low tide occurs during the afternoon, the nest is
exposed to the direct rays of the sun and the shallow water remaining in it
becomes warmer than that of the river, only to be abruptly cooled when the
river returns. Equally variable is the depth of water over the nests, varying

3301

Zoologica: New York Zoological Society [XXV :21 :1940]

from the four or five inches left in the nests at low tide to a maximum of five
feet at extreme high tide. Slightly variable also is the salinity of the river,
which, although normally fresh, becomes perceptibly brackish during any
prolonged dry weather.

The dates on which nesting began and ended were not determined as the
nests were present when these observations began, June 11, and were still
there on August 4. However, on returning to this locality September 7, only
the outlines of a few of the nests remained. A week later even they had dis-
appeared, so it is probable that nesting ceased during the latter part of
August. That these nests were successful under such conditions would seem
to be implied by the presence of young fish in several of the nests.

EXPLANATION OF THE PLATE.

Plate I.

Fig. 1. Nests of Lepomis auritus in the Chickahominy River, Virginia, exposed at
low tide.

Fig. 2. Nests of Lepomis auritus in the Chickahominy River, Virginia, along a
retaining wall, exposed at low tide.

*<✓/

RICHMOND.

PLATE I.

FIG. 2.

NESTING OF THE SUNFISH, LEPOMIS AURITUS (LINNAEUS), IN TIDAL WATERS.

Clark: Notes on Echinoderms

331

22.

Eastern Pacific Expeditions of the New York Zoological Society.
XXI. Notes on Echinoderms from the West Coast
of Central America.1

Hubert Lyman Clark

Museum of Comparative Zoology, Harvard University

(Plates I & II; Text-figures 1-4).

[This is the twenty-first of a series of papers dealing with the collections
of the Eastern Pacific Expeditions of the New York Zoological Society made
under the direction of William Beebe. The present paper is concerned with
specimens taken on the Eastern Pacific Zaca (1937-1938) Expedition, which
was made possible through the generosity of Mr. Templeton Crocker. For
data on localities, dates, dredges, etc., refer to Zoologica, Vol. XXIII, No. 14,
pp. 287-298.]

Contents.

Page

Introduction 332

ASTEROIDEA

Astropecten armatus Gray 332

Luidia asthenosoma Fisher 332

Luidia bellonae Liitken 332

Luidia foliolata Grube 333

Luidia ludwigi Fisher 333

Oreaster occidentalis Verrill 333

NidoreUia armata (Gray) 333

Patiria miniata (Brandt) 333

Linclcia colombiae Gray 334

Pharia pyramidata (Gray) 334

Phataria unifascialis (Gray) 334

Othilia aculeata Gray 334

Sclerasterias alexandri var. crassa var. nov. 335
OPHIUROIDEA

Ophiomyxa panamensis Liitken & Morten-

sen 336

Sigsbeia lineata Liitken & Mortensen 336

Amphiura arcystata H.L. Clark 336

Amphipholis geminata (Leconte) 336

Amphipholis platydisca Nielsen 337

Amphipholis squamata (delle Chiaje) 337

Amphichondrius granulosus Nielsen 337

Amphiodia sculptilis F. Ziesenhenne 337

Amphiodia urtica (Lyman) 338

Amphiodia vicina sp. nov 338

Ophiactis savignyi (Muller & Troschel) .... 339

Ophiactis simplex (Leconte) 340

Ophiothrix spiculata Leconte 340

Ophiothela gracilis Nielsen 340

Page

Ophionereis annulata (Leconte) 340

Ophionereis nuda Liitken & Mortensen .... 340

Ophiocoma aethiops Liitken 341

Ophiocoma alexandri Lyman 341

Ophiopteris papillosa (Lyman) 341

Ophioderma panamense Liitken 341

Ophioderma teres (Lyman) 342

Ophioderma variegatum Liitken 343

Diopederma danianum (Verrill) 343

Schizoderma diplax Nielsen 343

Ophiura lutkenii (Lyman) 343

Ophiozona pacifica (Liitken) 344

Genus Ophiolepis 344

Ophiolepis variegata Liitken 344

Ophiolepis julva sp. nov 345

Ophiolepis grisea sp. nov 346

ECHINOIDEA

Eucidaris thouarsii (Agassiz & Desor) 347

Stylocidaris dubia (H. L. Clark) 347

Astropyga pidvinata (Lamarck) 348

Arbacia incisa (A. Agassiz) 348

Lytechinus anamesus H. L. Clark 348

Lytechinus pictus (Veri-ill) 348

T oxopneustes roseus (A. Agassiz) 349

Strongylocentrotus purpuratus (Stimpson) 349

Echinometra vanbrunti A. Agassiz 349

Clypeaster europacificus H. L. Clark 349

Clypeaster ochrus H. L. Clark 350

Clypeaster speciosus Verrill 350

Moira clotho (Michelin) 350

Plagiobrissus pacificus sp. nov 351

Meoma grandis Gray 362

Lovenia cordiformis A. Agassiz 352

l Contribution No. 602, Department of Tropical Research, New York Zoological Society.

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Introduction.

Aside from the holothurians, which I have not examined, the collection
of Echinoderms which the Zaca made in 1937-38 consists of 822 specimens
of 57 species, of which 73 specimens are sea-stars, 645 are brittle-stars
and 104 are echini. Most of these are common forms and call for little
comment, but a few are of notable interest and deserve a critical report. At
least 4 species seems to be new to science, and 2 varieties are also treated as
new.

It is a pleasure to thank Dr. William Beebe for the opportunity of
studying this collection and for its deposit in the Museum of Comparative
Zoology.

Asteroidea.

Asfropecfen armatus Gray.

Gray, 1840. Ann. Mag. Nat. Hist., 6, p. 181.

There are 24 specimens of this common sea-star from a dozen stations
between Manzanillo, Mexico, and Hannibal Bank, Panama, in 2.5-60 fms.
of water. Most of them are small, R= 12-42 mm., but several have R exceed-
ing 100 mm. and a notable specimen from Port Culebi'a, Costa Rica, has
R=200-210 mm. This big one is dark yellowish-brown above and below, but
the adambulacral armature and the tips of the superomarginal spines are
conspicuously lighter. In this individual the superomarginal spines are fully
developed, a complete double series extending from the interradial line to
the tip of each ray. In no other adult specimen are both series complete,
and much diversity is shown in their development. One small specimen
(R=42 mm.) taken 14 miles south of Judas Point, Costa Rica, is very
nearly referable to A. calif ornicus, as the superomarginal spines are con-
fined to the interbrachial arcs and there is a single pair only in each arc, the
first superomarginal in each series having one small erect spinelet.

Geographical Distribution: San Pedro, California, to Punta Santa
Elena, Ecuador; low water to 80 fms.

Luidia asthenosoma Fisher.

Fisher, 1906. Proc. Wash. Acad. Sci., 8, p. 124.

This species was not taken by the Zaca in 1936 but there is a single
small specimen in the present collection, from Hannibal Bank, Panama, in
35-40 fms. The rays are only about 23 mm. long and 6 mm. wide. The color
above is pale brown, variegated with whitish ; beneath it is nearly white.

Geographical Distribution: Monterey, California, to Los Coronados
Islands, L.C., Mexico; 11-339 fms.

Luidia belionae Liitken.

Lutken, 1865. Vid. Med. Dansk. Nat. Foren. 1864, p. 133.

There are 2 small but typical specimens of this species; one with R=75
mm., from 16 fms. in Chamela Bay, Mexico, and one, with R=100 mm., from
30 fms. off Manzanillo, Mexico.

Geographical Distribution: Gulf of California to Iquiqui, Chile; also
Galapagos Islands; 2-30 fms.

1940]

Clark: Notes on E chinoderms

333

Luidia foliolata Grube.

Grube, 1866. Jahresber. schles. Ges. Vaterl. Cultur. 1865, p. 59.

There are a dozen specimens of this well-known Californian species.
The largest has R=140 mm. and br=27, but the others are much smaller
and several are very badly damaged. They were taken at various stations
between Tenacatita Bay, Mexico, and Corinto, Nicaragua, in 2-30 fms.

Geographical Distribution: Southeastern Alaska to Mazatlan, Mexico;
2-189 fms.

Luidia ludwigi Fisher.

Fisher, 1906. Proc. Wash. Acad. Sci., 8, p. 122.

This interesting species is represented by 2 specimens taken east of
Cedros Island, Lower California, in 45 fms. (St. 126 D-13, -14). In one
R=only 37 mm. but the larger one has R=100 mm. They are alike in color, a
reddish fawn, deepest along median area of each ray. It is interesting to
note that the Zaca in 1936 secured 4 species of Luidia but although in 1937-38
she again took 4 species of the genus, only 2 are common to both collections.

Geographical Distribution : Monterey Bay to San Pedro, California;
15-50 fms.

O reaster oeeidentalis Verrill.

Verrill, 1867. Trans. Conn. Acad., 1, p. 278.

There are 3 specimens of this common species, 1 from a trap at Guatulco,
Mexico, and 2 from Port Parker, Costa Rica. All are of moderate size with
R=100-110 mm.

Geographical Distribution: Lower California to Ecuador; also Cocos
and Galapagos Islands ; low water to 50 fms.

Midorellia armata (Gray).

Pentaceros armatus Gray, 1840. Ann. Mag. Nat. Hist., 6, p. 277.

Nidorellia armata Verrill, 1867. Trans. Conn. Acad., 1, p. 280.

Besides a normal 5-rayed specimen, with R=90 mm., from St. 200
D-28-30, at Corinto, Nicaragua, in 3 fms., there is a slightly smaller specimen
from the same station, which has 5 rays orally but from above seems to have
6 rays ; one ambulacral furrow when about 8 mm. from the mouth forks and
a ray has developed in association with each division. These 2 rays, how-
ever, are only about 50 mm. long, while the 4 other rays have R=83 mm.

Geographical Distribution: Guaymas, Mexico, to Zorritos, Peru; also
Galapagos Islands; low water to 40 fms.

Patiria miniata (Brandt).

Asterina miniata Brandt, 1835. Prodromus, p. 68.

Patiria miniata Verrill, 1914. Harriman Exp. Sea-stars, p. 264.

This northern species is represented by a single very small specimen
taken at St. 126 D-19, southeast of Cedros Island, Lower California, in 25
fms. It has 5 rays, with R=8 mm. and might well be called in the Asterina
stage.

Geographical Distribution: Sitka, Alaska, to Gulf of California; low
water to 165 fms.

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Zoologica: New York Zoological Society

Linckia colombiae Gray.

Gray, 1840. Ann. Mag. Nat. Hist., 1, p. 285.

A single small 6-armed specimen, scarcely 5 mm. across the disk but
with R=40 mm., more or less, represents this common species. It was taken,
with the preceding, at St. 126 D-19, southeast of Cedros Island, Lower Cali-
fornia, in 25 fms.

Geographical Distribution: San Pedro, California, to Colombia; also
Galapagos Islands; low water to 55 fms.

Pharia pyramidata (Gray).

Ophidiaster pyramidatus Gray, 1840. Ann. Mag. Nat. Hist., 1, p. 284.

Pharia pyramidata Sladen, 1889. “Challenger” Asteroidea, p. 784.

A half a dozen specimens with R ranging from 50 to 90 mm., were taken
at Tenacatita and Sihuatanejo Bays, Mexico, and Port Parker and Uvita Bay,
Costa Rica.

Geographical Distribution: Gulf of California to Zorritos, Peru; also
Galapagos Islands; low water to 10 fms.

Phataria unifascialis (Gray).

Linckia unifascialis Gray, 1840. Ann. Mag. Nat. Hist., 1, p. 285.

Phataria unifascialis Sladen, 1889. “Challenger” Asteroidea, p. 786.

Of the 14 specimens, with R ranging from 15 to 87 mm., few call for
comment. They were taken at Tenacatita and Sihuatanejo Bays, Mexico;
Cardon Isl. at Corinto, Nicaragua; Uvita Bay, Costa Rica, and Bahia Honda,
Panama. The specimen from Corinto is a very fine one with R=87 mm. and
br— 11. The one from Uvita Bay is very odd, for each ray has been bitten or
broken off at the tip and then more or less regenerated; the longest is only
46 mm. long but it is 12 mm. broad. The color is definitely yellow.

Geographical Distribution: Gulf of California to Zorritos, Peru; also
Galapagos Islands; low water to 10 fms.

Othilia ac is I eat a Gray.

Gray, 1840. Ann. Mag. Nat. Hist., 1, p. 281.

Plate I, figs. 1 & 2.

At Corinto, Nicaragua, St. 200 D-5,-6,-7, in 2-2.5 fms., the Zaca dredged
4 specimens of an echinasterid, which are apparently examples of this long
lost species. Although described almost a century ago it has never since been
taken and no specimens are known to be extant. It is therefore extraordi-
narily interesting that the Zaca should have secured these 4 individuals.
Gray’s specimens were from Guacamayo, Central America, fine sand, 13 fms.,
and apparently were larger than any of these from Costa Rica. As he gives
no measurements, however, and only a very brief and imperfect description,
much has to be inferred in regard to the types. It is worth while therefore
to give some details about the Zaca specimens.

The smallest specimen, now dry, has R=14 mm.; r=3 mm. and br is
the same. There are 6 relatively large spines on the disk, the central one
the smallest. On each ray there are about 7 such spines in three indefinite
series of 2 or 3 each. Many minute spinelets are scattered here and there,
mostly on the distal half of each arm. All the larger skeletal plates have, in
this dry specimen, a shagreen-like surface. Color light gray-brown with the
tips of the large spines and the lower surface pale yellow.

1940]

Clark: Notes on Echinoderms

335

A second specimen has R=21 mm. with r=5 mm. and br=4.5. The color
is as in the smaller specimen. There are about 5 big spines in each of three
series on each arm, and 4 or 5 similar but somewhat smaller spines on the
disk. There are very few minute spinelets except near tip of each arm.

The third specimen is nicely preserved in alcohol. It has R=28-30 mm.,
r=7, br=6. The body surface is very smooth with scarcely any small spine-
lets except at very tip of arm and here and there on oral surface of distal
part of arm. There are about 20 of the big spines on each ray, and 7 or 8
on the disk. On the rays there are marginal series of 5 or 6 spines, an
irregular median dorsal and an incomplete lateral on each side; these 3
series having 3-5 spines each. The color is quite different from the dried
specimens; it is more or less definitely brown on the sides of the arms, where
it is darkest, but dorsally it is fawn-color and orally nearly buff or yellowish ;
the spines are lightest at tip.

The largest specimen (Plate I, figs. 1 & 2) is dried from alcohol and
its color is dull yellow-brown with the papular areas a darker brown.
R=33-35 mm., r=8 and br is also 8. There are about 5 rather indefinite series
of big, nearly white, spines (2-5 mm. long) on each ray. Papulae are con-
spicuous; on the oral surface they chiefly occur singly; on sides of ray, there
are 2 or 3 in a group, while aborally there are up to 8-10 in each area.
Aborally there are a very few widely scattered minute spinelets; 6-8 of
these are on the carinal ridge at the very tip of each arm. On distal half of
arm there are a very few small spines scattered on the sides and oral sur-
face. Adambulacral spines in 3 more or less distinct series; innermost,
minute curved spinelets well up in furrow; on furrow margin a series of
pointed spinelets, a millimeter long; outside these is a series of similar but
frequently stouter spinelets, but this series is not so nearly complete as the
marginal one.

Although these specimens are probably all young, they are noticeably
distinct from any of the better known sea-stars of the Panamic region. The
largest was submitted to Dr. W. K. Fisher, the world’s authority on sea-
stars, who kindly had the best possible photographs made (by Mr. B. B.
Fisher) and permits me to publish them herewith. He believes it is justi-
fiable to refer these specimens to Gray’s long-lost species. Mr. Dilwyn John,
of the British Museum, assures me that Gray’s specimens are no longer
extant, although their fate is unknown. They are described as having 7 series
of large spines on the arms, indicating that they were considerably larger
than the largest of the Zaca quartette, as previously stated. All efforts to
locate “Guacamayo” have proved futile but as Gray states that Hugh
Cuming, Esq., was the collector, and Cuming is not known to have collected
on the eastern coast of Central America, we are justified in believing that
the original specimens of aculeata were from the western coast.

Geographical Distribution : Prior to Zaca collection, known only from
a single station on the western coast of Central America, the location of
which is unknown.

Sclerasterias alexandri var. crassa var. nov.

A single small sea-star from St. 126 D-19, southeast of Cedros Island,
Lower California, in 25 fms. caused me so much perplexity that I sent it to
Dr. Fisher who very kindly made a careful study of its important features
and wrote me: “I would list this as a variation of S. alexandri.” It differs
“from the type of alexandri in having coarser spines and constantly dipla-
canthid adambulacrals.” However “the coarseness of spines is usually quite
variable in Coscinasterinae.” “There is a well-developed web between outer
inferomarginal spines which in S. heteropaes is practically absent. The

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[XXV :22

crossed pedicellariae are like those of alexandri in size and shape.” In color,
“heteropaes is gray-green; alexandri, red-brown.”

This interesting specimen, which may be designated as the holotype of
var. crassa (M.C.Z. No. 3753) has a disk 6 mm. across and short, stout arms
only 20 mm. long, 4 mm. in actual diameter at base, but 7 mm. wide on the
actinal side, when the inferomarginal spines are included. As in typical
alexandri there are but 3 series of abactinal spines, a carinal and on each
side a superomarginal. These spines are shorter, stouter and blunter than in
typical alexandri. The skeleton is much coarser and the papular areas very
much smaller and less conspicuous than in the cotypes of alexandri with
which it has been compared. The color is a deep red-brown above, much
darker than these types. These differences combine to give it a markedly
different facies from that of Ludwig’s types. As there is but one specimen,
however, and that obviously young, it seems best to list it simply as a
variety, but more and larger specimens will probably justify its recognition
as a distinct species. It is strikingly different from heteropaes, the only other
Sclerasterias from the western coast of America.

Geographical Distribution: Bay of Panama; 52-210 fms. The variety
crassa is known only from the Zaca specimen.

Opiiiuroidea.

Ophiomyxa panatnensis Ltitken & Mortensen.

Liitken & Mortensen, 1899. Mem. M.C.Z., 23, p. 182.

At Station 126 D-19, off Cedros Island, Lower California, in 25 fms.
of water, 4 small adults of this brittle-star were taken, while a very young
one, only 5 mm. across the disk, was collected at St. 224 D-1,-2,-3, on Hannibal
Bank, Panama, in 35-40 fms.

Geographical Distribution: Gulf of California to Panama; also Cocos
and Galapagos Islands; 25-85 fms.

Sigsbeia lineata Liitken & Mortensen.

Liitken & Mortensen, 1899. Mem. M.C.Z. , 23, p. 183.

A single small adult was collected at St. 224 D-2,-3, on Hannibal Bank,
Panama, in 35 fms. It is nearly white now, but the brown lines on the disk
can still be detected.

Geographical Distribution: Cocos and Galapagos Islands; 30-100 fms.

Amphiura arcystata H.L. Clark.

H. L. Clark, 1911. Bull. 75, U.S.N.M., p. 145 (printed acrystata by
error ) .

A very small specimen, 4.5 mm. across the disk, was dredged in 38 fms.,
off Maldanado Point, Mexico, at St. 192 D-3. It is now white with the naked
skin of the disk, yellowish-brown.

Geographical Distribution: Japan and California to Gulf of California;
30-330 fms.

Amphipholis geminata (Leconte).

Ophiolepis geminata Leconte, 1851. Proc. Acad. Nat. Sci. Phila. 5, p. 317.
Amphipholis geminata Ljungman, 1867. Ofv. Kongl. Vet. Akad. Forh.,
23, p. 313.

1940]

Clark: Notes on Echinoderms

337

This very long-armed brittle-star* seems to be common in the Gulf of
Fonseca, El Salvador, and in the Gulf of Nicoya, Costa Rica. There are 27
specimens at hand from St. 199 D-l, -17-25, and 3 from St. 213 D 13-17.
A muddy bottom in shallow water is the chosen habitat.

Geographical Distribution : Panama (Pacific side) ; low water to 5 fms.

Amphipholis platydisea Nielsen.

Nielsen, 1932. Vid. Med. 91, p. 290.

There are 11 specimens of this fine big Amphiodia- like species, taken
at the same stations as the preceding; St. 199 D-7-16; St. 213 D-12-17; and
St. 214 D-l-4. Only 3 have the disk which is about 10 mm. across, still at-
tached to the mouth frame and the smallest of these is a typical example
of Nielsen’s variety microplax. It is odd that Nielsen did not realize that
his supposed variety is based on an individual with a regenerating disk.

Apparently platydisca is one of those amphiurids which shed the disk,
when mature, on very slight provocation. It may be that the shedding is" the
normal process of setting free ova and spermatozoa. But, at any rate, the
present series of specimens shows that at the time and place when they
were taken shedding was twice as frequent as retention. Nielsen’s material
was from Panama, so that the present series extends the range considerably
to the northward.

Geographical Distribution : Panama (Pacific side) ; low water to 15 fms.

Amphipholis squamata (delle Chiaje).

Asterias squamata delle Chiaje, 1828. Mem. Anim. sans Vert. Napoli,

3, p. 74.

Amphipholis squamata Verrill, 1899. Trans. Conn. Acad. 10, p. 312.

This small but wellnigh ubiquitous brittle-star is represented by a young
individual from St. 126 D-19 (Lower California, southeast of Cedros Island,
25 fms.) and a still smaller specimen, only 2 mm. across the disk from
Pocillopora coral, Jasper Island, Gulf of Nicoya, Costa Rica, 1 fm.

Geographical Distribution: Cosmopolitan; low water to 100 fms.

Amphichondrius granulosus Nielsen.

Nielsen, 1932. Vid. Med., 91, p. 304.

At St. 192 D-3, off Maldanado Point, Mexico, in 38 fms. on a muddy
bottom, 2 specimens of this interesting species were taken. The larger is
only 5 mm. across the disk and hence is little more than half grown but the
generic features are unmistakable.

Geographical Distribution: The species was known hitherto only from
the vicinity of La Jolla and San Diego, California.

Amphiodia sculptilis F. Ziesenhenne.

F. Ziesenhenne, 1940. Allan Hancock Pac. Exp., 8, No. 2, p. 24.

A single young specimen of this fine species was taken at St. 200
D-l, -3, -8, in 2-6.6 fms. at Corinto, Nicaragua, on a mangrove-leaves bottom.
It was previously known from various stations on the western coast of
Mexico. This individual is about 3.5 mm. across the disk and the arms are
some 25 mm. long. The general color is dark gray but the oral region and
the accompanying arm spines are cream. The single large tentacle scale is
very distinctive.

Geographical Distribution: Western coast of Mexico; 5-20 fms.

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Amphiodia urtiea (Lyman).

Amphiura urtiea Lyman, 1860. Proc. Boston Soc. Nat. Hist., 7, p. 258.
Amphiodia urtiea Verrill, 1899. Trans. Conn. Acad., 10, p. 313.

A single adult specimen, 6.5 mm. across the slightly elevated disk, was
taken at St. 196 D-14, in Tangola-Tangola Bay, Mexico, in 5 fms. on a shelly
bottom, thus extending the range considerably to the south.

Geographical Distribution : Shumagin Islands, Alaska, to Lower Cali-
fornia; 15-50 fms.

Amphiodia vieina2 sp. nov.

(Text-figures 1 & 2).

Disk 10 mm. in diameter, nearly circular and rather flat. Arms 5, 80-90
mm. long, rather stout basally, about 2 mm. wide regardless of the arm-
spines, tapering very gradually to the not very attenuate tip. Disk covered
by h coat of rather coarse overlapping scales without clearly indicated
primary plates; the largest ones more than half a millimeter across; very few
are even approximately circular, some are quite angular and some distinctly
elongated ; in each interradius there are 7-9 series separating the pairs of
radial shields. These shields are about 3 mm. long, 1.5 mm. wide distally;
they are in contact for one-half or more of their inner margins but proxim-
ally are separated by one large, long triangular scale or by 2 or more smaller
ones. Upper arm-plates wider than long; basally the width is almost thrice
the length and even far distally the length does not quite equal the width.
On the basal part of the arm, the distal margin of each plate is much longer
than the proximal, the resulting lateral angles being more or less rounded,
although often very acute, and more or less of the upper end of the side
arm-plates being conspicuous between them. On the basal part of the arm
there is a slight but usually rather distinct keel or ridge along the mid-line.

Text-figure 1. Text-figure 2.

Amphiodia vieina. Upper surface. Amphiodia vieina. Lower surface.

X 6. X 6. ^

Interbrachial areas closely covered, like the disk, by coarse overlapping
scales; the uppermost or marginal scales stand on edge and form a low
marginal “fence” around the disk, somewhat suggestive of Ophiophragmus ;
genital slits long, reaching nearly to the disk margin. First under arm-plate

2 4ucr7ms=neighboring, in reference to its nearness to A. psara, both structurally and
geographically.

1940]

Clark: Notes on Echinoderms

339

small, rounded, swollen, rather wider than long; second and succeeding plates
rounded pentagonal, wider than long; the second and third plates have the
lateral and distal margins concave, but this is not evident on the following
plates; these plates are not quite in contact, yet the side arm-plates seldom
meet in the mid-line; as a Consequence there is generally the indication of
a pit in the mid-line between the under arm-plates ; the basal plates, es-
pecially the second and third, have the margins thickened or even a little
swollen. Side arm-plates rather large and projecting but they do not actually
meet either above or below, even distally ; each carries 3 arm-spines, about a
millimeter long, subequal or the middle one a little the longest; slightly
swollen at base but tapering somewhat abruptly to a slender, if not acute,
point. Tentacle-scales 2, rather large, subequal; as usual in such cases one
is borne on the under arm-plate and one on the side arm-plate.

Oral shields spear-head shaped, longer than wide, with scarcely rounded
angles, the madreporite a trifle larger than the others and slightly swollen.
Adoral plates trigonal, slightly swollen, just meeting within, the inner
margin concave, the other two straight. Oral papillae 3 on each side; those
at the apex of the jaw are narrow and notably longer than wide, quite sepa-
rated from each other; the second papilla tends to show some diversity in
form, being flattened and somewhat angular, or rounded and scale-like, about
as wide as long; the distal papilla is largest and most scale-like and is car-
ried on the adoral plate. Teeth few, small and block-like.

Color of preserved specimens, now dry, light gray with a lavender tint,
variegated with light cream-color or white; many disk scales are white in
the holotype and in the smallest specimen but this is less evident in the
others; each radial shield has a more or less conspicuous blotch of white or
whitish on the distal half; oral surface very light; the under arm-plates
and arm spines nearly white.

Holotype, M.C.Z. No. 6026, from St. 213 D-13, off Ballenas Bay, Gulf of
Nicoya, Costa Rica, 35 fms., mud.

There are 12 paratypes from the same locality, (St. 213 D-13, -14, -15, -16;
35-45 fms.). One of these is remarkable for being symmetrically tetramerous;
as the arms are 100 mm. long, it is obviously a large adult, but as the disk
is missing little more can be said about it; the characters shown by arms
and mouth-parts are entirely like those of the holotype. Another specimen
worthy of mention is the smallest. It has a disk 7 mm. across with the scales
scarcely overlapping and hence smoother and flatter than in the large in-
dividuals.

This is a well-marked species for while it reminds one of psara (H. L.
Clark, 1935. Ann. Mag. Nat. Hist. (10) 15, p. 127) the difference in the
upper arm-plates is very striking, and the arm-spines and under arm-plates
are also different. Furthermore the oral papillae of psara are much stouter
and the distal one in particular is much larger than in vicina. Whether the
areas occupied by the two species overlap remains to be seen. So far as I
know, psara has not been taken south of Acapulco.

Geographical Distribution : Known only from the Zaca collection.

Ophiaetis savignyi (Muller & Troschel).

Ophiolepis savignyi Muller & Troschel, 1842. Sys. Ast. p. 95.

Ophiaetis savignyi Ljungman, 1867. Ofv. Kongl. Vet. Akad. Fork., 23,
p. 323.

This very common brittle-star is represented by 17 specimens, mostly
small, but several of those taken at St. 195 D-5, are adults 6 mm. across disk,
although all of this lot regardless of size have 6 arms. All of the material

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[XXV :22

was collected at St. 195 in Port Guatulco, Mexico, save 2 small specimens
found at Cardon Island, Corinto Bay, Nicaragua.

Geographical Distribution'. Tropicopolitan ; low water to 27 fms.

Ophiactis simplex (Leconte) .

Ophiolepis simplex Leconte, 1851. Proc. Acad. Nat. Sci. Phila., 5, p. 318.
Ophiactis simplex Liitken, 1859. Add. ad Hist. Oph. pt. 2, p. 130.

There are 4 very young specimens of this little Ophiactis from St. 126
D-19, east of Cedros Island, Mexico, in 125 fms.

Geographical Distribution : La Jolla, California, to Balboa, Canal Zone;
low water to 5 fms.

Ophiothrix spiculata Leconte.

Leconte, 1851. Proc. Acad. Nat. Sci. Phila., 5, p. 318.

There are 78 specimens of this very common brittle-star, the largest 12
mm. across the disk. Most of them are dull gray-blue or purplish but those
from St. 184 are very light colored, a pale grayish-brown, and those from
St. 126 are also very light but with a reddish tint, the disk quite rosy or in
the largest one a deep crimson in sharp contrast to the arms. This species
was taken at the following stations : 126 D-19, southeast of Cedros Island,
Mexico, 25 fms.; 184 D-2, oif Manzanillo, Mexico, 30 fms.; 199 D-7-16, La
Union, El Salvador, 5-6 fms.; 200 D-5-7, 28-30, Corinto, Nicaragua, 2-3
fms.; 218 D-5, Golfito, Costa Rica, 6 fms. There are 50 specimens taken
from Pocillopora coral, Jasper Island, Gulf of Nicoya, Costa Rica, 1 fm.

Geographical Distribution: Monterey Bay, California, to Bay of Sechura,
Peru, also Galapagos Islands ; low water to 45 fms.

Ophiothela gracilis Nielsen.

Nielsen, 1932. Vid. Med., 91, p. 256.

There are 3 small 5-armed Ophiothelas which I refer with some hesita-
tion to this species, on small fragments of a gorgonian from a tide-pool at
Uvita Bay, Costa Rica. The largest is only 2 mm. across the disk. The
genus needs critical study and revision.

Geographical Distribution: Panama; 4-5 fms.

Ophionereis annulate (Leconte).

Ophiolepis annulata Leconte, 1851. Proc. Acad. Nat. Sci. Phila. 5, p. 317.
Ophionereis annulata Lyman, 1860. Proc. Bost. Soc. Nat. Hist., 7, p. 203.

Half a dozen specimens were taken at Uvita Bay, Costa Rica, and a
single specimen at Bahia Honda, Panama.

Geographical Distribution: San Diego, California, to Panama; also
Galapagos Islands; low water to 20 fms.

Ophionereis nuda Liitken & Mortensen.

Liitken & Mortensen, 1899. Mem. M.C.Z., 23, p. 163.

A single small specimen, scarcely 5 mm. across the disk, was taken at
St. 224 D-l, -2, -3, on Hannibal Bank, Panama, in 35-40 fms.

1940]

Clark: Notes on Echinoderms

341

Geographical Distribution : Isabel Island, Mexico, to Gulf of Panama;
also Galapagos Islands; low water to 85 fms.

Ophiocoma aeihiops Lutken.

Lutken, 1859. Add. ad Hist. Oph., pt. 2, pp. 141, 145.

There is an adult specimen of this abundant Panamic brittle-star taken
from coral, Arriba Isthmus, Port Parker, Costa Rica. It is 20 mm. across
the light gray disk, which is irregularly speckled with deep brown. A small
adult with all arms broken off near the base but each obviously regenerating,
and 2 young specimens were collected in Bahia Honda, Panama. There are
9 specimens from St. 195 D-15, in Port Guatulco, Mexico, 1.5 fms., and there
is a single specimen from an unknown station.

Geographical Distribution : Lower California to Panama; also Cocos and
Galapagos Islands; low water to 10 fms.

Ophiocomss alexandri Lyman.

Lyman, 1860. Proc. Boston Soc. Nat. Hist., 7, p. 256.

This common long-armed brittle-star is represented by two lots of quite
different appearance. From an unknown station, there are 9 adults, 10-15
mm. across the disk, with the usual brown coloration, the arms banded with
a darker shade. There is no indication of green. The other lot was taken at
St. 195, D-15, in Port Guatulco, Mexico, 1.5 fms., and consists of 22 speci-
mens, ranging from 5 to 15 mm. across the disk with arms 7 times as much.
They are notable for their unusual coloration. In the smallest specimen, the
ground color is a very deep green, lightest near arm-tips, with more or less
indefinite markings of a light shade. In larger specimens the disk is more
brown and in the adults it is a definite uniform brown, (in one case almost
deep yellow ) with a green tinge around the margin. The arms are brown at
base but green distally, irregularly but definitely banded with a darker
shade. The shade and extent of the green show great diversity in different
individuals, but the arm-spines are in all cases quite green with the tips
often perceptibly lighter. The under arm-plates are green (or at base of
arms, brown) on the margins but a broad light area covers the middle; as a
result the under side of the arm appears to be brown (basally) or green,
with a broad longitudinal light stripe along the middle. This type of colora-
tion is frequent in specimens from the Galapagos Islands.

Geographical Distribution : Lower California to Panama; also Cocos and
Galapagos Islands; low water to 10 fms.

Ophiopteris papillose (Lyman).

Ophiocoma papillosa Lyman, 1875. Illus. Cat. M.C.Z., No. 8, pt. 2, p. 11.

Ophiopteris papillosa McClendon, 1909. Univ. Cal. Publ. Zool., 6, p. 49.

A young individual, scarcely 5 mm. across the disk and light grayish-
brown in color, was collected at St. 126, D 19, in 25 fms., southeast of Cedros
Island, Lower California.

Geographical Distribution : Carmel Peninsula, California, to Todos
Santos Bay, L.C., Mexico; low water to 50 fms.

Ophioderma panamense Lutken.

Lutken, 1859. Add. ad Hist. Oph., pt. 2, p. 91.

There are 12 specimens of this common brittle-star, ranging from 9 to

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19 mm. across the disk. They show some diversity in color from gray and
brown to more or less olive-green ; the arms are banded in all at least distally,
and this is the best distinction from teres. They were taken at St. 195 (Port
Guatulco, Mexico), at Cardon Island, Corinto Bay, Nicaragua, and at Port
Parker, Ballenas Bay and Uvita Bay, Costa Rica, always in very shallow
water. Two specimens were taken in Pocillopora coral, Jasper Island, Gulf of
Nicoya, Costa Rica, 1 fm.

U

Geographical Distribution : San Pedro, California, to Payta, Peru; also
Cocos and Galapagos Islands; low water to 10 fms.

Ophioderma teres (Lyman).

Ophiura teres Lyman, 1860. Proc. Boston Soc. Nat. Hist., 7, p. 198.

Ophioderma teres Meissner, 1901. Bronn’s Thier-reichs, 2, pt. 3, p. 915.

The examination of the Ophiodermas brought back by the Zaca revealed
the interesting fact that the 8 specimens to be referred to teres fall into two
easily distinguished groups. One of these is made up of unicolored individu-
als, usually very dark. The other consists of individuals having the dull
purplish upper arm-plates more or less conspicuously spotted with light buff
or cream-color; the disk is also more or less conspicuously spotted with the
same light shade. Moreover the lower surface of the arms, including the
lowest arm-spines, is white or pale yellowish, in marked contrast to the dull
interbrachial areas and sides of the arms. A re-examination of Lyman’s
original description shows that his type of teres was one of these spotted
individuals, which are strikingly different from panamense. Unfortunately
however the unicolored specimens are not so easily distinguished but seem to
intergrade with panamense. Nielsen (1932) suggested as the best distinc-
tion between them that in adult teres many upper arm-plates (usually
all or nearly all) are broken into two or more irregular fragments. This
fragmentation may occur in panamense but only in a few plates. Unfortu-
nately it is often insignificant in young teres. Usually panamense is grayish
or greenish or brown with more or less conspicuously banded arms. It is
never speckled as is typical teres, which also lacks any bands on the arms.
The unspeckled form of teres, which apparently seldom occurs with the
typical form, is often very dark brown and the lower surface of the arms
(except maybe near the disk) is not notably lighter. The arms are however
rarely banded distally and then only in small specimens; of course such
specimens are hard to tell from panamense but in teres the light band
consists of a short longitudinal white streak on each side of 3 or 4 segments
separated by a narrow dark line. In panamense the light bands are more
diffuse and variegated. On the whole the atypical form of teres needs a
distinctive name and may well be called variety unicolor. A specimen from
the Galapagos Islands, 35 mm. across the disk, M.C.Z. No. 114, may be
designated as the type. A very similar specimen, 31 mm. across the disk, is
in the present collection from Cardon Island, Corinto, Nicaragua, accom-
panied by 2 smaller specimens, one only 10 mm. across the disk.

The specimens of typical teres in the Zaca collection are from the Arriba
Isthmus and Abajo Isthmus, Port Parker, Costa Rica. The largest is 32
mm. across the disk. A small specimen was taken in Pocillopora coral from
Jasper Island, Gulf of Nicoya, Costa Rica, 1 fm.

Geographical Distribution: Gulf of California to Panama; also

Galapagos Islands ; low water to 10 fms.

1940]

Clark: Notes on Echinoderms

343

Ophioderma variegatum Liitken.

Liitken, 1856. Vid. Med., p. 21.

This slender-armed and often brightly colored species is easily recog-
nized by the naked adoral plates yet it is often confused with panamense.
There are 5 specimens at hand, not notable in any way. They are all dull
grayish-green above with banded arms and the oral area nearly or quite
white; the under surface of the arms is also nearly white in the largest
specimen, which is 13 mm. across the disk. In 2 of the specimens, a con-
spicuous light gray area covers more or less of the disk. This quintet were
taken at St. 200, Corinto, Nicaragua, on mangrove leaves, in 2-3 fms.

Notably different are 10 specimens from St. 195 D-17, -18, Port
Guatulco, Mexico, on sand in 6 fms. These are white beneath and strikingly
variegated above with green, rose-red and cream-color, and light and dark
dusky shades. There are 7 similar specimens from St. 196, Tangola-Tangola
Bay, Mexico, 7-10 fms., and a single large individual, 15 mm. across the disk,
from St. 182 D-4, Chamela Bay, Mexico, in 16 fms. on sand. There are also
2 small specimens from St. 213 D-6, -7, -8, -9, off Cedro Island, Costa Rica,
4-6 fms.

Geographical Distribution : San Diego, California, to Panama; also
Cocos Island; low water to 60 fms.

Diopederma danianum (Verrill).

Ophiura danianum Verrill, 1867. Trans. Conn. Acad., 1, p. 254.

Diopederma danianum H. L. Clark, 1913. Bull. Amer. Mus. Nat. Hist.,

32, p. 206.

This characteristic brittle-star of the western coast of Central America
is represented by 30 specimens, ranging from 9 to 26 mm. in disk diameter.
They were taken at the following stations: St. 183 D-2, Tenacatita Bay,
Mexico, 30 fms. ; St. 195 D-20, -21, Port Guatulco, Santa Cruz Bay, Mexico,
18-23 fms.; St. 196 D-6, Tangola-Tangola Bay, Mexico, 7 fms.; St. 197 D-2,
7 miles west of Champerico, Guatemala, 14 fms. ; St. 198 D-l, -2, La Libertad,
El Salvador, 13-14 fms.; St. 199 D-l, Gulf of Fonseca, El Salvador, 16 fms.

Geographical Distribution : Lower California and Cocos Island to
Panama; 3-25 fms.

Schizoderma diplax Nielsen.

Nielsen, 1932. Vid. Med., 91, p. 335.

This remarkable ophiuran was taken by the Zaca in 1938, only at St.
214 D-l, -2, -3, -4, 14 miles southeast of Judas Point, Costa Rica, on a
bottom of mud and shell in 42 fms. There are 19 specimens ranging from 6
to 11 mm. in disk diameter.

Geographical Distribution: Lower California to Panama; low water to
60 fms.

Ophiura liitkenii (Lyman).

Ophioglypha liitkenii Lyman, 1860. Proc. Boston Soc. Nat. Hist., 7, p.

197.

Ophiura liitkeni Meissner, 1901. Bronn’s Thier-reichs, 2, pt. 3, p. 925.
This northern brittle-star was found at St. 126 D-13, in 45 fms., east

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[XXV :22

of Cedros Island, Lower California, the southern limit of its range so far as
yet known. The 8 specimens are all small, only about 8 mm. across the disk.

Geographical Distribution-. Alaska to Lower California; 15-100 fms.

Ophiozona pacifica (Liitken).

Ophiolepis pacifica Liitken, 1856. Vid. Med., p. 22.

Ophiozona pacifica Lyman, 1865. Illus. Cat. M.C.Z., 1, p. 64.

There are 2 small adults of this species in the collection, both taken in
Pocillopora coral from Jasper Island, Gulf of Nicoya, Costa Rica, 1 fm.

Geographical Distribution : Previously known only from the Gulf of
Panama; low water to 10 fms.

Genus Ophiolepis.

This genus is represented by 318 specimens ranging from IV2 to 24 mm.
in disk diameter. As none of them can be properly referred to either crassa
or plateia, it would be natural to refer them to the third West Coast species,
the long-known variegata. But even a superficial examination shows this to
be an utterly erroneous procedure. Yet patient and long-continued study has
failed to discover a natural or satisfactory grouping. Aside from those more
or less adult specimens which are unquestionably variegata, there are two
groups, very unlike each other, which are so different from variegata, it
seems most natural to consider them distinct species. Unfortunately this
does not clear up the matter, as there are still many specimens, 9 mm. or less
in disk diameter, which cannot be referred definitely to any one of the 3
species. These specimens seem to be young variegata but most of them are
so unlike small specimens which are undoubtedly that species that one cannot
be wholly satisfied to call them variegata. It seems best to include them for
the present under that species, to which attention may now be given.

Ophiolepis variegata Liitken.

Liitken, 1856. Vid. Med. p. 23.

From St. 182 D-4, in Chamela Bay, Mexico, 16 fms., there are 7 very
good typical examples of this handsome species, 10-17 mm. across the disk;
the arms are 3 times the disk diameter. There are usually 3 arm-spines, the
lowest the smallest and the uppermost largest, but at the base of the arm
in the larger specimens there may be 4 spines and if so the uppermost may
be the smallest; the spines are usually quite sharply pointed but in the
larger specimens they may be thick clear to the tip and with a rather blunt
point.

From St. 184 D-2, at Manzanillo, Mexico, 30 fms., and from St. 195
D-17-20, in Port Guatulco, Mexico, 6-23 fms., there are 38 specimens, 7-15
mm. across the disk, very similar in all respects to those from St. 182, but
in the smallest there are but 2 arm-spines on most of the arm-segments near
the middle of the arm, the upper one the larger, while basally and distally
there are 3 as usual. From this same station, 195 D 17-20, there are 69
specimens, which I cannot determine satisfactorily but which I am for the
present considering young variegata. They differ obviously from typical
young individuals of the same size, in that the disk is very flat and thin
and nearly or quite circular, instead of being higher than the arm-bases
and definitely pentagonal. The arm-spines are usually 2 but on some part
of the arm, often far distal, there are 3 as in typical variegata. Orally there
are no tangible, constant differences. In color, the small flat individuals are
much more variegated than typical variegata, the disk and arm-plates them-

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Clark: Notes on Echinoderms

345

selves being blotched, mottled or speckled in marked contrast to the solidly
colored plates and scales of the typical form. The banding of the arms may
be almost entirely wanting or, at best, quite indistinct in these odd, youthful,
individuals, which are 6.5-9 mm. across the disk.

From St. 196 D-6, 13-15, in Tangola-Tangola Bay, Mexico, 5-10 fms.,
there are 8 specimens which are clearly variegata. The smallest is only 3
mm. across the disk and has only 2 arm-spines on each side arm-plate, but
the scaling of the disk and its coloring are quite typical. In 3 individuals
5-7 mm. across, the disk is very similar but there are 3 arm-spines on many
segments. The remaining 4 specimens are 12-16 mm. across the disk and
often have 4 arm-spines basally. From this same station there are 21 small
individuals, 4-10 mm. across the disk, which are like the perplexing lot
described above from St. 195. Although they are here included under
variegata, I am not satisfied with that identification.

Geographical Distribution: Gulf of California to Gorgona Island,
Colombia; low water to 60 fms.

Ophiolepis fu/vcF sp. nov.

Text-figure 3.

Disk 8 mm. in diameter, arms 21-23 mm. long. Disk flat, thin, rounded
pentagonal or quite circular, covered as in variegata by relatively large
plates, encircled by small ones; a notable peculiarity is that the central
group of 6 plates (a central surrounded by 5 radials) is rarely symmetrical
and never conspicuous; in many specimens it is quite indistinguishable. The
arms are somewhat widened at the base but not much flattened; the upper
arm plates are rounded hexagonal and, excepting the first 2 or 3, are as
long as wide or longer, with nearly straight distal margin ; the supplementary
plates are relatively conspicuous. The side arm-plates are very large,
longer than high, the distal margin not symmetrically semi-circular in out-
line but the lower half somewhat projecting and carrying 2 short, blunt,

Text-figure 3.

Ophiolepis fulva. Six arm seg-:
ments, seen from the side. X 14.

thick arm-spines, the upper one often the larger and longer. The under arm-
plates are longer than wide, except the first half dozen, and the distal margin
is definitely convex. Otherwise the oral surface is not evidently different
from that of variegata, although the arms at disk margin are definitely less
widened.

The color of the dorsal side is a mixture of browns and lighter shades,
inextricably mingled. The general impression is fulvous. The arms are
cross-banded on the upper surface, but there is no little diversity in the
number, width and distinctness of the bands. There is no indication of red.
The lower surface is tinted with yellow. Of course, the generally yellow-
tinted color is not the most important specific character but it is quite
obvious and as a rule is the most conspicuous feature. The side arm-plates
and arm-spines are distinctive and the irregular plating of the disk and the
narrow arm-bases are helpful characters.

Holotype, M.C.Z. No. 6049, from Zaca St. 195 D-13.

There are 152 paratypes, ranging from 1.5 to 8 mm. in diameter of disk.
All the material is from St. 195 D-13, Port Guatulco, Mexico, 8 fms., sandy

3 fulvus— brownish-yellow, tawny, in reference to the coloration.

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bottom. There is great diversity in the relative quantity of light and dark
browns, whites and creamy tints, some specimens being brighter or duller
than the holotype, but the generally fulvous tint is notably different from
the redder or grayer tints of other species.

Geographical Distribution : Known only from the Zaca 1937-38 collecting.

Ophiolepis grisea 4 sp. nov.

Text-figure 4.

Disk flat, pentagonal, 22 mm. in diameter. Arms 5, rather more than 100
mm. long or nearly 5 times the disk-diameter. In young individuals, 8-10
mm. across the disk, the arms are a trifle more than 4 times disk diameter.

The primary plates of the disk are very evident particularly in the
smaller specimens. The central plate is usually noticeably smaller than the
encircling ring of 5 radials. The arms are wide at the disk margin (6 mm. in
holotype) but are well arched and not at all flattened basally. Upper arm-
plates wider than long even far out on the arm; near the disk they are
very short, 3-4 times as wide as long; supplementary plates, at least 2 on
each side, extending far down on the sides of the arm. Side arm-plates high
and short, carrying 4 or often 5, and sometimes, close to disk, 6 spines.
These spines are relatively very small, flattened and sharp pointed; when
5 or 6 are present, the lowest and uppermost are smallest, the third from
the bottom the longest; when 4 only are present, the lowest is smallest while
the other 3 may be subequal, or the uppermost smallest. The longest scarcely
exceeds one-half the side arm-plate. In the young specimens from St. 213,

Text-figure 4.

Ophiolepis grisea. Six arm seg-
ments, seen from the side. X 7.

the arm-spines on the basal part of the arms are bristling, standing out at
right angles from the arm, but in adults, all the spines are closely appressed
to the side of the arms. Oral surface, even including the under arm-plates,
much as in variegata. Color definitely gray, more or less marked on both
disk and arms with lighter areas and with white spots or speckles. Under
the lens, there are many blackish spots and markings evident on both disk
and arms. Lower surface uniformly buffy white— not so near a real white as
in variegata. The line of demarcation between the upper and lower surfaces
is very sharply drawn on the sides of the arms, just above the uppermost
arm-spine. The arm-spines are therefore all white. The arms are not really
banded but at rather definite intervals there are transversely placed, some-
what elliptical areas, which distally become narrow light bands. Between
these areas there are widely separated small white spots, 4-7 on each arm.
In young individuals (less than 10 mm. across the disk) the banding of the
arms is more definite and in most such specimens there is a large irregular
light blotch on the disk, although this may be broken up into several small
spots.

Holotype, M.C.Z. No. 6050, from Zaca St. 199 D-5.

There are 4 paratypes from the same station, 199 D-5, which is off
Monypenny Point, Gulf of Fonseca, 7 fms., on a muddy bottom. These range
from 16 to 24 mm. across the disk but call for no comment. From St. 199

4 griseus=gray, in reference to the prevailing color.

1940]

Clark: Notes on Echinodei’ms

347

D-7, there are 3 specimens, each of which deserves mention. This station,
while in the Gulf of Fonseca, was in 6 fms. olf La Union, El Salvador, and
the bottom is described as “mud, mangrove leaves.” The smallest specimen
is 18 mm. across a square disk as it is symmetrically tetramerous. Its color
is like those from off Monypenny Point but the buffy shade of the lower
surface is deeper. A second specimen, 22 mm. across, has a similar pale buff
lower surface but the upper side is a light sandy gray; the blotches and
marks and arm-bands are as in the darker gray individuals. The third speci-
men, 23 mm. across, also has the pale buff lower surface, but the upper side
has a definitely reddish cast, which makes the lighter gray blotches and arm-
bands stand out more sharply than in any other specimen. One can but
wonder whether the mangrove leaves in the mud were a factor in causing the
brighter coloration.

From St. 213 D-6-9, off Cedro Island, Gulf of Nicoya, Costa Rica, 4-6
fms., mud, sand and shell, there are 15 specimens of Ophiolepis, 6-13 mm.
across the disk, which are apparently the young of this fine species. The gray
coloration with white or light gray markings on the disk and in narrow bands
on the arms is like that of the adults and even in the smallest individual
there are 4 arm-spines on some at least of the basal arm-segments, while in
variegata of such a size there are usually but 2. The arms are noticeably
slender and arched. The bristling arm-spines on the basal half of the arm
are probably a result of sudden immersion in alcohol, and not a normal
condition.

This large and handsome species was at first regarded as a form of
variegata but the character and arrangement of the arm-spines, the colora-
tion, and the longer arms preclude such a decision. The color suggests plateia
Ziesenhenne but the length and character of the arms set it off very decidedly
from that species. There seems to be no doubt therefore that it must be
recognized as a distinct species.

Geographical Distribution-. Known only from the Zaca 1937-38
collecting.

Echinoidea.

Eucidaris thouarsii (Agassiz & Desor).

Cidaris thouarsii Agassiz & Desor, 1846. Ann. Sci. Nat. (3), 6, p. 326.

Eucidaris thouarsii Doderlein, 1887. Jap. Seeigel, p. 42.

There are 10 specimens of this common urchin but only 2 are adult and
none is full grown. They were collected at the following places:

Mexico, Manzanillo, 30 fms., 2 young.

Mexico, Sihuatanejo, 2 adults and 2 young.

Panama, Bahia Honda, 30-50 fms., 3 young.

Geographical Distribution: Lower California to Panama; also Cocos
and Galapagos Islands; low water to 50 fms.

Stylocidaris dubia (H.L. Clark).

Tretocidaris dubia H.L. Clark, 1907. Bull. M.C.Z., 51, p. 204.

Stylocidaris dubia Mortensen, 1909. Ech. Deutsche Siidp. Exp., p. 52.

There are 4 very fine adults of this rather uncommon cidarid, 35-40
mm. in test diameter, from St. 214 D-4, 14 miles south by east off Judas
Point, Costa Rica, in 61 fms., and 41 young individuals, 5-25 mm. through
the test, from St. 224 D-l-3, on Hannibal Bank, Panama, in 35-40 fms.

Geographical Distribution: Gulf of Panama to Gorgona Island, Colom-
bia; 15-112 fms.

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Astropyga pulvinata (Lamarck).

Cidarites pulvinata Lamarck, 1816. Anim. s. Vert., 3, p. 59.

Astropyga pulvinata Agassiz & Desor, 1846. Ann. Sci. Nat. (3) , 6, p. 345.

There are 3 notable specimens of this very handsome sea-urchin. One
from St. 232, off Gorgona Island, Colombia, 2-8 fms., sand, is probably the
smallest specimen yet recorded ; it is scarcely 4 mm. in diameter, with bright
red interambulacra, brown-violet ambulacra, and white oral surface; the
slender, fragile spines are 5 mm. long, pale yellowish-green banded with
brown-violet. The other specimens are adult and were taken at St. 195 D-2,
in Port Guatulco, Mexico, in 3 fms. on sand; the smaller is 85 mm. across,
brown-violet above except for sharply defined, triangular areas on the distal
portion of each intrambulacrum, which are light reddish-brown, and fade
out into the very pale brown of the oral surface; the spines are pale green
on the aboral side of the test, more nearly white orally, but prettily marked
with numerous narrow rings (6-12) of dull violet. The larger specimen is 110
mm. across with the test colored much as in the smaller but the spines show
little banding except at the very base of a few; orally they are nearly white
but at the ambitus they become dull violet basally and dorsally most of the
spines are violet, although some are white at the tip. Evidently the green
tint which is so conspicuous on the spines of the young individuals tends to
disappear with age.

Geographical Distribution : Gulf of California to Panama; low water to
36 fms.

Arbacia incisa (A. Agassiz).

Echinocidaris incisa A. Agassiz, 1863. Bull. M.C.Z., 1, p. 20.

Arbacia incisa H.L. Clark, 1913. Bull. Amer. Mus. Nat. Hist., 32, p.

220.

There are 5 small specimens from St. 126 D-18, -19, southeast of Cedros
Island, Lower California, 20-25 fms., notable only for their uniformly black-
ish color. They range from 6 to 23 mm. in test diameter.

Geographical Distribution : Lower California to Zorritos, Peru; also
Galapagos Islands; low water to 29 fms.

Lytechinus anamesus H.L. Clark.

H.L. Clark, 1912. Mem. M.C.Z., 34, no. 4, p. 254.

A single small specimen, 18 mm. in diameter, taken at St. 126 D-13, off
Cedros Island, Lower California, in 45 fms., is the only representative of this
species, so characteristic of southern California.

Geographical Distribution: Santa Barbara, California, to Point San
Bartolome Bay, Lower California, 30-60 fms.

Lytechinus pictus (Verrill).

Psammechinus pictus Verrill, 1867. Trans. Conn. Acad., 1, p. 301.

Lytechinus pictus H.L. Clark, 1912. Mem. M.C.Z., 34, no. 4, p. 258.

This is another sea-urchin characteristic of southern California and
the west coast of Mexico. It is represented in the Zaca collection by 10
very young specimens, 3.5-13 mm. in diameter, taken at St. 126 D-18 off
Cedros Island, Lower California, in 20 fms. The relationship between pictus
and anamesus is very close and it is not unlikely that the latter is only a

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Clark: Notes on Echinoderms

349

long-spined form of pictus characteristic of deeper water, for it ranges from
20 down to 113 fms. while pictus is characteristic of the more strictly littoral
zone.

Geographical Distribution : Monterey, California, to Gulf of California;
also Cocos Island ; low water to 50 fms.

Toxopneustes roseus (A. Agassiz).

Boletia rosea A. Agassiz, 1863. Bull. M.C.Z. 1, p. 24.

Toxopneustes roseus Mortensen, 1903. “ Ingolf ” Ech., pt. 1, p. 136.

This large sea-urchin characteristic of the littoral zone of western Cen-
tral America is represented by 3 specimens 65 to 80 mm. in diameter. The
label says they were “flesh-colored” in life. They were taken at St. 222 D-3,
in Bahia Honda, Panama, in 8 fms.

Geographical Distribution : Mazatlan, Mexico, to Port Utria, Colom-
bia; low water to 30 fms.

Strongylocentrotus purpuratus (Stimpson).

Echinus purpuratus Stimpson, 1857. Crust. Ech. Pac. Shores N. Amer.,

p. 86.

Strongylocentrotus purpuratus A. Agassiz, 1872. Rev. Ech., pt. 1, p. 165.

A specimen 45 mm. in diameter is undoubtedly a representative of this
northern species. It was taken at Guadeloupe Island, off Lower California,
Nov. 8, 1937, but the Albatross took purpuratus in 1911 near Cedros Island,
which is somewhat further south.

Geographical Distribution : Puget Sound to Lower California; low water
to 10 fms.

Echinometra vanbrunti A. Agassiz.

A. Agassiz, 1863. Bull. M.C.Z. , 1, p. 21.

This well known Mexican sea-urchin was taken at the following places :

Mexico: Sihuatanejo Bay, 1 very small specimen, 9 mm. long.

Nicaragua: Corinto, Cardon Island, tide pools, 2 small adults.

Costa Rica: Uvita Bay, under rocks on reef, 1 small adult.

Geographical Distribution : Lower California to San Francisco Bay,
Ecuador; also Cocos and Galapagos Islands; low water to 10 fms.

Clypeasfer europacificus H.L. Clark.

H.L. Clark, 1914. Mem. M.C.Z., 46, p. 27.

There are 5 representatives of this fine clypeastroid in the collection.
Two large adults, 150 mm. long, yellow-green in color, and two young ones,
50 and 54 mm. long, dull purple above but nearly white underneath, were
taken at St. 214 D-l, 14 miles south by east of Judas Point, Costa Rica, in
42-61 fms. on a muddy bottom. The fifth specimen is a very young one,
only 14 X 13 mm. ; the upper surface is light dull purple, with pale greenish
spines, while the oral side is light greenish-yellow, almost white. It was
taken at St. 224 D-l, -2, -3, on Hannibal Bank, 35-40 fms.

Geographical Distribution : Gulf of California to Panama; also Gala-
pagos Islands; 7-56 fms.

350 Zoologica: New York Zoological Society [XXV:22

C lypeaster ochrus H.L. Clark.

H. L. Clark, 1914. Mem. M.C.Z., 46, p. 30.

There are 2 Clypeasters from St. 195 D-9, Port Guatulco, Mexico, 7 fms.,
green sand and shell, which are typical ochrus in form and tuberculation
of the test, but differ so markedly in color that I have hesitated over calling
them by that name. The test is reddish-white, most nearly white on the
dorsal interambulacral areas, most nearly dull rose-red on the poriferous
areas of the petals and on the oral surface. The spines are dirty white
or pale greenish but often have a rosy-tinge and the terminal portion may
be quite roseate or even dull-red. The smaller specimen, 59 X 55 X 15 mm.,
is more roseate than the larger, 80 X 73 X 20 mm. If this coloration is
natural, and there is not the slightest reason for thinking it artificial, and
should prove to be constant in any considerable group or area, it would
warrant a varietal name because it is so conspicuously different from
the usual color.

Geographical Distribution: Isabel Island, Mexico, to Port Utria, Colom-
bia; low water to 30 fms.

C lypeaster speciosus Verrill.

Verrill, 1870. Amer. Jour. Sci. (2) 49, p. 95.

There are 2 small Clypeasters from St. 184 D-2, off Manzanillo, Mexico,
30 fms., which are best referred to this species. They are obviously young but
in the form of the test and the character of the petals seem to be speciosus.
The larger is 65 X 55 X 17 mm., the smaller 44 X 39 X 11. They are yellow-
brown in color, more brown above, quite yellow orally, especially the smaller
one. The oral surface is rather flat but is distinctly depressed about the
mouth. The petals are relatively narrow but well opened at the tip. The
periproct is strictly marginal in both specimens but I have never seen any
other case in speciosus although I have examined a number of specimens of
which several were younger than these. Because of this position of the peri-
proct and the light yellow-brown color, I have hesitated to call these young
Clypeaster speciosus but I hesitate even more to consider them a “new”
species.

Geographical Distribution: Lower California, also Cocos and Galapagos
Islands; low water to 50 fms.

Moira clotho (Michelin).

Moera clotho Michelin, 1855. Rev. Mag. Zool., p. 247.

Moira clotho A. Agassiz, 1872. Rev. Ech., pt. 1, p. 147.

A superb series of this apparently rare spatangoid was taken at
St. 213 D-2, off Cedro Island, Gulf of Nicoya, Costa Rica, in 4 fms. on a
muddy bottom. There are 11 specimens ranging from 9 mm. X 8 X 6 to a
very fine adult, 50 X 45 X 37. The two smallest specimens are more or less
badly damaged but the others are in beautiful condition. All are pale fawn
color, the fascioles and petals somewhat darker, the enlarged tips of the
spines on the plastron, pure white.

Geographical Distribution: Lower California to Panama; low water to
10 fms.

1940]

Clark: Notes on Echinoderms

351

Pleagiobrissus pacificus5 sp. nov.

Plate II; figs. 3 & 4.

Test elongated and flattened, 36 mm. long, 25 mm. wide and 11-12 mm.
high. So similar to P. grandis of the West Indian region that it is difficult
to feel sure what characters are specific and what are merely due to youth-
fulness. The smallest specimen available of grandis is nearly twice as large,
63 X 45 X 23 mm. The Pacific species seems to be flatter and more rounded
anteriorly, but the only character shown which would seem to be surely
specific is in the ventral ambulacra. In the West Indian species these
ambulacra, on each side of the ventral plastron are extraordinarily narrow
and the plastron, like the lateral interambulacra, is densely covered with
tubercles; this is as true for the smallest as for the largest specimens. In
the specimen from the Pacific, these ambulacra are comparatively wide and
the plastron itself is narrower and less well-defined. This difference is best
shown by the actual measurements. In the smallest grandis, measured across
the middle of the ventral surface, we find the plastron is 14 mm. across, the
bare ambulacra on each side, only 3 mm., and the lateral interambulacra (to
test margin) 12.5 mm. In the type of pacificus the same measurements are
5, 3 and 7 mm. Thus in pacificus, the plastron is not equal in width to the
two ambulacra while in grandis it is more than double their combined
width; in pacificus, the ambulacra are almost half as wide as the lateral
tuberculated areas but in grandis they are scarcely one-fourth as much. As
a result of these differences the lower surface of pacificus looks quite unlike
that of grandis. One can only speculate as to how much of this difference
would persist in a pacificus 100 mm. long. The spines and pedicellariae in
the two species are similar but only a few pedicellariae are to be seen in the
specimen from the Pacific. One large tridentate with very narrow, widely
divergent valves was found and 2 or 3 of the “long globiferous” can be seen
near the posterior end of ambulacrum 5. A few very minute ophicephalous
pedicellariae are to be seen among the ventral spines. The peristome in
pacificus is relatively long and little depressed and the plates covering it are
notably longer than wide; but these are very likely youthful rather than
specific characters. The color of the specimen is very pale fawn, nearly white
— the test dorsally is white.

Holotype, M.C.Z. No. 7625, from St. 224 D-1,-2,-3, on Hannibal Bank,
Panama, in 35 fms. ; in view of the character of the specimen and the known
habits of the West Indian species, it is pretty certain that pacificus was
secured in the third haul of the dredge, which was on a bottom of sand,
shells and algae.

Besides the holotype, a second specimen was secured at the same time,
but is so much younger, it is of no help in defining the species. It measures
19 X 14.5 X 6.5 mm. and shows the same wide ventral ambulacra as the
holotype but even less well defined and more obviously youthful.

The discovery of this genus on the western coast of Central America is
of the greatest interest. It furnishes another link connecting the Panamic
with the West Indian fauna and is particularly interesting in showing that
striking novelties are still to be found in the Panamic region in spite of the
large amount of collecting and dredging that have been done there in the
past forty years.

Geographical Distribution-. Known only from the Zaca collection.

5 pacificus, in reference to the habitat, all other members of the genus occurring in the Atlantic
region or Mediterranean.

352

Zoologica: Neiv York Zoological Society

Meoma grandis Gray.

Gray, 1851. Ann. Mag. Nat. Hist. (2), 7, p. 132.

A single specimen of this spatangoid was taken at St. 195 D-16, in Port
Guatulco, Mexico, 10 fms., sand. It measures 98 X 85 X 48 mm. and the color
is a real brown.

Geographical Distribution : Gulf of California and west coast of Mexico,
20-60 fms.

Lovenia cordiformis A. Agassiz.

A. Agassiz, 1872. Bull. M.C.Z., 3, p. 57.

A small specimen, 35 X 28 X 14 mm., of this typically southern Cali-
fornian spatangoid, was taken at St. 126 D-19, southeast of Cedros Island,
Lower California, in 25 fms. The color is definitely light brown.

Geographical Distribution : Santa Barbara, California, to Guayaquil,
Ecuador; also Cocos, Galapagos and Hawaiian Islands.

EXPLANATION OF THE PLATES.
Plate I.

Fig. 1. Othilia aculeata Gray. Aboral surface. X 1.5.

Fig. 2. Oral surface. X 1.5.

Plate II.

Fig. 3. Plagiobrissus pacificus sp. nov. Aboral surface. X 2.
Fig. 4. Oral surface. X 2.

CLARK.

PLATE I.

FIG. 1.

FIG. 2.

NOTES ON ECH1NODERMS FROM THE WEST COAST OF CENTRAL AMERICA.

CLARK.

PLATE II.

FIG. 3.

FIG. 4.

NOTES ON ECHINODERMS FROM THE WEST COAST OF CENTRAL AMERICA.

Breder: Nesting Behavior of Eupomotis gibbosus

353

23.

The Nesting Behavior of Eupomotis gibbosus (Linnaeus)

In a Small Pool.

C. M. Breder, Jr.

New York Aquarium
(Plates I & II; Text-figures 1 & 2).

Introduction.

A general study of the nesting behavior of the family Centrarchidae
has already been given in this journal (Breder, 1936). The present con-
tribution is based on a continuation of these studies on one species, Eupo-
motis gibbosus (Linnaeus), and compared with other studies on a very
different type of fish, Ameiurus nebulosus (Le Sueur), under an identical
situation (Breder, 1939).

In the spring of 1940 fourteen mature Eupomotis were placed in a pool
on the writer’s property. This pool has been described in detail by Breder
(1939), and its general form may be seen in Text-fig. 1. Due to an unusually
hard freeze the Ameiurus which occupied it the previous year were all
frozen out, although the Rhodeus amarus which shared the pool with them
survived. Thus the pool, in addition to being physically similar, was also
alike in the organisms present in addition to those under study.

The Eupomotis were collected from the lake in Central Park, New York
City, at 59th Street, as part of a considerable number taken to the New
York Aquarium. These fish were in their second year and were 10 cm. long.
The chief purpose of the experiment was to note the number of nests con-
structed in a given body of water by a known number of fish and to compare
their behavior with that of the previously studied Ameiurus. It was impos-
sible to determine the numbers of each sex with certainty or to keep track
of each individual fish under the conditions of the experiment.

Reproductive Schedule.

Pre-spawning Behavior.

The fish were released on May 5 when the water was about 18.5° C. As
they began nesting on May 18, by which time it had reached 20°, there was
not much time for observation of the pre-spawning behavior. After a few
days of accommodation and exploration, the fish settled in a school in or over
the deepest part of the pool, near the water lilies in the north arm, rising
to near the surface on sunny days and staying in the deeper parts at night
and on dark days. Here they remained for the most part, with occasional
short sallies to other places until nest construction began. Those fish not
engaged in this activity continued to behave in a similar manner except
that as the season advanced they moved from the northwest part of the 24"

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[XXV :23

contour to the southeast. Much later a group of four or five was frequently
seen in the southwest extension of the 18" contour which reaches into the
southwest arm of the pool.

Nest Construction.

In all, seven nests were excavated, and since there were fourteen fish it
is tempting to assume that the sexes were equally divided and that each
male built one nest. It is doubtful if such is actually the case, however, as
there 'is reason to believe that some of the males built more than one nest,
as will be shown subsequently.

The nests are numbered in Text-figure 1 in the order of their construc-
tion, which gives a measure of order of choice of locality. Considered this
way, there were actually only three major choices, since nest 6 and nest 7

Text-figure 1.

Sketch map of the pool in which the present studies were made. The circled num-
bers, 1 to 7, indicate the nests of Eupomotis in the order constructed. The contour
lines indicate the depth of water in inches. See text for explanation.

1940]

Breder: Nesting Behavior of Eupomotis gibbosus

355

encroached on the older nest 3 and 2 respectively. While nest 5 did not en-
croach on nest 1, it closely approached it; but nest 5 was built while there
were eggs in nest 1. Nest 4 split the distance between nest 2 and nest 3, but
we have reason to believe that 2 and 4 were made by the same fish, as they
were alternately worked by one male. Thus it is apparent that the only avail-
able areas suited to these fish in this pool are in the upper tips of the two
arms. The contour lines show why this should be so, for at all other places
the shore line is too steep for sunfish to build nests, and apparently under
the conditions in this pool they seek water between 6" and 12" in depth.
The gravel bottom made nesting a simple matter of cleaning off such fine
detritus as might be present. Not much of a “dish” could be formed as the
large size of the gravel precluded much shifting by the efforts of these
small fish.

Text-figure 2.

The nests of Ameiurus of the previous year in the same pool. (After Breder,
1939). The circled numbers, 1 to 3, indicate the catfish nests in the order con-
structed. Differences in the symbols for aquatic plants, et cetera, indicate the
only changes in the pool between the two years. See text for explanation.

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[XXV : 23

TABLE I.

Schedule of Events in Reproducing Eupomotis.

Dates
May 5
18

19

20
21
22

23

24

25

26

27

28

29

30

31

June 1
2

3

4

5

6

7

8
9

10

11

12

13

14

15

Nest Number.

1 2 3 4 5

B

A A

B

(1)

Events.

14 fish placed in pool.
Nest just begun.

Nest begun.

Nest begun.

Temperature °C.
Air Water

23.0 20.0

25.3 20.0

RC A

Nest #2 reoccupied, courting
vigorously.

RE

(2)

RC

BC

E

L

Nest reoccupied, eggs.

Eggs hatched.

Nest reoccupied, courting.
Nest begun, courting.

2d eggs, young on nest.
Young left nest.

25.8 24.8

24.8 24.8

24.0 25.3

Absent from locality — observations suspended.

July

14

15

16

17

18

19

20
21
22

23

24

25

26

27

28

29

30

A A Y A A A A

L

A

RE

Young on nest #3. Nests
#5, 6 and 7 made during
absence and abandoned.

Young left nest #3. Nest
#7 reoccupied and spawning.

22.5 21.0

26.5 21.3

A Heavy storm this date.

A — nest abandoned. B — nest begun. C — courting. E — eggs present. H — -eggs
hatched. L — young fish left nest. R — nest reoccupied. Y — young on nest.

(1) — Plate I taken this day.

(2) — Plate II taken this day.

1940J

Breder: Nesting Behavior of Eupomotis gibbosus

357

If Text-figure 1 is studied closely it will be seen that the presence of
nest 1, because of the conformation of the shore, bottom and plants, pre-
cluded another nest in that entire arm except very close to it. The next nest,
2, was built as far from it as the pool allowed, being in the other arm. The
third nest, now that the other arm was occupied, also was built in the
southern arm and as far from nest 2 as the depth permitted. A considera-
tion of the bottom contours and the desire of these fishes to pick the sunniest
places well accounts for this and the preceding positions. With this restric-
tion, the only remaining place for a nest not in contact or encroaching on
another is that site on which nest 4 was constructed. After this there re-
mained no other place available in the pool in which a nest could be built
remote from one already constructed. In fact, the only remaining place
where overlapping was not necessary was that place selected for nest 5 which
was peripheral to nest 1.

When construction had reached this point no further nests could be
built in the spaces available except along the south shore of the south arm.
This place was apparently entirely out of the question because of shade.
The two remaining nests, 6 and 7, were consequently made so close to 3 and
2 respectively that they encroached on the latter as shown. Table I gives the
time sequence of these events.

Courting Activities.

An interesting note on the courting activities of these fish was made
June 8 on nest 2. Here a female approached and was very vigorous, entering
the nest voluntarily and circling the male and nudging the exposed gravel
with her snout. She was not feeding but seemed in a high state of sexual
excitement. It gave the impression that the activities of the male are largely
one of indicating his presence and willingness and of indicating the location
of the nest, the female actually doing the “selecting.” It would seem that
the chasing of females by males is of that nature rather than a matter of
“driving” the female over the nest as is usually reported. The latter would
seem to be an impossible procedure with all the open water normally avail-
able to these fishes. Usually the male is so willing to spawn that this feature
is not seen. For some reason this male did not seem to be ready or able to
perform his part and finally the female departed.

On June 9 a single male attended both nest 4 and nest 2. While he spent
most of his time on the new nest, he did give attention to both, and a ten-
tatively approaching female caused him to display over both, depending on
which one she happened to be nearer. At this time another male attempted to
reoccupy nest 3, presumably its maker which had abandoned it some time
before, but was driven off by the extreme vigor of the proprietor of nests 2
and 4. Nevertheless no eggs resulted from this activity.

Effects of Weather.

As is well known, the activity of sunfish is considerably controlled by
temperature and light. This was very evident as may be noted in Table I
by the frequent “Abandonments” and “Reoccupations” of nests. In late May
and early June there was an unseasonable amount of cloudy weather and
chilly days. It is reflected in the activity and lack of activity indicated in
this table. That these factors do not completely control sunfish activity is
indicated by the lack of exact synchronization of behavior. For example, on
May 24 two nests were abandoned and a third begun. Presumably these

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[XXV :23

vagaries in behavior have to do with variations in the physiological impetus
of the various individuals. When fish abandoned their nests they rejoined
the general school which would normally have been composed of females
alone.

The last nest was abandoned on July 30, on which date a heavy wind
and rainstorm did violent local damage, including the felling of a tree across
the pool.

Attention to Eggs and Young.

It is certain that at least three of the nests produced young, that is, nest
1 (two batches), nest 3 and nest 7. Later in the season young of four dif-
ferent size groups which corresponded to the spacing of these four hatchings
were evident. What happened during the period of personal absence is not
entirely clear but family reports indicate that there was no further egg pro-
duction.

Nest 1, the first to be established and perhaps therefore by the most
vigorous male, had at one time both eggs and young of very different ages
in the nest. The young left the nest the day after new eggs were deposited.
While it is known that sunfish will spawn on repeated days with a number
of females, the writer had not previously encountered or seen reports on
two batches of such a difference in age.

Discussion.

Compared with the nesting dates on this species recorded by Breder
(1936) for the same general region, the present dates agree very well. His
early date was May 11, the present May 18; his late date August 14, the
present July 30.

The depths selected by the size of fish involved agree perfectly with his
Figure 2. The diameters of the nests, about 20 cm., fall in the Pines Lake
group of that paper which also happens to be the closest geographically. It
will be noted that in the south arm of the pool, the only place where any
selection was possible, the nests are ranged along its northern edge, which
is what should be expected since the other side is shaded by the south bank.
Normally one expects sunfish nests on the north side of a small pond for
this reason, as has already been discussed in the earlier paper.

Compared with the catfish which occupied this pool the preceding year
(see Text-fig. 2), some curious differences and similarities appear. Sunfish
nest 1 occupied exactly the same place as catfish nest 3. In other words, the
first choice of the former was the last choice of the latter. The other two
catfish nests occupied places in which sunfish would surely not be expected to
nest, in a cavity and in a well-sheltered spot. This still leaves unexplained
why the one catfish nested in such a sunfish-like manner — at a place which,
as a matter of fact, was the first choice of the latter — a choice that might
have been guessed on a priori grounds. In this connection it should be men-
tioned that the pool suffered from no limitation of presumably choice cat-
fish nesting sites, but that they did nest as remote from each other as prac-
ticable. This wide selection of sites was not true in the case of the sunfish
which were strictly limited as to suitable sites, and indeed the pool was a
trifle too shaded to be ideal for such a species. Since the first catfish nest
occupied a position near the central part of the “U” formed by the pool, the
only thing that was left for the other two pairs was to retreat up each arm
as far as other factors permitted. Compared with the order of selection ex-
ercised by the sunfish, which is indicated in the body of the paper, we actu-
ally have a similar type of centrifugal influence expressed in two very dif-
ferent kinds of nest building fishes. These sunfish nests compared with those

1940]

Breder: Nesting Behavior of Eupomotis gibbosus

359

of Pines Lake (Breder, 1936) show thoroughly comparable selective reac-
tions. The remaining data in the body of the paper are confirmatory of
those in the earlier study.

Summary.

1. Earlier studies on Eupomotis are confirmed under semi-controlled con-
ditions.

2. A comparison is made between the nesting habits of Eupomotis and
Ameiurus under identical conditions in which it is seen that the first
choice of one was the last choice of the other.

3. The role of nesting males in pursuing females is indicated to be indica-
tive of his presence and willingness to mate, rather than an overpower-
ing effect.

4. Males may operate, intermittently, over two close nests.

5. Selection of nesting sites of Eupomotis and Ameiurus is shown to be
controlled by a centrifugal influence between nesting individuals, modi-
fied by the limitations of the availability of suitable nesting sites.

References.

Breder, C. M., Jr.

1936. The Reproductive Habits of the North American Sunfishes (Family
Centrarchidae) . Zoologica 21 (1) : 1-48.

1939. Variations in the Nesting Habits of Ameiurus nebulosus (Le Sueur).
Zoologica 24 (25) : 367-378.

360

Zoologica: New York Zoological Society

EXPLANATION OF THE PLATES.

Plate I.

Nest 3 on May 25. See Table I. This nest was begun the day previous. It was im-
possible to photograph the nest groups, due to difficult lighting conditions and
interfering vegetation.

Plate II.

Nest 1 on June 5. See Table I. This shows the nest on the day after the first
batch of eggs was laid.

BREDER.

PLATE I.

THE NESTING BEHAVIOR OF EUPOMOTIS GIBBOSUS (LINNAEUS)

BREDER.

PLATE II.

THE NESTING BEHAVIOR OF EUPOMOTIS GIBBOSUS (LINNAEUS)

Raney: Reproductive Activities of Hybrid Minnow

361

24.

Reproductive Activities of a Hybrid Minnow, Notropis
cornutus X Notropis rubellus.

Edward C. Raney

Department of Zoology, Cornell University, Ithaca, New York.

A considerable number of natural hybrids have been recognized in
recent years among American Cyprinidae. One of the most common is the
cross between the common shiner and the rosy-faced shiner, Notropis
cornutus (Mitchill) X Notropis rubellus (Agassiz). The subspecies of
Notropis cornutus involved appears to be unimportant and hybrids of
Notropis rubellus with subspecies cornutus, frontalis and chrysocephalus
have been found occasionally throughout their range. Hubbs & Brown
(1929, p. 37) have pointed out that hybrids of this combination were mis-
identified as Notropis pilsbryi by Forbes & Richardson (1909, p. 149).

It has generally been assumed that the hybridization in this as well as
in other minnow combinations is due to chance fertilization. The breeding
habits of Notropis cornutus and Notropis rubellus in some localities are
such that they are thrown into close contact at the spawning time. Both
species may utilize an unprepared gravel bottom in running water for
breeding but appear to prefer the pebble nests of the river chub, Nocomis
micropogon, and the horny-headed chub, Nocomis biguttatus, when these
are available. However, Notropis cornutus is quite adaptable and has been
observed (Raney, 1940, p. 6) digging a shallow nest in riffles. In the re-
ported observations of cornutus and rubellus spawning in close proximity,
it has been over a Nocomis nest. Hubbs & Brown (1929, p. 37) “observed
rubellus mating in midwater a few inches over the heads of breeding
cornutus.” They also noted that this hybrid combination was most common
below obstructions where a large number of cornutus and a considerably
smaller population of rubellus were present. This is in agreement with my
observations in Catatonk Creek at Candor, New York, (Susquehanna Drain-
age), where many hybrids have been collected and numerous others seen in
an area of gravel riffles below an impassable dam. At this locality practically
all of the spawning of rubellus and much of that of cornutus takes place
over the nests of Nocomis micropogon. The use of Nocomis nests by Notropis
cornutus or Notropis rubellus has been reported by Greeley (1929, p. 172,
and 1938, p. 67), Hankinson (1932, p. 417), Hubbs & Brown (1929, p. 37),
Hubbs & Cooper (1936, p. 49), Raney & Lachner (1939, p. 160) and Raney
(1940, p. 7). Hankinson (1932, p. 417) describes the breeding behavior of
rubellus as follows: “Spawning could be seen among these rosy-faced
minnows since it always took place high above the other fish of the aggre-
gation (mostly cornutus) and several inches above the gravel. The males
would collide with the females, striking them on or near the head with their
roughened snouts, and the two would contort themselves in a way that
suggested spawning.”

Several times in the past few years, during late May and early June,
both cornutus and rubellus and hybrids between them have been observed

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[XXV :24

over Nocomis nests. The hybrids of both sexes appeared to be in a condition
of sexual ripeness. The nuptial tubercles were well developed in the males,
and they showed the red colors that are apparent in the breeding season.
The females contained well developed eggs and some of them showed
red nuptial colors and tubercles. A suspicion that these hybrids were
capable of spawning was confirmed during the past spring (1939) when a
number were seen performing the breeding act.

Recognition in the Field.

The recognition of these hybrids in the field became relatively easy
after one had watched them for many hours. The hybrids were inter-
mediate in body shape between the parent species, cornutus and rubellus,
and could be distinguished by this character alone. The spawning hybrids
were slightly larger than any male rubellus seen at this locality. However,
the breeding color of the males furnished the best clue to their identification.
The following color characters (not apparent in preserved specimens) were
noted in breeding males when viewed under water under natural conditions.

Notropis c. cornutus
The mid-dorsal stripe and the
stripe on each side of the back
were conspicuous and varied
from a bright silver to a light
green against the dark back-
ground of the back. These stripes
were outstanding highlights and
could be seen at a considerable
distance. The snout was dusky.

Hybrid
The stripes
were easily
seen but
were red. The
snout was
slightly red.

Notropis rubellus
The stripes were ab-
sent or indistinct and
the anterior half of
the body was bright
red. The snout was
bright red.

The following brief color descriptions of males were taken after several
hours’ preservation in 10 per cent, formalin. In Notropis c. cornutus the
sides, the branchiostegal region and at times the lower part of the cheek
and opercle, the paired fins, and the outer or posterior border of the median
fins were red. The upper parts of the cheek and opercle were a slate blue
as was a narrow band immediately behind the opercle. The back was nearly
black, with the mid-dorsal and lateral light stripes mentioned above.

In the hybrid, Notropis c. cornutus X Notropis rubellus, the sides,
branchiostegal region, the lower cheek and opei’cle, the back at the base
of the dorsal fin, the pectoral and to a lesser extent the pelvic and anal fins
were red.

In Notropis rubellus the snout, chin, top of head, anterior half of body,
the base of the dorsal fin, the base of the pectoral fins and to a lesser degree
the base of the ventral fins were red.

The degree of development of breeding tubercles in the male hybrids
is interesting as they appear to have both the pattern and size of the
tubercles possessed by both the parent species. In ripe males of rubellus
many small tubercles cover the chin, snout, top and side of head, upper two-
thirds of the opercle, and the posterior edge of almost every scale on the
back and upper sides. All fins are covered with small tubercles. They are
best developed on the upper sides of the pectoral fin and least developed on
the anal fin. In males of cornutus there are fewer but larger and sharper
tubercles on the chin, snout, top of the head, and on the back in front of the
dorsal fin. A few minute tubercles are found on the upper sides. In male
hybrids there are many fine tubercles at approximately the same positions
on the body and fins as in the parent species. There are also several dozen
larger tubercles on the top of the head, as in cornutus.

1940] Raney : Reproductive Activities of Hybrid Minnow 363

Spawning of Hybrids.

Observations of spawning hybrids were made on June 10, 1939, in
Catatonk Creek, two miles south of Candor, New York. On this date
Notropis c. cornutus had about reached the end of its breeding season and
no spawning males of this species were seen (the spawning season had
reached its peak about June 1, when practically every Nocomis nest was
occupied by 5 to 25 cornutus males). Many ripe adults of Notropis rubellus
were seen over the same Nocomis nests, but were also seen as late as June
10 spawning over gravel shallows in riffles.

At 5 P.M. on June 10, when the water temperature was 69° F., the air
66° F., a group of hybrids, Notropis c. cornutus X Notropis rubellus, was
seen over a nest of Nocomis micropogon (the only species of Nocomis found
in Catatonk Creek). The nest, roughly two feet in diameter, consisted of
stones up to two inches in diameter piled to a height of four inches above
the general level of the gravelly creek bottom. It was located in moderately
swift water at a depth of 10 inches. The stream at this point was 70 feet
wide and the nest was about 25 feet from shore. Many Nocomis nests were
available in this area of Catatonk Creek but few were located in as fast a
riffle as this particular one and relatively few were occupied by spawning
fish at this late date.

At various times, from 10 to 16 male hybrids ranging in standard
length from 65 to 75 mm. (as determined by measuring captured specimens)
were seen facing upstream over the Nocomis nest. They attempted to
maintain a definite position and generally an individual did not move more
than five inches in any direction. They alternately moved to right and left
a few inches and, in general, their behavior was similar to that of spawning
rubellus, males of which hold positions facing upstream, shifting to either
side and striking an adjacent male from time to time but not moving far
from the original position. The rubellus and hybrid males differed in be-
havior from those of cornutus mainly in that they did much less fighting for
position and did not drive away predators slightly larger than themselves.

Many cornutus females were observed below the nest, occasionally
coming up over the males and, at times, continuing upstream in front of
the nest. A number of individuals of both sexes of rubellus were seen both
over the downstream half of the Nocomis nest and over the gravel below
the nest but they were not observed spawning although the males jockeyed
some for position. The rubellus males and females occupied a slightly
higher stratum than the cornutus females and the hybrids, which were
down over the pebbles of the nest so that the lower fins at times appeared
to be in contact with the bottom.

The spawning act was identical with that observed for cornutus as
recently reported in detail (Raney, 1940). A female cornutus came up-
stream over a hybrid male and dipped down to lie along side him. He threw
his caudal peduncle over hers and lifted his pectoral fin which was inserted
underneath the front end of her body. This activity forced her firmly
against the bottom of the nest, where she rested on her side with her vent
facing upstream. The eggs were forced out at the moment that the male’s
body was thrown into a curve. The breeding act was over in less than a
second. The female then swam downstream several feet but later returned
to spawn with the same or another male. The male hybrid remained at the
same place and was able to spawn again immediately. The breeding act
was noted a dozen times within 15 minutes, and the details were observed
clearly from a distance of about four feet. No female hybrids were observed
at this Nocomis nest.

Upon the completion of a spawning act several small fallfish, Leucoso-
mus corporalis, dashed to the spot and dug vigorously among the pebbles for
eggs. Several times male hybrids were also seen standing on their heads as

364 Zoologica: New York Zoological Society [XXV :24

if attempting to eat eggs. This behavior may, however, be a primitive type
of digging activity such as is developed to a greater degree in cornutus
males. A search revealed many eggs among and under the pebbles of the
nest. Eggs of both Notropis cornutus and Nocomis micropogon were
found, indicating that the nest had been utilized for some time. However,
it was impossible to isolate the eggs from a single mating under natural
conditions and, as yet, I have been unable to hatch cornutus eggs and raise
the young. With rare exceptions the cornutus females were larger than the
hybrid males with which they spawned. Males of cornutus probably spawn
occasionally with a larger female, but I have not seen it happen in many
observations over a period of years. After observing these spawning
hybrids for nearly an hour, as many as could be captured were preserved
and their identification checked.

On a previous occasion a male cornutus was seen spawning with a
female hybrid. This observation was also made in Catatonk Creek, just below
Candor, on May 31, 1939. At 2:30 P. M. a group of about 30 large cornutus
males were holding positions over a Nocomis nest. About a dozen individu-
als of rubellus were present over the lower side and below the nest. At one
time five hybrids were seen occupying the same area, mingling with rubellus.
A female hybrid which had some red coloration on the anterior third of the
body came up over a male cornutus, dipped down beside him and went
through a spawning act in a manner apparently identical to that followed
by a female cornutus. No repetitions of this spawning act took place during
several hours of observation, although cornutus spawned freely throughout
the afternoon.

Data prom Hybrids.

An examination of 18 cornutus X rubellus hybrids specimens collected
over a period of five years in a two-mile stretch of stream of Catatonk
Creek below Candor gives some interesting results. These specimens (in
the Cornell University Museum) were mostly taken from early May
through June, and they could easily be sexed by an examination of the
gonads. The sexes were equally divided among these 18 specimens, and all
but one small male (60 mm. in standard length) appeared to be capable of
producing sex elements. Dr. Carl L. Hubbs informs me that he, too, has
found a normal sex ratio in cornutus X rubellus hybrids, and that these
hybrids often show gonads which appear to be ripe and normally developed.

These data are interesting in the light of results obtained from a study
of aquarium-produced and natural hybrids of sunfishes as reported by
Hubbs & Hubbs (1933), who found several combinations of hybrid sun-
fishes to be predominantly males (81 to 95 per cent.). The evidence also
indicated that the resulting hybrids were sterile. Bailey & Lagler (1938, p.
605) also found a predominance of males (80 to 96 per cent.) in a sample of
natural hybrid sunfishes. However, a recently studied collection of a
hitherto unreported hybrid sunfish, Lepomis auritus X Lepomis cyanellus,
taken by Dr. A. H. Wright in a pond along the South Branch of the Potomac
River, one mile above Franklin, Pendleton Co., West Virginia, on August
14, 1931, shows an altered sex ratio in favor of the females. In this collec-
tion (examined by both Dr. Carl L. Hubbs and Dr. Reeve M. Bailey) there
were 6 male and 18 female hybrids (25 per cent, males). In this same
collection there were 17 males and 14 females of Lepomis auritus and 14
males and 17 females of Lepomis cyanellus. There also appeared to have
been some back-crossing, as some of the hybrids strongly approached
Lepomis auritus.

There is no direct proof that either males or females of the cornutus X
rubellus hybrids are fertile. However, as has already been mentioned, each
sex may perform the spawning act in nature with cornutus. Possibly the
hybrids may spawn with one another, although this act has not as yet been

1940] Raney: Reproductive Activities of Hybrid Minnow 365

observed. Male hybrids have been forced to emit milt upon pressure. The
fact that some hybrids appear to be closer in characters to one parent
species suggests that they may have resulted from a back cross. Hubbs &
Brown (1929, p. 36) reported “our large number of specimens (hybrids)
from numerous localities completely bridge over the wide gap in characters
between the two parent species.” These data also suggest that at times the
breeding act involving a hybrid and one of the parent species may be
successful.

There is considerable variation in the condition and position of the
eggs in the ovary of hybrids taken during May and June. In some the eggs
are all enlarged and appeared much as they do in either cornutus or rubellus
females which are ready for spawning. In a few ovaries the eggs are very
small and it appears that they would ripen much too late, if at all, for
the spawning season. In a few hybrids, for example one 73 mm. in standard
length taken on October 18, 1938, several unusually large eggs occur at
intervals among many small eggs. It seems that these fish may have
retained some eggs from the previous spring.

Table I.

Length-frequency distribution of mature hybrids, Notropis c. cornutus X
Notropis rubellus and Notropis rubellus taken during May and June
from Catatonk Creek, below Candor, New York.

Standard
Length
in mm.

Hybrids: Notropis c.
cornutus X Notropis
rubellus

Notropis

rubellus

Female

Male

Female

Male

44

2

46

4

48

i

5

50

3

4

52

4

1

54

1

2

56

1

58

2

i

60

i*

i

62

1

6

64

1

5

66

1

68

70

2

72

i

74

i

76

i

l

78

3

l

80

l

82

1

84

1

86

88

i

90

92

i

Number

9

9

25

19

Mean

80.89

70.44

58.00

48.95

Standard

Deviation

6.24

7.24

5.77

3.57

♦Immature.

366 Zoologica: New York Zoological Society [_XXV:24

Some of the female hybrids had tubercles and some red color on the
anterior part of the body but the majority had neither tubercles nor color.
Females as well as the males of rubellus are tuberculate at the spawning
time, and both develop red color. A female rubellus in nuptial color is
illustrated in Greeley (1930, plate 10). However the male rubellus is more
brightly colored and has more of the body covered with tubercles. In
cornutus the female is rarely colored and then usually only on the posterior
border of the caudal and anal fins, and she rarely, if ever, has breeding
tubercles.

The male hybrids taken during May and June had the breeding color and
breeding tubercles well developed. The one exception was a small individual,
60 mm. in standard length, which was sexed without question as a male but
was apparently immature. In the hybrids the lips were dusky as in rubellus,
rather than light colored as in cornutus.

As in rubellus, the females of the hybrids attain a greater length than
the males. Table I gives the length-frequency distribution of 18 hybrids
from Catatonk Creek below Candor, New York, collected from early May
through June over a period of five years. Length measurements of a random
sample of 44 specimens of Notropis rubellus in spawning condition, col-
lected at the same locality on June 10, 1939, are included. Seventeen addi-
tional hybrids were secured during October, 1938 and 1939, but since most
of these specimens were difficult to sex with accuracy they are not included
here. However all measurements of these October specimens fell within
the range as given in Table I. In cornutus the males attain a greater
length than the females as reported by Raney (1940, p. 5).

One large hybrid specimen placed in an aquarium in October, 1939,
proved to be extremely hardy and outlived several hundred specimens of
cornutus and rubellus placed in similar containers under similar conditions.
Not only did he outlive all of the specimens of the parent species, but a
dozen other species of cyprinids as well, and is thriving and active at the
present time (June, 1940).

Summary.

1. Hybrids between Notropis rubellus and several sub-species of Notropis
cornutus are fairly common in some localities, especially below dams and
over Nocomis nests during the spawning season in late May and early June.

2. The hybrids apparently result from a chance combination of sexual
elements as the parent species spawn at the same spot.

3. Hybrids can be recognized in the field by such characters as body
shape and coloration. Male hybrids have well developed tubercles and red
colors, as do some female hybrids.

4. Several male hybrids, Notropis c. cornutus X Notropis rubellus, were
observed spawning with female Notropis c. cornutus over a pebble nest of
Nocomis micropogon. On another occasion a female hybrid went through the
spawning process with a male Notropis c. cornutus.

5. The actual breeding behavior of male hybrids is identical with that
of Notropis cornutus although their behavior over a nest is similar to that of
rubellus. Eggs were deposited by cornutus. Many hybrids have seemingly
normal eggs and milt has been obtained from male hybrids upon stripping.

6. The sexes were evenly divided between 18 hybrids collected during
the breeding season over a period of five years.

7. Female hybrids attain a significantly greater maximum and mean
standard length than males. This is also true of female rubellus but in
cornutus the males are larger.

8. One hybrid kept in an aquarium over winter appeared to be an un-
usually hardy individual, outliving all specimens of parent species kept in
similar containers under similar conditions.

1940]

Raney: Reproductive Activities of Hybrid Minnow

367

Bibliography.

Bajley, Reeve M. & Lagler, Karl F.

1938. An analysis of hybridization in a population of stunted sunfishes in
New York. Pap. Mich. Acad. Sci., Arts and Letters, Vol. 23: 577-606,
figs. 1-5.

Forbes, Stephen Alfred & Richardson, Robert Earl

1909. The fishes of Illinois. Nat. Hist. Surv. Illinois, Vol. 3: i-cxxi, 1-357,
figs. 1-76, pis. 1-68.

Greeley, J. R.

1929. Fishes of the Erie-Niagara Watershed. In: A biological Survey of
the Erie-Niagara System. Suppl. 18th Ann. Rept., 1928: 150-179, pis.
1-8.

1930. Fishes of the Lake Champlain Watershed. In: A Biological Survey
of the Champlain Watershed. Suppl. 19th Ann. Rept. N. Y. Cons. Dept.,
1929: 44-104, pis. 1-16.

1938. Fishes of the area with annotated list. In: A Biological Survey of the
Allegheny and Chemung Watersheds. Suppl. 27th Ann. Rept. N. Y.
Cons. Dept., 1937 : 48-73, pis. 1-2.

Hankinson, Thomas L.

1932. Observations on the breeding behavior and habitats of fishes in south-
ern Michigan. Pap. Mich. Acad. Sci., Arts and Letters, Vol. 15: 411-
425, pis. 33-34.

Hubbs, Carl L. & Brown, Dugald E. S.

1929. Materials for a distributional study of Ontario fishes. Trans. Royal
Canadian Institute, Vol. 17 (Part I) : 1-56.

Hubbs, Carl L. & Cooper, Gerald P.

1936. Minnows of Michigan. Bull. Cranbrook Inst. Sci., No. 8: 1-95, pis. 1-10.

Hubbs, Carl L. & Hubbs, Laura C.

1933. The increased growth, predominant maleness, and apparent infertility
of hybrid sunfishes. Pap. Mich. Acad. Sci., Arts and Letters, Vol. 17 :
613-641, figs. 69-71, pis. 64-65.

Raney, Edward C.

1940. The breeding behavior of the common shiner, Notropis comutus
(Mitchill). Zoologica, Vol. 25 (Part I) : 1-14, fig. 1, pis. 1-4.

Raney, Edward C. & Lachner, Ernest A.

1939. Observations on the life history of the spotted darteT, Poecilichthys
maculatus (Kirtland). Copeia, (3): 157-165, figs. 1-2.

NEW YORK ZOOLOGICAL SOCIETY

General Office: 90 Broad Street, New York City

OFFICERS

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Secretary, David H. McAlpin
Chairman, Executive Committee, Laurance S. Rockefeller
Treasurer, Cornelius R. Agnew

General Director, Zoological Park and Aquarium, Allyn R. Jennings
Assistant General Director, Harry Sweeny, Jr.

SCIENTIFIC STAFF

Zoological Park

Raymond L. Ditmars, Curator of Mammals and Reptiles
Lee S. Crandall, Curator of Birds
Leonard J. Goss, Curator of Health
Claude W. Leister, Curator of Educational Activities
William Bridges, Editor and Curator of Publications

Aquarium

Charles M. Breder, Jr., Director
Christopher W. Coates, Aquarist
Ross F. Nigrelli, Pathologist
G. M. Smith, Research Associate in Pathology
Homer W. Smith, Research Associate in Physiology

Department of Tropical Research

William Beebe, Director
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Editorial Committee

Fairfield Osborn, Chairman

Allyn R. Jennings Charles M. Breder, Jr.

William Beebe William Bridges

ZOOLOGICA

SCIENTIFIC CONTRIBUTIONS
OF THE

NEW YORK ZOOLOGICAL SOCIETY

VOLUME XXV
Part 4

Numbers 25-34

PUBLISHED BY THE
THE ZOOLOGICAL PARK,

December 31, 1940

SOCIETY
NEW YORK

CONTENTS

PAGE

25. Eastern Pacific Expeditions of the New York Zoological

Society. XXII. Mollusks from the West Coast of Mexico
and Central America. Part I. By Leo George Hertlein,

& A. M. Strong. (Plates I & II) 369

26. On the Electric Powers and Sex Ratios of Foetal Narcine

brasiliensis (Olfers). By C. M. Breder, Jr., & Stewart
Springer 431

27. A Study of the Activities of a Pair of Galago senegalensis

moholi in Captivity, Including the Birth and Postnatal
Development of Twins. By Florence De L. Lowther.
(Plates I-VI) 433

28. Diets for a Zoological Garden: Some Results During a

Test Period of Five Years. By Herbert L. Ratcliffe 463

29. The Biology of the Smoky Shrew ( Sorex fumeus jumeus
Miller). By W. J. Hamilton, Jr. (Plates I-IV; Text-

figure 1) 473

30. Social and Respiratory Behavior of Small Tarpon. By

Arthur Shlaifer & C. M. Breder, Jr. (Plates I & II;
Text-figure 1) 493

31. New Observations on the Blood Group Factors in Simiidi

and Cercopithecidae. By P. B. Candela, A. S. Wiener

& L. J. Goss 513

32. Muscle Dystrophy in Tree Kangaroos Associated with

Feeding of Cod Liver Oil and Its Response to Alpha-
Tocopherol. By Leonard J. Goss 523

33. Mortality Statistics for Specimens in the New York Aquar-

ium, 1939. By Ross F. Nigrelli. (Plates I-III) 525

34. A Comparison of Some Electrical and Anatomical Char-

acteristics of the Electric Eeel, Electrophorus electricus
(Linnaeus). By R. T. Cox, W. A. Rosenblith, Janice
A. Cutler, R. S. Mathews & C. W. Coates. (Text-

figures 1-7) 553

Index to Volume XXV 563

Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 369

25.

Eastern Pacific Expeditions of the New York
Zoological Society. XXII.

Mollusks from the West Coast of Mexico and Central America.

Part I.1

Leo George Hertlein & A. M. Strong

California Academy of Sciences.

(Plates I & II).

[This is the twenty-second of a series of papers dealing with the collec-
tions of the Eastern Pacific Expeditions of the New York Zoological Society
made under the direction of William Beebe. The present paper is concerned
with specimens taken on the Templeton Crocker Expedition (1936) and the
Eastern Pacific Zaca Expedition (1937-1938). For data on localities, dates,
dredges, etc., refer to Zoologica, Vol. XXII, No. 2, pp. 33-46, and Vol. XXIII,
No. 14, pp. 287-298.]

Contents.

Page

Introduction 370

Class Pelecypoda 373

Order Prionodesmacea 373

Superfamily Solemyacea 373

Family Solemyidae 373

Genus Solemya Lamarck 374

Subgenus Acharax Dali 375

Solemya ( Acharax ) agassizii

Dali 375

Solemya ( Acharax ) johnsoni

Dali 376

Solemya ( Acharax ) macrodac-

tyla Mabille & Rochebrune. . . 376
Solemya ( Acharax ) patagonica

E. A. Smith 377

Subgenus Petrasma Dali 377

Solemya ( Petrasma ) panamensis

Dali 377

Solemya ( Petrasma ) valvulus

Carpenter 378

Superfamily Nuculacea

Family Nuculidae 378

Genus Nucula Lamarck 379

Nucula ( Nucula ) declivis Hinds 380
Nucula ( Nucula ) exigua

Sowerby 381

Subgenus Ennucula Iredale 382

Nucula ( Ennucula ) tenuis

Montagu 382

Nucula ( Ennucula ) cardara

Dali 383

Nucula (Ennucula?) linki Dali. 383
Subgenus Nucolopsis Woodring. . 383
Nucula (Nucolopsis) schencki

Hertlein & Strong, n. sp 384

Subgenus uncertain :

Nucula agujana Dali 384

1 Contribution No. 607, Department of Tropical Research, New York Zoological Society.

Page

Nucula chrysocoma Dali 385

Nucula colombiana Dali 385

Nucula grayi d’Orbigny 385

Nucula iphigcnia Dali 386

Nucula panamina Dali 386

Nucula paytensis A. Adams... 386

Nucula pigajcttae Dali 386

Nucula pisum Sowerby 387

Nucula savatieri Mabille &

Rochebrune 387

Nucula taeniolata Dali 387

Nucula tanneri Dali 388

Genus Acila H. & A. Adams 388

Subgenus Truncacila Schenck in

Grant & Gale 388

Acila (Truncacila) castrensis

Hinds 388

Family Nuculanidae 389

Genus Nuculana Link 390

Subgenus Saccella Woodring 392

Nuculana (Saccella) acuta

Conrad 392

Nuculana (Saccela) callimene

Dali 393

Nuculana (Saccella) elcnensis

Sowerby 393

Nuculana (Saccella) eburnea

Sowerby 395

Nuculana (Saccella) gibbosa

Sowerby 395

Nuculana (Sacce la) impar

Pilsbry & Lowe 396

Nuculana (Saccella) laeviradius

Pilsbry & Lowe 396

Nuculana (Saccella) taphria

Dali 397

JAM 6 19«

370

Zoologica: New York Zoological Society

[XXV :25

Contents (continued) :

Page

Subgenus Politoleda Hertlein &

Strong, n. subg 397

Nucuiana ( Politoleda ) polita

Sowerby 397

Subgenus Costelloleda Hertlein &

Strong, n. subg , .. 398

Nucuiana ( Costelloleda ) cos-

tellata Sowerby 398

Nucuiana ( Costelloleda ) marella
Hertlein, Hanna & Strong, n.

sp 399

Subgenus Thestyleda Iredale 400

Nucuiana ( Thestyleda ) hamata

Carpenter 400

Nucuiana ( Thestyleda ) cordyla

Dali 401

Subgenus Spinula Dali 401

Nucuiana ( Spinula ) calcar Dali 401

Nucuiana ( Spinula ) calcar ella

Dali 401

Subgenus Jupiteria Bellardi 402

Subgenus uncertain :

Nucuiana acrita Dali 402

Nucuiana agapea Dali 402

Nucuiana bicostata Sowerby... 403

Nucuiana crispa Hinds 403

Nucuiana cuneata Sowerby. . . . 403

Nucuiana excavata Hinds 404

Nucuiana hindsii Hanley 404

Nucuiana lobula Dali 404

Nucuiana loshka Dali 404

Nucuiana lucasana Strong &

Hertlein 405

Nucuiana ornata d’Orbigny.... 405

Nucuiana oxia Dali 405

Nucuiana peruviana Dali 405

Nucuiana pontonia Dali 406

Nucuiana rhytida Dali 406

Genus Adrana H. & A. Adams 406

Adrana arcuata Sowerby 408

Adrana crenifera Sowerby 408

Adrana elongata Sowerby 409

Adrana exoptata Pilsbry & Lowe 409
Adrana penascoensis Lowe. . . . 410
Adrana sowerbyana d’Orbigny. . 410
Adrana suprema Pilsbry &

Olsson 411

Adrana taylori Hanley 412

Adrana tonosiana Pilsbry &

Olsson 412

Genus Yoldia Moller 412

Subgenus Katadesmia Dali 413

Yoldia ( Katadesmia ) vincula

Dali 413

Subgenus Megayoldia Verrill &

Bush 414

Page

Yoldia ( Megayoldia ) martyria

Dali 414

Subgenus Orthoyoldia Verrill &

Bush 414

Yoldia ( Orthoyoldia ) panamen-

sis Dali 415

Subgenus Yoldiella Verrill & Bush 415
Yoldia ( Yoldiella ) cecinella Dali 416
Yoldia ( Yoldiella ) chilenica

Dali 416

Yoldia ( Yoldiella ) dicella Dali. 416
Yoldia ( Yoldiella ) granula Dali 416
Yoldia ( Yoldiella ) indolens Dali 417
Yoldia ( Yoldiella ?) infrequens

Dali 417

Yoldia ( Yoldiella ) leonilda Dali 417
Yoldia ( Yoldiella ?) mantana

Dali 417

Genus Cyrilla A. Adams 418

Cyrilla munita Carpenter in Dali 419
Subgenus N eopleurodon Hertlein &

Strong, n. subg 419

Cyrilla (N eopleurodon) subdolus

Strong & Hertlein 420

Genus Malletia Desmoulins 420

Malletia ( Malletia ) chilensis

Desmoulins 421

Subgenus Neilo H. & A. Adams. . . 422
Malletia (Neilo) goniura Dali. . 422

Subgenus Minormalletia Dali 422

Malletia ( Minormalletia ) arci-

formis Dali 423

Malletia ( Minormalletia ) ben-

thima Dali 423

Subgenus uncertain :

Malletia faba Dali 423

Malletia inequalis Dali 423

Ma letia magellanica E. A. Smith 424
Malletia patagonica Mabille &

Rochebrune 424

Malletia peruviana Dali 424

Malletia truncata Dali 424

Genus Tindaria Bellardi 425

Subgenus Tindariopsis Verrill &

Bush 425

Tindaria (Tindariopsis) sulcu-

lata Gould 425

Subgenus uncertain :

Tindaria atossa Dali 426

Tindaria compressa Dali 426

Tindaria j)anamensis Dali 426

Tindaria mexicana Dali 427

Tindaria salaria Dali 427

Tindaria smirna Dali 427

Tindaria thea Dali 427

Tindaria virens Dali 428

Introduction.

Two expeditions to the eastern tropical Pacific were conducted by Mr.
Templeton Crocker and Dr. William Beebe under the auspices of the New
York Zoological Society. The means of transportation for these trips was
Mr. Crocker’s yacht Zaca. The first of these left San Diego, California, on
March 19, 1936. Dredgings were made at many localities from a short dis-
tance south of San Diego to Cape San Lucas and into the Gulf of California,
north to Santa Inez Bay. Clarion Island in the Revillagigedo group was also
visited and some dredgings and shore collections made. At certain places
shore collecting was conducted but a large part of the mollusks obtained by
this expedition resulted from dredging, mostly from the southern part of
the Gulf of California. The depth varied from 1 to 600 fathoms but most of
the collection was obtained in less than 100 fathoms. This expedition re-
turned to San Diego, California, on May 25, 1936. A paper by Beebe2 con-
tains an itinerary, list of stations and nets and dredges used, and a general

2 Beebe, W. The Templeton Crocker Expedition. II. Introduction, Itinerary, List of Stations,
Nets and Dredges. Zoologica, Sci. Contrib, New York Zool. Soc., Vol. 22, pt. 1, April 5, 1937,
pp. 33-46, 8 text figs.

JAN 6 1941

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 371

account of the trip by the same author3, appeared in a book entitled “Zaca
Venture.”

The second expedition by Messrs. Crocker and Beebe left San Diego,
California, on November 6, 1937. This cruise took the Zaca along western
Lower California and the west coast of Mexico as far south as Gorgona
Island, Colombia. Dredging and shore collecting on this trip yielded much
fine research material. The Zaca returned to California and the party dis-
banded at Balboa on April 5, 1938. An account of the itinerary, list of sta-
tions, nets and dredges on this trip is contained in a paper by Beebe4.

The present collections, then, are almost entirely from tropical and sub-
tropical western North and Central America between 30° and 3° North
Latitude. Collecting over such an extensive area has naturally brought to
light many interesting occurrences, and a number of new species.

Several important extensions in range were discovered during the work.
Especially noteworthy are those species heretofore known from more north-
ern latitudes but now found to range south to Cape San Lucas. Also, numer-
ous species, previously known to occur in more southern latitudes were found
in the Gulf of California. An illustration of this is the discovery in the Gulf
of Cancellaria cumingiana Petit, heretofore known only from Ecuador. An-
other interesting occurrence is that of a single valve of Cardium magnum
Linnaeus, a species well known off Florida, which was found at Santa Inez
Bay. It seems possible that this may have been transported to the west coast
by some fortuitous means.

Through an arrangement with Dr. Beebe the mollusks were submitted
to the authors for identification and description. Types of new species and
figured specimens have been deposited in the type series of the California
Academy of Sciences; duplicates of some of the other species and certain
unique specimens were retained in the same institution for the purpose of
comparative studies of tropical west American mollusks now under way by
the authors. The bulk of the collection accompanied by identifications has
been returned to Dr. Beebe of the New York Zoological Society.

Numerous papers contain records of species inhabiting the Gulf of
California but the present collection revealed many additional ones not here-
tofore known to live in that region. There are also a very considerable num-
ber of forms previously unknown. Although it has been generally recog-
nized that the Gulf of California contains one of the richest known molluscan
faunas, the meticulous care taken by Messrs. Crocker and Beebe in record-
ing locality data makes this collection one of the most important ever made
in the area.

Scope and Plan of Present Report.

The preparation of a paper on the present collections offered problems
which have required considerable time. No checklist containing the majority
of the species known to occur between San Diego and Panama has been pub-
lished. Several papers dealing with the mollusks of this region exist but
any systematic work required consultation of many scattered references and
even in these many species have never been illustrated. After due consid-
eration and consultation the authors decided to prepare the paper with the
idea in mind that it should be useful not only as a checklist of the present
collection but also containing references and illustrations. We therefore
have prepared a systematic report with all pertinent references to each

3 Beebe, W. Zaca Venture (Harcourt Brace & Co., New York City, New York). 1938, 303
pp. + I-XVI. 23 illustrations.

4 Beebe, W. Eastern Pacific Expeditions of the New York Zoological Society, XIV. Introduction,
Itinerary, List of Stations, Nets and Dredges of the Eastern Pacific Zaca Expedition, 1937-1938.
Zoologica, Sci. Contrib. New York Zool Soc., Vol. 23, pt. 3, September 28, 1938, pp. 287-298, (2
maps pp. 290, 291) .

372

Zoologica: Neiv York Zoological Society

[XXV :25

species gathered together in the synonymy. The total lack of a complete
checklist of the tropical west American mollusks has necessitated a
compilation of all the species known to have been described or cited as
occurring in that region. Although it would not be possible in the time
available to monograph or even give a critical review of all the species, the
great utility of a complete catalogue justifies, in our opinion, the inclusion
herein of as complete a census as possible. In some cases we have included
additional species which are not known to occur in the tropics but which
are related to tropical forms, or which for systematic reasons it seems de-
sirable to include. Keys to the genera of certain of the families, and, where
information is adequate, keys to species, have been included. However, in
many cases, present knowledge does not justify an attempt to furnish a key.
The present unstable nomenclature5 of mollusks has led the authors to adopt
a conservative course in dealing with nomenclatorial problems in connection
with this work. Illustrations are planned for all species represented in the
collection which are heretofore unillustrated or of which the illustration is
not readily available. The range, type locality, repertory of the type where
known to us, collecting stations, notes on the species and descriptions of
new species are also included. At present it is planned that the entire work
will appear at intervals, each part to include one or more families, beginning
with the Pelecypoda. In the final part of the paper we hope to present gen-
eral observations on the collection as a whole and remarks regarding sig-
nificant features observed regarding distribution; and if feasible, a bibliog-
raphy of many of the important papers which it is necessary to consult in
a study of mollusks of tropical western North America.

At the beginning of this work it was planned that Dr. G. Dallas Hanna,
Curator of the Department of Paleontology of the California Academy of
Sciences, could collaborate in this work. Unfortunately the work had only
begun when duties called him to Alaska and elsewhere which then left the
preparation to the present authors. A few species were described during the
early stages of the work and in these we have happily been able to include
Dr. Hanna as a co-author.

Acknowledgements.

We wish here to thank Mr. Templeton Crocker, owner of the yacht
Zaca, for his generosity and interest which made this paper possible. His
collecting and careful recording of locality data have been of great help in
work on this collection. To Dr. William Beebe we extend our thanks for his
unfailing interest and cooperation throughout this work. His collecting and
recording of locality information is a model of its kind. To Dr. G. Dallas
Hanna we extend our thanks for his cooperation and unfailing help in ad-
vising us regarding many problems which have arisen during the prepara-
tion of the paper. His advice and help have been generously available at all
times and are greatly appreciated by the authors. Mr. A. G. Smith, Berkeley,
California, Dr. U. S. Grant IV, of the University of California at Los
Angeles, and Dr. H. G. Schenck and Dr. A. M. Keen of Stanford University,
have furnished information regarding certain of the species. The secre-
tarial work of Miss Winifred O’Neill and Miss Alta Holton and the prepara-
tion of photographs by Mr. Frank L. Rogers incident to the preparation of
this portion of the report is herewith acknowledged. This was done during
the course of Federal Works Progress Administration Project No. 8569.

5 Henderson, J. Our Unstable Biological Classification and Nomenclature. Presidential Address,
Fourth Annual Meeting of the American Malacological Union at Stanford University, June 26-28,
1934, pp. 1-13.

Other papers useful in dealing with nomenclatorial problems are : Procedure in Taxonomy,
including a Reprint of the International rules of Zoological Nomenclature with Summaries of
Opinions Rendered to date, completely Indexed. By E. T. Schenk and J. H. McMasters. Stanford
University Press, Stanford University, California, 1936, VII -j- 1-72. Terminology of Types. By
D. L. Frizzell. Amer. Midi. Nat., Vol. 14, no. 6, 1933, pp. 637-668.

1940] Hertlein & Strong: MollusJcs of Eastern Pacific Expeditions 373

Class Pelecypoda.

The pelecypods or bivalve shells are well known to collectors and sys-
tematists. Oysters and scallops are examples of edible pelecypods of eco-
nomic importance. Marine species occur in all oceans. The total number is
not exactly known but there are perhaps about 8,000 species. They are most
numerous on the coasts of continents or on land masses which have for-
merly been connected with continental land masses. About 500 species are
known to occur between Bering Sea and San Diego, California, and at least
an equal or greater number exist between San Diego and Peru. The largest of
all mollusks is the giant clam, Tridacna gigas Linnaeus0, which attains a
length of 54 inches and a weight of more than 500 pounds. Some excellent
manuals used in the classification and general arrangement of pelecypods are
those of Tryon7, Chenu8, Fischer9, Woodward10, Dali* 11 and Thiele12. In general,
although with some exceptions, we have followed the arrangement used by
Dali. The importance of the hinge in classification has been discussed by
Neumayr13 and by Dali14. The structure of shells has been discussed by
Boggild15 and others, and the coloration has been discussed by Bennett16. A
useful general bibliography of literature dealing with the Pelecypoda is
found in Dali’s article on Mollusks in Eastman’s Translation of Zittel’s Text-
book of Palaeontology and in recent papers on bivalves by Haas17. Additional

Order Prionodesmacea.

Superfamily Solemyacea.

Family Solemyidae.

The family Solemyidae is usually placed first in systematic arrange-
ment of the pelecypods. The shell characters are quite different from the
following group, the Nuculidae, and as pointed out by some authors, there is
little evidence of relationship between the two groups. The gill-structure of
the Solemyidae is believed by some writers to be due probably to specializa-
tion and not a remnant of a generalized style.

6 See Hedley, C. A Revision of the Australian Tridacna. Rec. Australian Museum , Vol. 13, no. 4,
April 12, 1921, pp. 163-172, pi. 27-34. See especially pp. 168-170. See also Mem. Australian Mus. 3,
pt. 8, 1899, p. 505, and E. A. Smith, Proc. Malacol. Soc. London , Vol. 3, 1898, p. 112.

7 Tryon, G. W. Structural and Systematic Conchology. Volume 3, 1884, Pelecypoda, pp. 116-353,
pis. 104-133.

8 Chenu, J. C. Manuel de Conchyliologie, Vol. 2, 1862, pp. 1-199, 1015 figs, in text.

9 Fischer, P. Manuel de Conchyliologie, 1880-1887, pp. 1-1369, 23 pis. Pelecypodes, pp. 897-1187,
pis. 16-23.

19 Woodward, S. P. Manual of the Mollusca. Reprint of Fourth Edition (1880), 1910, Conchifera,
pp. 393-507, pis. 16-23. Ap. to Manual of Mollusca by R. Tate, 1910, Conchifera, pp. 64-81.

11 Dali, W. H. Text-Book of Palaeontology by K. von Zittel. Edited by C. R. Eastman, Vol. 1,
1913. Pelecypoda, pp. 422-507, Figs. 637-836 in text.

12 Thiele, J. Handbuch der systematischen Weichtierkunde, (Verlag von Gustav Fischer. Jena),
Teil 3, 1934, Bivalvia, pp. 782-948, figs. 788-867. Also, Handworterbuch der Naturwissenschaften,
Zweite Auflage, Bd. 1, 1931, Bivalvia, pp. 996-1010, figs. 1-8. See also Bivalvia ( Palaontologie) by
E. Jaworski, pp. 1010-1026, figs. 1-43.

18 Neumayr, M. Zur Morphologie des Bivalvenschlosses. Sitzungsber. Akad. Wiss. Wien, Bd. 88,

Abt. 1, 1883, pp. 385-419, 2 pis. Beitrage zu einer Morphologischen Eintheilung der Bivalven.

Denkschr. Alcad. Wiss. Wien, Bd. 58, 1891, pp. 701-801.

Dali, W. H. Tertiary Mollusks of Florida. A new classification of the Pelecypoda, etc. Trans.
Wagner Free Inst. Sci., Vol. 3, pt. 3, March, 1895, pp. 486-566.

15 Boggild, G. B. The Shell Structure of the Mollusks. D. Kgl. Danslce Vidensk. Selsk. Skrifter
Naturbidensk. og Math. Afd., 9 Raekke, II, 2, 1930, pp. 233-325, 15 pis., 10 figs, in text. See also
Schenck, H. G., Literature on the Shell structure of Pelecypods. Bull. Mus. Hist. Nat. Belg., Vol.
10, no. 34, 1934, pp. 1-20.

16 Bennett, E. W. Coloration of Mollusca in relation to Light. Rec. Canterbury Mus., Vol. 3,
no. 3, November 23, 1928, pp. 185-197.

17 Haas, F. Bivalvia (Muscheln) H. G. Bronns Klassen und Ordnungen des Tier-reichs, Bd. 3,
Abt. Ill, 1 Lfrg., Leipzig, 1929, pp. 1-176 ; Bd. 3, Abt. Ill, 2 Lfrg., 1929, pp. 1-292. With bibli-
ography; Bd. 3, Abt. Ill, 3 Lfrg., 1931, pp. 177-384; Bivalvia. Dr. H. G. Bronns Klassen und
Ordnungen des Tier-reichs, Bd. 3, Abt. Ill, 4 Lfrg., Leipzig. 1933, pp. 385-544. Also pp. 12-141. Short
bibliography; Bd. 3, Abt. Ill, 1 Lfrg., 1937, pp. 1-208; Bd. 3, Abt. Ill, 2 Lfrg., 1938, pp. 209-466.

374

Zoologica: New York Zoological Society

[XXV :25

references which deal with the fauna upon which this paper is based will
be found in succeeding pages.

Genus Solemya Lamarck.

Soleviya Lamarck, Anim. s. Vert., Vol. 5, 1818, p. 488. Species in original

list: Solemya australis and Solemya mediterranea. Dali, Nautilus, Vol. 22,

no. 1, May, 1908, p. 1. “Type S. australis Lamarck.” — Dali, Bull. Mus. Comp.

Zool., Vol. 43, no. 6, October, 1908, p. 361. S. australis Lamarck considered to be

the type. Grant & Gale, Mem. San Diego Soc. Nat. Hist., Vol. 1, 1931, p.

109. Type: Solenomya mediterranea Lamarck =Solemya mediterranea Lamarcks
Solemya togata (Poli).

Solenimya Lamarck, Bowdich, Elem. Conch., Pt. 2, 1822, p. 8. “Solenimya.
( Solemya . Lam.) M.” Sole species: Solenimya australis cited as fig. 17.

Solenomya Lamarck, Menke, Synop. Meth. Moll., 1830, p. 119. “ {Solemya,

Lam.)” Sole species: Solenomya mediterranea Lamarck. Children, Quart.

Jour. Lit. Sci. and Arts, Vol. 14, January 1823, p. 300 (reprint p. 27). Reprint by
Kennard, Salisbury and Woodward, Smithson. Misc. Coll., Vol. 82, no. 17, July 11,
1931, p. 7. Type species: S. mediterranea Lamarck.

Type (designated by Children) : Solenomya mediterranea Lamarck.
[=Solemya mediterranea Lamarck, Anim. s. Vert., Vol. 5, 1818, p. 489. Ref. to
“Poli, test. 2, p. 42, et vol. 1. tab. 15, f. 20”; “Solen. Encycl. pi. 225,
f. 4.” “Habite la Mediterranee, dans le sable.” Referred to Solenomya togata
Poli by Bucquoy, Dautzenberg & Dollfus, Moll. Mar. Roussillon, Vol. 2, 1898,
p. 718, pi. 92, figs. 8, 9, 10. Mediterranean; Adriatic; Atlantic; Senegal;
Madeira].

Shell elongate, Scden-shaped, gaping at each end, epidermis dark, horny
and extending beyond the ventral margins; umbos posterior; ligament
amphidetic, chiefly internal; hinge edentulous with an obliquely inclined
chondrophore, below which is an internal rib on the inner surface of the
valve; pallial line obscure. Outer layer of shell composed of long prismatic
cells, nearly parallel with surface and mingled with dark cells as in Pinna;
inner layer also cellular.

Dali18' has published a revision of the Solemyidae.

Solemya has been recorded from Paleozoic to Recent (Carboniferous
and possibly Silurian to Recent). The genus has been cited from the Eocene
of California by Clark & Woodford10 and it also occurs from upper Oligocene
or lower Miocene to Recent in western North America. Solemya ( Solemya )
lomitensis Olsson20 has been described from the Oligocene of Peru, and
Marwick21 has recorded the genus from the lower Miocene of New Zealand. It
also occurs in the Cenozoic of various other parts of the world.

In Solemyarina I redale22 with the type S. velesiana Iredale “The median
rib is not curved, but is angulated posteriorly, while the anterior portion of
the ligament is small and linear and the posterior portion is small and sub-
linear, the posterior muscle scar free” (Iredale, 1939).

Zesolemya was proposed by Iredale23 for the New Zealand species
Solemya parkinsonii Gray.

is Dali, W. H., A Revision of the Solenomyacidae. Nautilus, Vol. 22, no. 1, May, 1908, pp. 1-2.
See also, Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908, pp. 361-366.

19 Solemya sp., Clark & Woodford, Univ. Calif. Publ. Bull. Dept. Geol. Sci., Vol. 17, no. 2,
December 31, 1927, p. 85, pi. 14, fig. 1. Univ. Calif. Loc. 3162. North side of Deer Valley, north of
Mt. Diablo, Mt. Diablo Quadrangle, California. Meganos formation ; Eocene.

20 Solemya (Solemya) lomitensis Olsson, Bull. Amer. Paleo., Vol. 17, no. 63, June 5, 1931,
p. 127 (31), pi. 15 (3), fig. 5. “Lomitos,” Peru; Oligocene.

21 Marwick, J., Geol. Surv. New Zealand, Palaeo. Bull. No. 13, 1931, p. 48, pi. 1, fig. 1.

22 Solemyarina Iredale, Rec. Austral. Mus., Vol. 18, no. 4, June 29, 1931, pp. 202, 231. “I intro-
duce Solemyarina for these small species, designating velesiana as type.” Sydney, Australia.

• Iredale, Brit. Mus. (Nat. Hist.) Great Barrier Reef Exped. 1928-29. Sci. Repts., Vol. 5, no.

6, Moll. Pt. 1, 1939, p. 232. ‘‘Orthotype: Solemya velesiana Iredale.”

23 Zesolemya Iredale, Brit. Mus. (Nat. Hist.) Great Barrier Reef Exped. 1928-29. Sci. Repts.,

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 375
Key to the Species of Solemya.

A. Ligament internal (subgenus) Petrasma

a. Anterior end with 8 or 9 obscure rays panamensis

aa. Anterior end with fine radial striae only valvula

B. Ligament external (subgenus) Acharax

a. Dorsal margin with a heavy callous patagonica

aa. Dorsal margin without accessory callous

b. Entire surface with subequally spaced radiating rays johnsoni
bb. Surface with central area smooth or with obsolete rays

c. Posterior end evenly rounded agassizii

cc. Posterior end meeting hinge line and basal margin at
rounded right angles macrodactyla

Subgenus Acharax Dali.

Acharax Dali, Nautilus, Vol. 22, no. 1, May, 1908, p. 2. “Type S. johnsoni
Dali.” Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908, p. 364.

Type (by original designation) : Solemya johnsoni Dali.

Ligament opisthodetic, wholly external, visible internally only where it
crosses the gap between the margins of the valves. Nymphs without props
(Dali).

Acharax has been recorded from Oligocene to Recent.

Solemya (Acharaxl agassizii Dali.

S[olemya], agassizii Dali, Nautilus, Vol. 22, no. 1, May, 1908, p. 2. “Off Tilla-
mook Bay, Oregon, south to Aguja Point, Peru, in 1036-1800 fathoms.”

Solemya ( Acharax ) agassizii Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 365, pi. 16, fig. 10. U. S. S. Albatross station 3360, “Gulf of
Panama, in 1672 fathoms, sand, bottom temperature 36°. 4 F.” Also at other sta-
tions from 1588, 1740, 1772, and 1793 fathoms; off Aguja Point, Peru, in 1036

fathoms; range; Gulf of California to off Aguja Point, Peru. Zetek, Rev.

Nueva, Nos. 1 & 2, 1918, p. 37. Panama. — I, S. Oldroyd, Stanford Univ., Publ.
Univ. Ser. Geol. Sci., Vol. 1, 1924, p. 9, pi. 40, fig. 11 (under subgenus Acharax) .
Original records cited.

Acharax agassizii Dali, Dali, U. S. Nat. Mus., Bull. 112, 1921, p. 9. Tillamook,
Oregon, to Point Aguja, Peru.

Type Locality : Gulf of Panama, in 1672 fathoms. Type No. 106,885 U. S.
Nat. Mus.

Range : Off Tillamook, Oregon, to Aguja Point, Peru, in 1036 to 1800
fathoms.

The shell of Solemya agassizii is ornamented by about five or six an-
terior radial channels rather than from nine to twelve on S. johnsoni. S.
agassizii differs from S. macrodactyla Mabille & Rochebrune, a more south-
ern species, in the presence of a smooth middle area on the valves, as well
as the proportionately shorter posterior end, and much larger size. S. grandis
Verrill & Bush-4, an Atlantic species, has a shorter and wider shell.

Solemya ( Acharax ) dalli Clark25 from the lower Miocene or upper Oligo-

24 Solemya grandis Verrill & Bush, Proc. U. S. Nat. Mus., Vol. 20, [No. 1139.], 1898, p. 885,
pi. 86, figs. 1 and 2. “Two good specimens and some fragments, at four stations, between N. lat.
39° 58' 30", W. long. 70° 30', and N. lat. 37° 24', W. long. 74° 17', in 300 to 1,600 fathoms, 1880-1884."

25 Solemya dalli Clark, Univ. Calif. Publ. Bull. Dept. Geol. Sci., Vol. 15, no. 4, January 5, 1925,

p. 73, pi. 8, fig. 2. L.S.J.U. Loc. NP. 120. "Shales outcropping in sea cliffs west of West Twin

River for distance of % mile. Twin, Washington.” Tegland, Univ. Calif. Publ. Bull. Dept.

Geol. Sci., Vol. 23, no. 3, October 11, 1933, p. 103, pi. 4, figs. 1, 2, 3, 4, 5, 6, 7, 8, 9, 10. Type

locality. Also from Restoration Point, Blakeley formation, near Seattle, Washington. Upper
Oligocene.

376

Zoologica: New York Zoological Society

[XXV :25

cene of Washington is a somewhat similar species and Solemya tokunagai
Yokoyama20 and S. yessoensis Kanehara27, from the Tertiary of Japan, are
members of the same group.

Solemya ( Acharax ) belenensis Olsson28 has been described from the
Oligocene of Peru.

Solemya l Acharax) johnsoni Dali.

Solemya johnsoni Dali, Proc. U. S. Nat. Mus., Vol. 14, 1891, p. 189. “U. S.
steamer Albatross, station 3010, off coast of Lower California, in 1,005 fathoms.”

Dali, Proc. U. S. Nat. Mus., Vol. 17, 1894, p. 712, pi. 25, fig. 1. Coast of

Ecuador in 1,740 fathoms; Gulf of Panama, in 1,672-1,793 fathoms; Gulf of

California in 1,000-1,588 fathoms; Straits of Fuca, in deep water. — Keep,

West Amer. Shells (The Whitaker & Ray Co.,) San Francisco, 1904, p. 19, fig. 8.

Deep water as far north as Puget Sound. Dali, Nautilus, Vol. 22, no. 1, 1908,

p. 2. Puget Sound to Panama Bay in 60 to 1740 fathoms. Keep, West Coast

Shells, (The Whitaker & Ray-Wiggin Co.) San Francisco, 1911, p. 24, fig. 10.
Same locality as cited 1904. Edit, by Baily, 1935, p. 45, fig. 20 (as Solemya

(Acharax) johnsoni). Oregon to Panama. Zetek, Rev. Nueva, Nos. 1 & 2,

1918, p. 37. Panama. 1. S. Oldroyd, Stanford Univ. Publ. Univ. Ser. Geol.

Sci., Vol. 1, 1924, p. 9, pi. 14, fig. 1 (under subgenus Acharax) . Type locality cited.
Range: Oregon to Panama.

Solenya [Typ. error] johnsoni [Dali], Pilsbry, Nautilus, Vol. 5, no. 4, August,
1891, p. 47.

Solenomya johnsoni Dali, Rogers, The Shell Book, (Doubleday, Page & Co.,
Garden City, New York), 1913, p. 368. Puget Sound Southward.

Acharax johnsonii Dali, Dali, U. S. Nat. Mus., Bull. 112, 1921, p. 9 (under
genus Solemya). Oregon to Panama.

Type Locality : Off the coast of Lower California, in 1,005 fathoms.

Range-. Puget Sound to Panama.

There are nine to twelve anterior radial channels on the shell of Solemya
johnsoni, while there are only five or six such channels on the similar S.
agassizii.

Solemya l Acharax) macrodactyla Mabille & Rochebrune.

Solemya macrodactyla Mabille & Rochebrune, Miss. Sci. Cap Horn, Vol. 6,

Zool., pt. 2, 1891, p. H109, pi. 8, fig. 4. “Hab. Baie Orange.” Dali, Nautilus,

Vol. 22, no. 1, May, 1908, p. 2. Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,

1908, p. 364. Orange Harbor, Patagonia, north to Chiloe Island, southwest Chile,
in 20 to 369 fathoms.

Type Locality : Orange Harbor, Patagonia.

Range-. Orange Harbor and north to Chiloe Island, southwest Chile, in
20 to 369 fathoms.

This southern species, Solemya macrodactyla, bears a resemblance to
Solemya agassizii which occurs farther north, but the posterior end is less
rounded than it is in that species. Mabille & Rochebrune compared S', mac-

26 Solemya tokunacjai Yokoyama, Jour. Coll. Sci. Imp. Univ. Tokyo. Vol. 45, Article 5, March
21, 1925, p. 31, pi. 6, figs. 1, 2, 3. “Shiogu in Tatsuta ; Tsuchibashi in Kadono ; Enamura Quarry.”

Pliocene. Kanehara, Jap. Jour. Geol. & Geogr., Vol. 14, nos. 3 & 4, 1937, p. 156, pi. 15, figs.

10, 11 (as Solemya ( Acharax ) tokunagai). “Near Morae, Atsuta-gun, Ishikari.” Oiwake Series;
upper Miocene to Pliocene.

27 Solemya ( Acharax ) yessoensis Kanehara, Jap. Jour. Geol. & Geogr., Vol. 14, nos. 3 & 4,
October, 1937, p. 155, pi. 15, fig. 12. “Near Morai, Atsuta-gun, Ishikari.” Oiwake Series ; upper
Miocene to Pliocene.

2$ Solemya ( Acharax ) belenensis Olsson, Bull. Amer. Paleo., Vol. 17, no. 63, June 5, 1931,
p. 127 (31), pi. 15 (3), fig. 7. “Heath formation, Lobos, Pajarabobo.” Peru; Oligocene.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 377

rodactyla to S. borealis Totten which occurs from Halifax, Nova Scotia, to
Connecticut, and to S. parkinsonii Gray29 of New Zealand. Dali has pointed
out that Solemya macrodactyla may ultimately be relegated to the synonymy
of S. patagonica E. A. Smith.

Solemya lAcharaxI patagonica E. A. Smith.

Solemya patagonica E. A. Smith, Sci. Res. Voy. Challenger, Zool., Vol. 13,

1885, pp. 24, 208, pi. 11, figs. 1, la. “ Habitat . Station 311, off west coast of

South Patagonia, in 245 fathoms.” — Dali, Nautilus, Vol. 22, no. 1, May,

1908, p. 2. Dali, Bidl. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908, p. 364.

West coast of Patagonia.

Type Locality : Off west coast of Chile, in 245 fathoms.

Range'. West coast of Chile.

At the time of description of Solemya patagonica, E. A. Smith com-
pared the snecies to S. parkinsonii Gray of New Zealand. Dali considered it
likely that S. macrodactyla Mabille & Rochebrune might be relegated to the
synonymy of S. patagonica, but stated that the type of S. patagonica ap-
peared to be abnormally callous dorsally.

E. A. Smith {Ann. & Mag. Nat. Hist., Ser. 7, vol. 18, October, 1906,
p. 253) has cited S. patagonica from off India, and Melvill & Standen (Proc.
Zool. Soc. London, November 13, 1906, p. 793) have cited it from the Gulf
of Oman in the Arabian Sea. We have not seen specimens from that region.

Subgenus Petrasma Dali.

Petrasma Dali, Nautilus, Vol. 22, no. 1, May, 1908, p. 2. ‘‘Type S. borealis
Totten.” Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908, p. 363.

Type (by original designation) : Solemya borealis Totten. [ Amer . Jour.
Sci.,. Ser. 2, Vol. 26, July, 1834, p. 366, pi. [unnumbered], fig. 1. h, i. “In-
habits the coast of Rhode Island.” Also illustrated by M. Smith, East Coast
Mar. Shells (Edwards Bros., Ann Arbor, Michigan), 1937, p. 25, fig. 29a.
Nova Scotia to Connecticut],

Ligament internal but not exposed internally in front of the chondro-
phore.

Solemya f Petrasma! panamensis Dali.

S[olemya], panamensis Dali, Nautilus, Vol. 22, no. 1, May, 1908, p. 2. “extends
from off Santa Barbara, Cal., to Panama Bay.”

Solemya (Petrasma) panamensis Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 366. “U. S. S. ‘Albatross,’ station 2799, Panama Bay, in 2914>
fathoms, mud.” Also off Santa Barbara, California, in 68 fathoms.

Solemya panamensis Dali, Zetek, Rev. Nueva, Nos. 1 & 2, 1918, p. 37. Panama.

1. S. Oldroyd, Stanford Univ. Publ. Univ. Ser. Geol. Sci., Vol. 1, 1924, p. 10,

pi. 9, fig. 4. Original record cited. Pilsbry & Lowe, Proc. Acad. Nat. Sci.

Philadelphia, Vol. 84, 1932, p. 141. “On mud flats at extreme tide, rare. La Paz.”

Lowe, Trans. San Diego Soc. Nat. Hist., Vol. 8, no. 6, 1935, p. 27. Punta

Penasco, Sonora, Mexico, dredged in 10 fathoms. Strong, Proc. Calif. Acad.

Sci., Ser. 4, Vol. 23, no. 12, 1937, p. 192. San Martin Island, Lower California,
Mexico.

29 Solemya parkinsonii Gray, Voy. Erebus & Terror, 1874, p. 6, pi. 3, fig. 1 Suter, Man.

New Zealand Moll. (Wellington, New Zealand), 1913, p. 830. pi. 58, fig. 1. Throughout New Zealand,
in mud about 6 inches below the surface ; Kermadec Islands ; Milford Sound, in 100-120 fathoms.

378 Zoologica: Neiv York Zoological Society [XXV :25

Petrasma panamensis Dali, Dali, U. S. Nat. Mus., Bull. 112, 1921, p. 9. Santa
Barbara, California, to Panama.

Type Locality: Panama Bay, 29^2 fathoms. Type No. 110,678 U. S. Nat.

Mus.

Range: Santa Barbara, California, to Panama.

This is more expanded in front and less sharply truncate in front than
S. agassizii of the same length, and they may be separated at once by the
difference in the hinges. S. volvulus Carpenter is a much smaller species and
has no anterior prop to the chondrophore (Dali).

Solemya I Petrasma) volvulus Carpenter.

Solemya volvulus Carpenter, Ann. & Mag. Nat. Hist., Ser. 3, Vol. 13, April,
1864, p. 311. Cape St. Lucas. Reprint in Smithson. Misc. Coll., No. 252, 1872, p.

210. Carpenter, Rept. Brit. Assoc. Adv. Sci. for 1863 (issued August,

1864), p. 618. Reprint in Smithson. Misc. Coll., No. 252, 1872, p. 104. Dali,

Nautilus, Vol. 22, no. 1, May, 1908, p. 2. San Pedro, California, to the Gulf of

California. Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908, p.

364. San Pedro, California, to the Gulf of California. 1. S. Oldroyd, Stanford

Univ. Publ. Univ. Ser. Geol. Sci., Vol. 1, 1924, p. 11, pi. 40, fig. 10 (under subgenus
Petrasma) . Type locality cited. Range: San Pedro, California, to the Gulf of

California. Lowe, Trans. San Diego Soc. Nat. Hist., Vol. 8, no. 6, 1935, p.

27. Punta Penasco, Sonora, Mexico, dredged in 10 fathoms.

Solemya ( Petrasma ) volvulus Carpenter, Lamy, Journ. de Conchyl., Vol. 57,
no. 3, 1909, p. 208. Gulf of California.

Petrasma volvulus Carpenter, Dali, U. S. Nat. Mus., Bull. 112, 1921, p. 9. San
Pedro, California, to the Gulf of California.

Type Locality: Cape San Lucas, Lower California.

Range: San Pedro, California, to Punta Penasco, Sonora, Mexico.

The shell of Solemya volvulus is small, thin, transparent and attains a
length of about 20 mm. It is ornamented by radial slender brownish lines
which are finely striate posteriorly, distantly spaced on the medial portion
of the shell, and short, broad, and bifurcate anteriorly. The chondrophore is
without props.

Superfamily Nuculacea.

Family Nuculidae.

The family Nuculidae is an ancient one and has representatives in the
Paleozoic. Praenucula Pfab30 assigned to this family has been described
from the Silurian of Bohemia. In the Nuculidae the pallial impression is en-
tire, due to the lack of development of siphons. The shells are entirely
closed, not gaping, and the inner shell-layer is pearly. The family occurs in
both shallow and deep water. A few occur in tropical waters but most of
the species occur in temperate and boreal waters.

Two papers by Schenck31 on the classification of nuclid pelecypods may
be consulted regarding the nomenclature of members of this family. Dr.
Schenck has kindly examined a number of specimens of Nucula and Nucul-
ana in the present collection and has furnished us notes on certain of the
species.

30 Praenucula Pfab, Palaeontogr., Bd. 80, Abt. A, Lief. 4-6, April, 1934, p. 234. "Typus des
Genus: P. dispar expansa n. var.” p. 235, pi. 3, figs. 10, 11, 15a, 15b. Osek.

siSchenek, H. G. Classification of Nuculid Pelecypods. Bull. Mus. Roy. d’Hist. Nat. Belgique,

Vol. 10, no. 20, June, 1934, 78 pp„ 5 pis. Schenck, H. G. Revised Nomenclature for some

Nuculid Pelecypods. Jour. Paleo., Vol. 13, no. 1, pp. 21-41, 4 pis., January, 1939.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 379

Genus Nucula Lamarck.

Nucula Lamarck, Mem. Soc. Hist. Nat. Paris, Vol. 1, 1799, p. 87. Sole

species: Area nucleus Linnaeus. Schenck, Bidl. Mus. Roy. d’Hist. Nat.

Belg., Vol. 10, no. 20, 1934, p. 18, pi. 1, fig. 8; pi. 3, fig. 2; pi. 4, figs. 4, 4a,
4b; pi. 5, figs. 1, la. “(Type by monotypy: Area nucleus Linne.)”

Type (by monotypy) : Area nucleus Linnaeus. [Syst. Nat., Ed. 10, 1758,
p. 695. “Habitat in Europa.” Illustrated by Schenck, Bull. Mus. Roy. d’Hist.
Nat. Belg., Vol. 10, no. 20, 1934, p. 18, pi. 1, fig. 8; pi. 3, fig. 2; pi. 4, figs. 4,

4a, 4b; pi. 5, figs. 1, la. Schenck, Proc. Malacol. Soc. London, Vol. 21,

pt. 4, March, 1935, fig. 1 (p. 260)].

Shell closed, not gaping; profile ovate-trigonal; a “pouting” of the
escutcheonal area, which the radial ribs do not cross; beaks opisthogyrate,
appressed; prodissoconch unornamented; radial ribs faint, low, wide and
flat, often difficult to see on the middle part of the shell, but they are more
distinct near the ventral margin where they form the “pectinate margin” ;
interspaces narrow, about one-tenth the width of the ribs ; interior nacreous ;
pallial line simple; two subequal adductor muscle scars and additional
muscle scars; longer (anterior) row of teeth arched, with 16 ± to 24 ±
teeth; the shorter (posterior) row straight with 7 ± to 11 ±; axis of
chondophore forms an arc of a circle of which the arcuate dorsal margin is
apart (Schenck).

Key to the Species of Nucula32.

A. Outer surface smooth

a. Periostracum olive

b. Periostracum dark olive

c. Length less than 10 mm

d. Shape subcordate, umbos weak, length 8 mm., height

4 mm tenuis

dd. Shape subtrigonal, umbos prominent, length 6 mm.,

height 5 mm linki

cc. Length over 20 mm., solid, thick panamina

bb. Periostracum pale olive

e. Length less than 10 mm., shape, ovate colombiana

ee. Length 14-21 mm. savatieri

aa. Periostracum greenish or black

f. Periostracum greenish

g. Anterior hinge teeth 8, posterior 18 cardara

gg. Anterior hinge teeth 5, posterior 11 grayi

ff. Periostracum nearly black agujana

B. Outer surface sculptured

a. Concentric sculpture only

b. Length less than 10 mm. pisum

bb. Length over 20 mm. tanneri

aa. Radial sculpture present

c. Radial sculpture faint

d. Length less than 5 mm.

e. Anterior teeth less than 15

32 In the case of species which are not illustrated and which we have not seen, the key is based
upon descriptions.

380 Zoologica: New York Zoological Society [XXV :25

Key to the Species of Nucula (continued) :

f. Sculpture definitely reticulate, concentric strong,
anterior teeth 9, posterior 5 schencki

ff. Sculpture not reticulate, concentric weak, an-
terior teeth 10, posterior 6 chrysocoma

ee. Anterior teeth 17, posterior 9 paytensis

dd. Length over 10 mm.

g. Anterior hinge teeth 6, posterior 10 taeniolata

gg. Anterior hinge teeth 7, posterior 15-17 pigafettae

cc. Radial sculpture strong

h. Length less than 10 mm.

i. Concentric sculpture heavier, regular rugae ..exigua

ii. Concentric sculpture fine, irregular declivis

hh. Length over 30 mm. iphigenia

Nucula I Nucula I declivis Hinds.

Plate I, Figures 1, 2, 3, 6, 7.

Nucula declivis Hinds, Proc. Zool. Soc. London, December, 1843, p. 98.

“Hab ?” Hinds, Zool. Voy. Sulphur, Moll. Pt. 3, 1844 (January,

1845, on cover of Pt. 3), p. 63, pi. 18, fig. 8. “Inhab ?” Hanley,

Thes. Conch., Vol. 3, 1860, Nucula, p. 154, pi. 230 ( Nucididae , pi. 5), fig. 147.

“Hab ?” Sowerby, Conch. Icon., Vol. 18, Nucula, Octouer, 1810,

sp. 31, pi. 4, fig. 31. “Hab ?” Dali, Proc. U. S. Nat. Mus., Vol. 31,

1909, p. 250. “Panama to Magellan Straits.’’ — Zetek, Rev. Nueva, Nos. 1 & 2,

1918, p. 37. Panama. Lowe, Trans. San Diego Soc. Nat. Hist., Vol. 8, no.

6, 1935, p. 27. Punta Penasco, Sonora, Mexico. Strong & Hertlein, Allan

Hancock Pacific Exped. (Univ. South. Calif.), Vol. 2, no. 12, 1939, p. 183. Bahia
Honda, and off Taboga Island, Panama.

Type Locality. Original locality not known. Type locality here desig-
nated as off Taboga Island, Panama, in 3-9 fathoms.

Range : Punta Penasco, Sonora, Mexico, to Panama. [?] Panama to
Magellan Straits (Dali).

Collecting Stations: Mexico: Santa Inez Bay, Gulf of California (145-
D-1,-3). 4-15 fathoms, sand; Manzanillo Bay (184-D-2). 30 fathoms, gravelly
sand bottom; Nicaragua: Corinto (200-D-19 and beach). 12-13 fathoms,
mangrove leaves; Costa Rica: (203-D-l,-3) . 12-15 fathoms, sand and shelly
mud bottom.

This species has a small, smooth, oblique shell ornamented by very fine
radial striations. In the original description it is stated that the ventral mar-
gin is crenulate, and that the species is “a still more oblique shell than N.
pisum, to which it is closely allied.” From the original illustrations alone it
would be difficult to separate N. declivis from N. pisum, Sowerby although
Sowerby’s species appears to be less oblique. N. pisum was originally de-
scribed from Valparaiso, Chile. Hanley and Dali have considered N . semiornata
d’Orbigny33 to be synonymous with N. pisum. Specimens of N. declivis from
the Gulf of California resemble, in general features, N. crenulata Adams34
from Florida.

33 Nucula semi-ornata d'Orbigny, Voy. Amer. Merid., Vol. 5, 1846, p. 624, pi. 84, figs. 27. 28, 29.
“Nous l’avons recueillie dans la baie de San Bias, au nord de la Patagonie, ou elle vit au niveau
des plus basses marees. Elle y est tres-rare. M. Darwin l'a recontree fossile a la baie Blanche.”
3 i Nucula crenulata A. Adams, Proc. Zool. Soc. London, 1856, p. 52. “Hab. Guadaloupe Mus

Cuming.” Hanley Thes. Conch., Vol. 3, 1860, p. 154, pi. 229, ( Nuculidae , pi. 4), figs. 134, 135.

“Guadaloupe.” Sowerby, Conch. Icon., Vol. 18, Nucula, October, 1870, sp. 25, pi. 4, fig. 25.

“Hab. Guadeloupe.”

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions

381

Nucula (Nucula) exigua Sowerby.

Plate I, Figures 4, 5.

Nucula exigua Sowerby, Proc. Zool. Soc. London for 1832 (issued March 13,
1833), p. 198. “Hab. ad Columbian-) Occidentalem (Bay of Caraccas).” [Ecuador
Lat. 0° 35'S.] “A single specimen found in sandy mud at nine fathoms depth.”

Sowerby, Conch. Illustr., 1833, Nucidae, p. 6, pi. 16, figs. 24, 24*. “Bay of

Caraccas.” Muller, Synop. Test. Viv., 1836, p. 192. Type locality cited.

Hanley, Cat. Rec. Bivalve Shells, p. 172, 1843, p. 376, 71856, Suppl. pi. 20, fig. 13,

1856. W. Colombia. d’Orbigny, Voy. Amer. Merid., Vol. 5, 1846, p. 625.

Original locality cited. C. B. Adams, Ann. Lyceum Nat. Hist. New York,

Vol. 5, 1852, p. 479. Panama. Also earlier records cited. Carpenter, Cat.

Mazatlan Shells, February, 1856, p. 145. (as Nucula ? exigua). Mazatlan, Mexico.

Also earlier records cited. Carpenter, Rept. Brit. Assoc. Adv. Sci. for 1856

(issued 1857), pp. 249, 277, 311. Gulf of California; Central America; Panama;

Bay of Caraccas, Ecuador. Hanley, Thes. Conch., Vol. 3, 1860, p. 154, pi.

229, (Nuculidae, pi. 4), fig. 136. “Bay of Caracas. W. Columbia.” Carpen-

ter, Proc. Zool. Soc. London, 1863, p. 364. Reprint in Smithson. Misc. Coll., No.

252, 1872, p. 200. C. B. Adams’ record cited. — Sowerby, Conch. Icon., Vol.

18, Nucula, October, 1870, sp. 26, pi. 4, fig. 26. “Hab. West Columbia.” Dali,

Bull. Mus. Comp. Zool., Vol. 43, no. 6, 1908, p. 370. Bay of Panama (Cuming) ;
off Acapulco, Mexico, in 660 fathoms; near the Galapagos Islands, in 812
fathoms; off Aguja Point, Peru, in 1036 fathoms; west cost of Patagonia, in 122

and 194 fathoms. Dali, Proc. U. S. Nat. Mas., Vol. 37, 1909, p. 250.

Acapulco, Mexico, to Ecuador and to the Magellanic region. Zetek, Rev.

Nueva, Nos. 1 and 2, 1918, p. 51. Zoogeographical province of Panama. Ols-

son, Nautilus, Vol. 37, no. 4, 1924, p. 126. Salinas, Ecuador. Common. E.

K. Jordan, Contrib. Dept. Geol. Stanford Univ., Vol. 1, no. 4, 1936, p. 112.
Magdalena Bay, Lower California, Pleistocene.

Nucula ( Nucula ) exigua Sowerby, Schenck, Jour. Paleo., Vol. 13, no. 1, 1939,
p. 36, pi. 6, figs. 1-8, 11. Type loc. cited. Also western North America.

Type Locality: Bay of Caraccas, Ecuador, in 9 fathoms, in sandy mud.
Type in Brit. Mus. (Nat. Hist.).

Range: San Bartolome Bay [Turtle Bay], Lower California, and Gulf of
California to Ecuador. [?] Magellanic region (Dali).

Collecting Station: Nicaragua: Corinto (200-D-19). 12-13 fathoms,

mangrove leaves.

The shell of Nucula exigua is ornamented by concentric and radial
ornamentation but the concentric is much the stronger. It is a small shell
averaging not over 5 mm. in length. A strongly projecting lunular area
just under the beaks appears to be a characteristic feature of the species.
These differences are well shown on photographs of the type of N. exigua
shown to us by H. G. Schenck.

Some authors have considered N . exigua to be identical with N. supra-
striata Carpenter (in Arnold)35 but Schenck has given reasons for consider-
ing them to be distinct species. The radial ribbing on N. suprastriata is not
well developed and usually is present only ljear the ventral margin. The
lunule and escutcheon is more deeply impressed on N. suprastriata and it is
a larger shell averaging 5.5 to 6 mm. in length. Nucula calmitensis 01sson3fa
from the Miocene of Costa Rica was compared to N. exigua by Olsson.

Nucula paytensis A. Adams, bears considerable similarity to N. exigua.
Photographs of the type of the species described by Adams show a more
rounded outline and broader projecting lunular area but we are not certain
whether these differences are constant characters which can be used to
separate the two species.

35 Nucula ( Nucula ) suprastriata Carpenter in Arnold, Mem. Calif. Acad. Sci., Vol. 3, 1903, p.
96, pi. 18, tig. 6. “Upper San Pedro series, Los Cerritos” Upper Pleistocene. Also at Spanish
Bight, San Diego, California, upper Pleistocene. Rare in lower San Pedro of Deadman Island, lower
Pleistocene, and in the upper San Pedro Series of San Pedro, California, upper Pleistocene.

Schenck, Jour. Paleo. , Vol. 13, no. 1, 1939, p. 36, pi. 6, figs. 9, 10, 12, 13. Various localities

in the Pleistocene of southern California.

36 Nucula cahuitensis Olsson, Bull. Amer. Paleo., Vol. 9, Bull. No. 39, June 21, 1922, p. 343
(171), pi. 21 (18), figs. 21-24. ‘‘Gatun Stage: Zone G, Saury Creek,” Costa Rica. Miocene.

382

[XXV :25

Zoologica: New York Zoological Society

Subgenus Ennucula Iredale.

Ennucula Iredale, Rec. Australian Mus., Vol. 18, no. 4, June 29, 1931, pp. 202,

231. “Type Nucula obliqua Lamarck. Schenck, Bull. Mus. Roy. d’Hist. Nat.

Belg., Vol. 10, no. 20, 1934, p. 37, pi. 3, figs. 4, 4a, 4b; pi. 4, figs. 3, 3a, 3b. Type:

Nucula obliqua Lamarck. Cotton & Godfrey, Moll. South Australia, Pt. 1.

Handbook of Flora and Fauna of South Australia, issued by South Australian

Branch of Brit. Sci. Guild. Adelaide, 1938, p. 40. “Genotype E. obliqua

Lamarck.” Iredale, Brit. Mus. (Nat. Hist.) Great Barrier Reef Exped. 1928-

29 Sci. Repts. Vol. 5, no. 6, Moll., pt. 1, 1939, p. 235. Orthotype: Nucula obliqua
Lamarck.

Type (by original designation) : Nucula obliqua Lamarck. [Anim. s.
Vert., Vol. 6, 1819, p. 59. “Habite les mers australes, au cap aux Huitres.”

Illustrated by Schenck, Bull. Mus. Roy. d’Hist. Nat. Belg., Vol. 10,

no. 20, 1934, p. 37, pi. 3, figs. 4, 4a, 4b; pi. 4, figs. 3, 3a, 3b. “Cap aux Huitres,
Nouvelle Hollande.”]

The type species of Nucula is nucleus Linne, a European species which
differs appreciably from antipodean shells so classed, the latter having a
notably oblique chondrophore, above which the teeth become much smaller,
and the angle of opposition of the two rows of teeth is scarcely marked;
further, the edge of the European shell is strongly denticulate, whereas
ours is practically smooth (Iredale).

Nucula I Ennucula I tenuis Montagu.

Area tenuis Montagu, Test. Brit., Suppl., 1808, p. 56, pi. 29, fig. 1. “on the
shore near Dunbar,’' England.

Nucula tenuis Montagu, Forbes & Hanley, Hist. Brit. Moll., Vol. 2, 1853, p.
223, pi. 47, fig. 6; and (animal) pi. P, fig. 5. England and North Atlantic, Massa-
chusetts. Also Pliocene. Hanley, Thes. Conch., Vol. 3, 1860, p. 161, pi. 229

(Nuculidae, pi. 4), figs. 140, 141. Britain, W. Europe. Sowerby, Conch.

Icon., Vol. 18, Nucula, October, 1870, sp. 20, pi. 3, fig. 20. “Hab. Britain, Europe.”

1. S. Oldroyd, Stanford Univ. Publ. Univ. Ser. Geol. Sci., Vol. 1, no. 1, 1924,

p. 13, pi. 5, fig. 12; pi. 37, fig. 4. Point Barrow, Alaska, to Coronado Islands, Cali-
fornia. Circumboreal. Pliocene at San Pedro and San Diego, California.

Antevs, For. Geol. Foren. i Stockholm, Bd. 50, Heft 4, 1928, p. 493.

Nucula (Nucula) tenuis Montagu, Grant & Gale, Mem. San Diego Soc. Nat.
Hist., Vol. 1, 1931, p. 111. Earlier records cited. “Miocene” [Pliocene] of St. Paul
Island, and Nushag-ak Island, Alaska (Dali), and Recent. Circumboreal.

Nucula (Ennucula) tenuis Montagu, Schenck, Jour. Paleo., Vol. 13, no. 1,
January, 1939, p. 33, pi. 8, figs. 1-4, 9, 12, 14, 15. Earlier records cited.

Not Pronucula tenuis Powell, Rec. Canterbury Mus., Vol. 3, no. 2, December
14, 1927, p. 119, pi. 22, figs. 1, 2. Southwest of Otago, New Zealand, in 100
fathoms.

Type Locality. Near Dunbar, England, on shore.

Range: Northern Europe. Northeastern North America. Circumboreal.
Point Barrow, Alaska, to Cedros Island, Lower California.

Collecting Station : Mexico: East of Cedros Island (126-D-12). 45 fath-
oms, crushed shell and mud bottom.

One specimen of Nucula tenuis was dredged in Lat. 28° 20' N., Long.
115° 10 30" W., one mile east of Cedros Island, Lower California. Heath37
has recently studied the soft parts of this species.

This appears to be a wide-ranging form. According to Schenck some
of the specimens dredged off British Columbia agree exactly with specimens
of the species from the North Sea. Schenck has placed Nucula tenuis in the
subgenus Ennucula Iredale. The bathymetric range of the species given by
Antevs is 2 to 1,200 meters.

37 Heath, H., Mem. Mus. Roy. d'Hist. Nat. Belgique , Ser. 2, Fasc. 10, 1937, pp. 4, 9, etc., pi. 3,
figs. 19, 20, 21, 22, 26; pi. 4, figs. 31, 33, 34; pi. 5, fig. 44; pi. 9, fig. 81. Behring Sea, in 36 fathoms.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 383

Nucula lEnnuculal cardara Dali.

Nucula cardara Dali, Proc. V. S. Nat. Mus., Vol. 52, December 27, 1916, p.

394. “Station 5673, in 1090 fathoms, mud, off San Diego, California.” Dali,

U. S. Nat. Mus., Bull. 112, 1921, p. 10. Monterey, California, to Lower California,

in deep water. 1. S. Oldroyd, Stanford Univ. Publ. Univ. Ser. Geol. Sci.,

Vol. 1, 1924, p. 12. Type locality and range same as cited by Dali. Keen,

Check List West North Amer. Mar. Moll. (Stanford Univ. Press), 1937, p. 23.
Lat. 30°-37°N.

Nucula ( Ennucida ) cardara Dali, Schenck, Jour. Paleo., Vol. 13, no. 1,
January, 1939, p. 34, pi. 5, figs. 12, 14, 18, 21. Type locality cited.

Type Locality : Off San Diego, California, in 1090 fathoms, mud. Type
No. 265905 U. S. Nat. Mus.

Range : Monterey, California, to Lat. 30° N., Lower California, in deep
water.

Shell elongate-ovate, thin, polished, light olive green in color; hinge
with 8 anterior and 18 posterior teeth; margins of valves smooth. Length,
16 mm.; height, 11.5 mm.; diameter, 8 mm.

Nucula lEnnucula?) linki Dali.

Nucida linki Dali, Proc. U. S. Nat. Mus., Vol. 52, December 27, 1916, p. 394.

“Station 3034, 24 fathoms, mud; off Point Fermin, Lower California.” Dali,

U. S. Nat. Mus., Bull. 112, 1921, p. 10. Queen Charlotte Sound to Guaymas,

Mexico. 1. S. Oldroyd, Stanford Univ. Publ. Univ. Ser. Geol. Sci., Vol. 1,

1924, p. 12. Type locality and range same as cited by Dali. Keen, Check List

West North Amer. Mar. Moll. (Stanford Univ. Press), 1937, p. 23, Lat. 23°-55° N.

Nucula {Ennuculai) linki Dali, Schenck, Jour. Paleo., Vol. 13, no. 1, January,
1939, p. 33, pi. 5, figs. 2, 6, 11, 15. Type locality cited.

Type Locality: Off Point Fermin, California, in 24 fathoms, mud. [Ac-
cording to Schenck, not from off Point Fermin, Lower California, as orig-
inally cited by Dali]. Type No. 107649 U. S. Nat. Mus.

Range: Queen Charlotte Sound, British Columbia, to Guaymas, Sonora,
Mexico.

Shell small, inflated, very inequilateral, subtriangular and of a dark
olive color; hinge with six anterior and eleven posterior teeth; valve mar-
gins entire. Length, 6 mm.; height, 5 mm.; diameter, 3.6 mm.

Subgenus Nucolopsis Woodring.

Nucolopsis Woodring, Carnegie Inst. Washington, Publ. No. 366, May, 1925,
p. 14. [Cited in caption only. ? Typ. error.].

Nucidopsis Woodring, Carnegie Inst. Washington, Publ. No. 366, May, 1925,

p. 14. “Type. Nucula ( Nuculopsis ) hilli new species.” Schenck,

Bull. Mus. Roy. d’Hist. Nat. Belg., Vol. 10, no. 20, June, 1934, p. 33. Type: Nucula
hilli Woodring.

Not Nuculopsis Girty, Ann. New York Acad. Sci., Vol. 21, 1911, p. 133. “Type
species, Nucula ventricosa Hall.” Pennsylvanian series.

Not Nuculopsis Rollier, Abh. Schweiz. Palaont. Gesell., Vol. 38, Part 2, 1912,
p. 64. Renamed Isoleda by Rollier, Rev. Crit. Paleozool., Vol. 27, no. 1, January,
1923, p. 67. “(G.-T. Nuc. Palmae Sow.).” Jurassic.

Type (by original designation) : Nucula, ( Nuculopsis ) hilli Woodring,
Carnegie Inst. Washington, Publ. No. 366, May, 1925, p. 14, pi. 1, figs. 2
and 3. Bowden, Jamaica, Miocene.

Shell medium-sized, subelliptical, inequilateral ; sculpture consisting of
strong concentric rugae; chondrophore long, narrow, oblique, deeply exca-
vated ; anterior series of teeth more than twice as long as posterior series ;

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anterior teeth reduced in size toward chondrophore, posterior teeth not re-
duced ; interior of valve subnacreous ; lower inner margin of valve smooth
(Woodring) .

At the present time the subgenus “Nucolopsis” is represented by the
type species Nucula hilli, in the Miocene of Bowden, Jamaica, and by one
species, N. schencki, in the Recent fauna of western Mexico.

At the time of description of this subgenus Woodring spelled it Nucolop-
sis in the caption but following that on the same page it is consistently
spelled “Nuculopsis.” There is a prior Nuculopsis described by Girty and
also one by Rollier. Dr. Schenck is working on the family Nuculidae and is
of the opinion that Nucolopsis may be regarded as a typographical error for
Nuculopsis. We have used the original spelling used by Woodring but the
name may not be valid, and if not it will be rectified in Schenck’s forthcom-
ing work on the Nuculidae.

Nucula I Nucolopsis) schencki Hertlein & Strong, sp. nov.

Plate I, Figures 8, 9, 10.

Shell thin, polished; rather compressed; beaks small; dorsal margin
anterior to beaks elevated and broadly rounded; anterior end more acutely
rounded; ventrai margin and posterior end broadly rounded; sculpture con-
sists of rather c.oseiy spaced concentric ribs crossed by faint radiating
striae which are weaker on the umbos ; iunular area not depressed, area of
escutcheon marked only by a very short thin line. Length 2.0 mm., height
1.7 mm., thickness botn valves l.U mm.

Interior of paratypes reveals a smooth, glossy surface with faint crenu-
lations along the ventral margin corresponding to the exterior striae;
chondrophore relatively large, deeply set, elongate, subtriangular, the axis
oblique ; teeth well developed, about 9 in the anterior series and 5 in the
posterior series.

Holotype, from Sta. 195-D-9, dredged in 7 fathoms (12.6 meters) Lat.
15° 44' 28" N., Long. 96° 07' 51" W., Port Guatulco, Mexico, gray sand and
crushed shell bottom.

Nucula schencki sp. nov. resembles Nucula {“Nuculopsis”) hilli Wood-
ring38 described from the Miocene of Jamaica more closeiy than any Recent
west American shell. The more elongate shape, lack of radial striae exte-
riorly, noncrenulate lower interior margin and greater number of teeth
on the Jamaican shell easily separate it from the present species. We have
had available photographs of the type specimen of Nucula hilli for compari-
son with our new species. The photographs were furnished by Dr. H. G.
Schenck who received them from Dr. Paul Bartsch of the U. S. National
Museum.

The new species has been referred to the subgenus Nucolopsis because
it resembles that group in most ot its general characters although it does
have very faint crenmations on the interior of the ventral margins.

This species is named for Dr. Hubert G. Schenck of Stanford University
in recognition of his work on the Nuculidae.

The subgeneric position of the following species of Nucula is uncertain.

Nucula agujana Dali.

Nucula agujana Dali, Bull. Mils. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 370, pi. 10, figs. 6, 7. U. S. S. Albatross station 4654, “twenty-four miles N.
68° W. from Aguja Point, Peru, in 1036 fathoms, mud, bottom temperature 37°. 3
F.”

38 Nucula ( Nuculopsis ) hilli Woodring, Carnegie Inst. Washington, Puhl. No. 366, May, 1925,
p. 14, pi. 1, figs. 2 and 3. Bowden, Jamaica, Miocene.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 385

Type Locality: 24 miles northwest of Aguja Point, Peru, in 1,036 fath-
oms, mud. Type No. 110,571 U. S. Nat. Mus.

Range : Known only from the type locality.

Shell small, triangular, inequilateral, polished, nearly black with paler
olivaceous umbos; hinge with 15 anterior and 9 posterior teeth; margins
of the valves smooth. Length, 11 mm.; height, 8 mm.; diameter, 5.75 mm.

Nucula chrysocoma Dali.

Nucula chrysocoma Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 370, pi. 18, figs. 3 and 4. U. S. S. Albatross Station 4656, “off the coast of
Peru in S. Lat. 6°55' and W. Long. 83°34', in 2,222 fathoms, green mud, bottom
temperature 35°. 2 F.” Also off Manta, Ecuador, in 401 fathoms, and off Acapulco,
Mexico, in 660 fathoms.

Type Locality : Off the coast of Peru, Lat. 6° 55' S., Long. 83° 34' W., in
2,222 fathoms, green mud. Type No. 110,572 U. S. Nat. Mus.

Range : Off Acapulco, Mexico, to Peru, in deep water.

Shell small, plump, solid, subtriangular, polished, and of a light yellow-
ish-olive color; sculpture of irregularly spaced impressed concentric lines
and fine slightly raised, close-set radial lines; hinge with 10 anterior and 6
posterior teeth; margins of valves entire. Length, 5.0 mm.; height, 4.5 mm.;
diameter 3.0 mm.

Nucula colombiana Dali.

Nucula colombiana Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 371. “West coast of Colombia, in Panama Bay at station 2799, in 29% fathoms.”
Also in Panama Bay, in 51 fathoms ; off Manta, Ecuador, in 401 fathoms ; southern
Chile, in 122 and in 194 fathoms; off west coast of Patagonia, Lat. 51° 12' S., in

258 fathoms. Dali, Proc. U. S. Nat. Mus., Vol. 37, 1909, p. 250. Panama to

Patagonia.

Type Locality : West coast of Colombia, in Panama Bay, in 29y2 fath-
oms. Type No. 110,686 U. S. Nat. Mus.

Range: Panama to west coast of Patagonia in Lat. 51° 12' S.

Shell small, very inequilateral, ovate, white with a. pale olive perios-
tracum, smooth and brilliantly polished; hinge with 14 anterior and 7 poste-
rior teeth ; margins of valves smooth. Length, 4.5 mm. ; height, 3.0 mm. ;
maximum diameter, 2.2 mm.

Nucula grayi d’Orbigny.

Nucula obliqua Lamarck, Sowerby, Conch. Illustr., Nuculae, 1833, p. 5, pi. 16,

fig. 21. “Valparaiso, Mr. Cuming.” Hanley, Cat. Recent Bivalve Shells, p.

171, 1843, p. 376, 1856?, Suppl. pi. 20, fig. 9, 1856. “Valparaiso, South Seas.”

Not Nucula obliqua Lamarck, Hist. Nat. Anim. s. Vert., Vol. 6, 1819, p. 59.
“Habite les mers australes, au cap aux Huitres.”

Nucula grayi d’Orbigny, Voy. Amer. Merid., Vol. 5, 1846, p. 625. “Cette
espece, la plus grande que nous connaissions dans les mers du Chili, a ete recueillie
par nous pres de Valparaiso, ou elle est rare.” New name for “ Nucula obliqua,

Gray (non Lamarck).” Hanley, Thes. Conch., Vol. 3, 1860, p. 15 7, pi. 229

( Nuculidae , pi. 4), fig. 126. Chili. Hupe, Hist, de Chile, Zool., Vol. 8, 1854,

p. 304. Sowerby, Conch. Icon., Vol. 18, Nucula, October, 1870, sp. 13, pi. 2,

fig. 13. “Hah. New Zealand.” [Not New Zealand], Dali, Proc. U. S. Nat.

Mus., Vol. 37, 1909, p. 250. Valparaiso, Chile.

Type Locality: Valparaiso, Chile.

Range: Known only from the vicinity of the type locality.

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Shell thin, oval, much longer than broad, with a polished greenish-olive
periostracum; margins entire.

Nucula iphigenia Dali.

Nucula iphigenia Dali, Proc. U. S. Nat. Mus., Vol. 18, April 23, 1896, p. 15.
“U. S. Fish Commission station 3396, in 259 fathoms, Gulf of Panama; tempera-
ture, 47.4°F.” Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,

p. 369, pi. 7, figs. 1 and 4. “Gulf of Panama, in 259 fathoms, hard bottom;
temperature, 47.4° F.”

Type Locality : Gulf of Panama, in 259 fathoms, hard bottom. Type No.
122,895 U. S. Nat. Mus.

Range: Known only from the type locality.

This fine shell is one of the largest known nuculas, and peculiar from
its elongated shape and posterior attenuation. The periostracum seems to
have been thin, dull, and yellowish (Dali). The margins of the shell are
denticulate. The hinge with about 30 anterior and 15 posterior teeth. Length,
35 mm.; height, 22.5 mm.; diameter, 16 mm.

Nucula panamina Dali.

Nucula panamina Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 368, pi. 6, fig. 11. U. S. S. Albatross, station 3360, “Gulf of Panama, in 1672
fathoms, sand, bottom temperature 42°F.”

Type Locality: Gulf of Panama, in 1,672 fathoms, sand. Type No.
122,894 U. S. Nat. Mus.

Range: Known only from the type locality.

Shell large, thick, solid, rather donaciform with dark polished olivaceous
periostracum; interior more or less radially striate, but valve margins
smooth and entire; hinge with 20 anterior and 10 posterior teeth. Length
22 mm.; height, 16 mm.; diameter, 9 mm.

Nucula paytensis A. Adams.

Nucula paytensis A. Adams, Proc. Zool. Soc. London, 1856, p. 51. “ Hab . Payta,

Peru. Mus. Cuming.” Sowerby, Conch. Icon., Vol. 18, Nucula, October, 1870,

sp. 23, pi. 3, fig. 23. “Hab. Payti, Peru.” Dali, Proc. U. S. Nat. Mus., Vol.

37, 1909, p. 250. Paita, Peru.

[ Nucula crenulata ] var.? N. paytensis A. Adams, Hanley, Thes. Conch., Vol.
3, 1860, p. 154, pi. 230 (Nuculidae, pi. 5), figs. 160 and 161. [No locality cited].

Type Locality : Paita, Peru.

Range: Known only from the type locality.

A very oblique, ovate, gibbose species, concentrically grooved and
decussately striated (A. Adams).

Nucula pigafettae Dali.

Nucula pigafettae Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 369. U. S. S. Albatross, station 2780, “Magellan Straits, in 369 fathoms, mud,
bottom temperature 47 °F.”

Type Locality : Magellan Straits, in 369 fathoms, mud. Type No. 96,243
U. S. Nat. Mus.

Range: Known only from the type locality.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 387

Shell small, inequilateral, with blackish-brown or dark olivaceous peri-
ostracum; irregularly concentrically striated and sometimes very minutely
radiately striated but not visibly reticulate; hinge with 15-17 anterior and
7 posterior teeth; margins of valves entire. Length, 15 mm.; height, 10.5
mm.; maximum diameter, 7.5 mm.

Nucula pisum Sowerby.

Nucula pisum Sowerby, Proc. Zool. Soc. London, 1832 (issued March 13,
1833), p. 198. “Hab. ad Valparaiso.” ‘‘Dredged in coarse sand and gravel, at

various depths from seven to forty fathoms.” — Sowerby, Conch. Illustr.,

Nuculae, 1833, p. 6, pi. 16, fig. 23. Valparaiso. Hanley, Cat. Rec. Biv.

Shells, p. 172, 1843, p. 376, 1856?, Suppl. 20, fig. 12, 1856. Valparaiso. Hupe,

Hist, de Chile, Zool., Vol. 8, 1854, p. 304. Valparaiso, etc. Hanley, Thes.

Conch., Vol. 3, Nuculidae, 1860, p. 153, pi. 229 ( Nuculidae , pi. 4), fig. 133. Val-
paraiso, Chile. Philippi, Los Fos. Terc. i. Cuart. Chile, 1887, p. 190, pi. 41,

fig. 25. Recent on coast of Chile, also “fosil en la hacienda de La Cueva.”

Dali, Proc. U. S. Nat. Mus., Vol. 37, 1909, p. 250. Valparaiso to San Bias, Chile.

Type Locality: Valparaiso, Chile, in 7 to 40 fathoms, sand and gravel.

Range: Valparaiso to San Bias, Chile.

Reference to the illustration of Nucula pisum given in Reeve’s39 Con-
chologia Iconica has been omitted by us from the synonymy of this species
because von Ihering40, stated that the illustration is definitely that of N.
semiornata d’Orbigny, an Atlantic species.

Nucula savatieri Mabille & Rochebrune.

Nucula savatieri Mabille & Rochebrune, Miss. Sci. Cap Horn, Vol. 6, Zool., pt.

2, 1889, p. H 112, pi. 8, figs. 2a, 2b, 2c. "Hab. Canal du Beagle. Baie

Orange.” Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908, p. 367,

pi. 18, fig. 11. Gulf of Panama, in 322 fathoms; 24 miles off Aguja Point, Peru,
in 1036 fathoms; on the west coast of Patagonia in 122 fathoms; Straits of
Magellan, in 77 fathoms.

Type Locality: Beagle Canal and Orange Bay, Chile.

Range: Straits of Magellan, Patagonia, to the Gulf of Panama.

Shell compressed-ovate, very inequilateral, pale olive periostracum;
valves with smooth entire margins ; hinge with 24 anterior and 10 posterior
teeth. Length, 14-21 mm. ; height, 10-15 mm. ; diameter, 5-8 mm.

Nucula taeniolata Dali.

Nucula taeniolata Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 368, pi. 7, figs. 3 and 5. U. S. S. Albatross, station 3417, “off Acapulco, Mexico,

in 493 fathoms, mud, bottom temperature 40.6° F.” Heath, Mem. Mus. Roy.

d’Hist. Nat. Belg., Ser. 2, Fasc. 10, 1937, pp. 4, 6, 17, pi. 3, fig. 24 (Anatomy).
Acapulco, Mexico, in 493 fathoms.

Type Locality: Off Acapulco, Mexico, in 493 fathoms, mud. Type No.
122,897 U. S. Nat. Mus.

Range : Known only from the type locality.

Shell elongate, almost rostrate, thin, polished, and with dark olive-col-
ored periostracum, more or less minutely irregularly wrinkled with a few
very faint almost microscopic irregularly distributed radial striae ; margins
of valves simple, smooth; hinge with 10 anterior and 6 posterior teeth.
Length 17 mm.; height, 11 mm.; diameter, 6.5 mm.

39 Nucula pisum Sowerby, Sowerby, Conch. Icon., Vol. 18, Nucula, October, 1870, sp. 24, pi. 4,
fig. 24.

40 von Ihering, H., An. Mus. Nac. Buenos Aires, Ser. 3, Vol. 7, 1907, p. 227.

388

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Nucula tanneri Dali.

Nxicula tanneri Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908, p.
367. “Straits of Magellan at station 2780, in 369 fathoms, mud, bottom tempera-
ture 47°F.” Also on the west coast of Patagonia, in 61 to 348 fathoms, mud,
temperature 48° to 54°.

Type Locality: Straits of Magellan, in 369 fathoms, mud. Type No.
96,243 U. S. Nat. Mus.

Range : Straits of Magellan and west coast of Patagonia.

Shell oval, inequilateral, brownish periostracum with concentric zones
of darker and lighter shade; sculpture of faint concentric ripples; margins
smooth; hinge with 14-17 anterior and 7-9 posterior teeth. Length, 22.5 mm. ;
height, 16.5 mm.; diameter, 10.0 mm.

Genus Acila H. & A. Adams.

Acila H. & A. Adams, Gen. Rec. Moll., Vol. 2, January, 1858, p. 545.

Stoliczka, Mem. Geol. Surv. India, Palaeont. Indica, Cret. Fauna South India,

Vol. 3, 1871, p. 325. “N. divaricata, Hinds, is the type.” Schenck, Geol. Soc.

America, Spec. Publ. No. 4, 1936. See especially pp. 7-44.

Type (designated by Stoliczka) : Nucula divaricata Hinds. [ Proc . Zool.

Soc. London, 1843, p. 97. China Sea; from eighty-four fathoms. — Hinds,

Zool. Voy. Sulphur, Moll. pt. 3, 1844 [January, 1845 on cover of pt. 3], p. 62,

pi. 18, fig. 4. “Inhab. China Sea. From eighty-four fathoms.” Sower-

by, Conch. Icon., Vol. 18, Nucula, October, 1870, sp. 29, pi. 4, fig. 29. “Hab.

Chinese Seas.” Schenck, Geol. Soc. America, Spec. Publ. No. 4, 1936,

p. 90, pi. 15, figs. 1-10; text fig. 8 (1, 2). Earlier records cited].

The species of the genus Acila are characterized by the presence of
divaricate sculpture on the shell.

A recent monograph by Schenck41 of the members of this group should
be consulted for information regarding the species of Acila. The genus is
known from Cretaceous to Recent.

Subgenus Truncacila Schenck in Grant & Gale.

Truncacila Schenck in Grant & Gale, Mem. San Diego Soc. Nat. Hist., Vol. 1,

November 3, 1931, p. 115. Type: Nucula castrensis Hinds. Schenck, Geol.

Soc. Amer., Special Publ. No. 4, July 18, 1936, p. 23. Type: Nucula castrensis
Hinds.

Type (by original designation ) : Nucula castrensis Hinds.

Shell without the shallow sinus characteristic of the typical section of
Acila, the posterior end being truncated, as a rule, nearly at right angles
(Grant & Gale).

Acila I Truncacila I castrensis Hinds.

Nucula castrensis Hinds, Proc. Zool. Soc. London, 1843, p. 98. "Hab. Sitka,
Northwest America. A single specimen was dredged in the harbour, from seven

fathoms, sand.” Hinds, Zool. Voy. Sulphur, Moll., Pt. 3, 1844 [Jan. 1845 on

cover of Pt. 3 ] , p. 63, pi. 18, fig. 5. “Inhab. Sitka, North-west America. A single

specimen was dredged in the harbour, from seven fathoms, sand.” Sowerby,

Conch. Icon., Vol. 18, Nucula, October, 1870, sp. 32, pi. 4, fig. 32. "Hab. North-
West America.”

Nucida lyalli Baird, Proc. Zool. Soc. London, 1863, p. 71. "Hab. Esquimalt
Harbour, Vancouver Island; dredged by Dr. Lyall in 8 to 10 fathoms.”

Acila castrensis Hinds, Ashley, Proc. Calif. Acad. Sci., Ser. 2, Vol. 5, 1895, pp.
327, 344. Purisima and Tunitas Creek, San Mateo County, California, Pliocene.

41 Schenck, H. G. Nuculid Bivalves of the genus Acila. Geol. Soc. America, Special Publ. No. 4,
July, 1936, 149 pp., 18 pis.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 389

Also Miocene to Recent. Frizzell, Nautilus, Vol. 44, no. 2, 1930, pp. 50-53.

Friday Harbor, Puget Sound, Washington, Recent. Haas, Dr. H. G. Bronns

Klassen u. Ordnung des Tierreichs, Moll., Bivalvia, Bd. 3, Lief. 4, 1933, pp. 529-
531, fig. 258.

Nucula ( Acila ) castrensis Hinds, I. S. Oldroyd, Stanford Univ. Publ. Univ.
Ser. Geol. Sci., Vol. 1, no. 1, 1924, p. 14, pi. 5, fig. 11; pi. 37, figs. 1, 2. Bering Sea to

San Diego, California, also Miocene, Pliocene and Pleistocene. Grant &

Gale, Mem. San Diego Soc. Nat. Hist., Vol. 1, 1931, p. 116, pi. 1, figs. 6a, 6b.
Earlier records cited. Miocene to Recent.

Acila (Truncacila) castrensis Hinds, Schenck, Geol. Soc. America, Special
Publ. No. 4, 1936, p. 96, pi. 10, figs. 1-15; pi. 14, figs. 2, 3; text figure 7 (2-5).
Pliocene to Recent. Earlier records cited with discussion of the species.

Type Locality. Sitka, Alaska, dredged in harbor in 7 fathoms.

Range: Sitka, Alaska, to Cedros Island, Lower California, 40 to 262
fathoms.

Collecting Station: Mexico; East of Cedros Island (126-D-10). 60
fathoms, crushed shell and eel grass bottom.

A single valve of Acila castrensis was dredged about lfi miles east of
Cedros Island. This is a southern extension in the range of the species.
Heath42 has recently discussed the soft parts of A. castrensis and Schenck
has discussed the species in detail in his work on Acila.

Family Nuculanidae.

The shells of the Nuculanidae are somewhat similar to those of the
Nuculidae but the posterior end is elongate and rostrate; the ends of the
shell are sometimes partly gaping; a ligament is present, often behind the
beaks; the interior of the shell is porcellanous or sub-nacreous; the pallial
line is sinuate due to the presence of siphons.

Most of the species of the Nuculanidae dwell in muddy portions of the
bottom of the sea.

No representatives of Yoldia, Cyrilla, Malletia or of Tindaria are pres-
ent in the collections made by Crocker and Beebe from west American waters.
This may be explained in part by the fact that most of the species of such
genera as Yoldia and Tindaria in the region covered by these expeditions
occur at depths considerably greater than those from which most of the
collections were secured.

Key to the Genera of Nuculanidae.

A. No lateral teeth on hinge line

a. Ligament internal Nucidana

aa. Ligament external

b. Shell pointed posteriorly

c. Shell veneriform, thickened Tindaria

cc. Shell elongate

d. Shell lanceolate, length more than double width

Adrana

dd. Shell broader, length about one third greater than

width Yoldia

bb. Shell rounded posteriorly Malletia

B. A well developed lateral tooth in right valve with corresponding depres-
sion in left; shell very small Cyrilla

42 Heath, H. Mem. Mus. Roy. d’Hist. Nat. Belg., Ser. 2, Fasc. 10, 1937, pp. 4, 9, qtc. pi. 4,
figs 30, 36; pi. 6, fig. 50; pi. 9, figs. 76, 77. See also Hilton, W. A., Jour. Entomol. & Zool., Vol. 11,
no. 4, 1919, p. 77, figs. 1-7. Monterey Bay, California, in 60-80 fathoms, and south of Alaskan
Peninsula, in 483 fathoms.

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Genus Nuculana Link.

Nuculana Link, Beschreib. Nat.-Samml. Univ. Rostock, Abt. 3, May 17, 1807,

p. 155. Sole species: “N. rostrata” [Chemnitz]. Grant & Gale, Mem. San

Diego Soc. Nat. Hist., Vol. 1, 1931, p. 118. Type: Area rostrata Chemnitz.

Leda Schumacher, Essai Nouv. Syst. Test., 1817, pp. 55, 172, 173. Type:
Area rostrata Chemnitz. Illustrated by Schumacher on pi. 19, fig. 4.

Type (by monotypy) : Area rostrata Chemnitz. [Neues Syst. Conchyl.-
Cab., Bd. 7, 1784, p. 206, pi. 55, figs. 550, 551. Recent, northern coasts of
Europe. Montagu, Suppl. Test. Brit., 1808, p. 55, pi. 27, fig. 7. Nor-

wegian and Baltic Seas, also Scotland].

The genus Nuculana and closely related forms originated in the Paleo-
zoic. Praeleda Pfab45 has been described from the Silurian of Bohemia and
has been recorded from the Ordovician of Belgium. The genus Nuculana is
widely distributed but the majority of the species occur on muddy bottoms
in cold water. Verrill & Bush44 published a revision of the genera of the
Nuculanidae of the Atlantic coast of the United States. Stewart45 has dis-
cussed some of the superspecific groups in the Nuculanidae as has Iredale46
and Prashad47.

Key to the Species of Nuculana.

A. Umbos distinctly anterior

a. Surface smooth or with very feeble sculpture

b. Hinge with 15 anterior and 22 posterior teeth loshka

bb. Hinge with 15 anterior and 38 posterior teeth peruviana

aa. Surface with distinct concentric sculpture

c. Umbos at about the anterior fourth

d. Posterior dorsal margin nearly straight hamata

dd. Posterior dorsal margin strongly concave cordyla

cc. Umbos at about the anterior third

e. Anterior end with a distinct radiating ray or shallow
groove

f. Size small, length about 6 mm acuta

ff. Size large, length about 20 mm. or more

g. Concentric sculpture fine, weak marella

gg. Concentric sculpture strong, distant riblets

costellata

ee. Anterior end without a radiating ray or groove or if pres-
ent, very weak

43 Praeleda Pfab, Palaeontogr., Bd. 80, Abt. A, Lief. 4-6, April, 1934, p. 231. “Typus des genus :
P. compar (Barr.).”

44 Verrill, A. E., & Bush, H. J. Revision of the genera of Ledidae and Nuculidae of the Atlantic

Coast of the United States. Amer. Jour. Sci., Ser. 4, Vol. 3, 1897, pp. 51-63, 22 figs, in text. —

Proc. U. S. Nat. Mus., Vol. 20, 1898, pp. 775-901.

Yonge ( Phil . Trans. Roy. Soc. London, Ser. B, Biol. Sci., No. 566, Vol. 230, August 9, 1939,
pp. 79-147, 1 pi., 39 figs, in text) has discussed the anatomy of the protobranchiate mollusks.

45 Stewart, R. B., Acad. Nat. Sci. Philadelphia, Spec. Publ. No. 3, 1930, pp. 48-64.

46 Iredale, T„ Rec. Austral. Mus., Vol. 17, no. 4, 1929, pp. 158-159. — Brit. Mus. (Nat.

Hist.) Great Barrier Reef Exped. 1928-29. Sci. Repts. Vol. 5, no. 6, Moll. Pt. 1, 1939, pp. 238-241.
In this paper three new genera of Nuculanidae are proposed. Eptoleda, ‘‘Type: Leda darwini
Smith”; Zygonoleda, "Type: Z. corbuloides minutalis subsp. nov.” ; Tepidoleda, “Type: T. lata
orion subsp. nov.”

Powell (Proc. Malacol. Soc. London, Vol. 21, pt. 4, March, 1935, pp. 252-255, pi. 27), has de-
scribed some Recent and Tertiary Nuculanidae from New Zealand.

47 Prashad, B., Siboga Exped., Vol. 53c, Lamell., 1932, pp. 18-19.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 391

h. Anterior end without radiating ray or groove

i. Without deeply impressed lines near base calcar

ii. With 6 to 8 impressed lines near base, not coinci-
dent with lines of growth calcarella

hh. Anterior end with a very weak radiating ray or
groove cuneata

B. Umbos central or subcentral

a. Sculpture faint or obsolete over most of the shell

b. Striae obsolete on central portion of shell; no radials

c. Size large, conspicuous oblique striae posteriorly, length

about 35 mm. polita

cc. Size smaller, no oblique posterior striae, length about 15

mm. pontonia

bb. Striae present on basal margin, radially rippled acrita

aa. Sculpture strong, continuous, concentric

d. Posterior dorsal area defined by two ridges or angles
e. Anterior end with a radial ray or groove
f. Concentric ridges evenly spaced

g. Radiating lyrae well developed

h. Radiating lyrae over entire shell hindsii
hh. Radiating lyrae confined to posterior end

crispa

gg. Radiating lyrae, if present, very fine, confined to
the umbonal region

i. Anterior groove shallow, barely discernible .

eburnea

ii. Anterior groove very conspicuous ornata

ff. Concentric ridges not evenly spaced

j. Ridges widely spaced toward umbos, close toward

margin impar

jj. Ridges irregularly spaced elenensis

ee. Anterior end without a radial ray or groove

k. Posterior end obtuse (60°) , shell thin, fragile agapea

kk. Posterior end acute, shell solid bicostata

dd. Posterior dorsal area with not more than one ridge or angle

1. size small, length less than 25 mm.

m. Dorsal area defined by one ridge, posterior end attenu-
ated.

n. Concentric sculpture extending at least to posteri-
or dorsal ridge

o. Size about 15 to 20 mm.

p. Anterior hinge teeth 26, posterior 20

callimene

pp. Anterior hinge teeth 12, posterior 12

taphria

oo. Size smaller, length 10 mm. or less

q. Length about 10 mm. excavata

qq. Length about 5 mm oxia

nn. Shell smooth, at least in front of posterior dorsal
ridge laeviradins

392 Zoologica: Netv York Zoological Society [XXV :25

Key to the Species of Nuculana (continued) :

mm. Posterior dorsal area without ridge or angle, both
ends rounded

r. Anterior end without a radiating ray or groove

s. Posterior end attenuated but not acute

rhytida

ss. Both ends evenly rounded lobula

rr. Anterior end with a radiating ray or groove

lucasana

11. Size large, shell heavy, length over 30 mm. gibbosa

Subgenus Saccella Woodring.

Ledina Sacco, Moll. Terr. Terz. Piemonte e Liguria, Pt. 26, December, 1898,
p. 53. “(tipo L. fragilis (Chemnitz.)).”

Not Ledina Dali, Trans. Wagner Free Inst. Science, Vol. 3, pt. 4, April, 1898,
p. 580. Type: Leda eborea Conrad, 1860 (not L. eborea Conrad, 1846) =L. smirna
Dali, Eocene of Gulf States of United States.

Saccella Woodring, Carnegie Inst. Washington, Publ. No. 366, May 20, 1925,
p. 15. Type: Area fragilis Chemnitz Leda commutata Philippi). New name for
Ledina Sacco not Ledina Dali.

Type (by original designation) : Area fragilis Chemnitz. [Neues Syst.
Conchyl.-Cab., Bd. 7, 1784, p. 199, pi. 55, fig. 546. “Mittellandischen Meeres.”

; — Bucquoy, Dautzenberg & Dodfus, Moll. Mar. Roussillon, Vol. 2, Fasc.

5 (Pelecypoda, Fasc. 18), 1891, p. 215, pi. 37, figs. 26, 27, 28, 29, 30, 31.
Mediterranean, and Atlantic Ocean. ( =Leda commutata Philippi). Miocene
to Recent in the Mediterranean region.].

Saccella resembles Lembidus 4S but the former has concentric instead of
diagonal sculpture and has a narrower and shallower posterior groove. Sac-
cella occurs Recent and fossil in Europe, eastern and western North and
Central America and elsewhere. It has been recorded fossil in the Eocene of
California40 and Tertiary of New Zealand00.

Nuculana I Saccella) acuta Conrad.

Plate I, Figure 11.

Nucula acuta Conrad, Amer. Mar. Conch., 1832, p. 32, pi. 6, fig. 3. North
Carolina; Recent.

Leda acuta Conrad, Verrill, Trans. Connecticut Acad. Arts and Sci., Vol. 6,

1884, p. 259, pi. 30, fig. 15. New England Coast. Dali, Bull. Mus. Comp.

Zool., Vol. 12, no. 5, 1886, p. 251, pi. 7, figs. 3a, 3b, 8 [Not all the synonymy].

New England; Florida; Jamaica. Dali, Trans. Wagner Free Inst. Sci.,

Vol. 3, pt. 4, 1898, p. 592. Oligocene to Recent on Atlantic Coast. Also Recent on

Pacific Coast of California. Dali & Simpson, Bull. U. S. Fish Comm., Vol.

20, pt. 1, for 1900 (issued separately November 29, 1901), p. 458. Mayaguez

Harbor, Porto Rico. Dali, Proc. U. S. Nat. Mus., Vol. 37, 1909, p. 250.

California, the Gulf of Panama, and south to Valparaiso, Chile. Also Atlantic.

— : Gardner, V . S. Geol. Surv., Prof. Paper 142-A, 1926, p. 12, pi. 2, figs. 13-16.

Eastern North America; Miocene to Recent.

48 Lembulus Risso, Hist. Nat. l’Europ. Merid., Vol. 4, 1826, p. 319. Type (designated by Gray,
1847): Lembulus rossianus Risso ( —Area pella Linnaeus). Recent, Mediterranean Sea. Illustrated
by Risso, 1826, pi. 11, fig. 166, and by Bucquoy, Dautzenberg & Dollfus, Moll. Mar. Roussillon, Vol.
2, Fasc. 5 (Pelecypoda, Fasc. 18), 1891, p. 218, pi. 37, figs. 32, 33, 34, 35. Mediterranean; Adriatic;
Atlantic Ocean.

49 Vokes, H. E., Ann. New York Acad. Sci.', Vol. 38, 1939, pp. 41-43.

50 See Nuculana ( Saccella ) duplicarina Laws, Trans, and Proc. Roy. Soc. Neiv Zealand, Vol.

68, pt. 4, Match, 1939, p. 468, pi. 62, fig. 2. Pakaurangi Point, Kaipara Harbor, New Zealand.
Tertiary. — & See also, Marwick, Geol. Surv. New Zealand, Palaeo. Bull. No. 13, 1931, pp. 50, 51.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions

393

Nuculana acuta Conrad, M. Smith, East Coast Mar. Shells (Edwards Bros.,
Ann Arbor, Michigan), 1937, p. 26, pi. 2, fig. 2. Off Martha’s Vineyard to the
West Indies.

Type Locality. North Carolina.

Range: Off Martha’s Vineyard, Massachusetts, to the West Indies.

Nuculana acuta has been recorded from various localities on the west
coast ranging from Alaska to Chile. Dali in 1909 cited the range as from
California south to Chile. In a later work51 he cited it as occurring from
Nazan Bay, Atka Island, Aleutian Islands, Alaska, to the Gulf of California.
The species is well known on the Atlantic coast from Massachusetts to the
West Indies, but its occurrence in Western America can be considered very
doubtful until definitely proved.

Nuctilcma l Saccella l callimene Dali.

Plate I, Figure 13.

Leda ( Jupiteria ) callimene Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 372, pi. 17, figs. 3 and 4. “U. S. S. ‘Albatross,’ Station 3396,
Gulf of Panama, 259 fathoms, mud, bottom temperature 47.4° F.” Also at Tome,
Chile, in 14 fathoms.

Leda callimene Dali, Dali, Proc. U. S. Nat. Mus., Vol. 37, 1909, p. 250. Gulf

of Panama to Tome, Chile. Zetek, Rev. Nueva, Nos. 1 & 2, 1918, p. 37.

Panama.

Type Locality: Gulf of Panama, in 259 fathoms, mud. The type is No.
122,910 U. S. Nat. Mus.

Range: Gulf of Nicoya to Tome, Chile.

Collecting Station: Costa Rica: 3 miles south of Blanco Island, Gulf of
Nicoya. 100 fathoms.

A single specimen of Nuculana callimene was dredged in the Gulf of
Nicoya. Compared to N. taphria Dali it is more elongate, less inflated, and
the exterior surface is less polished.

Specimens somewhat similar to Nuculana callimene have been recorded
by E. K. Jordan52 from the Pleistocene of Magdalena Bay, Lower California.

Nuculana I Saccella I elenensis Sowerby.

Plate I, Figures 12, 14, 15, 16, 17, 18, 19, 22.

Nucula elenensis Sowerby, Proc. Zool. Soc. London, 1832 (issued March 13,
1833), p. 198. “ Hab . ad Sanctam Elenam.” “Dredged in sandy mud at a depth of

six fathoms.” Recent. -Sowerby, Conch. Illustr., Nuculae, 1833, p. 4, pi. 15,

fig. 14. “St. Elena, Mr. Cuming.” Muller, Synop. Test. Viv., 1836, p. 191.

Type locality cited. Reeve, Conch. Syst., Vol. 1, 1841, p. Ill, pi. 85, fig. 14.

Hanley, Cat. Rec. Bivalve Shells, 1843, p. 169, Suppl. pi. 19, fig. 56, 1846.

“W. Colombia.” C. B. Adams, Ann. Lyceum Nat. Hist. New York, Vol. 5,

1852, p. 478. Panama. Also earlier records cited.

Leda ? elenensis Sowerby, Carpenter, Cat. Mazatlan Shells, February, 1856,
p. 145. Mazatlan, Mexico. Also earlier records cited.

Leda elenensis Sowerby, d’Orbigny, Voy. Amer. Merid., Vol. 5, 1846, p. 545.
Santa Elena, Ecuador. — — — Hanley, Thes. Conch., Vol. 3, 1860, Leda, p. 121,

pi. 228, [ Nuculidae , pi. 3], figs. 70, 71, 72. Panama. -Sowerby, Conch. Icon.,

Vol. 18, Laeda, September, 1871, sp. 32, pi. 6, figs. 32a, 32b. “Hab. St. Elena.

51 Dali, w. H., U. S. Nat. Mus.. Bull. 112, 1921, p. 10.

52 Leda sp. aff. callimene Dali (young),.. E. K. Jordan, Bull. South . Calif. Acad. Sci., Vol. 23,
pt. 5, September-October (issued October 25), 1924, p. 148. Quaternary at Magdalena Bay, Lower

California. E. K. Jordan, Contrib. Dept. Geol. Stanford Univ., Vol. 1, no. 4, 1936, p. 112.

Previous record cited.

394

Zoologica: Neiv York Zoological Society

[XXV :25

Panama.” Dali, Proc. U. S. Nat. Mus., Vol. 37, 1909, p. 251. Santa Elena,

Ecuador. Olsson, Nautilus, Vol. 37, no. 4, 1924, p. 126. Salinas, Ecuador.

Leda elenensis Sowerby var. gibbosa Hanley, Thes. Conch., Vol. 3, 1860, p.

121.

Leda elenensis Sowerby var. media Hanley, Thes. Conch., Vol. 3, 1860, p. 121.

Leda elenensis Sowerby var.? pyriformis Hanley, Thes. Conch., Vol. 3, 1860,
p. 121.

Leda ( Jupiteria ) elenensis Sowerby, Dali, Bull. Mus. Comp. Zool., Vol. 43,
no. 6, p. 374, October, 1908. In 52 fathoms in Panama Bay.

Leda ( Saccella ) acapulcensis Pilsbry & Lowe, Proc. Acad. Nat. Sci. Phila-
delphia, Vol. 84, May 21, 1932, p. 107, pi. 17, figs. 1 and 2. “Mexico: Acapulco,
20 fathoms.”

Nuculana acapulcensis Pilsbry & Lowe, Strong, Hanna & Hertlein, Proc.
Calif. Acad. Sci., Ser. 4, Vol. 21, no. 10, December 21, 1933, p. 118. Acapulco,
Mexico.

Nuculana elenensis Sowerby, Hertlein, Bull. South. Calif. Acad. Sci., Vol. 33,
pt. 2, May-August (issued August 31), 1934, p. 65. San Ignacio Lagoon, Lower

California; Pleistocene. E. K. Jordan, Contrib. Dept. Geol. Stanford Univ.,

Vol. 1, no. 4, 1936, pp. 112, 118. Magdalena Bay, Lower California; Pleistocene.
Recent, Panama and Santa Elena, Ecuador.

Nuculana elenense Sowerby, Strong & Hertlein, Allan Hancock Pacific Exped.
(Univ. South. Calif.), Vol. 2, no. 12, 1939, p. 183. Bahia Honda, Veragua, Panama;
off Taboga Island, Panama.

Type Locality : Santa Elena, Ecuador, in 6 fathoms, sandy mud.

Range : Santa Inez Bay, Lower California, to Salinas, Ecuador.

Collecting Stations: Mexico: Arena Bank, Gulf of California (136-D-
13,-18). 40-45 fathoms, mud and Area conglomerate bottom; Santa Inez Bay,
Gulf of California (144-D-2). 2^2 fathoms, sand, weed and rock bottom;
also (145-D-l,-3, and shore). 4-13 fathoms, sand bottom; Tenacatita Bay
(183-D-3). 40 fathoms, sandy mud bottom; Manzanillo (184-D-2). 30
fathoms, gravelly sand bottom; Acapulco (189-D-3). 13 fathoms, mud bot-
tom; 4 miles south-southwest of Maldanado Point (192-D-3). 38 fathoms,
mud bottom; Port Guatulco (195-D-19). 17 fathoms, gray mud and crushed
shell bottom; Tangola-Tangola Bay (196-D-17). 23 fathoms, mud bottom;
Costa Rica: Port Parker (203-D-i,-3) . 12-15 fathoms, shelly mud bottom.

Nuculana elenensis Sowerby is a very variable species and as pointed
out by Jordan neither the form nor sculpture is constant in a series. Sowerby
gave the varietal names, gibbosa, media and pyriformis, to forms of
this species. With a very large series of shells from many localities we can
find no constant distinguishing feature for separation of Pilsbry & Lowe’s
species acapulcensis. They compared it with Leda agapea Dali which we
have not seen. The specimens here referred to elenensis have a strong groove
on the anterior end. This is not shown on the figures of acapulcensis Pilsbry
& Lowe. We have studied a series of several hundred specimens from Sta.
203-D-3. These agree well with Hanley’s figure 71 (Thes. Conch., Vol. 3,
pi. 228, Nuculidae, pi. 3). Almost none of our specimens show the nearly
smooth area near the ventral margin as shown by Pilsbry & Lowe on their
pi. 17, fig. 1. One specimen in the collection shows a tendency toward this
type of sculpture and in many of the larger specimens there is a definite
line beyond which the sculpture is somewhat finer.

Nuculana bicostata Sowerby appears to be distinguished by the pres-
ence of two pronounced rostral ribs while there is but one rib on the posterior
part of N. elenensis. Nuculana acuta Conrad is more truncate posteriorly and
is more compressed than N. elenensis. The shorter and more contracted
posterior portion of the shell serves to separate N. elenensis from young
specimens of N. peruviana Dali from Peru.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 395

Nueulana l Saccella I eburnea Sowerby.

Plate II, Figures 1, 2, 3.

Nucula eburnea Sowerby, Proc. Zool. Soc. London, 1832 (issued March 13,
1833), p. 198. "Hah. ad oras Columbiae Occidentalis.” “Found in sandy mud, at

a depth of from seven to nine fathoms, in the Bay of Caraccas.” Sowerby,

Conch. Illustra., Nuculae, 1833, pp. 4, 6, pi. 15, fig. 10. Bay of Caraccas.

Muller, Synop. Test. Viv., 1836, p. 191. Type locality cited. Reeve, Conch.

Syst., Vol. 1, 1841, p. Ill, pi. 85, fig. 10. — — Catlow, Conch. Nomencl., 1845,

p. 53_ Hanley, Cat. Rec. Bivalve Shells, p. 169, 1843, 376, 71856, Suppl. pi.

19, fig. 57, 1846. W. Colombia.

Leda eburnea Sowerby, d’Orbigny, Voy. Amer. Merid., Vol. 5, 1846, p. 546.
“M. Cuming l’a decouverte dans la baie de Caracas, sur la cote occidentale de

la republique de la Equateur.” Carpenter, Rept. Brit. Assoc. Adv. Sci. for

1863 (issued 1864), p. 560. Reprint in Smithson. Misc. Coll., No. 252, 1872, p. 46.

Panama; Bay of Caraccas. Hanley, Thes. Conch., Vol. 3, 1860, p. 124,

pi. 228 ( Nuculidae , pi. 3), fig. 90. Panama; Bay of Caraccas. Sowerby,

Conch. Icon., Vol. 18, Laeda, September, 1871, pi. 5, fig. 29. “Hab. Bay of Panama.”

Dali, Proc. U. S. Nat. Mus., Vol. 37, 1909, p. 250. “Gulf of Panama to the

Bay of Caraques, Ecuador.” Zetek, Rev. Nueva, Nos. 1 & 2, 1918, p. 37.

Panama.

Nucula lyrata Hinds, Proc. Zool. Soc. London, 1843, p. 100. “Hab. Panama;

from thirty fathoms.” Hinds, Zool. Voy. Sulphur, 1844 (January, 1845 on

the cover of Moll., Pt. 3), p. 64, pi. 18, fig. 12. Original locality cited.

Type Locality : Bay of Caraccas, Ecuador, in 7 to 9 fathoms, sandy

mud.

Range : Meanguera Island, Gulf of Fonseca, El Salvador, to Bay of
Caraccas, Ecuador.

Collecting Stations: El Salvador: Meanguera Island, Gulf of Fonseca
(199-D-l). 16 fathoms, sandy mud and crushed shell bottom; Nicaragua:
Corinto (200-D-19). 12-13 fathoms, mangrove leaves on bottom; Costa Rica:
14 miles southeast of Judas Point (214-D-l to 4). 42-61 fathoms, mud, shell
and rocky bottom.

The shell of Nueulana eburnea Sowerby has a smooth or appressed
area anterior to the posterior keel somewhat resembling N. laeviradius
Pilsbry & Lowe. It differs from that species in the great size, more acuminate
posterior end and in lacking the elevated ridge on the anterior end which
runs from the beaks to the anterior ventral margin on N. laeviradius. Speci-
mens of N. eburnea in the present collection average about 12.6 mm. in
length. Nueulana lyrata Hinds, originally described from Panama, was con-
sidered to be a synonym of N. eburnea by Hanley and Carpenter. From an
inspection of the figure given by Hinds it appears likely that N. lyrata can be
considered synonymous with N. eburnea.

Nueulana I Saccella I gibbosa Sowerby.

Plate II, Figures 5, 8.

Nucula gibbosa Sowerby, Proc. Zool. Soc. London, 1832 (issued March 13,
1833), p. 198. “Hab. ad littora Peruviae.” “Found in soft mud, at a depth of five
fathoms, at Tumbez in Peru.” “A variety, with a less elevated ridge in the centre
of the anterior dorsal margin, and of much smaller size, was found in mud, at

twelve fathoms, in the Gulf of Nocoiyo.” Sowerby, Conch. Illustr., Nuculae,

1833, p. 4, pi. 15, fig. 9. “Gulf of Nocoiyo. Mr. Cuming.” Deshayes, ed. of

Lamarck’s Anim. s. Vert., Vol. 6, 1835, p. 508. (as Nucula gibossa. Cuming, but
in index it is cited as Nucula gibbosa). “Habite les mers du Perou, pres des

rivages.” Muller, Syn. Test. Viv., 1836, p. 191. Original locality cited.

Reeve, Conch. Syst., Vol. 1, 1841, p. Ill, pi. 85, fig. 9. Hanley, Cat. Rec.

Bivalve Shells, p. 169, 1843, p. 376, 71856, Suppl. pi. 19, fig. 55, 1846. Peru.

396

Zoologica: Neiv York Zoological Society

[XXV :25

Leda gibbosa Sowerby, d’Orbigny, Voy. Amer. Merid., Vol. 5, 1846, p. 545.

“Cote de Payta (Perou).” Hanley, Thes. Conch., Vol. 3, Nuculidae, 1860,

p. 120, pi. 228 ( Nuculidae , pi. 3), fig. 79. Peru. Sowerby, Conch. Icon.,

Vol. 18, Laeda, November, 1871, sp. 51, pi. 8, fig. 51. “ Hab . Peru.” — Dali,

Proc. U. S. Nat. Mus., Vol. 37, 1909, p. 251. Gulf of Panama to Paita, Peru.

— - — — Zetek, Rev. Nueva, Nos. 1 & 2, 1918, p. 37. Panama. Strong, Hanna

& Hertlein, Proc. Calif. Acad. Sci., Ser. 4, Vol. 21, no. 10, 1933, p. 118. Acapulco,
Mexico.

Leda (Jupiteria) gibbosa Sowerby, Dali, Bull. Mus. Comp. Zool., Vol. 43, no.
6, 1908, p. 372. Panama Bay in 26 to 47 fathoms, mud.

Nuculana gibbosa Tomlin, Jour. Conch., Vol. 18, no. 7, 1928, p. 189. Coiba
Island, Panama, in 10 fathoms. Gorgona Island, Colombia, in 20 fathoms.

Type Locality : Tumbez, Peru, in 5 fathoms, soft mud.

Range: Acapulco, Mexico, to Tumbez, Peru.

Collecting Stations: Mexico: Tangola-Tangola Bay (196-D-19). 30
fathoms, mud bottom; Costa Rica: off Ballena Bay, Gulf of Nicoya (213-D-
11-17). 35-45 fathoms, mud bottom.

This is a thick, elongate, posteriorly pointed shell with fairly gibbous
beaks. It is ornamented by strong concentric somewhat upturned ribs which
are separated by smooth bottomed grooves.

Nuculana ISaccellal impar Pilsbry & Lowe.

Plate II, Figure 6.

Leda impar Pilsbry & Lowe, Proc. Acad. Nat. Sci. Philadelphia, Vol. 84,
May 21, 1932, p. 106, pi. 17, figs. 3, 4, 5, 6. “Mexico: Guaymas, 20 fathoms.”

Hertlein, Bull. South. Calif. Acad. Sci., Vol. 33 pt. 2, May-August (issued

August 31), 1934, p. 62. Maria Magdalena Island, Ties Marias Islands; Pleis-
tocene. Lowe, Trans. San Diego Soc. Nat. Hist., Vol. 8, no. 6, 1935, p. 27.

Punta Penasco, Sonora, Mexico.

Type Locality: Guaymas, Mexico, in 20 fathoms. Type No. 155636 Acad.
Nat. Sci. Philadelphia.

Range: Punta Penasco, Sonora, Mexico, to Port Parker, Costa Rica.

Collecting Stations: Mexico: Santa Inez Bay, Gulf of California (145-D-
1,-3). 4-13 fathoms, sand bottom; Costa Rica: Port Parker (203-D-3). 12
fathoms, shelly mud bottom.

The sculpture of this species is characteristic. This consists of con-
centric ridges, which are fine and close on the beaks, then widely spaced,
then again becoming closer and about equal to their intervals from about
the middle of the valves to the ventral margin.

Nuculana ISaccellal laeviradius Pilsbry & Lowe.

Plate II, Figures 4, 7.

Leda laeviradius Pilsbry & Lowe, Proc. Acad. Nat. Sci. Philadelphia, Vol. 84,
May 21, 1932, p. 106, pi. 17, fig. 7. “Mexico: Guaymas, 20 fathoms.”

Leda leviradius Pilsbry & Lowe, Lowe, Trans. San Diego Soc. Nat. Hist.,
Vol. 8, no; 6, 1935, p. 27. Punta Penasco, Sonora, Mexico.

Type Locality: Guaymas, Mexico, in 20 fathoms. Type No. 155635 Acad.
Nat. Sci. Philadelphia.

Range: Punta Penasco, Sonora, Mexico, to Port Parker, Costa Rica.
Collecting Stations: Mexico: Santa Inez Bay, Gulf of California (145-
D-1,-3). 4-13 fathoms, sand bottom; Costa Rica: Port Parker (203-D-l,-3) .
12-15 fathoms, sandy mud, crushed shell and shelly mud bottom.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 397

The presence of a smooth band preceding the posterior ridge and the
strong hinge in proportion to the size of the shell are characteristic features
of this species. In some specimens a considerable portion of the exterior of
the shell is smooth.

Nuculana (Sac cello J taphria Dali.

Nucula caelata Hinds, Proc. Zool. Soc. London, December, 1843, p. 99. “Hab.
California, between 38° 18' and 34° 24' north latitude; namely, at Russian
Bodegas, San Francisco, and Santa Barbara, in from six to ten fathoms.” “Cab.

Belcher.” Hinds, Zool. Voy. Sulphur, Moll., pt. 3, 1844 [January, 1845 on

cover of pt. 3], p. 64, pi. 18, fig. 13. Original locality cited.

Not Nucula coelata Conrad, Amer. Jour. Sci., Vol. 23, no. 2, January, 1833,
p. 343. Claiborne, Alabama. London Clay. [Eocene],

Leda caelata Hinds, Hanley, Thes. Conch., Vol. 3, 1860, p. 129, pi. 228

( Nuculidae , pi. 3), figs. 95, 96. California. Sowerby, Conch. Icon., Vol. 18,

Laeda, February, 1871, sp. 6, pi. 2, figs. 6a, 6b. “Hab. California.”

Leda taphria Dali, Nautilus, Vol. 10, no. 6, Oct., 1896, p. 70. New name for
Nucula caelata Hinds, not Nucula caelata Conrad, 1833. Claiborne, Eocene.

Dali, Nat. Hist. Soc. Brit. Columbia, Bull. No. 2, January, 1897, p. 7

(in text), pi. 2, figs. 6 and 8. Bodega Bay, California to San Diego. Arnold,

Proc. U. S. Nat. Mus., Vol. 32, 1907, p. 544, pi. 48, figs. 5, 5a. Pliocene to Recent

in the California Province. 1. S. Oldroyd, Stanford Univ. Publ. Univ. Ser.

Geol. Sci., Vol. 1, no. 1, 1924, p. 16, pi. 37, figs. 7, 7a, 8. Bodega Bay, California
to Lower California. Also Miocene to Recent in California.

Nuculana taphria Dali, Grant & Gale, Mem. San Diego Soc. Nat. Hist., Vol. 1,
1931, p. 121, pi. 1, figs. 8 and 9. Earlier records cited. Miocene to Recent.

Hertlein, Bull. South. Calif. Acad, Sci., Vol. 33, pt. 2, May-August (issued

August 31), 1934, p. 62. Maria Magdalena Island, Tres Marias Islands; Pleis-
tocene. E. K. Jordan, Contrib. Dept. Geol. Stanford Univ., Vol. 1, no. 4,

1936, p. 112. Magdalena Bay, Lower California (Dali) ; Pleistocene. Willett,

Trans. San Diego Soc. Nat. Hist., Vol. 8, no. 30, 1937, p. 386. About 2 miles NE. of
Playa del Rey, Baldwin Hills, Los Angeles County, California; Upper Pleistocene.

Type Locality : Bodega Bay, California, in 6 to 10 fathoms.

Range : Bodega Bay, California, to Punta Arena, Lower California.

Collecting Stations : Mexico: East of Cedros Island (126-D-12). 45
fathoms, crushed shell and mud bottom; Arena Bank, Gulf of California
(136-D-22). 45 fathoms, mud bottom.

The concentric sculpture on this species is uniform and prominent. The
beaks are nearly central and the rostrum is rather bluntly pointed.

Heath53 has studied the soft parts of this species.

Subgenus Politoleda Hertlein & Strong, nov.

Type: Nucula polita Sowerby.

Shell elongate, posteriorly pointed. Sculpture consists of very fine con-
centric lines of growth, and on the lower half of the shell fine wavy in-
cised lines which on the whole posterior dorsal part of the shell are nearly
parallel to the posterior dorsal margin.

Nuculana (Politoleda! polita Sowerby.

Plate II, Figure 9.

Nucula polita Sowerby, Proc. Zool. Soc. London, 1832 (issued March 13,
1833), p. 198. “Hab. ad Panamam.” “A single specimen of this very beautiful
species was dredged up in sand from a depth of seven fathoms.” -Sowerby,

53 Heath, H., Mem. Mus. Roy. d’Hist. Nat. Belgique, Ser. 2, Fasc. 10, 1937, pp. 5, 6, etc., pi.
7, figs. 59, 60.

398

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[XXV :25

Conch. Illustr., Nuculae, March 8, 1833, p. 4, pi. 15, fig. 11. “Panama. Mr. Cuming.”

Deshayes, ed. of Lamarck’s Anim. s. Vert., Vol. 6, 1835, p. 507. Panama.

Hanley, Cat. Bivalve Shells, p. 169, 1843, Suppl. pi. 19, fig. 53, 1846.

Panama. Miiller, Syn. Test. Viv., 1836, p. 190. “Legit H. Cuming ad Pan-

amam”. Reeve, Conch. Syst., Vol. 1, 1841, p. Ill, pi. 85, fig. 11.

Leda polita Sowerby, Hanley, Thes. Conch., Vol. 3, Leda, 1860, p. 125, pi. 228

( Nuculidae , pi. 3), fig. 68. Panama. Sowerby, Conch. Icon., Vol. 18, Laeda,

September, 1871, sp. 16, pi. 3, fig. 16. “Hab. Panama.” Zetek, Rev. Nueva,

Nos. 1 & 2, 1918, p. 37. Panama. Pilsbry & Lowe, Proc. Acad. Nat. Sci.

Philadelphia, Vol. 84, p. 141, 1932. On the beach at Old Panama.

Nuculana polita Sowerby, Strong & Hertlein, Allan Hancock Pac. Exped.
( Univ . South. Calif.), Vol. 2, no. 12, 1939, p. 183. Beach near Panama Vieja
(Old Panama).

Type Locality. Panama, in 7 fathoms, sand.

Range: Champerico, Guatemala, to Panama.

Collecting Stations: Guatemala: 7 miles west of Champerico (197-D-
1-2). 14 fathoms, mud bottom; El Salvador: La Libertad (198-D-2). 14 fath-
oms, mud bottom; Meanguera Island, Gulf of Fonseca (199-D-l). 16 fathoms,
sand, mud, and crushed shell bottom; Panama: Gulf of Chiriqui (221-D-1-5).
35-40 fathoms, sandy mud bottom.

The shell of Nuculana polita is oblong, anteriorly beaked, ornamented
by subdued concentric striae and by oblique striae on the lower part and
over the whole posterior portion of the shell. Young specimens 15 mm. in
length lack the oblique striae. Specimens reach a length of 45 mm. Nuculana
guppyi Dali"'4 from the Miocene of Trinidad bears some similarity to N.
polita but has a more attenuated shell and the beaks are more anterior.

Subgenus Costelloleda Hertlein & Strong, nov.

Type: Nucula costellata Sowerby.

Shell elongate and with strong concentric sculpture. Costelloleda is
somewhat similar in general appearance to Poroleda I redale55 and Thestyleda
Iredale.

Nuculana I Costelloleda I costellata Sowerby.

Plate II, Figure 10.

Nucula costellata Sowerby, Proc. Zool. Soc. London, 1832 (issued
March 13, 1833), p. 198. “Hab. ad Panamam.” “Two specimens were taken in

sandy mud at a depth of ten fathoms.” Sowerby, Conch. Illustr., Nuculae,

1833, p. 4, pi. 15, fig. 8. “Panama.” Muller, Syn. Test. Viv., 1836, p. 191. “Legit

H. Cuming ad Panamam.” Reeve, Conch. Syst., Vol. 1, 1841, p. 110, pi.

85, fig. 8. Hanley, Cat. Rec. Bivalve Shells, p. 169, 1843, p. 376, 71856,

Suppl. pi. 19, fig. 54, 1846. “Panama.”

Leda costellata Sowerby, Hanley, Thes. Conch., Vol. 3, 1860, p. Ill, pi. 228
D-1,-4). 25-29 fathoms, mud, crushed shell and sand bottom; Cape San Lucas,

September, 1871, sp. 24, pi. 4, fig. 24. “Hab. Panama.” Zetek, Rev. Nueva,

Nos. 1 & 2, 1918, p. 37. Panama. Strong, Hanna & Hertlein, Proc. Calif.

Acad. Sci., Ser. 4, Vol. 21, no. 10, 1933, p. 118. Acapulco, Mexico.

Leda (Leda) costellata Sowerby, Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
p. 375, 1908. Gulf of California to Panama. Sta. 2823, in 26 fathoms.

54 See Leda guppyi Dali, Maury. Bidl. Amer. Paleo., Vol. 10, Bull. No. 42, p. 174 (22), pi. 23
(12), fig. 3, 1925. Manzanilla, Trinidad; Miocene.

5 5 Poroleda Tate, Jour. Roy. Soc. New South Wales, Vol. 27, 1893, p. 186. “Examples:

Scaphula ( ?) elongata, Hutton, Trans. N. Z. Inst., XVII., p. 332, 1885. Poroleda lanceolata, Tate.”

Cotton & Godfrey, Moll. South Australia, Pt. 1. Handbook of Flora and Fauna of South

Australia, issued by South Australian Branch Brit. Sci. Guild. Adelaide, 1938, p. 45. Type:
Poroeda lanceolata Hutton [Tate, 1893, p. 186, pi. 12, fig. 6. Gellibrand River, Australia; Eocene].

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 399

Leda rostellata Sowerby, Pilsbry & Lowe, Proc. Acad. Nat. Sci. Philadelphia,
Vol. 84, 1932, p. 141. Acapulco, Mexico, dredged in 20 fathoms.

Type Locality: Panama, in 10 fathoms, sandy mud.

Range: Santa Inez Bay, Lower California, to Panama.

Collecting Stations: Mexico: Santa Inez Bay, Gulf of California (143-
D-1,-4) . 25-29 fathoms, mud, crushed shell and sand bottom; Cape San
Lucas, Lower California; El Salvador: Meanguera Island, Gulf of Fonseca
(199-D-l) . 16 fathoms, sand, mud and crushed shell bottom; Costa Rica: 14
miles southeast of Judas Point (214-D-1-4). 42-61 fathoms, mud, shell and
rocks on bottom.

Oblong, thin, covered with small concentric acute riblets ; anteriorly
beaked and acuminated, with two dorsal approximated and crenulated ribs
(Hanley) .

Several specimens of Nuculana costellata were collected, usually only
one or two at a locality. Most of these specimens are very large compared
to those which have appeared previously in the literature, but they do not
differ otherwise.

Nuculana flexuosa56, described from the lower Miocene of Florida, was
compared to N. costellata by Heilprin but judging from the figures it does
not appear to be very closely related to the west coast species.

Nuculana ICosfelloledcsl marella Hertlein, Hanna & Strong, sp. nov.

Plate II, Figures 12, 13.

[?] Leda cestrota Dali, Pilsbry & Lowe, Proc. Acad. Nat. Sci. Philadelphia,
Vol. 84, 1932, p. 141. “Dredged at 20 fathoms. Acapulco.”

Not Leda cestrota Dali, Proc. U. S. Nat. Mus., Vol. 12, 1889, p. 255, pi. 13,
fig. 7. “Hab. Station 2145, near Colon (Aspinwall), in 25 fathoms, mud.”

Shell elongate, slender, compressed, thin, narrowly rostrate, white under
a thin light brown epidermis ; beaks low, at about the anterior third, pos-
terior dorsal margin only moderately concave and the anterior slightly con-
vex; sculptured with numerous, fine, close spaced concentric laminae;
lunular depression very narrow, smooth; escutcheon long, narrow, micro-
scopically striated, the valve margins distinctly elevated, bounded on each
valve with two rays separated by a shallow depression running from the
beaks to the end of the rostrum, on which the concentric laminae are stronger
but flattened; anterior end with a broad, shallow depression running from
the beaks to the basal margin; interior polished, hinge with 32 small teeth
on the anterior side and about 50 on the posterior side of the central pit.
The type measures: longitudinal diameter, 32 mm.; vertical diameter, 11.3
mm. ; thickness of the two valves, 5.0 mm.

Holotype, a single specimen, is without definite locality although it
probably came from the Gulf of California because it was collected during
the expedition to the Gulf of California in 1936. About a dozen single valves
and one specimen with both valves were collected at Sta. 221-D-1-5. Lat 7°
54' 15" to 7° 52' 30" N., Long. 82" 02' 45" to 82° 01' 00" W., Gulf of Chiriqui.
Panama, 35-40 fathoms, sandy mud bottom.

This species bears a slight resemblance to Nuculana costellata Sowerby,
but is larger, more rostrate and possesses finer concentric sculpture. The
new species bears some resemblance to Nuculana cestrota Dali, a species de-
scribed from the Caribbean and reported by Pilsbry & Lowe from Acapulco,
Mexico. The shell of N. marella has a greater altitude in proportion to the

56 Leda flexuosa Heilprin, Trans. Wagner Free Inst. Sci., Vol. 1, 1887, p. 119, pi. 16, fig. 66.

From the “Silex-bearing marl (Miocene) of Ballast Point, Hillsboro Bay,” Florida. Dali, U. S.

Nat. Mus., Bull. 90, 1915, p. 117, pi. 23, figs. 8 and 10.

Zoologica: New York Zoological Society

400

[XXV :25

length and is therefore not so slender, and the anterior and posterior dorsal
slopes are straighter than those of the species described by Dali.

Subgenus Thestyleda Iredale.

Thestyleda Iredale, Rec. Australian Mus., Vol. 17, no. 4, September 4, 1929,
pp. 158, 187. “Type Leda ramsayi E. A. Smith.”

Type (by original designation) : Leda ramsayi E. A. Smith. [Sci. Rept.
Voy. Challenger, Zool., Vol. 13, 1885, p. 241, pi. 20, figs. 3, 3a. “off Sydney,
New South Wales, in 950 fathoms; green mud.”]

Shell of medium size, thin, elongate, umbos slightly tumid, acutely
rounded anteriorly, rostrate posteriorly; anterior dorsal slope short, pos-
terior dorsal slope more than twice as long, concave in anterior half then
almost straight; ventral margin convexly rounded anteriorly then con-
tinuing in an almost straight line to the posterior end which is squarely
truncated ; ornamented by well marked regular concentric rugae which turn
upward at almost a right angle upon crossing the lower keel; two radiating
ridges form keels along the rostrum; hinge, consisting of anterior and pos-
terior series of chevron shaped teeth; chondrophore large.

The west American species Nuculana hamata and N. cordyla appear to
be referable to the subgenus Thestyleda.

Nuculana IThestyledal hamata Carpenter.

Plate II, Figure 14.

Leda hamata Carpenter, Rept. Brit. Assoc. Adv. Sci. for 1863 (issued 1864),
pp. 612, 644. “Santa Barbara; Cat. Is., 20-60 fm.; common.” Reprint in Smithson.

Misc. Coll., No. 252, 1872, pp. 98, 130. Carpenter, Proc. Calif. Acad. Sci.,

Vol. 3, February, 1866, p. 210. Coast of California. Cooper, Geol. Surv.

Calif., Geogr. Catalog. Moll., 1867, p. 12. “Santa Barbara 16-28 fms. Catalina I.?”

Sowerby, Conch. Icon., Vol. 18, Laeda, November, 1871, sp. 56, pi. 9, fig. 56.

“Hab. Catalina Islands.” Dali, Proc. U. S. Nat. Mus., Vol. 24, 1902, p. 558,

pi. 40, fig. 9. Near Catalina Island, California, in about 50 fathoms. Arnold,

Mem. Calif. Acad. Sci., Vol. 3, 1903, p. 97, pi. 17, fig. 4. Pliocene at Deadman
Island, San Pedro, California. Pleistocene of San Pedro and San Diego, California.
Recent from Santa Barbara to Santa Catalina Island. 1. S. Oldroyd, Stan-

ford Univ. Publ. Univ. Ser. Geol. Sci., Vol. 1, no. 1, 1924, p. 23, pi. 6, fig. 4;
pi. 37, fig. 5. “Type locality, Santa Barbara, California. Range: Puget Sound to
Panama Bay. In the Pliocene and Pleistocene of California.”

Nuculana hamata Carpenter, Grant & Gale, Mem. San Diego Soc. Nat. Hist.,
Vol. 1, 1931, p. 125, pi. 1, figs. 14 and 15. Earlier records cited. Pleistocene and
Recent.

Not Zealeda hamata. Marwick, Proc. Malacol. Soc. London, Vol. 16, no. 1,

April, 1924, p. 25, figs. 1 and 2 ( p. 26). “ Locality . Awatere Beds (Pliocene),

east shore of Lake Grassmere, Marlborough, New Zealand.”

Type Locality : Santa Barbara, California.

Range : Puget Sound to Panama Bay.

Collecting Station: Mexico: East of Cedros Island (126-D-12). 45
fathoms, crushed shell and mud bottom.

The shell of Nuculana hamata is extended into a long rostrum pos-
teriorly which is squarely truncated at the end. The exterior of the shell is
ornamented by strong concentric sculpture. The sculpture of the subspecies
Nuculana hamata limata Dali57 is very variable.

57 Leda hamata Carpenter variety limata Dali, Proc. U. S. Nat. Mus., Vol. 52, December 27, 1916,

p. 397. “off Santa Rosa Island, California, in 50 fathoms.” 1. S. Oldroyd, Stanford Univ. Publ.

Univ. Ser. Geol. Sci., Vol. 1, 1924, p. 24 (as Leda hamata limata). Type locality cited. Range:
Santa Barbara to San Diego, California.

1940] Hertlein & Strong : Mollusks of Eastern Pacific Expeditions 401

Nuculana ( Thestyleda ) elaborata Prashad58 described from the East
Indies, is a similar species.

Nuculana (Thestyledal cordyla Dali.

Leda ( Leda ) cordyla Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October,
1908, p. 375, pi. 6, figs. 6 and 7. U. S. S. Albatross Station 3354, “Gulf of Panama,
in 322 fathoms, mud, bottom temperature 46°F.” Also at Station 2792, off Manta,
Ecuador, in 401 fathoms.

Type Locality. Gulf of Panama, in 322 fathoms, mud. Type No. 122,915
U. S. Nat. Mus.

Range: Gulf of Panama to Manta, Ecuador, in deep water.

Shell, small, rostrate, with 14 anterior and 19 posterior hinge teeth.
It is somewhat similar to N. hamata.

Subgenus Spinula Dali.

Spinula Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908, p. 377.

“Type, Leda calcar Dali.” Thiele, Handbuch syst. Weichtierkunde, Teil 3,

1934, p. 789. L. (S.) calcar Dali, cited.

Type (by original designation ) : Leda calcar Dali.

Shell rostrate, acute behind, smooth, with a well-developed short amphi-
detic ligament, an internal resilium supported by triangular chondrophores,
a defined lunule and escutcheon ; a long, slender, completely united siphon,
no papal tentacles; pallial sinus obsolete (Dali).

Ledaspina Marwick50 from the Tertiary of New Zealand “differs from
Spinula Dali in being almost equilateral and not drawn out posteriorly.”

Nuculana I Spinula I calcar Dali.

Leda ( Spinula ) calcar Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October,
1908, p. 378, pi. 10, figs. 1, 10. U. S. S. Albatross station 4658, “off the Peruvian
coast, in S. Lat. 8° 30', W. Lon. 85° 36', in 2370 fathoms, green mud, bottom
temperature 35°. 3 F.”

Type Locality: Off the coast of Peru, in Lat. 8° 30' S., Long. 85° 36' W.,
in 2370 fathoms, green mud. Type No. 110,573 U. S. Nat. Mus.

Range: Known only from the type locality.

Shell compressed and acutely rostrate; posterior dorsal margin nearly
straight, posterior end acutely angular, with a wide shallow sulcus below,
setting off the rostrum.

Nuculana ISpinulal calcarella Dali.

Leda ( Spinula ) calcarella Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October,
1908, p. 378. U. S. S. Albatross station 4656, “off the coast of Peru, in S. Lat. 6° 55'
and W. Lon. 83° 34', in 2222 fathoms, green mud, bottom temperature 35°. 2 F.”

Type Locality: Off the coast of Peru, in Lat. 6° 55' S., Long. 83° 34' W.,
in 2,222 fathoms, green mud. Type No. 110,575 U. S. Nat. Mus.

Range: Known only from the type locality.

Shell smaller than Nuculana calcar and near the base there are six or
eight lines not coincident with the lines of growth.

58 Nuculana (Thestyleda) elaborata Prashad, Siboga Exped., Monogr. 53c, Lamell., 1932, p. 22,
pi. 1, figs. 32, 33, 34. “Holotype and an odd valve brought up in a deep-sea trawl at St. 52 (9° 3' 4
S., 119° 56' .7 E., 959 metres).”

59 Marwick, J., Geol. Surv. New Zealand, Palae Bull. No. 13. 1931. p. 53. “Type (original in-
troduction) : Ledaspina stimulea n. sp. Oligocene, New Zealand,” p. 53, pi. 2, figs. 21, 24. From
Loc. 1240 “Ihungia mudstone. Island Creek, Tutamoe, S. D.” Oligocene.

402

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[XXV :25

Subgenus Jupiteria Bellardi.

Jupiteria Bellardi, Mon. Nuculidi Terr. Terz. Piemonte e Liguria, 1875, p. 20.

- — -Dali, Trans. Wagner Free Inst. Sci., Vol. 3, pt. 4, April, 1898, p. 579. “Type

L. concava Bronn.” Sacco, Moll. Terr. Terz. Piemonte e Liguria, Pt. 26,

December, 1898, p. 56. “(Tipo J. concava (Brn.)).” Woodring, Carnegie

Inst. Washington, Publ. 366, 1925, p. 18. Type: Nucula concava Bronn.

Type (designated by Dali) : Leda concava Bronn. [= Nucula concava
Bronn, Ital. Tert. Gebilde, 1831, p. 110. Italy. From “Ta. c.” [=“ T abbiano ,
links zwischen Cq. and Parma” in the “Jungerer Grobkalk, blau, thonig, in

den Apenninen.”] Sacco, Moll. Terr. Terz. Piemonte e Liguria, Pt. 26,

December, 1898, p. 56, pi. 12, figs. 1, 2, 3. Several localities in the Pliocene
of Italy],

Small corbuloid, strongly inflated shells; blunt and barely ridged ros-
trum; shallow sinus.

A few of the tropical west American species have been placed under
Jupiteria by Dali but some of those are now believed to be referable to
Saccella. It is probable however that some of the west American species
may be referable to Jupiteria.

Jupiteria occurs in the Miocene and Pliocene of the Mediterranean
region, in the Midway, early Tertiary of southeastern United States, and
from Miocene to Recent in the Caribbean region. It has been recorded
from the upper Eocene of California60. Marwick & Finlay61 cited Jupiteria
from the Danian, upper Cretaceous of New Zealand, Marwick02 cited it from
the Medial Tertiary, and Powell63 described a species which he referred to
Jupiteria, from the waters of New Zealand.

An Australian species, “Leda fortis 6i,” was believed by Hedley to be
referable to Jupiteria but Iredale considered the species to belong to his
genus Teretileda.

The subgeneric position of the following species of Nuculana is uncer-
tain.

Nueulana acrita Dali.

Leda (Jupiteria) acrita Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October,
1908, p. 374. (J. S. S. Albatross Station 2799, “Panama Bay, in 29% fathoms,
mud.” Also other localities in Panama Bay, in 14-62 fathoms.

Type Locality: Panama Bay, in 29% fathoms, mud. Type No. 110,690
U. S. Nat. Mus.

Range: Panama Bay.

Shell small, white, subequilateral, inflated and acutely rostrate pos-
teriorly; hinge with about 15 anterior and 13 posterior teeth. Length, 6.2
mm.; height, 3.8 mm.; diameter, 3.0 mm.

Nuculana agapea Dali.

Leda (Jupiteria) agapea Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October,
1908, p. 373, pi. 6, figs. 4 and 5. U. S. S. Albatross station 3360, “Gulf of Panama,
in 1672 fathoms, sand, bottom temperature 42°F.” Also off the coast of Ecuador
in 1573 fathoms.

60 Jupiteria ( Ledina ) duttonae Vokes, Ann. New York Acad. Sci., Vol. 38, January 4, 1939,
p. 44, pi. 1, fig. 6. From Loc. 1817 (Univ. Calif.), S. W. % of N. W. ^4 of Sec. 15, T. 18
S., R. 14 E., 100 feet up the fourth small draw from west end of ridge; immediately opposite the
place where Urruttia Canyon enters Salt Creek. Domengine formation. Eocene.

61 Finlay, H. J., and Marwick, J., Geol. Surv. New Zealayid, Palaeo. Bull. No. 15, 1937, p. 16.

62 Marwick, J., Geol. Surv. New Zealand , Palaeo. Bull No. 13, 1931, pp. 51-53.

63 Nuculana ( Jupiteria ) manawatawhia Powell, Discovery Repts., Vol. 15, 1937, p. 164, pi. 45,
fig. 9. “ Habitat : Off Three Kings Islands” New Zealand.

64 Leda fortis Hedley, Rec. Australian Mus., Vol. 6, no. 5, July 18, 1907, p. 362, pi. 66, figs. 2
and 3. Dredged from 800 fathoms, 35 miles east of Sydney, Australia, in the Tasman Sea.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 403

Type Locality. Gulf of Panama, in 1,672 fathoms, sand. Type No.
122,911 U. S. Nat. Mus.

Range : Gulf of Panama to coast of Ecuador, in deep water.

This species is most nearly related to L. pontonia Dali, but has the beaks
more anterior and the sculpture coarser and more deeply incised (Dali).

Nuculana bicostata Sowerby.

Laeda bicostata Sowerby, Conch. Icon., Vol. 18, Laeda, November, 1871, sp.
37, pi. 6, fig. 37. “Hab. Panama.”

Type Locality : Panama.

Range: Known only from the type locality.

Shell arched, semi-ovate, compressed, covered with a pale epidermis,
sculptured with nearly concentric ridges undulated at the back and beaded
on the ribs-; anterior side roundly sub-acuminated with ventral margin
roundly convex; posterior side rayed with two crenulated arched ribs,
dorsal margin depressed, concave, acute at the end; umbos rounded, ele-
vated (Reeve).

The arched form and distinct elevated and curved posterior ribs seem
to distinguish this shell from the varieties of Laeda elenensis. (Reeve).

Nuculana crispa Hinds.

Nucula crispa Hinds, Proc. Zool. Soc. London, December, 1843, p. 100. “Hab.

Gulf of Nicoya; from thirty-six fathoms.” Hinds, Zool. Voy. Sulphur, Moll.,

Pt. 3, 1844 [January, 1845 on cover], p. 64, pi. 18, fig. 14. Original locality cited.

Leda crispa Hinds, Hanley, Thes. Conch., Vol. 3, 1860, p. 134, pi. 229 ( Nuculi -

dae, pi. 4), figs. 107, 108. Gulf of Nicoya. Sowerby, Conch. Icon., Vol. 18,

Laeda, November, 1871, sp. 55, pi. 9, fig. 55. “Hab. Gulf of Nocoyo.”

Type Locality: Gulf of Nicoya, in 36 fathoms.

Range : Known only from the type locality.

The original illustration of this species shows a small rostrate form
with strong concentric sculpture. There is a strong ridge on the rostrum and
anterior to this is a narrowly elongate triangular depression separating the
ridge from the remainder of the shell.

Nuculana cuneata Sowerby.

Plate I, Figures 20 and 21.

Nucula cuneata Sowerby, Proc. Zool. Soc. London, 1832 (issued March 13,
1833), p. 198. “Hab. ad Valparaiso.” “Dredged in coarse sand and gravel, at

various depths, from fourteen to forty-five fathoms.” — Sowerby, Conch.

Illustr., Nuculae, 1833, p. 4, pi. 15, fig. 15. Valparaiso. New Guinea. Reeve,

Conch. Syst., Vol. 1, 1841, p. Ill, pi. 85, fig. 15. Philippi, Reise durch die

Wueste Atacama, 1860, pp. 176-177. Valparaiso; Paposo, Chile.

Leda cuneata Sowerby, Hupe, Hist, de Chile, Zool., Vol. 8, 1854, p. 307. “Habita

Valparaiso y otras partas de la Republica.” Hanley, Thes. Conch., Vol. 3,

1860, p. 128, pi. 228 ( Nuculidae , pi. 3), figs. 92, 93. Sowerby, Conch. Icon.,

Vol. 18, Laeda, September, 1871, sp. 35, pi. 6, figs. 35a, 35b. “Hab. Valparaiso,
New Guinea.”

Type Locality: Valparaiso, Chile, in from 14 to 45 fathoms, sand and
gravel.

Range: Known with certainty only from the type locality.

404

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[XXV :25

Shell small, rostrate-ovate, with an ashy or greenish-olive periostracum,
without conspicuous indented ray, and with concentric costellated sculpture.

Nuculana cuneata has been recorded from New Guinea but that record
is believed to have been based on some similar but different species. Accord-
ing to Hanley65, “Leda inornata” A. Adams06, which was described from
New Guinea, is identical with Nuculana cuneata. Dali (1909), however,
considered N. cuneata to be identical with N. acuta.

Nuculana excavata Hinds.

Nucula excavata Hinds, Proc. Zool. Soc. London, December, 1843, p. 100. “Hab.

Panama; dredged among mud in thirty fathoms.” Hinds, Zool. Voy. Sulphur,

Moll. Pt. 3, 1844 [January, 1845 on cover], p. 64, pi. 18, fig. 17. Original locality
cited.

Leda excavata Hinds, Hanley, Thes. Conch., Vol. 3, Nuculidae, 1860, p. 135, pi.

229 ( Nuculidae , pi. 4), figs. 104, 105. Panama. Sowerby, Conch. Icon., Vol.

18, Laeda, September, 1871, sp. 38, pi. 6, figs. 38a, 38b. “Hab. Panama.”

Type Locality : Panama, in 30 fathoms, mud.

Range : Known only from the type locality.

The original illustration of this species shows an ovate-rostrate shell
with a rostral ridge, anterior ray, and concentric sculpture.

Nuculana hindsii Hanley.

Leda hindsii Hanley, Proc. Zool. Soc. London, 1860, p. 440. “Hab. Nicoyae fre-
tum?” “Mr. Hinds referred this shell to his Nucula crispa; it is very distinct,

however, from the delineated type.” Hanley, Thes. Conch., Vol. 3, Nuculidae,

1860, p. 135, pi. 229 ( Nuculidae , pi. 4), fig. 102. Gulf of Nicoya? Sowerby,

Conch. Icon., Vol. 18, Laeda, November, 1871, sp. 60, pi. 9, fig. 60. “Hab. Gulf of
Nicoya?”

Type Locality: ? Gulf of Nicoya, Costa Rica.

Range : Known only from the type locality.

A small, pointed, ovate-oblong shell with a conspicuous anterior in-
dented ray and fold, and with concentric sculpture.

Nuculana lobula Dali.

Leda ( Jupiteria ) lobula Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October,
1908, p. 375. U. S. S. Albatross station 3422, “in 141 fathoms, mud, off Acapulco,
Mexico, bottom temperature 53°. 5.”

Type Locality: Off Acapulco, Mexico, in 141 fathoms, mud.

Range: Known only from the type locality.

Shell small, olivaceous, oval with regular rounded concentric threads;
hinge with about 13 anterior and 9 posterior teeth.

Nuculana loshka Dali.

Lecla (Leda) loshka Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 376, pi. 17, fig. 2. U. S. S. Albatross station 3392, “Gulf of Panama, in 1270
fathoms, hard bottom, temperature 36°. 4 F.”

Type Locality: Gulf of Panama, in 1,270 fathoms, hard bottom. Type
No. 122,916 U. S. Nat. Mus.

65 Hanley, S., Thes. Conch., Vol. 3, Nuculidae, 1860, explanation to figs. 92 and 93, pi. 228
(Nuculidae, pi. 3).

66 Leda inornata A. Adams, Proc. Zool. Soc. London, June 16, 1856, p. 48. “Hab. New Guinea.
Mus. Cuming.”

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 405

Range : Known only from the type locality.

Shell thin, very inequilateral and posteriorly produced in a long rostrum;
disk smooth but under a lens showing fine concentric and a few irregular
faint radial striations.

Nuculana lucasana Strong & Hertlein.

Nnculana lucasana Strong & Hertlein, Proc. Calif. Acad. Sci., Ser. 4, Vol. 22,
no. 6, December 31, 1937, p. 160, pi. 34, figs. 9, 12, 13. “Lat. 23° 03' to 23° 06' N.,
Long. 109° 36' to 109° 31' W., dredged about 10 miles due east of San Jose del
Cabo, Lower California, Mexico, in 20 to 220 fathoms.”

Type Locality. East of San Jose del Cabo, Lower California, Mexico, in
20 to 220 fathoms. Type No. 6966 Calif. Acad. Sci. Paleo. Type Coll.

Range : Known only from the type locality.

Shell differing from Nucidana lobula Dali in the greater diameter in
proportion to the height, and in the presence of an anterior radial de-
pression, a feature not mentioned in the original description of Dab’s
species.

Nuculana ornata d’Orbigny.

Leda ornata d’Orbigny, Voy. Amer. Merid., Vol. 5, 1846, p. 546, pi. 82, figs.

4-6. “rencontree dans du sable de fond, recueilli a Payta (Perou).” — Hanley,

Thes. Conch., Vol. 3, Nuculidae, p. 128, pi. 228 ( Nuculidae , pi. 3), figs. 97, 98.

Payta, Peru. Sowerby, Conch. Icon., Vol. 18, Laeda, September, 1871, sp. 36,

pi. 6, figs. 36a, 36b, “Hab. Payta, Peru.” Wolf, Geogr. & Geol. Ecuador

(Leipzig), 1892, p. 631. Pliocene of Uimbi, Ecuador. Recent in region about
Paita, Peru. Dali, Proc. U. S. Nat. Mus., Vol. 37, 1909, p. 251. Paita, Peru.

Type Locality : Paita, Peru, sand.

Range : Known only from the type locality.

Shell ovate-elongate, rostrate, subventricose, with fine even concentric
sculpture and a strong indented anterior ray.

Nuculana oxia Dali.

Leda oxia Dali, Proc. U. S. Nat. Mus., Vol. 52, December 27, 1916, p. 395.
“Station 2901, off Santa Rosa Island, California, in 48 fathoms, muddy bottom.”

Dali, U. S. Nat. Mus., Bull. 112, 1921, p. 11. Santa Rosa Island, California,

to the Gulf of California. 1. S. Oldroyd, Stanford Univ. Publ. Univ. Ser.

Geol. Sci., Vol. 1, 1924, p. 18. Type locality and range same as cited by Dali.

— E. K. Jordan, Proc. Calif. Acad. Sci., Ser. 4, Vol. 15, no. 7, 1926, p. 244.

San Quintin, Lower California; Pleistocene.

Nuculana oxia Dali, Grant & Gale, Mem. San Diego Soc. Nat. Hist., Vol. 1,

1931, p. 125. Earlier records cited. Keen, Check List West North Amer.

Mar. Moll. (Stanford Univ. Press), 1937, p. 23. Lat. 23°-34°N.

Type Locality: Off Santa Rosa Island, California, in 48 fathoms, mud.
Type No. 214,448 U. S. Nat. Mus.

Range: Santa Rosa Island, California, to the Gulf of California.

Shell minute, very acute posteriorly, beaks subcentral; sculpture of
regular equal concentric ridges with subequal interspaces; a depressed ray
from beak to base anteriorly; hinge with about eight teeth on either side
of the resilifer.

Nuculana peruviana Dali.

Leda ( Leda ) peruviana Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October,
1908, p. 377. “dredged in 1036 fathoms, mud, off Aguja Point, Peru, at station
4654.”

406

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[XXV :25

Type Locality: Off Aguja Point, Peru, in 1,036 fathoms, mud.

Range: Known only from the type locality.

Shell large, slender, rostrate; surface mostly smooth but with a few
coarse irregular wrinkles on basal half of disk anteriorly; hinge with about
15 anterior and 38 posterior teeth.

Nuculana pontonia Dali.

Leda pontonia Dali, Proc. U. S. Nat. Mus., Vol. 12, 1889, p. 257, pi. 13, figs. 5,
5b. “Stations 2807, and 2808, in 812 and 634 fathoms, mud and sand, near the

Galapagos Islands, Pacific Ocean; temperatures 38°. 4 and 40°F.” Dali,

U. S. Nat. Mns., Bull. 112, 1921, p. 12. Santa Barbara Islands, California, to Peru

and the Galapagos Islands, in deep water. 1. S. Oldroyd, Stanford Univ.

Publ. Univ. Ser. Geol. Sci., Vol. 1, 1924, p. 20, pi. 1, figs. 4 and 5. Locality and

range same as cited by Dali. Heath, Mem. Mus. Roy. d'Hist. Nat. Belg.,

Ser. 2, Fasc. 10, 1937, pp. 5, 17, 20. Off San Diego, California, in 822 fathoms.

Leda ( Jupiteria ) pontonia Dali, Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 374. Off Galapagos Islands; Gulf of Panama; off San Diego,
California; in 634 to 1672 fathoms.

Type Locality: Near the Galapagos Islands, in 634-812 fathoms, mud
and sand.

Range: Santa Barbara Islands, California, to Peru, and the Galapagos
Islands, in deep water.

Shell stout, strong, inflated with recurved pointed, posterior end; con-
centric sculpture of fine regular threads which become obsolete on the
umbos; hinge with 16 anterior and 13 posterior teeth.

Nuculana rhytida Dali.

Leda {Leda) rhytida Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October,
1908, p. 376. U. S. S. Albatross station 3422, “off Acapulco, Mexico, in 141 fathoms,
mud, bottom temperature 53°. 5 F.”

Type Locality: Off Acapulco, Mexico, in 141 fathoms, mud. Type No.
122,918 U. S. Nat. Mus.

Range: Known only from the type locality.

Shell small, elongate-ovate, translucent white, with olive-gray periostra-
cum; posteriorly attenuated but not acute; sculpture of concentric riblets
which become obsolete on upper posterior portion of shell; hinge with 11
anterior and nine posterior teeth.

Genus Adrana H. & A. Adams.

Adrana H. & A. Adams, Gen. Rec. Shells, Vol. 2, 1858, p. 547. — Stoliczka,

Mem. Geol. Surv. India. Palaeont. Indica. Cret. Fauna South India, Vol. 3, 1871,
p. 320. “Type Nuc. [ Adrana ] lanceolata, Lam.”

Type (designated by Stoliczka) : Nucula lanceolata Lamarck. [Anim. s.
Vert., Vol. 6, 1819, p. 58. No locality cited. According to d’Orbigny (Voy.
Amer. Merid., Vol. 5, 1846, p. 545), “Le veritable N. lanceolata de Lamarck
est propre a la Nouvelle-Hohand.” According to Hanley (Thes. Conch., Vol.
3, 1860, explanation to pi. 227 ( Nuculidae , pi. 2, fig. 35) it is Leda taylori
from Guatemala. Chenu (Man. de Conchyl., Vol. 1, 1862, p. 180, fig. 904)
gave an illustration of a shell cited as “A. lanceolata, Lamarck,” but no
locality was cited. The figure given by Chenu and the same figure given by
Tryon but cited as Leda ( Adrana ) sowerbyana (Struct, and Syst. Conch.,
Vol. 3, 1884, pp. 249, 443, pi. 126, fig. 32. “Xipixapi”) somewhat resembles

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 407

that of Adrana electa A. Adams (. Proc . Zool. Soc. London, 1856, p. 48 (as

Leda electa ) “Hab. Santos, Brazil ( Capt . Martin). Mus. Cuming.” — •

Hanley, Thes. Conch., Vol. 3, 1860, p. 109, pi. 227 ( Nuculidae , pi. 2), figs.

40, 41 (as Leda electa) . Santos, Brazil. Sowerby, Conch. Icon.,

Vol. 18, Laeda, February, 1871, sp. 2, pi. 1, figs. 2a, 2b. “Hab. Santos, Brazil”),
and so also does that of G. B. Sowerby cited as Nucula lanceolata Lamarck
(Gen. Shells, pi. 102, fig. 1, which Hanley referred to Leda electa), and that
of G. B. Sowerby II cited as Laeda sowerbyana (Conch. Icon., Vol. 18,
Laeda, pi. 1, fig. la)].

The shells referred to this genus are very elongate, lanceolate, and
often have nearly straight doi’sal margins, the beaks scarcely projecting;
the sculpture is fine and composed of concentric or somewhat oblique riblets.

Nine species of Adrana have been described from tropical west Ameri-
can waters. Three of these, Adrana elongata Sowerby, A. sowerbyana
d’ Orbigny and A. suprema Pilsbry & Olsson, have large shells with a nearly
straight dorsal border. The others are smaller and the dorsal border slopes
somewhat in each direction from the beak. No species of Adrana has been
described from the Tertiary of western North America but several have
been recorded from Eocene67 to Recent in the southeastern part of the
United States and from Miocene88 to Recent in the Caribbean region. The
genus has been recorded from the Oligocene of Peru. At the present time
the genus Adrana appears to be entirely confined to the New World. One
Recent species “Leda metcalfii,” cited from the Philippine Islands by Hanley
and by Sowerby was originally described without information as to locality.
We have not noticed it cited in the literature as present in recent collections
from the Philippine Islands.

Key to the Species op Aclrana.

A. Anterior and posterior dorsal margins forming a nearly straight line

a. Umbos a little anterior to the center

b. Ventral margin scarcely concave toward ends elongata

bb. Ventral margin distinctly concave toward ends producing more

attenuate terminations sowerbyana

aa. Umbos central suprema

B. Anterior and posterior dorsal margins forming an obtuse angle at the

umbos

a. Beaks nearly central

b. Branching riblets on anterior dorsal area tonosiana

bb. Anterior dorsal area without branching riblets arcuata

aa. Beaks slightly anterior to center

c. Surface fully sculptured

d. Anterior ventral margin narrowly constricted taylori

dd. Anterior ventral margin not constricted crenifera

cc. Posterior dorsal area smooth

e. Concentric sculpture of coarse riblets exoptata

ee. Concentric sculpture fine penascoensis

07 Adrana aldrichiana Harris, Proc. Acad. Nat. Sci. Philadelphia, Vol. 47, April 9, 1895,
p. 47, pi. 1, fig. 6 (under the genus Leda). "Brazos River, Tex., one and one-half miles below

Mosley’s Perry.” "Lower Claiborne. Eocene.” Harris, Bull. Amer. Paleo., Vol. 6, No. 31,

June 30, 1919, p. 71, pi. 25, figs. 20, 21.

68 See Olsson, A. A., Bull. Amer. Paleo., Vol. 9, Bull. No. 39, 1922, pp. 346-347 (174-175).

Maury, Serv. Geol. e Miner. Brazil, Monogr. 4, 1924, p. 405. Mansfield, Florida State

Geol. Surv., Bull. No. 8, 1932, p. 36.

408

[XXV :25

Zoologica: New York Zoological Society

Adrana arcuata Sowerby.

Laeda arcuata Sowerby, Conch. Icon., Vol. 18, Laeda, September, 1871, sp. 20,
pi. 4, fig. 20. “Hab ?”

Nuculana arcuata Sowerby, Pilsbry & Olsson, Nautilus, Vol. 48, no. 4, 1935,
p. 118, pi. 6, figs. 2 and 3. Point Ancon, Santa Elena, Ecuador. Also Ecuadorian
and Peruvian coasts.

Type Locality : Not originally cited. Point Ancon, Santa Elena, Ecuador,
here designated as type locality.

Range: Coasts of Ecuador and Peru.

Shell elongate with arcuate ventral margin ; ornamentation of regular
thread-like ribs over the entire surface.

Adrana crenifera Sowerby.

Nucula crenifera Sowerby, Proc. Zool. Soc. London, 1832 (issued March 13,
1833), p. 19 7. “Hab. ad Xipixapi.” “It was dredged in sandy mud, at a depth of

nine fathoms.” Sowerby, Conch. Illustr., Nuculae, 1833, pp. 3, 6, pi. 14,

fig. 3. “Xipixapi, Mr. Cuming.” Reeve, Conch. Syst., Vol. 1, 1841, p. 110, pi.

84, fig. 1. Hanley, Cat. Rec. Biv. Shells, p. 167, 1843, p. 376, 1856?, Suppl.

19, fig. 51, 1846. Xipixapi.

Leda crenifera Sowerby, Hanley, Thes. Conch., Vol. 3, Leda, 1860, p. 110, pi.

227 ( Nuculidae , pi. 2), figs. 37 and 38. Xipixapi. Sowerby, Conch. Icon.,

Vol. 18, Laeda, September, 1871, sp. 21, pi. 4, figs. 21a, 21b. “Hab. Xipixapi.”

Yoldia ( Adrana ) crenifera Sowerby, Dali, Proc. U. S. Nat. Mus., Vol. 37,
1909, p. 251. Jipijapa, Ecuador.

Type Locality: Xipixapi, Ecuador, in 9 fathoms, sandy mud.

Range: Known only from the type locality.

Transversely elongated, lanceolate, smooth, with very fine transverse
striae, the dorsal margin carinated, the keel crenulated (Hanley).

Adrana janeiroensis E. A. Smith69 from off Rio de Janeiro, Brazil, bears
some resemblance to A. crenifera. A species of Adrana somewhat similar
to A. crenifera has been mentioned by Olsson70 as occurring in the Oligocene
of Peru, and a species compared to A. crenifera was cited by Wolf71 from
the Pliocene of Ecuador. “Nucula dorbignyi” described by Philippi72 from
the Tertiary of Chile was compared to Adrana crenifera.

The species “Leda metcalfi” Hanley is very similar if not identical
with A. crenifera. It was described by Hanley73 without locality information.
Later, Hanley74 cited it from the Philippine Islands as did Sowerby75. Except
for citation of the species in the paper by Faustino76 we have not noticed
a record of it from the Philippine Islands in any recent papers on the fauna
of that region.

69 Adrana janeiroensis E. A. Smith, Brit. Mus. (Nat. Hist.) Brit. Ant. (“Terra Nova”)
Exped., 1910. Nat. Hist. Kept. Zool., Vol. 2, no. 4, Moll. pt. 1, p. 98, pi. 2, fig. 16, 1913: “Off Kio
de Janeiro in 40 fathoms. Lat. 22° 56' S., Long. 41° 34' W.”

70 Olsson, A. A., Bull. Amer. Paleo., Vol. 17, no. 63, 1931, p. 134 (38).

71 Wolf, T., Geogr. & Geol. Ecuador, (Leipzig), 1892, p. 631. Pliocene of Uimbi, Ecuador.

72 Nucula dorbignyi Philippi, Los Fos. Terc. i. Cuart. Chile, 1887, p. 188, pi. 41, fig. 10. “Es
de Levu i esta sobre un gran pedazo de roca dura.”

73 Hanley, S., Proc. Zool. Soc. London, 1860, p. 370. l,Hab. ? ’

74 Hanley, S., Thes. Conch., Vol. 3, 1860, expl. to pi. 227 ( Nuculidae , pi. 2), fig. 34.

75 Sowerby, II, G. B., Conch. Icon., Vol. 18, Laeda, September, 1871, sp. 18, pi. 4, fig. 18.

76 Faustino, L. A., Bur. Sci. Philippine Islands, Monogr. 25, October 30, 1928, p. 18.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 409

Adrana elongata Sowerby.

Plate II, Figure 16.

Nucula elongata Sowerby, Proc. Zool. Soc. London, 1832 (issued March 13,
1833), p. 197. “Hab. in Columbia Occidental!. ’' “Dredged in sandy mud, at a depth

of twelve fathoms, at Xipixapi.” Sowerby, Conch. Illustr., Nuculae, 1833,

pp. 3, 6, pi. 14, fig. 2. “Salango, Mr. Cuming.” Muller, Synop. Test. Viv.,

1836, p. 190. Original record cited. Reeve, Conch. Syst., Vol. 1, 1841, p. 110,

pi. 84, fig. 2. Catlow & Reeve, Conch. Nomencl., 1845, p. 53. — Hanley,

Cat. Rec. Bivalve Shells, p. 167, 1843, p. 376, ?1856, Suppl. pi. 19, fig. 50, 1846.

[Not accurate figure.]. West Colombia. d’Orbigny, Voy. Amer. Merid.,

Vol. 5, 1846, 545. Original record cited.

Leda elongata Sowerby, Hanley, Thes. Conch., Vol. 3, 1860, p. 106, pi. 227

( Nuculidae , pi. 2), fig. 39. “Salango, W. Columbia.” Sowerby, Conch. Icon.,

Vol. 18, Laeda, 1871, sp. 4, pi. 1, fig. 4. "Hab. Salango, West Columbia.”

Not Leda elongata Treat, Ann. de Paleo., Vol. 22, fasc. 2, 1933, p. 52, pi. 6, fig.
3. “Environs d’Ankitohazo ; niveau inferieur.” Madagascar. Permo-Triassic.

Yoldia ( Adrana ) elongata Sowerby, Dali, Proc. U. S. Nat. Mus., Vol. 37, 1909,

p. 251. Coast of Ecuador. Zetek, Rev. Nxieva, Nos. 1 & 2, 1918, p. 51. Zoo-

geographical province of Panama.

Nucidana ( Adrana ) soiverbyana d’Orbigny, Strong, Hanna & Hertlein, Proc.
Calif. Acad. Sci., Ser. 4, Vol. 21, no. 10, 1933, p. 118. Acapulco, Mexico.

Not Nacida ( Adrana ) sowerbyana d’Orbigny.

Type Locality: Xipixapi, Ecuador, in 12 fathoms, sandy mud.

Range: Acapulco, Mexico, to Ecuador.

Collecting Stations: Guatemala: 7 miles west of Champerico (197-D-l-
2). 14 fathoms, mud bottom; El Salvador: La Libertad (198-D-2). 14
fathoms, mud bottom.

Specimens from three localities appear referable to Adrana elongata
Sowerby. The application of the specific name elongata to this species is some-
what unfortunate because the species is not as elongate in form as is Adrana
sowerbyana, a similar species. The beaks are more central and the ends
much less tapering on A. elongata than is the case in A. sowerbyana. Another
large species of the genus is Adrana suprema Pilsbry & Olsson with a length
of 107 mm. It differs from A. elongata Sowerby in that the beaks are nearly
central rather than somewhat anterior as in the species described by Sowerby.

Adrana exoptafa Pilsbry & Lowe.

Plate II, Figure 11.

Leda ( Adrana ) exoptata Pilsbry & Lowe, Proc. Acad. Nat. Sci. Philadelphia,
Vol. 84, p. 107, pi. 17, figs. 8 and 9, May 21, 1932. “Mexico: Guaymas, 20 fathoms.”

Type Locality: Guaymas, Mexico, in 20 fathoms. Type No. 155633 Acad.
Nat. Sci. Philadelphia.

Range: Guaymas, Sonora, Mexico, to Tangola-Tangola, Oaxaca, Mexico.

Collecting Stations: Mexico: 4 miles south-southwest of Maldanado
Point (192-D-l). 26 fathoms, mud bottom; Santa Cruz Bay (195-D-21). 18
fathoms, mud bottom; Tangola-Tangola Bay (196-D-17). 4 fathoms, sand
bottom.

The shell of this elongate form possesses a smooth band anterior to
the posterior carina. The concentric sculpture on the anterior part of the
shell becomes obsolete and grades into the smooth area. Some of the speci-
mens in the present collection are slightly upturned at the posterior ex-

410

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[XXV :25

tremity of the shells but others barely show this feature and seem to agree
almost exactly with Adrana exoptata Pilsbry & Lowe. Adrana newcombi
Angas77 described from the east side of the isthmus of Panama appeals to
be a very similar species.

Adrana penascoensis Lowe.

[ l]Yoldia apparently electa A. Adams, Dali, Amer. Jour. Conch., Vol. 7, no. 2,
November 2, 1871, p. 154. From the head of the Gulf of California.

Leda ( Yoldia ) lanceolata Lamarck, Stearns, Proc. U. S. Nat. Mus., Vol. 17
1894, p. 147. Gulf of California.

Leda ( Adrana ) penascoensis Lowe, Trans. San Diego Soc. Nat. Hist., Vol. 8,
no. 6, March 21, 1935, p. 18, pi. 1, fig. 8. “Punta Penasco, Sonora, dredged 10
fathoms.”

Type Locality : Punta Penasco, Sonora, Mexico, in 10 fathoms. Type
No. 11393 Lowe Coll, in San Diego Soc. Nat. Hist.

Range : Gulf of California.

Shell white, with a straw-colored glossy periostracum; strongly com-
pressed beaks much nearer the anterior end. Dorsal line nearly straight,
ventral margin curved, anterior and posterior ends about equally angular.
Dorsal edges of both valves slightly crenate the entire length. Sculpture of
fine concentric lines of growth over the entire surface of both valves, except
a narrow portion bordering the posterior dorsal margin, which is entirely
smooth. Diameter 37.5 mm., altitude 9.4 mm. (Lowe).

Adrana sowerbyana d’Orbigny.

Nucula layiceolata Lamarck, Sowerby, Conch. Illustr., Nuculae, 1833, p. 3,

pi. 14, fig. 1. “Xipixapi.” Deshayes, ed. of Lamarck’s Anim. s. Vert., Vol.

6, 1835, p. 504. [No locality cited], Reeve, Conch. Syst., Vol. 1, 1841,

p. 110, pi. 84, fig. 1. Hanley, Cat. Bivalve Shells, Suppl. pi. 19, fig. 49, 1846.

Not Nucula lanceolata J. Sowerby, Miner. Conch., Vol. 2, 1817, p. 178, pi.
180, fig. 1. “At Bawdsey, Suffolk, in Crag.”

Not Nucula lanceolata Lamarck, Anim. s. Vert., February-June, 1819, p. 58.
“Habite ”

Leda sowerbyana d’Orbigny, Voy. Amer. Merid., Vol. 5, 1846, p. 544. Ref-
erence cited “Nucula lanceolata, Sow., 1831, Gen. of Shells, no. 17, fig. 1 (non
Layiceolata, Lamk., 1819).” “Elle a ete pechee par M. Cuming a la cote de
Xipixapi, republique de l’Equateur.”

[?] Nucula elongata Sowerby, Valenciennes, Voy. Venus, Zook, Moll., 1846,
pi. 23, fig. 4 (two figs.).

Leda soiuerbiana d’Orbigny, Hanley, Thes. Conch., Vol. 3, 1860, p. 107, pi.
227 (Nuculidae, pi. 2), fig. 33. Xipixapi.

Laeda sowerbyana d’Orbigny, Sowerby, Conch. Icon., Vol. 18, Laeda, Novem-
ber, 1871, sp. 1, pi. 1, fig. lb (only). “Hab. Xipixapi.”

Yoldia (Adrana) sowerbyana d’Orbigny, Dali, Proc. U. S. Nat. Mus., Vol.

37, 1909, p. 251. “Jipijapa, Ecuador.” Zetek, Rev. Nueva, Nos. 1 & 2, 1918,

p. 51. Panamanian zoogeographical province.

Type Locality: Xipixapi, Ecuador.

Range : Panama to Ecuador.

A few specimens in the collection appear to be intermediate between
Adrana sowerbyana and A. elongata. The shell of Adrana sowerbyana
d’Orbigny is similar to Adrana elongata Sowerby but is narrower and with
more tapering extremities. Hanley believed that the specimen figured by

77 Leda (Adrana) newcombi Angas, Proc. Zool. Soc. London, March 5, 1878, p. 314, pi. 18, figs.
16, 17. “ Hab . Dredged in Navy Bay, Aspinwall, in 8 fathoms.”

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 411

Valenciennes as “Nucula elongata” Sowerby might probably be referred to
sowerbyana. The figure given by Valenciennes is somewhat less tapered at
the ends than sotverbyana but otherwise it is quite similar. Stearns cited
Adrana sowerbyana (as L. lanceolata ) from the Gulf of California but Lowe
described Adrana penascoensis from that region and Pilsbry & Lowe in dis-
cussing Adrana exoptata stated that A. elongata and A. sotverbyana occur
much further south. A. scaphoides Rehder'8 from eastern Colombia is a
similar species.

The name Adrana sotverbyana presents some complications in regard to
nomenclature. Nucula lanceolata J. Sowerby was originally described as a
fossil from the Pliocene of England in 1817. Lamarck also described under
the same name in 1819, a Recent shell without locality information. G. B.
Sowerby in the Genera of Shells (No. 17, 1825, pi. 102, fig. 1) illustrated a
species under the name of Nucula lanceolata Lamarck but without informa-
tion as to locality, while in 1833 G. B. Sowerby Jr. (Conch. Illustr., Nuculae,
p. 3, pi. 14, fig. 1) illustrated a shell as Nucula lanceolata Lamarck with the
locality as Xipixapi, which is in western Colombia, and also cited a reference
to the figure in the Genera of Shells. d’Orbigny in 1846 noticed that the
name Nucula lanceolata of Lamarck was preoccupied by the same combina-
tion of names used by J. Sowerby, hence d’Orbigny proposed for Lamarck’s
species the new name Leda sowerbyana and stated that it was collected at
Xipixapi by Cuming. This evidently refers to the shell and locality in
Conchological Illustrations although in the synonymy d’Orbigny cited the
reference to the illustration in the Genera of Shells. Hanley in 1860 de-
scribed “Leda” taylori from Guatemala and in the synonymy cited the refer-
ence to Nucula lanceolata of Lamarck and to the figure of Nucula lanceolata
given by Sowerby in the Conchoiogical Illustrations while the illustration of
N. lanceolata in the Genera of Shells is referred by Hanley to Leda electa
A. Adams from Brazil. Sowerby in 1871 referred the illustration, of N.
lanceolata Lamarck in the Genera of Shells to “Leda” sowerbyana.

Although d’Orbigny stated that the true Nucula lanceolata Lamarck
came from New Holland, it does not appear to have been recognized subse-
quently in that region. The figure in the Genera of Shells, which has a dis-
tinctly curved anterior dorsal margin and rounded anterior ventral margin,
appears to be distinct from the one in the Conchological Illustrations which
has an almost straight dorsal margin and is attenuated at both ends. We
therefore follow Hanley in referring this figure (1833) to the west American
sowerbyana while the one in the Genera of Shells (1825) may be referred to
the east American “Leda” electa A. Adams.

“Nucula darwini1 9” described by Philippi from the Tertiary of Chile
was compared to “N. lanceolata Sow.”

Adrana suprema Pilsbry & Olsson.

Nuculana (Adrana) suprema Pilsbry & Olsson, Nautilus, Vol. 48, no. 4, April,
1935, p. 117, pi. 6, fig. 1. “Beach between Port Guanico and the mouth of the
Guanico river, Los Santos province, Panama.”

Type Locality. Beach between Port Guanico and the mouth of the
Guanico river, Los Santos province, Panama.

Range : Known only from the type locality.

Shell resembling Adrana elongata but with the beaks nearly central.

78 Adrana scaphoides Rehder, Nautilus, Vol. 53, no. 1, July, 1939, p. 17, pi. 6, fig. 5. “Near
Cartagena, Colombia.”

79 Nucula darwini Philippi, Los Fos. Terc. i Cuart. Chile, 1887, p. 188, pi. 41, fig. 17, From
same locality as Nucula darwini, that is, “Es de Levu i esta sobre un gran pedazo de roca dura.”

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Adrana taylori Hanley.

Leda taylori Hanley, Proc. Zool. Soc. London, 1860, p. 370. “Hab. Guatemala.”

Hanley, Thes. Conch., Vol. 3, 1860, p. 109, pi. 227 ( Nuculidae , pi. 2), fig.

35. “Guatemala.” Sowerby, Conch. Icon., Vol. 18, Laeda, September, 1871,

sp. 17, pi. 4, fig. 17. “Hab. Guatemala.”

Type Locality : Guatemala.

Range : Known only from the type locality.

Sowerby (1871) in discussing Adrana arcuata stated that “In Leda
taylori, which resembles this species, there is peculiar narrowness and almost
a constriction in the ventral margin at the anterior end not observable in
Laeda arcuata.” Adrana taylori is similar to A. crenifera Sowerby and A.
electa A. Adams80.

Adrana tonosiana Pilsbry & Olsson.

Plate II, Figure 15.

Nuculana ( Adrana ) tonosiana Pilsbry & Olsson, Nautilus, Vol. 48, no. 4,
April, 1935, p. 117, pi. 6, fig. 4. “Estero Bucaru, mouth of the Tonosi River,
Los Santos, Panama.”

Type Locality: Estero Bucaru, Mouth of Tonosi River, Los Santos Pro-
vince, Panama. Type No. 164025 Acad. Nat. Sci. Philadelphia.

Range: Off Mazatlan, Sinaloa, Mexico, to Panama.

Collecting Stations: Guatemala: 7 miles west of Champerico (197-D-
1-2). 14 fathoms, mud bottom; El Salvador: La Libertad (198-D-1-2). 13-14
fathoms, mud bottom; Nicaragua: Corinto (200-D-19). 12-13 fathoms, man-
grove leaves on bottom; Panama: Gulf of Chiriqui (221-D-1-5). 35-40
fathoms, sandy mud bottom.

A large series of specimens dredged off Central America have been
referred to Adrana tonosiana Pilsbry & Olsson. There seems but little dif-
ference between these and young Adrana arcuata Sowerby. According to
Pilsbry & Olsson tonosiana is a smaller form, their specimens not exceeding
26 mm. Some of the specimens in the present collection attain a length of
33 mm. We have assigned these to tonosiana chiefly due to the strong
and sometimes branching ribbing on the anterior dorsal margin. This was
considered by Pilsbry & Olsson to be a characteristic feature of tonosiana
and not present on A. arcuata Sowerby. If this is constantly absent on A.
arcuata it would seem best to leave tonosiana as a valid species, otherwise
they are extremely similar. A few irregular radial striae are present on the
exterior of the shells in the present collection but these were not mentioned
in the description of Pilsbry & Olsson. A single somewhat worn right valve
from Sta. 221-D-1-5 has also been assigned to A. tonosiana because of the
branching ribbing on the anterior dorsal edge. The shell is larger than the
type of A. tonosiana and has a more pointed posterior end.

Adrana crenifera Sowerby is somewhat similar in form but the dorsal
margin has a greater slope from the beak and the edge of the posterior
ridge is said to be angular and decidedly crenulated.

Genus Yoldia Moller.

Yoldia Moller, Naturhist. Tidsskrift, IV (1), 1842, p. 91, also Index Moll.
Groenlandiae, 1842, p. 18. Species cited: “Y. arctica, Nucula arctica Gray” and

“Y. angularis nob., Nuc. myalis Couth.?” S. A. Miller, North Amer. Geol.

and Palaeo., (Cincinnati, Ohio), 1889, p. 516. “Type Y. myalis,” fig. 935.

80 Leda electa A. Adams, See Hanley, Thes. Conch., Vol. 3, 1860, p. 109, pi. 227 ( Nuculidae , pi.

2), figs. 40 and 41. “Santos, Brazil.” Sowerby, Conch. Icon., Vol. 18, Laeda, February, 1871,

sp. 2, pi. 1, figs. 2a, 2b. "Hab. Santos Brazil.”

1940] Hertlein & Strong : Mollusks of Eastern Pacific Expeditions

413

Verrill & Bush, Proc. U. S. Nat. Mus., Vol. 20, June 15, 1898, p. 858.

“Type. Yoldia hyperborea Torrell= Yoldia arctica Moller (not Gray).”

Cossmann, Act. Soc. Linn. Bordeaux, Vol. 66 (Conch. Neog. Aquit., Vol. 2,

Livr. 1), 1912, p. 113. Genotype: L. hyperborea Loven. Gardner, Geol. Surv.

Maryland. Upper Cretaceous, 1916, p. 518. “Type. Yolida arctica Gray.”

Woodring, Carnegie Inst. Washington, Publ. 366, 1925, p. 21. Type: “Nucula

arctica Moller not Gray ( = Nucula myalis Couthouy).” Stewart, Acad. Nat.

Sci. Philadelphia, Spec. Publ. No. 3, 1930, p. 59. Type designation of Verrill & Bush

accepted. Grant & Gale, Mem. San Diego Soc. Nat. Hist., Vol. 1, 1931, p. 126.

Type : Nucula arctica Gray.

Type (designated by Gardner) : Nucula arctica Gray. [Suppl. to Ap.

Parry’s First Voy., 1819-1820, p. 241, 1824. Also illustrated by

Grant & Gale, Mem. San Diego Soc. Nat. Hist., Vol. 1, 1931, pi. 1, figs. 10, 11
(figures copied from Mossevieh, Acad. Sci. Union Republ. Soviet Socialist.,
Livr. 19, 1928, pi. 1, figs. 1 and 3). Circumboreal] .

Somewhat similar to Nuculana but with thinner, subovate shell, slightly
rostrate, usually gaping posteriorly; exteriorly sculptured by growth-lines or
concentric striae; hinge consisting of two subequal series of small chevron-
shaped teeth; resilium-pit large, symmetrically underlapping both rows of
teeth; pallial sinus deep and wide, the apex broadly U-shaped.

There is considerable disagreement among various authors as to the
designation of the type species of Yoldia. Grant & Gale have given a discus-
sion of the problem. The designation of Yoldia myalis Couthouy by Miiler
in 1889 is not strictly valid due to the fact that in the original list of species
under Yoldia it was cited as “Nuc. myalis Couth.?” and is thus a species
inquirenda and not acceptable under the present rules of nomenclature. The
interpretation of the type of Yoldia as indicated by Grant & Gale appears to
be the correct one, and the type here accepted is Nucula arctica Gray. Yoldia
probably ranges from Eocene to Recent, although it has been recorded from
the Carboniferous. Several species of the genus Yoldia have been described
from the Eocene of Western North America.

Microyoldia Verrill & Bush81, with the type M. regularis Verrill, has not
been recorded from tropical west American waters.

Subgenus Katadesmia Dali.

Katadesmia Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1898, p.

379. Sole species: Yoldia ( Katadesmia ) vincula Dali. Thiele, Handbuch

Syst. Weichtierkunde, Teil 3, 1934, p. 790. Y. ( K .) vincula Dali, cited.

Type (by original designation and by monotypy) : Yoldia ( Katadesmia )
vincula Dali.

Southern species with well-defined functional opisthodetic ligament.

Yoldia l Katadesmia! vincula Dali.

Yoldia ( Katadesmia ) vincula Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 379, pi. 5, fig. 5. U. S. S. Albatross station 3360, “in 1672
fathoms, sand, Gulf of Panama, bottom temperature 42° F.” Also at stations 3354
and 3361, in 322 fathoms, and in 1471 fathoms.

Yoldia vincula Dali, Zetek, Rev. Nueva, Nos. 1 & 2, 1918, p. 38. Panama.

Type Locality: Gulf of Panama, in 1,672 fathoms, sand. Type No.
122,903 U. S. Nat. Mus.

Range: Gulf of Panama.

S'l Microyoldia Verrill & Bush, Amer. Jour. Sci., Ser. 4, Vol. 3, January, 1897, p. 56. “Type M.

regularis (V.),“ figs. 5 and 6 (p. 60). Verrill & Bush, Proc. U. S. Nat. Mus., Vol. 20, 1898,

p. 860. Type: Microyoldia regularis Verrill, p. 860, pi. 78, figs. 5 and 6.

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The species is notable for its pale color and brilliant polish (Dali).
Shell with the general form of Sanguinolaria rosea; hinge with 15 anterior
and about 25 posterior teeth.

Subgenus Megayoldia Verrill & Bush.

Megayoldia Verrill & Bush, Amer. Jour. Sci., Ser. 4, Vol. 3, January, 1897,

p. 55. “Type M. thraciaef ormis (Storer),” fig. 17. Verrill & Bush, Proc.

U. S. Nat. Mus., Vol. 20, i898, p. 859. Type: Megayoldia thraciaef ormis (Storer),

Type (by original designation): Megayoldia thraciaef ormis (Storer).
[=Leda thraciaef ormis Storer, Boston Jour. Nat. Hist., Vol. 2, 1838, p. 122,

text fig. “. . . from the stomach of a Platessa the dentata of Mitchill

called by our fishermen, Sand dab: this fish was caught off

Provincetown, Cape Cod, in about thirty fathoms of water.” 1. S.

Oldroyd, Stanfoid Univ. Publ. Univ. Ser. Geol. Sci., Vol. 1, 1924, p. 27, pi. 5,
fig. 1. Ai'ctic Ocean to Oregon and Puget Sound. Also Atlantic].

Broad, compressed form, with a very blunt, indefinite, postero-dorsal
rostrum, and with a low radial ridge, ending in a postero-ventral marginal
lobe. The chondrophore is remarkably large and strong, concave, striated
within, and projects much within the margin of the hinge-plate. The
pallial sinus is large and deep. Ligament external, strongly developed (from
Verrill & Bush) .

Megayoldia is known in the Oligocene and probably occurs from Eocene
to Recent. Yoldia emersoni Dali82 described from the Eocene of Alaska may
belong to this subgenus. Makiyama83 has discussed the relationships of
several Japanese species belonging to Megayoldia.

Yoldia IMegayoldial martyria Dali.

Yoldia martyria Dali, Nat. Hist. Soc. Brit. Columbia, Bull. No. 2, January, 1897,
p. 9, pi. 2, fig. 15. “Puget Sound, 35-135 fms.; Gulf of California off the Island of
San Pedro Martir, in 65 fms.; U. S. Fish Commission steamer Albatross.”

Dali, Checklist Rec. Biv. Moll. (Southwest Museum), 1916, p. 11. Kasa-an

Bay, Alaska, to Gulf of California. Dali, U. S. Nat. Mus., Bull. 112, 1921,

p. 13. Kasaan Bay, Alaska, to Gulf of California. 1. S. Oldroyd, Stanford

Univ. Publ. Univ. Ser. Geol. Sci. Vol. 1, 1924, p. 29, pi. 39, fig. 11. Type locality and

range same as cited by Dali. Keen, Check List West North Amer. Mar.

Moll. (Stanford Univ. Calif.), 1937, p. 27. Range: Lat. 23°-55° N.

Type Locality: (According to I. S. Oldroyd), off San Pedro Martir
Island, Gulf of California. Type in U. S. Nat. Mus.

Range: Kasaan Bay, Alaska, to the Gulf of California.

Shell olive-greenish, smooth, nearly equilateral; posterior end roundly
pointed, compressed and slightly recurved; hinge with 21 anterior and 17
posterior teeth.

Subgenus Orthoyoldia Verrill & Bush.

Orthoyoldia Verrill & Bush, Amer. Jour. Sci., Ser. 4, Vol. 3, 1897, p. 55.

“Type Orthoyoldia scapina (Dali).” Verrill & Bush, Proc. U. S. Nat. Mus.,

Vol. 20, 1898, p. 859. Type: Orthoyoldia scapina (Dali).

Type (by original designation): Orthoyoldia scapina (Dali). \=Yoldia
scapama Dali, Proc. U. S. Nat. Mus., Vol. 12, 1889, p. 254, pi. 13, fig.

82 Yoldia emersoni Dali, Harriman Alaska Exped., Vol. 4, 1910, p. 104, pi. 9, fig. 8. “Upper
beds, 3373.” Beds near the camp on the eastern side of Chichagof Cove. Upper beds of Stepovak
Series. Eocene.

83 Makiyama, J., Mem. Coll. Sci. Kyoto Imper. Univ., Ser. B, Vol. 10, no. 2, 1934, pp. 130-132.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 415

6. U. S. Fish Commission station 2762, “east of Rio Janeiro, in 59 fathoms,
mud; bottom temperature 57°F.”].

Shell oblong, gaping, blunt or rounded at both ends, without a distinct
rostrum; no carina. Pallial sinus large and broad. Teeth numerous in both
series (Verrill & Bush).

The shells of Orthoyoldia are soleniform or elongate-elliptic in shape.
Orthoyoldia is known from the Miocene84 of the Caribbean region. It is
now living in that region and is represented by at least one species in tropical
west American waters.

Yoldia fOrthoyoldial panamensis Dali.

Yoldia ( Orthoyoldia ) panamensis Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 380. U. S. S. Albatross station 3354, “Gulf of Panama, in 322
fathoms, mud, bottom temperature 46° F.” Also at station 3355, in 182 fathoms.

Yoldia panamensis Dali, Zetek, Rev. Nueva, Nos. 1 & 2, 1918, p. 38. Panama.

Type Locality : Gulf of Panama, in 322 fathoms, mud. Type No. 122,900
U. S. Nat. Mus.

Range : Gulf of Panama.

The shell of this species is small, narrow, elongate, and very inequilater-
al ; resilium small, vertical, triangular and wholly internal ; no visible external
ligament; hinge with 18 anterior and 52[?25]-27 posterior teeth.

Subgenus Yoldiella Verrill & Bush.

Yoldiella Verrill & Bush, Amer. Jour. Sci., Ser. 4, Vol. 3, January, 1897,

p. 55. “Type Yoldiella lucida (Loven),” figs. 3, 4, 11, 14. Verrill & Bush,

Proc. U. S. Nat. Mus., Vol. 20, 1898, p. 861. Type: Yoldiella lucida (Loven).

Type (by original designation): Yoldiella lucida (Loven). [= Yoldia
lucida Loven, K. Vet. Akad. Fork., 1846 (Index Moll. Lit. Scandinaviae occid.
Habit.), p. 34. “Boh.-Finm.” Illustrated as Portlandia lucida Loven, by Sars,
Bid. Kundsk. Norg. Ark. Fauna. I. Moll. Reg. Arct. Norvegiae, 1878, p. 37,
Tab. 4, figs. 8a, 8b. West Coast of Norway in 20-650 fathoms. Also in

Arctic and north Atlantic. — Verrill & Bush, Proc. U. S. Nat. Mus., Vol.

20, 1898, p. 861, pi. 77, fig. 2; pi. 80, fig. 3 (as Yoldiella lucida) ].

This group includes a large number of small, mostly deep-sea species,
with glossy, iridescent, ovate, and usually wedge-shaped shells, nearly always
having a slight postero-ventral sinuosity, which feebly defines an obscure,
blunt, rostral region, without any definite carination. The shells do not gap,
but close tightly, except that at the rostral angle of some species there may
be a slight divergence. The internal cartilage, which is often relatively
large, occupies a simple notch, which interrupts the hinge margin more or
less completely and generally shows externally in a dorsal view; the notch
usually terminates within on the inner or inferior surface of the hinge-plate,
and is often bounded within by a slight ridge. A weak external ligament is
present on the postero-dorsal margin. A relatively small, pallial sinus has
been observed in several of the species, but is usually indistinct. The siphon
tubes, as observed in a few of the species, are slender and united for more
than half their length. (Verrill & Bush).

84 See Yoldia ( Orthoyoldia ) ovalis Gabb, Woodring, Carnegie Inst. Washington, Publ. 366,
1925, p. 22, pi. 1, figs. 20, 21. Bowden, Jamaica; Miocene. Yoldia bocasensis Olsson (Bull. Amer.
Paleo., Vol. 9, Bull. No. 39, 1922, p. 348 (176), pi. 31 (28), figs. 21, 22. “Gatun Stage: Bocas del
Toro”) from the Miocene of Panama, is a similar species.

416

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Zoologica: New York Zoological Society

Yoldia (Yoldiella) eecinella Dali.

Yoldia eecinella Dali, Proc. U. S. Nat. Mus., Vol. 52, December 27, 1916, p.
399. “Station 2823, off La Paz, Gulf of California, in about 26 fathoms.”

Dali, U. S. Nat. Mus., Bull. 112, 1921, p. 14. Aleutian Islands to Gulf of

California. 1. S. Oldroyd, Stanford Univ. Publ. Univ. Ser. Geol. Sci.,

Vol. 1, 1924, p. 34. Type locality and range same as cited by Dali. Keen,

Check List West North Amer. Mar. Moll. (Stanford University Press), 1937,
p. 27. Lat. 23°-55° N.

Type Locality : Off La Paz, Lower California, Mexico, in about 26
fathoms. Type No. 211,424 U. S. Nat. Mus.

Range : Aleutian Islands to the Gulf of California.

Shell minute, polished, smooth, nearly equilateral, the margins arcuate
above and below, the anterior end rounded, the posterior more pointed and
slightly longer; beaks low and inconspicuous, with neither lunule nor
escutcheon indicated; hinge with 6 or 7 minute teeth on each side of the
subumbonal, very small resilifer. Length, 5 mm. ; height, 2.6 mm. ; diameter,
1.5 mm. (Dali).

Yoldia I Yoldiella I chilenica Dali

Yoldia ( Yoldiella ) chilenica Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 380. U. S. S. Albatross station 2781 “on the southern coast of
Chile (Lat. 51° 52' S. ) , in 348 fathoms, mud, bottom temperature 50° F.” Also
at station 2782, in 258 fathoms.

Type Locality : Southern coast of Chile (Lat. 51° 52' S.), in 348 fathoms,
mud. Type No. 96,923 U. S. Nat. Mus.

Range: Southern coast of Chile.

This species has not been illustrated but according to Dali it has some-
what the appearance of young Yoldia thraciaeformis. Hinge with 19 anterior
and 17 posterior teeth.

Yoldia I Yoldiella l dicella Dali.

Yoldia ( Yoldiella ) dicella Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 382. U. S. S. Albatross station 3418, “off Acapulco, Mexico, in
660 fathoms, sand, bottom temperature 39° F.”

Type Locality: Off Acapulco, Mexico, in 680 fathoms, sand. Type No.
122,917 U. S. Nat. Mus.

Range: Known only from the type locality.

This is said to be a “spade-shaped” little species quite distinct from
others of the genus in west American waters. Hinge with about 10 anterior
and 15 posterior teeth.

Yoldia I Yoldiella ) granula Dali.

Yoldia ( Yoldiella ) granula Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 382. U. S. S. Albatross station 2778, “in Magellan Straits, in 61
fathoms, bottom temperature 47°. 9 F.”

Type Locality: Magellan Straits, in 61 fathoms. Type No. 110,693 U. S.
Nat. Mus.

Range: Known only from the type locality.

Shell minute, oval, equilateral, moderately tumid, smooth, covered with
a polished straw-colored periostracum; interior porcellanous, with a very
slight pallial sinus and smooth margins; hinge line with about six teeth on

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 417

either side of a well-developed resilium, the most anterior tooth of all a
little more distant and notably larger than any of the others. Length, 1.7 ;
alt. 1.2; max. diam. 1.0 mm. (Dali).

Yoldia lYoldiella} indolens Dali.

Yoldie[a] ( Yoldiella ) indolens Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1898, p. 381. U. S. S. Albatross station 2784, “on the southern coast of
Chile, in 194 fathoms, mud, bottom temperature 51°. 9 F.” Also at station 2785,
in 122 fathoms.

Type Locality : Southern coast of Chile, in 194 fathoms, mud. Type No.
122,740 U. S. Nat. Mus.

Range : Southern coast of Chile.

Apparently no illustration has been given of this species. According to
Dali it resembles Yoldia chilenica but is more inflated. Hinge with about
a dozen teeth on each side of the pit for the resilium.

Yoldia lYoldiella?) infrequens Dali.

Yoldia (Yoldiella?) infrequens Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 381. U. S. S. Albatross station 2784, “in 194 fathoms, mud, off
the coast of southern Chile, bottom temperature 51°. 9 F.” Also at station 2783,
in 122 fathoms.

Type Locality : Off the coast of southern Chile, in 194 fathoms, mud.
Type No. 110,692 U. S. Nat. Mus.

Range : Off the southern coast of Chile.

Shell very small, callistaeform, tumid; both ends of shell evenly rounded;
concentrically striated ; hinge with about 9 anterior and 13 posterior teeth.
Length, 4.4 mm. ; height, 3.5 mm. ; maximum diameter, 2.0 mm.

Yoldia lYoldiella) leonilda Dali.

Yoldia ( Yoldiella ) leonilda Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 382. U. S. S. Albatross station, 3360, “Gulf of Panama, in 1672
fathoms, sand, bottom temperature 42° F.”

Yoldia leonilda Dali, Zetek, Rev. Nueva, Nos. 1 & 2, 1918, p. 38. Panama.

Type Locality: Gulf of Panama, in 1,672 fathoms, sand. Type No.
122,909 U. S. Nat. Mus.

Range : Gulf of Panama.

Shell small, oval, smooth, white ; hinge with 12 anterior and 14 posterior
teeth.

Yoldia lYoldiella?) mantana Dali.

Yoldia ( Yoldiella ?) mantana Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 381. U. S. S. Albatross station 2792, “in 401 fathoms, mud,
off Manta, Ecuador, bottom temperature 42°.9 F.”

Type Locality : Off Manta, Ecuador, in 401 fathoms, mud. Type No.
122,756 U. S. Nat. Mus.

Range: Known only from the type locality.

Shell small, elongate-oval, white, nearly equilateral, with a pale yellowish
periostracum ; surface mostly smooth but with a few irregular concentric
striae; hinge with about 9 anterior and 10 or 11 posterior teeth. Length,
5.0 mm. ; height, 3.2 mm. ; maximum diameter, 2.5 mm.

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Genus Cyrilla A. Adams.

Huxley ia A. Adams, Ann. & Mag. Nat. Hist., Ser. 3, Vol. 5, April, 1860, p.
303. Sole species: Huxleyia sulcata A. Adams. See also p. 432.

Not Huxleyia Bowerbank. Dali (1898), cited the use of the name Huxleyia
by Bowerbank but we have not noticed a citation to the original reference.

Not Huxleya Dyster, Quart. Jour. Micr. Soc., Vol. 6, 1858, p. 260.

Cyrilla A. Adams, Ann. & Mag. Nat. Hist., Ser. 3, Vol. 5, June, 1860, p. 478.
“In a former paper I gave the name Huxleyia to a new genus of bivalve Mollusca.
Should, however, this name have been already used, which is not improbable,

Cyrilla might be substituted for it.” A. Adams, Ann. & Mag. Nat. Hist.,

Ser. 3, Vol. 9, 1862, p. 295. A. Adams, Journ. cle Conchyl., Vol. 16, 1868,

p. 41. Dali, Trans. Wagner Free hist. Sci., Vol. 3, Pt. 4, 1898, p. 601.

Type: H. sulcata A. Adams.

Type (by monotypy) : Huxleyia sulcata A. Adams. [Ann. & Mag. Nat.
Hist., Ser. 3, Vol. 5, lb60, p. 303. “Hab. 16 miles from Mino-Sima, Straits

of Korea; dredged from 63 fathoms.” A. Adams, Journ. de Conchyl.,

Vol. 16, 1868, p. 42, pi. 4, fig. 2. Original locality cited.

Shell suborbicular, minute, exteriorly smooth or with simple sculpture;
pallial line entire; fossette for ligament enlarged and rounded; anterior
cardinal teeth reduced, posterior series of teeth well developed; interior of
shell not nacreous.

Nucinella Wood85 with the type Nucula miliar is Deshayes from the
Eocene of the Paris Basin, is a related genus. The name Pleurodon Wood
was replaced by Nucinella Wood due to the earlier use of Pleurodon by
Harlan. Pleurodon microdus Boettger, P. cinctus von Koenen, P. dobergensis
Lienenklaus and P. zinndorfi Zilch8'" all referred to Pleurodon, have been
described from the Oligocene of Germany and the genus has been cited from
Eocene to Recent. Pleurodon gunteri Mansfield87 has been described from
the Miocene, and P. woodii Dali88 from the Miocene and Pliocene of Florida,
and P. adamsii occurs in the waters off the coast of Florida. E. A. Smith80
recorded a species living off the Cape of Good Hope, under the name of
Nuculina ovalis Wood. It is probably a distinct species from the one originally
described under that name from the Crag, Pliocene of England. Jeffreys90
cited Nucinella ovalis Wood from the Strait of Korea in 40 fathoms but
probably that record can be referred to some similar species inhabiting the
North Pacific. Lamy described “ Nucinella Serrei ”91 from Brazil, and Cotton
& Godfrey have recently added a new species, “Pleurodon hedleyi,” to the
fauna of South Australia. Species of Cyrilla are also known off the coasts of
New Zealand92 and Japan as well as off North and Central America.

S5 Nucinella Wood, Palaeontogr. Soc., Monogr. Crag. Moll., Pt. 2, 1850, p. 72. Species cited:
Nucula miliaris Deshayes. In the synonymy of which was included Pleurodon ovalis S. Wood, 1840.
Nucinella to replace Pleurodon Wood, 1840 [not Pleurodon Harlan, Jour. Acad. Nat. Sci. Philadel-
phia, Vol. 6, 1831, p. 284]. Schenck, Jour. Paleo., Vol. 13, no. 1, January, 1939, p. 39.

“ Nucinella Wood, 1850 (genotype, N. miliaris (Deshayes), from the Eocene of the Paris Basin)”
[=Nucula miliaris Deshayes, Descript. Coq. Foss. env. Paris, Vol. 1, Livr. 15, 1829, p. 235, pi. 36,
figs. 7, 8, 9],

86 Pleurodon zinndorfi Zilch, Arch. f. Molluslcenkunde, Bd. 69, No. 5-6, November, 1937, p. 250,
figs, la, lb. “In den mitteloligozanen Meeressanden des Mainzer Beckens, am Welschberg bei Wald-
bockelheim.”

87 Pleurodon gunteri Mansfield, Florida State Geol. Surv., Bull. No. 8, 1932, p. 38, pi. 2, figs. 4
and 6. Vaughan Creek, Walton County, Florida. Upper Middle Miocene.

88' Pleurodon woodii Dali, Trans. Wagner Free Inst. Sci., Vol. 3, pt. 4, 1898, p. 600, pi. 24,

fig. 10. “Pliocene marls of the Caloosahatchie, Florida.” Mansfield, Florida State Geol. Surv.,

Bull. No. 8, 1932, p. 37, pi. 2, figs. 1 and 3. Cancellaria zone — borrow pit, Jackson Bluff, Leon
County, Florida ; upper Miocene.

89 Nuculina ovalis (S. Wood), E. A. Smith, Sci. Rept. Voy. Challenger, Zook, Vol. 13, 1885,
p. 230, pi. 19, figs. 1, lb. “Simon’s Bay, Cape of Good Hope, in 15 to 20 fathoms.”

90 Jeffreys, J. G., Jour. Linn. Soc. London , Vol. 14, 1879, p. 420.

91 Nucinella serrei Lamy, Bull., Mus. Nat. d’Hist. Nat. (Paris), Vol. 18, no. 7, 1912, p. 432,
figs. 1, 2, 3. Brazil.

92 Pleurodon maorianus Hedley, Rec. Australian Mus., Vol. 5, no. 2, January 28, 1904, p. 87,
fig. 14. “from 5 fathoms off Anchor Island, Dusky Sound, New Zealand.”

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 419

Two generic names Cyrillona and Cyrillista have been proposed by
Iredale03 for austral species formerly referred to Cyrilla or Nuculina. Dia-
bolica Jousseaume9,1 is referable to Cyrilla as indicated by Thiele.

Lamy95 has reviewed many of the species referred to Pleurodon and
Nucinella.

Key to the Species of Cyrilla.

A. Lateral tooth short, ligamentary pit large munita

B. Lateral tooth long, ligamentary pit small subdolus

Cyrilla munita Carpenter in Dali.

Cyrilla munita Carpenter, Dali, Trans. Wagner Free Inst. Sci., Vol. 3, pt. 4,
April. 1*98. p. 602. “From thirty fathoms off Catalina Island, California.”

Dali, U. S. Nat. Mus., Bull. 112, 1921, p. 14 (under the genus Pleurodon).

Santa Barbara Islands to the Gulf of California. Schenck, Jour. Paleo.,

Vol. 13, no. 1, January, 1939, p. 39, pi. 6, figs. 14 and 15. Type localities of
Cyrilla munita and Nucula petriola cited.

Nucula petriola Dali, Proc. U. S. Nat. Mus., Vol. 52, December 27, 1916, p.

395. “Off Santa Rosa Island, California, in 53 fathoms, mud.” 1. S. Oldroyd,

Stanford Univ. Publ. Univ. Ser. Geol. Sci., Vol. 1, 1924, p. 12. Type locality cited.

Nuculina munita Carpenter, Lamy, Bull. Mus. Nat. d’Hist. Nat. {Paris),
Vol. 18, no. 7, 1912, pp. 430, 431. Earlier records cited.

Pleurodon munitum (Carpenter) Dali, I. S. Oldroyd, Stanford Univ. Publ.
Univ. Ser. Geol. Sci., Vol. 1, 1924, p. 36. Locality and range same as cited by
Dali.

Nucinella munita [Carpenter in Dali], Keen, Check List West North Amer.
Mar. Moll., (Stanford Univ., Calif.), 1937, p. 23, Lat. 23°-34° N.

Type Locality. Off Catalina Island, California, in 30 fathoms. Type of
Nucula petriola, No. 271,416 U. S. Nat. Mus.

Range : Santa Rosa Island, California to ? Lat. 34° S.

A small rounded-quadrate shell with a large fossette.

Subgenus Neopleurodon Hertlein & Strong, nov.

Type : Pleurodon subdolus Strong & Hertlein.

Shell minute, suborbicular ; exteriorly smooth or with simple sculpture;
pallial line entire; hinge plate well developed with teeth forming a gentle
arc; a long lateral tooth present in right valve; interior not nacreous.

Neopleurodon differs from Cyrilla in the more even arcuate series of
teeth and in the very small area for the ligament, and longer lateral tooth.

Judging from the illustration, Pleurodon ovalis Wood00 from the Pliocene
of England may belong to this subgenus, as may “Pleurodon hedleyi” Cotton
& Godfrey87.

93 Iredale, T., Rec. Australian Mus., Vol. 17, no. 4, September 4, 1929, pp. 160, 188. Cyrillona ,
“Type Cyrilla dalli Hedley.” Mem. Australian Mus., Mem. 4, Vol. 1, pt. 5, July 29, 1902, p. 296,
fig. 44. “off Port Kembla, in 63-75 fathoms, and off Cape Three Points in 41-50 fathoms.” Cyrillista,
“Type Nuculina concentrica Verco.” Illustrated by Cotton & Godfrey, Moll. South Australia, Pt. 1,
1938, p. 56, fig. 38.

9 ± Diabolic a diabolica Jousseaume, Le Naturaliste, Ann. 19, (Ann. 11, Ser. 2), no. 257, Novem-
ber 15, 1897, p. 265. Sole species: Diabolica diabolica Jousseaume. No locality cited. Lamy,

Mem. Inst. Egypt, Vol. 37, 1938, p. 8, fig. 1 (as Nucinella ( Cyrilla ) diabolica) . Gulf of Akaba.

95 Lamy. E., Sur le Genre Pleurodon ou Nucinella S. Wood, avec description d’une espece
nouvelle. Bull. Mus. Nat. d' Hist. Nat. (Paris), Vol. 18, no. 7, 1912, pp. 429-433. See also Lamy, E.,
Joura. de Conchyl., Vol. 60, no. 2, December 15, 1912, pp. 108, 109 (footnote).

96 Pleurodon ovalis Wood, Mag. Nat. Hist., New Ser., Vol. 4, 1840, p. 230, Suppl. pi. 13, fig. 1.

“Coralline crag Ramsholt and Sutton.” See illustration by Wood, Palaeontogr. Soc., Monogr.

Crag. Moll., Pt. 2, 1850, p. 72, pi. 10, figs. 4a, 4b, 4c.

9” Pleurodon hedleyi Cotton & Godfrey, Moll. South Australia, Pt. 1. Handbook Flora and Fauna of
South Australia, issued by South Australian Brit. Sci. Guild, Adelaide, 1938, p. 57, fig. 39. “100
fathoms, 40 miles south of Cape Wiles, South Australia.”

420

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[XXV :25

Cyrilla I Neopleurodon) subdolus Strong & Hertlein.

Pleurodon subdolus Strong & Hertlein, Proc. Calif. Acad. Sci., Ser. 4, Vol. 22,
no. 6, December 31, 1937, p. 162, pi. 35, figs. 14, 18, 19. “Lat. 23° 12' N., Long.
106° 29' W., dredged in 12 fathoms, about five miles west of Mazatlan, Sinaloa,
Mexico.”

Type Locality: Off Mazatlan, Mexico, in 12 fathoms. Type No. 6,970
Calif. Acad. Sci. Paleo. Type Coll.

Range : Known only from the type locality.

The more regularly oval shell, persistant dark epidermis, and different
hinge are features separating Cyrilla subdolus from C. munita (Carpenter)
Dali. Cyrilla adamsi Dali" from the Straits of Florida is a somewhat similar
species but the hinge is different.

Genus Malletia Desmoulins.

Malletia Desmoulins, Act. Soc. Linn. Bordeaux, Vol. 5, February 15, 1832, p.

85. Stoliczka, Mem. Geol. Surv. India. Palaeont. Indica. Cret. Fauna South.

India, Vol. 3, 1871, pp. XX, 321. “Type, M. chilensis, Desm.” Verrill &

Bush, Amer. Jour. Sci., Ser. 4, Vol. 3, January, 1897, p. 56. “Type Malletia

chilensis Desmoulins.” Verrill & Bush, Proc. U. S. Nat. Mus., Vol. 20,

1898, p. 873. Type: Malletia chilensis Desmoulins. Dali, Trans. Wagner

Free Inst. Sci., Vol. 3, pt. 4, 1898, p. 581. “Type M: norrisii (Sby.)=M. chilensis

Desm.” Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908, p. 383.

Type: M. chilensis Desmoulins. Cossmann & Peyrot, Act. Soc. Linn. Bor-

deaux, Vol. 66 (Conch. Neog. Aquit., Vol. 2, Livr. 1), 1912, p. 121. Genotype:

M. chilensis Desmoulins. Grant & Gale, Mem. San Diego Soc. Nat. Hist.,

Vol. 1, 1931, p. 132. Type: M. chilensis Desmoulins. Prashad, Siboga Exped.,

Mon. 53c, Lamell., 1932, p. 25. Type: Malletia chilensis Desmoulins.

Solenella Sowerby, Proc. Zool. Soc. London, for 1832 (issued March 13, 1833),
p. 197. Sole species: Solenella norrisii. Valparaiso, Chile, in soft mud, at from
14 to 45 fathoms.

Pseudomalletia Fischer, Man. de Conchyl., 1886, p. 987. Species cited: P.
obtusa Sars and P. transversa Ponzi.

Type (designated by Stoliczka) : Malletia chilensis Desmoulins.

Shell oval, compressed, smooth or concentrically furrowed, epidermis
olive; ligament external, elongated, prominent; hinge with an anterior and
posterior series of fine sharp teeth; interior subnacreous; pallial sinus large
and deep; anterior adductor scar giving off a long oblique pedal line (Tryon:
Struct, and Syst. Conch.)

The shell of Malletia s.s. is oblong elliptical, blunt posteriorly and
without a definite rostrum or carination. The carinated and rostrate species
with strong concentric sculpture can in many cases be referred to Neilo
H. & A. Adams. The absence of a lunule, escutcheon and distinct rostrum
serve to separate Malletia from Yoldia. A characteristic feature of the hinge
of Malletia is that the posterior series of teeth are often more than twice as
numerous as the anterior series.

Malletia has been recorded from upper Cretaceous" to Recent. It is
primarily a Magellanic genus which is said to have reached the northern

9 & Pleurodon adamsi Dali, Trans. Wagner Free Inst. Sci., Vol. 3, pt. 4, April, 1898, p. G01,
pi. 24, fig. 9. “Dredged seven miles east of Fowey Rocks, Straits of Florida.”

99 Malletia gracilis Wilckens, Wiss. Ergeb. Schwed. Siidpolar-Exped., 1901-1903, Bd. 3, Lief. 12,
1910, p. 25, Taf. 2, fig. 4. Seymour Island, Antarctica ; Cretaceous. The shape of this species is
similar to that of M. chilensis Desmoulins.

Malletia ornata Sowerby has been reported from Patagonia in beds referred to the upper
Cretaceous, also in beds referred to the Eocene by von Ihering (An. Mus. Nac. Buenos Aires, Ser.
3, Vol. 7, 1907, pp. 4, 228). The species bears a resemblance to M. australis Quoy & Gaimard of
New Zealand. Powell (Shellfish of New Zealand, 1937, p. 55), referred M. australis to Neilo which
he considered to be of generic status.

1940] Hertlein & Strong : Mollusks of Eastern Pacific Expeditions 421

hemisphere during the Oligoeene or Miocene, but the genus appears to be
present in the Eocene of California.

One of the largest species of Malletia is M. gigantea100 which occurs at
Kerguelen Island in the south Atlantic. The size is noteworthy, the type
specimen is 62 mm. in length. The posterior end of M. gigantea is expanded
in a manner somewhat resembling Yoldia thraciaeformis. E. A. Smith men-
tioned that the animal of Yoldia subaequilateralis 101 “resembles that of
Solenella gigantea in all respects excepting that the edge of the foot is
bluntly serrated or scalloped.”

Malletia I Malletia I chilensis Desmoulins.

Malletia chilensis Desmoulins, Act. Soc. Linn. Bordeaux, Vol. 5, January 15,
1832, p. 85, pi. 1, figs. 1-8. Valparaiso, Chile, in 14 to 45 fathoms (according to

Dali). H. & A. Adams, Gen. Rec. Moll., Vol. 2, 1858, p. 549, pi. 126, figs.

6, 6a. Chenu, Man. de Conchyl., Vol. 2, 1862, p. 181, fig. 913. Kobelt,

Illustr. Conchylienbuch, Bd. 2, Lief. 10-11, 1881, p. 372, pi. 109, fig. 3. Chile.

Tryon, Struct, and Syst. Conch., Vol. 3, 1884, p. 249, pi. 126, fig. 34 (on

expl. to pi. 126 as Yoldia ( Malletia ) chilensis). Valparaiso, Chile. Fischer,

Man. de Conchyl., Fasc. 10, 1886, p. 987, pi. 17, fig. 22. Verrill & Bush,

Amer. Jour. Sci., Ser. 4, Vol. 3, 1897, p. 56, fig. 9 (on p. 60). Stempell,

Zool. Jahrb., Suppl. Bd. 4 (Fauna chilensis, Bd. 1), Heft 2, July 1, 1898, p. 343
(and following pages), pi. 22, figs. 2, 3, 4, 9, 12; pi. 23, figs. 13, 14, 15, 16, 17;

pi. 24, fig. 32 (anatomy). Stempell, Zool. Jahrbuch., Suppl. Bd. 5 (Fauna

Chilensis, Bd. 2), 1902, p. 219. Harbor of Coquimbo; Talcahuano, in 10 fathoms;

Calbuco, in 10 fathoms. -Dali, Proc. U. S. Nat. Mus., Vol. 37, 1909, p. 251.

Coquimbo, south to Talcahuano Bay, Chile. Heath, Mem. Mus. Roy. d’Hist.

Nat. Belg., Ser. 2, Fasc. 10, 1937, p. 5, pi. 10, fig. 87 (Anatomy). Talchano
[?Talcahuano], Chile.

Solenella norrisii Sowerby, Proc. Zool. Soc. London for 1832 (issued March
13, 1833), p. 197. “Hub. ad Valparaiso.” “Discovered by Mr. Cuming, in soft mud,
at from fourteen to forty-five fathoms depth.” — — - — Sowerby, Gen. Shells, Vol.
2, pi. 237 (four figs.), 71833. “Dredged by Mr. H. Cuming at Valparaiso.”

Reeve, Conch. Syst., Vol. 1, 1841, p. 48, pi. 30, figs. 1-4. Hanley,

Cat. Rec. Biv. Shells, p. 17, 1843, p. 337, 18567, Suppl. 10, figs. 8, 1843.

d’Orbigny, Voy. Amer. Merid., Vol. 5, 1846, p. 543. Dredged in 10 fathoms in the

Bay of Valparaiso, Chile. Woodward, Man. Moll., 1851-1856, p. 270, pi.

17, fig. 22. Ed. 2, 1868, p. 429, pi. 17, fig. 22. Reprint of Ed. 4 (1880), 1910, p.

429, pi. 17, fig. 22. Valparaiso, Chile. Hupe, Hist, de Chile, Zool., Vol. 8,

1854, p. 306. Various parts of the coast of Chile. Deshayes, Traite Elem.

Conchyl., Vol. 2, 1839-1857 [-1858], p. 270, pi. 34, figs. 5, 6, 7. Valparaiso, Chile.

Hanley, Thes. Conch.., Vol. 3, 1860, p. 164, pi. 226 ( Nuculidae pi. 1), figs. 1

and 2. Valparaiso, Chile.

Solenella norrisii Sowerby var. brevior Hanley, Thes. Conch., Vol. 3, 1860,
p. 164, pi. 226 ( Nuculidae , pi. 1). Valparaiso, Chile.

Type Locality : Valparaiso, Chile, in 14 to 45 fathoms, mud.

Range : Coquimbo to Talcahuano Bay, Chile.

Shell subovate, somewhat compressed, with polished olivaceous epider-
mis; beaks anterior to the middle; unornamented except by lines of growth;
hinge with teeth very fine and very numerous posteriorly but very few
anteriorly.

100 Solenella gigantea E. A. Smith, Ann. & Mag. Nat. Hist., Ser. 4, Vol. 16, 1875, p. 72. “Hab.

Royal Sound, Kerguelen’s Island.” E. A. Smith, Phil. Trans. Roy. Soc. London, Vol. 1 6s!

1879, p. 187, pi. 9, fig. 19. Royal Sound, both at Observatory Bay and near the eastern shore of
Swain’s Harbour, on mud in about 10 fathoms.

Malletia gigantea E. A. Smith, E. A. Smith, Sci. Res. Voy. Challenger, Zool., Vol. 13, Lamell.,
1885, p. 244. Betsy Cove, Kerguelen Island, in shallow water ; also at Balfour Bay, in 20 to 60

fathoms. Lamy, Bull. Mus. Nat. d’Hist. Nat. (Paris), Vol. 12, 1906, p. 203. Kerguelen

Islands. Lamy, Bull. Mus. Nat. d’Hist. Nat. (Paris), Vol. 21, no. 2, 1915, p. 76. Several

localities in Kerguelen Islands.

101 Yoldia subaequilateralis E. A. Smith, Ann. & Mag. Nat. Hist., Ser. 4, Vol. 16, 1875, p. 73.

“Hab. Swain’s Bay, Kerguelen’s Island.” E. A. Smith, Phil. Trans. Roy. Soc. London, Vol!

168, 1879, p. 187, pi. 9, fig. 18. Swain's Bay. Dredged in 7-10 fathoms in very sheltered water.

422

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Subgenus Neilo H. & A. Adams.

Neilo A. Adams, Proc. Zool. Soc. London, 1852, p. 93. Sole species: Neilo

cumingii A. Adams. “It is from the shores of New Zealand.” H. & A,

Adams, Gen. Rec. Moll., Vol. 2, 1858, p. 549. Verrill & Bush, Amer. Jour .

Sci., Ser. 4, Vol. 3, 1897, p. 57. Type: N. cumingii Adams. Verrill & Bush,

Proc. U. S. Nat. Mus., Vol. 20, 1898, p. 877. Type: Neilo cumingii Adams.

Dali, Trans. Wagner Free Inst. Sci., Vol. 3, pt. 4, 1898, p. 581. Type: N. australis

Quoy & Gaimard (+A. cumingii A. Adams). Suter, Man. New Zealand

Moll., 1913, p. 837. Type: N. cumingii A. Adams =australis Quoy & Gaimard.

Type (.by monotypy) : Neilo cumingii A. Adams. [Proc. Zool. Soc. Lon-
don, 1852, p. 93. “It is from the shores of New Zealand.” H. & A.

Adams, Gen. Rec. Moll., Vol. 2, 1858, p. 549, pi. 126, figs. 7, 7a, 7b.

Chenu, Man. de Conchyl., Vol. 2, 1862, p. 181, fig. 914. Suter, Man.

New Zealand Moll., 1913, p. 837, pi. 58, figs. 2, 2a (as Malletia australis
Quoy & Gaimard). New Zealand],

The subgenus Neilo differs from Malletia in the ark-like form and in that
the surface of the valves is usually ornamented by concentric sculpture.

Neilo has been recorded from late Cretaceous to Recent in Patagonia,
from the Helvetian, Miocene, of France, and from upper Oligocene to Recent
in New Zealand. The species described as Nucula abrupta by Conrad from
the Miocene of Astoria, Oregon, was later referred to Neilo, but, there is
some doubt as to the generic assignment of the species. One species assigned
to Neilo has been described from deep water off tropical western North
America.

Spineilo Finlay & Marwick102, with the type Malletia elongata Marshall
from the Wangaloan (Danian) of New Zealand, was proposed as a subgenus
of Neilo, the latter considered by them as of generic status.

Malletia INeilol goniura Dali.

Malletia goniura Dali, Proc. U. S. Nat. Mus., Vol. 12, 1889, p. 251, pi. 10, fig.
10. U. S. Fish Commission station 2793, “off the coast of Ecuador in 741 fathoms,
mud; bottom temperature 38°. 4 F.”

Malletia ( Nelro ) [typ. error for Neilo ] goniura Dali, Dali, Bull. Mus. Comp.
Zool., Vol. 43, no. 6, October, 1908, p. 386, pi. 18, fig. 6. U. S. S. Albatross station
3360. Gulf of Panama, in 1672 fathoms, sand, bottom temperature 42° F. Also
original locality cited.

Type Locality : Off the coast of Ecuador, in 741 fathoms, mud.

Range : Gulf of Panama to off the coast of Ecuador, in deep water.

This is a thin fragile shell on which two ridges radiate from the beak
to the truncate posterior margin. Hinge of type described as possessing 19
anterior and 25 posterior teeth. Another specimen described by Dali from
the Gulf of Panama apparently had 14 anterior and over 30 posterior teeth.

Subgenus Minormalletia Dali.

Minormalletia Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908, p.
385. Sole species: Malletia ( Minormalletia ) arcijormis Dali.

Type (by monotypy) : Malletia ( Minormalletia ) arciformis Dali.

Shell small, blunt, plump, with amphidetic ligament, no resilium, the
pallial sinus large, no radial depressions or sculpture (Dali).

Marwick103 has recorded Minormalletia from the medial Tertiary of New
Zealand.

102 Spineilo Finlay & Marwick, Geol. Surv. New Zealand, Palaeo. Bull. No. 15, May 20, 1937, p. 17.
“Genotype: Malletia elongata Marshall, Wangaloan (Danian), New Zealand,” p. 17, pi. 1, figs. 4
and 5. “Wangaloa (Type), common; Boulder Hill, common.” Danian, upper Cretaceous.

103 Marwick, J., Geol. Surv. New Zealand , Palaeo. Bull. No. 13, 1931, p. 55.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 423

Malletia IMinormalletial arciformis Dali.

Malletia (Minormalletia) arciformis Dali, Bull. Mus. Comp. Zool., Y ol. 43,
no. 6, October, 1908, p. 385, pi. 15, figs. 5 and 6. U. S. S. Albatross , station 3417,
“off Acapulco, Mexico, in 493 fathoms, mud, bottom temperature 40°.6 F.”

Type Locality : Off Acapulco, Mexico, in 493 fathoms, mud.

Range : Known only from the type locality.

The pallial sinus is notable in that the greater part of its basal scar is
identical with the pallial line itself, whereas in most of the Malletinae the
sinus tends to slope obliquely upward (Dali).

Dali pointed out that the profile of this species, although more rounded
below and behind, is somewhat similar to that of Area adamsii (Shuttleworth
(MS) ) E. A. Smith, a Caribbean species.

Malletia IMinormalletial benthima Dali.

Malletia ( Minormalletia ) benthima Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6,
October, 1908, p. 386, pi. 15, figs. 1 and 2. U. S. S. Albatross station 3417, “off
Acapulco, Mexico, in 493 fathoms, mud, bottom temperature 40°. 6 F.”

Type Locality: Off Acapulco, Mexico, in 493 fathoms, mud. Type No.
122,927 U. S. Nat. Mus.

Range : Known only from the type locality.

Shell small, thin, very inequilateral; hinge with 12-13 anterior and
17-18 posterior teeth.

The subgeneric position of the following species of Malletia is uncertain.

Malletia faba Dali.

Malletia faba Dali, Nat. Hist. Soc. Brit. Columbia, Bull. No. 2, January, 1897,
p. 10, pi. 2, fig. 10. “Off Sea Lion Rock, coast of Washington, in 477 fms.; off
Queen Charlotte Islands, British Columbia, in 876 fms.; off Tillamook in 786

fms.; off San Diego, Cal., in 822 fms., U. S. Fish Commission.” Dali,

Checklist Rec. Bivalve Moll., (Southwest Museum), 1916, p. 13. Queen Charlotte

Islands, Brit. Columbia, to Lower California. Dali, U. S. Nat. Mus., Bull.

112, 1921, p. 14. Queen Charlotte Islands, British Columbia, to Lower California.

1. S. Oldroyd, Stanford Univ. Publ. Univ. Ser. Geol. Sci., Vol. 1, 1924, p. 37,

pi. 13, fig. 16. Range same as cited by Dali. Keen, Check List West North

Amer. Mar. Moll., (Stanford Univ. Press), 1937, p. 22. Range: Lat. 30°-54° N.

Type Locality: Off Sea Lion Rock, Washington, in 477 fathoms. Type
No. 122,569 U. S. Nat. Mus.

Range : Queen Charlotte Islands, British Columbia, to Lower California.

Shell thin, ovate, inflated, smooth, slightly inequilateral; hinge with
about 9 small posterior and about 32 anterior teeth.

Malletia inequalis Dali.

Malletia inequalis Dali, Bidl. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 383. U. S. S. Albatross station 2772, “off Cape Virgins, in 31 fathoms, sand,
and 2778, Straits of Magellan, in 61 fathoms, mud, bottom temperature 47°. 9 F.”

Type Locality: Off Cape Virgins, in 31 fathoms, sand, and Straits of
Magellan, in 61 fathoms, mud. [Type locality here designated as off Cape
Virgins, in 31 fathoms, sand].

Range: Off Cape Virgins, and Straits of Magellan.

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[XXV :25

The truncation behind, the coalescent pallial sinus, and the compressed
form separate it from the other species (Dali). Hinge with 3 anterior and
24 posterior teeth.

Malletia magellanica E. A. Smith.

Malletia magellanica E. A. Smith, Proc. Zool. Soc. London, 1881, p. 39, pi. 5,
figs. 3, 3a. “Hab. Station 3, Mayne Harbour, 9 fathoms, greenish mud; and Cockle

Cove, 2-32 fathoms, mud.” Mabille & Rochebrune, Miss. Sci. Cap. Horn,

Moll., Vol. 6, Zool., Pt. 2, 1889, p. H 114. Mayne Harbor; Cockle Cove (E. Smith).

Type Locality : Mayne Harbor and Cockle Cove, Chile.

Range : Straits of Magellan and Magellanic region.

Malletia patagonica Mabille & Rochebrune.

Malletia patagonica Mabille & Rochebrune, Miss. Sci. Cap. Horn, Vol. 6, Zool.,
pt. 2, 1889, p. H 114, pi. 8, fig. 1. “Hab. Punta- Arenas.”

Malletia hyadesi Mabille & Rochebrune, Miss. Sci. Cap. Horn, Vol. 6, Zool.,
pt. 2, 1889, p. H 114. “Hab. Punta- Arenas.”

Malletia magellanica Mabille & Rochebrune, Dali, Bull. Mus. Comp. Zool.,
Vol. 43, no. 6, October, 1908, p. 383 (at least in part). Punta Arenas. Also off
Cape Virgins, in 31% fathoms, and Magellan Straits, in 77 fathoms.

Not Malletia magellanica E. A. Smith.

Type Locality. Punta Arenas, Chile.

Range: Southern Chile in the Magellanic region.

Malletia peruviana Dali.

Malletia peruviana Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 384, pi. 10, figs. 3 and 5. U. S. S. Albatross station 4654, “N. 68° W., twenty-
four miles from Aguja Point, Peru, in 1036 fathoms, mud, bottom temperature
37°. 3 F.”

Type Locality : Twenty-four miles N. 68° W., from Aguja Point, Peru,
fathoms, ooze. Type No. 122,906 U. S. Nat. Mus.

Range : Known only from the type locality.

This species is remarkable for its almost blackish color and extremely
shallow pallial sinus, which does not extend in front of the posterior adductor
scar; the anterior end is unusually short and almost pointed (Dali). Hinge
with about 10 anterior and 33 posterior teeth.

Malletia truncata Dali.

Malletia truncata Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 384, pi. 17, fig. 1. U. S. S. Albatross station 3374, “southwest of Malpelo
Island, Gulf of Panama, in 1823 fathoms, ooze, bottom temperature 36°.4 F.” Also
at station 3361, in 1471 fathoms, and station 3381, in 1772 fathoms.

Type Locality: Southwest of Malpelo Island, Gulf of Panama, in 1,823
fathoms, ooze. Type No. 122,906 U. S. Nat. Mus.

Range: Known only from the type locality.

Remarkable for its light yellow color, its surface devoid of radial sculp-
ture or indented margin, and its bluntly truncate posterior end. In the latter
character and its subequal division of the hinge teeth it recalls Neilo, but is
without the rostration of that form (Dali). Hinge with 20 anterior and
about 27 posterior teeth.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 425

Genus Tindaria Bellardi.

Tindaria Bellardi, Monogr. Nuculidi Terr. Terz. del. Piemonte e Liguria, 1875,

p. 28. Sole species: Tindaria arata Bellardi. Verrill & Bush, Proc. U. S. Nat.

Mus., Vol. 20, 1898, p. 880. Type: Tindaria arata Bellardi. Dali, Trans.

Wagner Free Inst. Sci., Vol. 3, pt. 4, April, 1898, p. 581. Type: T. arata Bellardi.

Sacco, Moll. Terr. Terz. Piemonte e Liguria, Pt. 26, December, 1898, p.

64. “(Tipo T. arata Bell.).” Cossmann & Peyrot, Act. Soc. Linn. Bordeaux,

Vol. 66 (Conch. Neog. Aquit., Vol. 2, Livr. 1), 1912, p. 125. Genotype: Tindaria

arata Bellardi. Woodring, Carnegie Inst. Washington, Puhl. 366, 1925,

p. 23. Type: Tindaria arata Bellardi.

Type (by monotypy) : Tindaria arata Bellardi. [Mon. Nuculid. Terr.

Terz. Piemonte e Liguria, 1875, p. 28, fig. 27. Sacco, Moll, Terr. Terz.

Piemonte e Liguria, Pt. 26, December, 1898, p. 64, pi. 12, figs. 55, 56. Hel-
vetian and Tortonian of Italy].

Shell small, veneriform, strongly inflated; beaks turned forward; sculp-
tured by concentric rugae of varying development; ligament external, opis-
thodetic ; hinge uninterrupted below umbo, posterior series of teeth strongly
curved, the anterior series of teeth are heavier, shorter and straighten;
pallial sinus moderately developed, narrowly triangular.

Tindaria is known from the Miocene of France and Italy and Miocene
to Recent in the Caribbean region. It is not known to occur with certainty
in the Tertiary of the western Americas but several species have been de-
scribed from west American waters between Sitka, Alaska, and Cape Horn.

Subgenus Tindariopsls Verrill & Bush.

Tindariopsis Verrill & Bush, Amer. Jour. Sci., Ser. 4, Vol. 3, 1897, p. 59.
“Type T. agathida (Dali).”

Type (by original designation) : T. agathida (Dali). \—Malletia ( Tin-
daria) agathida Dali, Proc. U. S. Nat. Mus., Vol. 12, 1889, p. 252, pi. 13, fig.
10. South from St. Kitts, in 687 fathoms, east from Tobago, in 880 fathoms].
This subgenus was proposed for species with ovate shells, with a short
rostrum, defined by a radial ridge and a furrow.

Tindaria ITindariopsisI sulculata Gould.

Nucula sulculata Couthouy, Gould, U. S. Explor. Exped., Vol. 12, 1852, p. 424,
Atlas, 1856, pi. 37, figs. 539a-e. Orange Harbor, Patagonia.

Leda sulculata Couthouy, Hanley, Thes. Conch., Vol. 3, 1860, p. 129 footnote.

Mabille & Rochebrune, Miss. Sci. Cap. Horn. Vol. 6, Zool., Pt. 2, 1889, p.

H 113. Cape Horn. Orange Bay. Stempell, Sitzungsber. Kon. Preuss. Akad.

Wiss. Berlin, Jahrg. 1897, pp. 17-28. Stempell, Zool. Jahrb., Bd. 4 (Fauna

Chilensis, Bd. 1), Heft 2, July 1, 1898, jd. 343 (and following pages), pi. 22, figs.
1, 5, 6, 7, 8, 10, 11; pi. 23, figs. 19, 21; pi. 24, figs. 23, 24, 25, 26, 28, 29, 30, 31;
pi. 25, figs. 33, 34, 35, 36, 37, 38, 40, 41, 43 (anatomy).

Not Lembulus sulculatus Risso, Hist. Nat. Europ. Merid., Vol. 4, 1826, p. 320.

Leda lugubris A. Adams, Proc. Zool. Soc. London, 1856, p. 49. “Hab ?

Mus. Cuming.” E. A. Smith, Proc. Zool. Soc. London, 1881, p. 39. Port

Rosario in 2-30 fathoms; Wolsey anchorage, in 17 fathoms. Mabille &

Rochebrune, Miss. Sci. Cap. Horn, Vol. 6, Zool., Pt. 2, 1889, p. H 113. Port
Rosario (E. Smith).

Leda orangica Mabille & Rochebrune, Miss. Sci. Cap. Horn, Vol 6 Zool
Pt. 2, 1889, p. H 113, pi. 8, fig. 3. “Hab. Baie Orange.”

Tindaria (Tindariopsis) sulculata Gould, Dali, Bull. Mus. Comp. Zool., Vol.
43, no. 6, October, 1908, p. 390. Straits of Magellan. Also earlier records.

Type Locality: Orange Harbor, Chile.

426

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[XXV :25

Range : Cape Horn and Straits of Magellan.

The subgeneric position of the following species of Tindaria is uncertain.

Tindaria atossa Dali.

Tindaria atossa Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 388, pi. 15, figs. 3 and 4. U. S. S. Albatross, station 3392, “Gulf of Panama, in
1270 fathoms, hard bottom, temperature 36°. 4 F.” Also at station 3393, Gulf of
Panama, in 1020 fathoms.

Type Locality : Gulf of Panama, in 1,270 fathoms, hard bottom. Type
No. 122,920 U. S. Nat. Mus.

Range : Gulf of Panama.

Shell small, olivaceous, moderately polished, finely concentrically striated
all over, with the beaks slightly anterior, the posterior end blunGy pointed,
the anterior rounded; ligament small, amphidetic; anterior teeth six, pos-
terior ten, the dorsal slopes gently, the basal margin roundly arcuate; in-
terior white, margin entire. Measurements of the type: Length, 6.5 mm.;
height, 4.5 mm.; maximum diameter, 3.0 mm. (From Dali).

Tindaria atossa resembles T. panamensis but the whole surface is sculp-
tured and the posterior end is less elongated and pointed.

Tindaria compressa Dali.

Tindaria compressa Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October,
1908, p. 387, pi. 15, figs. 7 and 8; pi. 17, figs. 15 and 16. U. S. S. Albatross station
3360, “in 1672 fathoms, sand, bottom temperature 42° F.” Also at station 3414,
“southwest of the isthmus of Tehuantepec, in 2232 fathoms, green mud, bottom
temperature 38°. 5 F.”

Type Locality : Lat. 6° 17' 00" N., Long. 82° 5' 0" W., in 1,672 fathoms,
sand. Type No. 122,921 U. S. Nat. Mus.

Range : Southwest of the Isthmus of Tehuantepec to the Gulf of Panama,
in deep water.

Shell small, thin, cythereiform, inequilateral, concentrically striated ;
hinge with 10-12 anterior and 21-25 posterior teeth.

Tindaria amabilis Dali104 of the Antilles is a somewhat similar species.

Tindaria panamensis Dali.

Tindaria panamensis Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October,
1908, p. 388, pi. 17, figs. 10 and 12. U. S. S. Albatross, station 3392, “Gulf of
Panama, in 1270 fathoms, hard bottom, temperature 36°. 4 F.”

Type Locality : Gulf of Panama, in 1,270 fathoms, hard bottom. Type
No. 122,922 U. S. Nat. Mus.

Range : Known only from the type locality.

Surface with fine concentric threads which become much finer on the
posterior third of the shell; hinge with 7 anterior and about 13 posterior
teeth. Dimensions of type; length, 5.5 mm.; height, 4.3 mm.; maximum
diameter 2.8 mm.

The more greenish color and dull surface of this species are said to be
characteristic in comparison to similar species.

104 Malletia ( Tindaria ) amabilis Dali, Bull. Mus. Comp. Zool., Vol. 18, June, 1889, p. 438,

pi. 40, fig. 8. Dali, Proc. U. S. Nat. Mus., Vol. 12, 1889, p. 252. In 607 fathoms, off St. Kitts,

and in 880 fathoms east of Tobago.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 427

Tindaria sp105., cited by Woodring from the Miocene of Bowden,
Jamaica, is said to possess a similar shell but it is shorter and the posterior
end is more inflated.

Tindaria mexicana Dali.

Tindaria mexicana Dali, Bull. Mus. Camp. Zool., Vol. 43, no. 6, October, 1908,
p. 389, pi. 17, figs. 11 and 14. U. S. S. Albatross, station 3418, “off the Mexican
coast in 660 fathoms, sand, bottom temperature 39° F.”

Type Locality: Off the Mexican coast, in 660 fathoms, sand. Type No.
122,925 U. S. Nat. Mus.

Range: Known only from the type locality.

Very similar to T. smirna, but with more prominent beaks; shorter and
blunter, in proportion to height, and decidely less pointed and produced
behind (Dali). Hinge with 11 anterior and about 21 posterior teeth.

Tindaria solaria Dali.

Tindaria salaria Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 387. “Dredged in Mid-Pacific, off Salar y Gomez Island, at U. S. S. Albatross,
station 4693, in 1142 fathoms, manganese nodules, bottom temperature 35°. 4 F.”

Type Locality: Off Sala y Gomez, West of Chile, in 1,142 fathoms,
manganese nodules.

Range: Known only from the type locality.

The shell of this species, in its general exterior characteristics, is said
to resemble that of Nucida proxima Say.

Tindaria smirna Dali.

Tindaria smirna Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 389, pi. 17, figs. 6 and 7. U. S. S. Albatross, station 3360, “Gulf of Panama, in
1672 fathoms, sand, bottom temperature 42° F.”

Type Locality: Gulf of Panama, in 1,672 fathoms, sand. Type No.
122,919 U. S. Nat. Mus.

Range: Known only from the type locality.

Shell small, polished, subtriangular, of a blackish olive tint; smooth over
two thirds of the surface, but near the basal margin there are about 10
concentric grooves crossed by very fine radial striae; hinge with about 8
anterior and 12 posterior teeth. Length 5.5 mm.; height, 4.0 mm.; maximum
diameter, 2.7 mm.

Close to T. atossa, but much more brilliantly polished, the shell shorter,
more turgid, more nearly equilateral, and with the ligament distinctly and
subequally amphidetic, while in T. atossa it appears, to the naked eye, to
be entirely opisthodetic, though on opening the valves a small portion is seen
to pass in front of the beaks (Dali).

Tindaria thea Dali.

Tindaria thea Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 390. U. S. S. Albatross, station 4654, “N. 68° W., twenty-four miles from’Aguja
Point, Peru, in 1036 fathoms, mud, bottom temperature, 37°. 3 F.”

105 Tindaria (Tindaria) sp., Woodring, Carnegie Inst. Washington, Publ. 366, May, 1926, p. 23,
pi. 1, figs. 22, 23. Bowden, Jamaica ; Miocene.

428

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[XXV :25

Type Locality. 24 miles northwest of Aguja Point, Peru, in 1,036 fath-
oms, mud. Type No. 110,577 U. S. Nat. Mus.

Range : Known only from the type locality.

Just about the color of dark, wet, tea-leaves after they have been steeped.
Longer and more pointed, and more attenuated behind, than T. smirna,
which has very similar sculpture but a more yellowish color. In looking from
below, the series of hinge teeth seems uninterrupted, but the black color of
the semi-internal resilium can be seen through the interstices, as it is situ-
ated above the tooth-line (Dali). Hinge with 8 anterior and 14 posterior
teeth.

Tindaria virens Dali.

Malletia ( Tindaria ) virens Dali, Proc. U. S. Nat. Mus., Vol. 12, 1889, p. 254,
pi. 13, fig. 3. “In 122 to 449 fathoms, mud, on the west coast of Patagonia; tem-
peratures 47° to 50° F.”

Tindaria virens Dali, Bull. Mus. Comp. Zool., Vol. 43, no. 6, October, 1908,
p. 389. Coast of Southern Chile and western Patagonia, in 122 to 449 fathoms.

Type Locality: West coast of Chile, in 122 to 449 fathoms, mud.

Range: Coast of southern Chile, in 122 to 449 fathoms.

Lighter colored, with alternating zones and with sharper and deeper
concentric sulci, than those exhibited by T. atossa of the Panama fauna,
wrhich is the most nearly related species (Dali, 1908).

EXPLANATION OF THE PLATES.

Plate I.

Fig. 1. Nucula (Nucula) declivis Hinds. Length 5 mm., height 4 mm., thickness
(both valves) 2.9 mm. Hypotype, left valve, from Station 203-D-l,
dredged in 15 fathoms (27 meters), in Lat. 10° 56' 05" N., Long. 85°
49' 25" W., Port Parker, Costa Rica. P. 380.

Fig. 2. Nucula ( Nucula ) declivis Hinds. Length 4.8 mm., height 3.8 mm., thick-
ness 1.6 mm. Hypotype, right valve, from Station 145-D-l, dredged in
13 fathoms (24 meters) in Lat. 26° 52' N., Long. 111° 53' W., Santa Inez
Bay, east coast of Lower California. View of the interior. P. 380.

Fig. 3. Nucula ( Nucula ) declivis Hinds. Length 4.6 mm., width 3.4 mm., thick-
ness (both valves) 2.7 mm. Hypotype, left valve, from the same locality
as the specimen shown in Fig. 2. P. 380.

Fig. 4. Nucula ( Nucula ) exigua Sowerby. Length 5.7 mm., height 5.4 mm.,
thickness (both valves) 3.9 mm. Hypotype, left valve, dredged at Loc.
27595 (C.A.S.), San Bartolome (Turtle Bay), Lower California. P. 381.

Fig. 5. Nucula (Nucula) exigua Sowerby. View of posterior end of specimen
shown in Fig. 4.

Fig. 6. Nucula (Nucula) declivis Hinds. View of posterior end of specimen shown
in Fig. 1.

Fig. 7. Nucula (Nucula) declivis Hinds. View of specimen shown in Fig. 1
but with less magnification.

Fig. 8. Nucula (Nucolopsis) schencki Hertlein & Strong, sp. nov. Length 2.0 mm.,
height 1.7 mm., thickness (both valves), 1.0 mm. Holotype, left valve,
from Station 195-D-9, dredged in 7 fathoms (12.6 meters), Lat. 15° 44'
28" N., Long. 96° 07’ 51" W., Port Guatulco, Mexico. P. 384.

Fig. 9. Nucula (Nucolopsis) schencki Hertlein & Strong, sp. nov. Length 2.8
mm., height 2.5 mm. Paratype, right valve, from the same locality as the
holotype shown in Fig. 8. View of the interior. P. 384.

Fig. 10. Nucula (Nucolopsis) schencki Hertlein & Strong, sp. nov. Dorsal view
of the holotype shown in Fig. 8.

1940] Hertlein & Strong: Mollusks of Eastern Pacific Expeditions 429

Fig. 11. Nucnlana ( Saccella ) acuta Conrad. Length 8.6 mm., height 4.8 mm.,
thickness (both valves) 3.4 mm. Hypotype, left valve, dredged at a depth
of two fathoms at Cedar Keys, Florida, H. Hemphill collector (Loc. 5710).
P. 392.

Fig. 12. Nuculana (Saccella) elenensis Sowerby. Length 13.6 mm., height 7.0 mm.,
thickness (both valves) 5 mm. Hypotype, left valve, from Station 196-
D-17, dredged in 23 fathoms (42 meters) in Lat. 15° 45' N., Long. 96°
05' 34" W., Tangola-Tangola Bay, Mexico. P. 393.

Fig. 13. Nuculana ( Saccella ) callimene Dali. Length 14.5 mm., height 8.9 mm.,
thickness (both valves) 6.5 mm. Hypotype, left valve, from 3 miles south
of Blanco Island, Gulf of Nicoya, Costa Rica. P. 393.

Fig. 14. Nuculana ( Saccella ) elenensis Sowerby. Length 16 mm., height 8.5 nun.,
thickness (both valves) 6.5 mm. Hypotype, left valve, from the same
locality as the specimen shown in Fig. 2. P. 393.

Fig. 15. Nuculana ( Saccella ) elenensis Sowerby. Dorsal view of specimen shown
in Fig. 14.

Fig. 16. Nuculana ( Saccella ) elenensis Sowerby. Length 13.3 mm., height 6.8 mm.,
thickness (both valves) 4.8 mm. Hypotype, left valve, from the same
locality as the specimen shown in Fig. 12. P. 393.

Fig. 17. Nuculana (Saccella) elenensis Sowerby. Reproduction of figure of Leda
elenensis Sowerby given by Hanley, Thes. Conch., Vol. 3, 1860, pi. 228
(Nuculidae, pi. 3), fig. 70. Panama.

Fig. 18. Nuculana (Saccella) elenensis Sowerby. Reproduction of figure of Leda
elenensis Sowerby given by Hanley, Thes. Conch., Vol. 3, 1860, pi. 228
(Nuculidae, pi. 3), fig. 71. Panama.

Fig. 19. Nuculana (Saccella) elenensis Sowerby. Reproduction of figure of Leda
elenensis Sowerby given by Hanley, Thes. Conch., Vol. 3, 1860, pi. 228
(Nuculidae, pi. 3), fig. 72. Panama.

Fig. 20. Nuculana cuneata Sowerby. Reproduction of figure of Leda cuneata
Sowerby given by Hanley, Thes. Conch., Vol. 3, 1860, pi. 228 (Nuculidae,
pi. 3), fig. 93. “Valparaiso.” [Not the record “New Guinea!”]. Dorsal
view. P. 403.

Fig. 21. Nuculana cuneata Sowerby. Reproduction of figure of Leda cuneata
Sowerby given by Hanley, Thes. Conch., Vol. 3, 1860, pi. 228 (Nuculidae,
pi. 3), fig. 92. Right valve. Same locality as cited for Fig. 20.

Fig. 22. Nuculana (Saccella) elenensis Sowerby. Length 15.5 mm., height 8.1 mm.

Hypotype, left valve, from the same locality as the specimen shown in
Figs. 12 and 16. View of the interior. P. 393.

Figs. 12, 14-19, 22, illustrate the variability of Nuculana elenensis. Fig. 14.

is a typical N. acapulcensis Pilsbry & Lowe but in view of the variability of

the species we have considered these all to be referable to N. elenensis.

All the specimens illustrated on the plate except those shown in figures

17, 18, 19, 20, 21, are in the type collection of the Department of Paleontology

of the California Academy of Sciences.

Plate II.

Fig. 1. Nucidana (Saccella) eburnea Sowerby. Length 12.4 mm., height 6.5 mm.

Hypotype, right valve, from Station 199-D-l, dredged in 16 fathoms
(29 meters) in Lat. 13° 18' N., Long. 87° 43' W., Meanguera Island,
Gulf of Fonseca, El Salvador. P. 395.

Fig. 2. Nuculana (Saccella) eburnea Sowerby. Length 11.2 mm., height 6 mm.,
thickness (both valves) 4.4 mm. Hypotype, left valve, from the same
locality as the specimen shown in Fig. 1. P. 395.

Fig. 3. Nuculana (Saccella) eburnea Sowerby. Length 13.3 mm., height 6.6 mm.

Hypotype, left valve, from the same locality as the specimens shown
in Figs. 1 and 2. View of the interior. P. 395.

Fig. 4. Nuculana (Saccella) laeviradius Pilsbry & Lowe. Length 7.8 mm., height
4.4 mm., thickness (both valves) 3.6 mm. Hypotype, right valve, from

430

Zoologica: New York Zoological Society

Station 145-D-l, -3, dredged in 4-13 fathoms (7. 5-2. 4 meters) in Lat. 26°
52' N., Long., 111° 53' W., Santa Inez Bay, east coast of Lower California.
P. 396.

This illustration shows a comparatively smooth specimen in contrast
to the one with strong concentric sculpture shown in Fig. 7.

Fig. 5. Nuculana ( Saccella ) gibbosa Sowerby. Length 36.8 mm., height 18.9 mm.,
thickness (one valve) 8.2 mm. Hypotype, left valve, from Station 213-
D-ll-17, dredged in 35 fathoms (63.7 meters) in Lat. 9° 44' 52" — 9° 42'
N., Long. 84° 51' 25"— 84° 56' W., off Ballenas Bay, Gulf of Nicoya,
Costa Rica. P. 395.

Fig. 6. Nuculana ( Saccella ) impar Pilsbry & Lowe. Length, 14.6 mm., height, 7.6
mm., thickness (both valves), 6.3 mm. Hypotype, right valve, from Loc.
23802 (C.A.S.), San Luis Gonzaga Bay, east coast of Lower California.
P. 396.

The character of the ribbing, closely spaced near the beaks, widely
spaced on the middle of the shell, then closely spaced near the base,
is characteristic of this species.

Fig. 7. Nuculayia ( Saccella ) laeviradius Pilsbry & Lowe. Length 6.0 mm., height
3.3 mm. Hypotype, right valve, from Loc. 23,802 (C.A.S.), San Luis
Gonzaga Bay, east coast of Lower California. P. 396.

Fig. 8. Nuculana ( Saccella ) gibbosa Sowerby. Length 37.5 mm., height 19.8 mm.,
thickness (single valve) 7.9 mm. Hypotype, right valve, from the same
locality as the specimen shown in Fig. 5. View of the interior. P. 395.

Fig. 9. Nuculana ( Politoleda ) polita Sowerby. Length 29.4 mm., height 14.0 mm.,
thickness (both valves, approximately) 9.6 mm. Hypotype, left valve,
from Station 197-D-l, dredged in 14 fathoms (25 meters) in Lat. 14° 16'
N., Long. 92° 03' W., 7 miles west of Champerico, Guatemala. P. 397.

Fig. 10. Nuculana ( Costelloleda ) costellata Sowerby. Length 22 mm., height 8.0
mm., thickness (both valves) 4.4 mm. Hypotype, left valve, from Station
143-D-l, dredged in 29 fathoms (53 meters) in Lat. 26° 58' 30" N.,
Long. 111° 57' 30", Santa Inez Bay, east coast of California. P. 398.

Fig. 11. Adrana exoptata Pilsbry & Lowe. Length 20.0 mm., height 6.0 mm.,
thickness (both valves, approximately) 3.0 mm. Hypotype, left valve,
from Station 195-D-21, dredged in 18 fathoms (33 meters) in Lat. 15°
44' 45" N., Long. 96° 06' 55" W., Santa Cruz Bay, Mexico. P. 409.

Fig. 12. Nuculana ( Costelloleda ) marella Hertlein, Hanna & Strong, sp. nov.

Length 32 mm., height 11.3 mm., thickness (both valves), 5.0 mm. Holo-
type, right valve, apparently from the Gulf of California. P. 399.

Fig. 13. Nuculana ( Costelloleda ) marella Hertlein, Hanna & Strong, sp. nov.
View of interior of left valve of holotype.

Fig. 14. Nuculana ( Thestyleda ) hamata Carpenter. Length, 12.0 mm., height, 7.2
mm. Hypotype, left valve, from Station 126-D-12, dredged in 45 fathoms
(82 meters) in Lat. 28° 20' N., Long. 115° 10' 30" W., east of Cedros
Island, Lower California, Mexico. P. 400.

Fig. 15. Adrana tonosiana Pilsbry & Olsson. Length 33.1 mm., height 9.3 mm.,
thickness (both valves), 5.4 mm. Hvpotype, left valve, from the same
locality as the specimen shown in Fig. 9. P. 412.

Fig. 16. Adrana elongata Sowerby. Length 49.8 mm., height 12.0 mm., thickness
(both valves) 5.5 mm. Hypotype, left valve, from Station 197-D-2, dredged
in 14 fathoms (25 meters) in Lat. 14° 13' N., Long. 92° 02' W., 7 miles
west of Champerico, Guatemala. P. 409.

All the specimens illustrated on this plate are in the tvne collection of the

Department of Paleontology of the California Academy of Sciences.

HERTLEIN & STRONG.

PLATE I.

MOLLUSKS FROM THE WEST COAST OF MEXICO AND CENTRAL AMERICA.

HERTLEIN & STRONG

PLATE II.

MOLLUSKS FROM THE WEST COAST OF MEXICO AND CENTRAL AMERICA.

Breder & Springer: Foetal Narcine brasiliensis

431

26.

On the Electric Powers and Sex Ratios of Foetal Narcine
brasiliensis (Olfers) .

C. M. Breder, Jr.

New York Aquarium

&

Stewart Springer

Bass Biological Laboratories

Although the small electric rays of the West Indies and Florida, Narcine
brasiliensis (Olfei's), are reasonably well known and recorded as giving elec-
tric shocks of some force, little seems to be known about their uterine
embryos. Bean & Weed (1911) describe and figure the young of a 17-inch
specimen which contained 14 foetuses and quote the reports of trammel-net
fishermen along the Florida Keys regarding the electric powers of adults.

Trammel-net fishermen operate near the New York Aquarium’s field
station on Palmetto Key, which is in the vicinity of Boca Grande; see Breder
(1939). They report that this ray is frequently taken in their gear. The
shocking powers of these fish are sufficient to cause the operators to leave
them in the trammel pockets when they clear their nets. Here they are left
to die, being removed later when the fishermen reach home and spread their
nets for drying.

One of these groups, the Spearing family, has been very kind in bring-
ing interesting specimens to the laboratory. On July 8, 1940, they brought
in a female Narcine of 295 mm. over all. It had aborted two premature
young in their bucket and subsequently died. Manipulation caused it to
extrude six more embryos. These were still in a living state and made the
usual feeble efforts of ray embryos to swim. Nothing exceptional was noted
in the manipulation incident to the extrusion, but when the young were
handled four of them were noted to be capable of giving electric shocks.
Although in a much weakened stage and not nearly ready to be born, this
shock was sufficient to cause the senior writer to drop them, although this
result was perhaps as much from surprise as from the current. The fine
tingling sensation was typical of an interrupted or alternating current. The
young were not all in a single stage of development. The range of their
condition may be estimated by the following figures of length of embryo
and diameter of yolks. The two largest, dead when received, and the two
smallest, showed no reaction. Whether the two smallest had not yet de-
veloped the ability or whether they were so weakened as to be unable to dis-
charge is not clear. The over-all lengths of the embryos were : male, 65, 67 ;
female, 65, 73, 76, 78, 78 and 80 mm. The diameters of the yolks, respec-
tively, were: 16, 14, 5, 6, 7, 8, 9 and 7 mm. The smallest female was ab-
normal in that the yolk sac appeared deflated and the ventral fins were
slightly asymmetrical.

432

Zoologica: New York Zoological Society [XXV:26 :1940]

One of us, Springer, handling similar material, detected no electric ef-
ects from the young, but in this case the operator is particularly insensitive
to electric shocks, while on the contrary the other one of us is more than
usually sensitive to such effects. In addition, moreover, Mr. L. A. Krumholz,
working at the same place with the latter, also felt the discharge.

The prenatal development of this ability to give off electric shocks sug-
gests interesting complications in the internal circuits of the mother and
young and causes one to wonder what the characteristics of the exterior
total current might be like. Also the question of why no discharge was felt
while the young were being forced out is of note. Suppression of or im-
possibility of discharge before birth, insulating or dissipating qualities in
the parent, suggest themselves as possibilities.

The following data were taken from material obtained from a trammel-
net at Punta Gorda Beach, July 7, 1939, giving total length and width in mm.

Males Females

Gravid

Non-gravid

T. L.

W.

T. L.

W.

T. L. W.

300

160

310

173

375 308

295

164

284

165

271 152

228

124

272

153

The one gravid female of 310 mm. in length contained four males 82 X
50, 79 X 42, 81 X 43, 81 X 43, and one female 84 X 45. The other of 284
mm. contained one male and three females of 78 to 80 mm. in length.

Sex Ratio of Embryos.

Length of Mother

Male

Female

% Male

284

1

3

25

295

2

6

25

310

4

1

80

432

9

5

64

16

15

52

It is apparent that while there seems to be a tendency for the young
to be predominantly of one sex in a given female, the young from so few as
four females taken at widely separated times and places, when combined,
closely approximate a 1-to-l ratio. It would also appear that small mothers
tend to produce more females while the large tend to a preponderance of male
offspring, although of course, these specimens are too few for a basis of
generalization.

References.

Bean, B. A. & Weed, A. C.

1911. An electric ray and its young from the west coast of Florida. Proc.
U. S. Nat. Mtis. 40: 231-232, pi. 10 and 11.

Breder, C. M., Jr.

1939. On the trail of the Tarpon. Bull. N. Y. Zool. Soc. 42 (4) : 98-110, 24
Illustrations.

Lowther: Study of Galago senegalensis moholi

433

27.

A Study of the Activities of a Pair of Galago senegalensis moholi
in Captivity, Including the Birth and Postnatal
Development of Twins.

Florence De L. Lowther

Barnard College, Columbia University

(Plates I-VI).

Contents.

Page

Introduction 433

Description of Galago senegalensis moholi. . 434
History and Classification — Paleontology. . . . 434

Distribution 437

Conditions of Observation 437

Functioning of the Structures Characteristic

of the Galago 438

The Head 438

a. Facial Expression, b. Lips. c. Nose. . 438

d. Eyes 440

e. Ears 440

f. Jaws. Teeth and Tongue 441

The Limbs 442

a. The Hand and its Action 442

b. The Foot and its Action 443

c. Volar and Solar Pads 443

d. The Hind Limb and its Action,

aa. Climbing, bb. Moistening of
Pads. cc. Jumping 443

The Reproductive Organs 445

Disposition 446

Page

Behavior of the adults 446

Feeding 446

Grooming 447

Calls 448

Sex Behavior 448

Nesting Habits 449

Breeding Season and Period of Oestrus.... 449

Galago activities 449

Comparison with Breeding Habits of

Other Lemuroidea 450

Discussion of Environmental Causes of

Oestrus Change 451

Period of Gestation 452

Birth and Number of Young 452.

Post-natal Condition of the Young 453

Nursing and Care of the Young 455

Parental Attitudes 455

Development of the Young Galago 456

Sleep 457

Acknowledgments 457

Bibliography 458

Introduction.

Galagos, small, elusive lemuroids of arboreal and nocturnal habits, are
found exclusively in Africa, where they are well known and widely distrib-
uted. Though the anatomical features of the many varieties of these animals
have been studied, little seems to have been reported concerning the ways in
which their structures function, nor is there any sufficient record of galago
habits and family life. The account is spotty and incomplete, partially at
least, because their nocturnal and tree-dwelling habits render observation
difficult in the field and in captivity. In a study of the galago in captivity,
an effort must be made to reproduce in some measure certain elements of its
native environment, such as perching structures, space for jumping and dark
places for retirement during daylight hours. The nocturnal habits of the
group necessitate the observer’s continuous occupancy of the animals’ night
quarters.

For more than three years, beginning September, 1937, I have made a
close study of a pair of Galago senegalensis moholi A. Smith, a subspecies

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[XXV :27

of galago found in abundance throughout the savanna country of central
and southern Africa. The opportunity to study these unique and interesting
forms was enlarged by the fortunate birth of twins in captivity, making a
family of four. Thus it has been possible to add, to the story of the behavior
and disposition of the adults, a record of the birth and development of a
male and female in captivity. While in the following discussion I have
emphasized observations of their habits and family life, I have also de-
scribed the functioning of those structures which are characteristic of the
galago.

Description of Galago senegalensis moholi.

The Galago s. moholi is a small, round-headed animal with a shortened
face; large, almost naked, membraneous ears with transverse ridges; round,
wide-open eyes with vertical pupils, and a short rhinarium situated high on
the face (PI. I, Fig. 1).

The body length including the head of the adult male is 6V2 inches and
the long, non-prehensile tail, which is used for balancing, 8V2 inches. The
front limbs are shorter than the long hind limbs. The foot, which is highly
specialized for jumping, has elongated calcaneum and navicular bones, pro-
viding adequate leverage for spectacular jumps.

The opposable huge hallux and less well developed pollex are primate
characteristics possessed by the galago, as are the small, and in this species
flattened, nails found on all the digits with the exception of the index toe.
Here a claw, characteristic of the Lemuroidea, is found.

The dental formula of the Galago s. moholi, like that of the true le-
murs, is

2 i
2

1 c 3_p
1 3

= 36 total

O

The two pairs of procumbent lower incisors are flanked by modified
canines, making the typical lemuroidean “tooth-comb.”

The neck of the galago is short. The pelage is soft, heavy and almost
chinchilla-like. The color of this subspecies is a mottled slate gray and silver
with a light brown wash on the dorsal surface of the posterior half of the
body. The ventral surface is largely white with slate gray on the deeper
portions of the fur, seen only when the fur is separated.

On the inner surface of both arms and legs and across the chest, the
pelage is clear yellow. There are white markings on the nose, around the
neck and lower jaw. Through the usual gray of the dorsal pelage, longer
black hairs protrude sparsely (PI. IV, Fig. 9). There are black markings
around the eye and along either side of the nose from the eye toward the
upper lip (PI. I, Fig. 1). The pelage is shorter near the hands and feet, and
more silvery in color.

There are no well defined sex differentiations. The adult male is slightly
larger than the female and the yellow of the pelage perhaps more marked.
The length of the tails in this group varies between 8% and 10 inches. (The
tail of the adult female under discussion is unnaturally short because of an
injury to the tip sustained at capture (PI. V, Fig. 12).

History and Classification — Paleontology.

In 1796 Etienne Geoffroy-St. Hilaire first described one of the “quad-
rumana” which the natives called “Galago,” found in western Africa in the
region of Senegal. A skull of this form was brought to him at the Museum
in Paris by Michael Adanson who had spent some years in exploration of

1940]

Lowther: Study of Galago senegalensis moholi

435

the Senegal, and a skin was later provided by the Due de Nivernais. The
addition of pictures and descriptions furnished by Adanson led Geoffroy
to the conclusion that this was a new type of animal which resembled in
many respects the makis (lemurs), the lorises, and tarsier. It differed from
each of these, however, to such an extent that he felt justified in calling this
a new species intermediate, perhaps, between the Loris and the Tarsius. He
determined to adopt for it the native name “Galago” to which he attached
“du Senegal,” perhaps to distinguish it from two other forms (one larger
and one smaller) which Adanson had reported to be present in that region.
He described1 it as possessing hind legs longer than the body, built for jump-
ing; the front legs short; the tail longer than the body; ears large and
membranous with transverse ridges.

Like the lemurs there were nails on hands and feet with the exception
of the second toe which had a claw. The hind foot was very long due to the
elongation of the calcaneum and scaphoid (navicular). The fur was grayish-
tan on the dorsal surface and white with yellow on the under surface. It
was also shorter on the hands and head while the under side of the hands and
feet and the ears and nose were naked. The animal was described as arboreal
and insectivorous. Geoffroy incorrectly stated that there were but one pair
of upper incisors.

In 1836 Sir Andrew Smith described a galago of similar size which he
found near the Limpopo River, Bechuanaland, South Africa, which closely
resembled Geoffroy’s senegalensis, but differed from it in two features.

1. On the ventral surface of the body the fur was white on the tips of
the hair only; near the body it was slate gray.

2. There were two pair of incisors on the upper jaw instead of one pair
as described for senegalensis.

On the basis of these differences he felt compelled to consider this a
new species and called it Galago moholi, which is the Bechuana name for
the animal in that locality.

“Had the upper jaw on this species not been furnished with four cutting
teeth I should have been disposed to have considered it as identical with Galago
senegalensis of Geoffroy.” (Page 42).

For some years thereafter, the few species of galagos known were
classified in accordance with the number of upper incisor teeth described
for each. When, however, in 1851 Isadore Geoff roy-St. Hilaire reported that
all galagos normally possessed two pair of upper incisors, the main reason
for separating Galago senegalensis from Galago moholi disappeared.

“Les Galagoides de M. Smith sont les especes qui n’auraient que deux
incisives superieures, et par consequent trente-quatre dents en tout. Tels seraient,
suivant lui les Galago senegalensis et demidoffii. Cette characteristique est
erronee; ces especes ont normalement quatre incisives superieures comme les
autres; seulement deux sont caduques, comme il arrive si souvent chex les
Lemurides.” (Page 80.)

The Galago moholi is now recognized as a sub-species of Galago senegal-
ensis. According to W. L. Sclater, 1900, (Page 23),

“This little Lemur is closely allied to the West African form with which it
has frequently been confounded, but an examination of the two species ( Galago
senegalensis in Paris and Galago moholi in London) shows that they are really
distinct.”

According to E. Schwarz (1931), the Galaginae fall naturally into two
genera: (1) Euoticus with pointed nails; (2) Galago with flat nails. Euoticus
is represented by a single species {elegantulus) . Members of the genus

1 The dimensions were given as follows :

Body length 6 inches 10 lines

Tail length 8 inches 4 lines

Head length 1 inch es 8 lines

Anterior limbs 3 inches 4 lines

Posterior limbs including feet 6 inches 11 lines

436 Zoologica: New York Zoological Society [XXV:27

Galago, however, are numerous and Schwarz divides them provisionally into
four species:

1. Galago crassicawdatus

2. Galago alleni

3. Galago demidoffii

4. Galago senegalensis 2

The Galaginae, because of their long tails and ability to jump, are called
“African long tailed Lemurs;” other anatomical details, however, relate them
more closely to the short tailed, climbing, loris-like forms of Asia and Africa.
The current classification of the Lemuroidea provides for two main divi-
sions, the Lemuriformes and the Lorisiformes (W. K. Gregory, 1916), sub-
divided as follows :

Suborder Lemuroidea3

Division A. Lemuriformes — all inhabitants of Madagascar
Family 1. Lemuridae
Family 2. Indrisidae
Family 3. Chiromydae

Division B. Lorisiformes — -Inhabitants of Asia and Africa
Family 1. Lorisidae4

Subfamily 1. Lorisinae

Arctocebus — West Africa
Perodicticus — West Africa
Nycticebus — N. India to Philippine Is.

Loris — S. India, Ceylon
Subfamily 2. Galaginae

Euoticus — Tropical and subtropical Africa
Galago — Tropical and subtropical Africa

Palaeontologically the time and place of origin of the Lorisiformes, un-
like those of the Lemuriformes and the Tarsioids, are obscure because of
total lack of fossil evidence. Dr. George Pinkley of the American Museum of

2 R. W. Hayman (1937) further emphasized the difference in nail structure among the Galaginae.
He demonstrated that crassicaudatus has concave nail ends while demidoffii Fischer, alleni Water-
house and senegalensis Geoffroy all have nails which are bluntly rounded at the ends with the
exception of Galago senegalensis inustus Schwarz. Because this form possesses pointed and keeled
nails he is convinced that inustus Schwarz should have the status of a new species.

3 The relation of the Lemuroidea to other primates is not the function of this article ; however,
a brief outline of recent trends in Primate classification may be pertinent.

Classification of
Pocock — Zuckerman
(1918) (1933)

Order Primates

Series 1. Strepsirhini
Suborder Lemuroidea
Div. A. Lemuriformes
Div. B. Lorisiformes
Series 2. Haplorhini
Suborder Tarsioidea
Suborder Pithecoidea

Dr. George G. Simpson is now (1940) revising his classification, the main outline of which, exclusive
of fossil forms, is as follows :

Order Primates
Suborder Prosimii

Infraorder Lemuriformes

Superfamily Tupaioidea (Tree or squirrel shrews)

Superfamily Lemuroidea (Malagasy lemurs and lemuroids)

Infraorder Lorisiformes (Lorises, pottos, galagos — no fossil forms)

Infraorder Tarsiiformes (Tarsius and fossil tarsioids)

Suborder Anthropoidea

While the relation of the Lemuriformes and the Lorisiformes is not changed, the term Lemuroidea is
relegated to that of a superfamily instead of the more inclusive one of suborder, and the term
Lemuriformes includes the primitive Tupaiodea as well. The Tarsioids, as in the case of the Allen-
Coolidge modification, are put back once more with the Prosimii.

4 Some authors rank the two subfamilies as families, Lorisidae and Galagidae, respectively. In
that case no subfamilies are indicated.

Allen-Coolidge modification of Pocock,
Schwarz. Zuckerman, as outlined by R. M.
and A. W. Yerkes, 1935
Order Primates
Division 1. Prosimiae
Suborder 1. Lemuroidea
Suborder 2. Tarsioidea
Division 2. Anthropoidea

1940]

Lowther: Study of Galago senegalensis moholi

437

Natural History, in the following memorandum to the author (1940) elabo-
rates this statement as follows :

“The occurrence of fossil specimens of the lower primates indicates that in
the geologic past the geographic distribution of these forms was singularly dif-
ferent from that of today. Specimens representing the Lemuriformes and Tarsi-
oids (but not the Lorisiformes — lorises, pottos, and galagos), are found in fossili-
ferous deposits in both Europe and western North America. They are first known
from middle Paleocene formations in America, seem to have spread over holarctic
regions and appear in late Paleocene deposits of Europe. They range through the
Eocene, disappear in the late Eocene in Europe but not until the early Oligocene
in America. Lower primates are then quite unknown in the fossil record until the
appearance of Lemuroids in the late Pleistocene of Madagascar, where they are
now confined. And Tarsius, living today in the Malay archipelago, is the only
surviving representative of the Tarsioids. This leaves a hiatus in the fossil record
of nearly two-thirds of Cenozoic time — about forty million years.

“It is furthermore remarkable that the paleontological history of the Lorisi-
formes is entirely unknown although living forms are widely distributed through
tropical Africa and the Indo-Malayan region. It is true that the French Eocene
forms Pronycticebus and Pseudoloris were formerly believed to represent fossil
Lorisoids, but more careful examinations indicate that they represent a Lemuroid
and a Tarsioid, respectively. There is good reason, from comparative anatomical
studies, to believe that the ancestry of Lorisiformes is approximately as ancient
as is that of Lemuriformes. It therefore seems probable that during the time when
the fossil record of Lemuriformes and Tarsioids was accumulating in limited
areas in Europe and western North America, early Lorisoids were living in some
other part of the world — possibly Asia or Africa.”*

Distribution.

Euoticus, Galago alleni and G. demidoffii are forest dwellers, while the
six varieties of crassicaudatus and ten varieties of senegalensis are found
only in the savanna country, a type of environment defined by botanists as
“grassland with scattered shrubs and medium-sized trees.” (J. Chapin,
1933, page 103). According to Shortridge (1934), typical senegalensis and
other races of the species range through Africa to as far north as Senegal,
Nigeria, French Equatorial Africa, the Southern Soudan, Gallaland, and
Somaliland and as far south as Inhambane in Portuguese East Africa, but
apparently not south of the Southern Transvaal. According to Sclater
(1900), Schwarz (1931) and Shortridge (1934), the subspecies moholi is
widely distributed over eastern, southeastern, southern and southwestern
Africa, in the north at least as far as the Tabora district in Tanganyika
Territory, possibly extending further northwest.

Conditions of Observation.

For the first two winters, I shared the same room with the adult pair
of galagos. During the day, the animals were confined in a cage 6X4X4
feet. On a shelf in the cage was a small sleeping box which provided the
needed darkness. At night for periods of two to five hours, they were lib-
erated and given the freedom of the room for exercise. A shaded light made
observation possible without distress to the animals. By placing food on
their feeding shelf, they were finally coaxed back into the cage. While shar-
ing a room with a pair of galagos had obvious disadvantages to the observer,
nevertheless, it offered the surest method of obtaining detailed data through-
out their active period. During the exercise period, all damageable articles
were covered with sheets. At frequent intervals the walls of the room were
washed down.

* According to Dr. W. K. Gregory the removal of Pronycticebus from the Lorisidae to the
Adapidae by Dr. G. G. Simpson (1940)’ deprives the Lorisoids of any known early Tertiary repre-
sentative.

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From June to September, the galagos were removed to the country,
where they enjoyed the complete freedom of a screened-in porch 10X14 feet,
adjoining my room. As usual, they slept in a sleeping box on a shelf. Sev-
eral times each night their activities were checked.

The birth of the young galagos occurred in the second spring, April,
1939, in the apartment room, under the most favorable conditions for con-
tinuous observation.

The third winter, when the young animals were half-grown, the incon-
venience of caring for so large a family made advisable their transfer to a
10X7 feet partitioned section of a heated greenhouse on the roof of Barnard
College, Columbia University. While conditions now prevent a constant
watch, some record of activity is made at varying intervals, at least three
times each night, twice by myself and once by the night watchman.

In the daytime the animals sleep fairly continuously. A darkened and
elevated recess in the cage, greenhouse or porch protects them from the
sunlight. Their quarters are kept scrupulously clean by frequent scrubbing.

In both the greenhouse and the screened-in porch, small growing trees
planted in tubs have been provided, as well as horizontal and vertical round,
wooden, exercise bars. One side of the porch is screened by half-inch mesh-
ing, large enough to admit moths, the galagos’ favorite fare. The moths and
other insects are attracted by a light with a reflector situated in the en-
closure.

Full-sized doors leading into both the greenhouse enclosure and porch
make it convenient for visitors to enter and watch at close range the behavior
of the galago family.

The Galago s. moholi seems to be hardy. It can tolerate a wide range of
temperatures from summer heat of more than 90 degrees in their sleeping
box to as low as 40 degrees on a cold winter night.

Functioning of the Structures Characteristic of the Galago.

It is my purpose at this point to supplement the outline of the known
features of the adult Galago s. moholi by further details based on my own
observations of the functioning of the outstanding structures which differ-
entiate this form. From time to time when possible I have attempted to
compare these structures with those reported of closely related lemuroids,
as well as with Tarsius, which, while not so closely related, displays many
interesting similarities and differences.

The Head.

a. Facial Expression, b. Lips. c. Nose.

The galagos, like the Lemurs, are extraordinarily expressionless — “no-
toriously blank,” as S. Zuckerman has expressed it. They cannot (A. Smith
to the contrary) make grimaces or even wrinkle the brow. They can merely
stare or droop their lids, move the ears and open the mouth wide enough
to expose the teeth.

This lack of facial expression is due in large measure to the immobility
of the upper and lower lips. There are two reasons for this condition: 1. the
middle of the upper lip is bound by the frenulum to the underlying premaxil-
lary area; 2. the lack of development of the labial and nasal muscles of the
facial field. Ernest Huber 1931, (page 23) who has made an exhaustive study
of facial muscles of mammals and primates, states that:

“The facial muscles of the Lemur play no role as musculature of facial
expression. Indeed we can hardly speak of facial expression in the Lemur.”

1940]

Lowther: Study of Galago senegalensis moholi

439

In common with all other Lemuroidea and many mammals, the galago
has a moist, glandular rhinarium with crescent-shaped nostrils, which E.
Geoffroy-St. Hilaire (1812) and R. I. Pocock (1918) call strepsirhine. This
structure extends into the labial area and becomes the phiitrum, or median,
attached part of the upper lip. Maxillary portions of the upper lip which
are covered with fur may terminate on either side of the naked phiitrum
as in the case of the lemurs, or, as in the Galago s. moholi (Pocock, 1918,
and J. D. Boyle, 1932) may meet in front of (over) the labial portion of
the rhinarium, a deep groove remaining to show the line of union. In this
way the upper lip is completely bounded by fur (PI. I, Fig. 1).

When the Galago s. moholi is angry and prepared for defense or offense,
it opens the mouth so wide that the labial area is stretched and the canines
and premolars exposed. There is no ability to draw up the lip in a snarl or
to make grimaces of any sort. The enlarged photograph (PI. I, Fig. 1) of
the face of one of the galagos shows the preliminary stage of rage. The
mouth is partially opened though not wide enough to show the teeth. The
eyes are focused on the cause of irritation located just above its head.

The sense of smell of the galago as of other Lorisiformes is well de-
veloped and still serves a more important function than in the higher
primates.

Le Gros Clark (page 177) states that:

“In general there is a close correlation between the degree of complexity
of the turbinate system and the acuity of smell.” . . . “In Lorisiformes the first
ethmo-turbinal is very large and actually covers over the maxillo-turbinal while
in Lemuriformes it is much smaller.”

The presence of a relatively large olfactory lobe in the brain is addi-
tional evidence that the galago and the lemurs, in common with the lower
mammals, retain their dependence on the sense of smell.

The galago places great dependence upon its sense of smell. Curiosity
is satisfied through this means. Any strange person, or object, is first thor-
oughly smelled, or food is approached nose first unless it is food-on-the-wing,
when it is seized first and smelled afterwards. The facial tactile vibrissae, or
“whiskers,” around the nose are said to be closely associated with the sense
of smell. Though they are not as well developed as in the genus Lemur, they
are still evident in some degree. The mystacial, mental and genal, are pres-
ent though more delicate and less obvious (PI. I, Fig. 1) than in Lemur.
According to Ernst Huber (pages 22-23), among the Lemurs:

“The mystacial and mental vibrissae jointly with the rhinarium, a highly
sensitive patch of mucous membrane at the tip of the snout, subserve the
tactile sense of the latter, which is closely associated with the sense of smell. . . .
This sense, which plays an eminent role in the life of the primitive, terrestrial
mammals, is of subordinate importance in the primates and has already become
deteriorated in the Lemuroidea. Undoubtedly in connection with their adaptation
to arboreal life, the sense of touch, with the aid of the facial tactile vibrissae, is,
however, of the greatest usefulness to these primitive primates in their life amidst
the branches of trees. Indeed, the sense of touch together with the highly devel-
oped sight and keen hearing are the guiding senses of the lemur.”

In my judgment the sense of smell should be added as an important guiding
sense.

The mouth and nose of the Tarsius are strikingly different from those
of the galago or other lemuroids. There are no crescent-shaped nostrils with
moist rhinarium extending into the upper lip. Like higher primates, the
nostrils are oval and there is only a remnant of the moist area about these
apertures, a condition which Pocock (1918) considers of such fundamental
importance that in the classification of the primates he uses this feature
as a basis for primary subdivision. He calls the lemuroids “Strepsirhini”
and all other primates including Tarsius, Haploi'hini.

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Zoologica: New York Zoological Society

[XXV :27

d. Eyes.

The large, round eye bulges forward, giving a conspicuously rounded
appearance to the surface. In the daylight the entire surface of the eye is
colored a homogeneous light brown. Since the whole visible portion of the
eye is covered by the rounded, transparent cornea, no portion of the sclera
can be seen. The pupil, as A. D. Bartlett (1863) observed long ago in the
case of Galago crassicaudatus montieri, is vertical (PI. I, Fig. 1), a condition
common to Lemuroidea and to many nocturnal mammals. Tarsius, however,
differs from the galago in that the pupil is horizontal. A recent photograph
which appeared in Collier’s Magazine (August, 1939) of Professor John
Fulton’s two living specimens at Yale University, shows the pupil in the
light as a horizontal slit, although in their other respects the eye is strik-
ingly similar.

Under the hand lens, the vertical pupil of the galago eye appears to be
slightly rounded at each end and somewhat wider in the center. Fine dark
lines radiate from the rim of the slit into the iris. The spacing of the lines
seems to vary with the intensity of the light. When the light is bright and
the slit narrow the lines seem crowded together. In a dimmer light, the slit
is larger and the radiating lines farther apart. A slight scalloping of the
edge of the slit between the lines is apparent. At night in a dim light the
iris opens to a prodigious extent, leaving only a small portion of the brown
visible as a rim. By reflected light the pupil is brilliant orange (like the
lemur). The eyelids, which are kept wide open, have lashes on the rims
visible on the upper lid of the right eye in the enlargment (PI. I, Fig. 1).
It is difficult to see the lid except when the animals are sleeping. We suc-
ceeded in getting a photograph of a form with extended lid after it had
been temporarily blinded by a flash-light bulb which it was examining when
the light was discharged. The photograph clearly shows the upper lid cov-
ering the major portion of the bulging eye (PI. II, Fig. 3).

The eye is so large that the extrinsic muscles are unable to move it. In
compensation the head moves flexibly as a whole in quick motions from side
to side and sometimes in an elipse. The head may be bent to the side at such
an angle that one eye is placed almost vertically above the other. It may also
turn practically 180 degrees so that the nose is directly over the spine and,
as Cooke (1939) has noted in the case of the Tarsius, the animal frequently
moves its head in this manner to look upward. The galago does this when it
is clinging to some vertical or inclined branch or when clinging upside down
to a horizontal branch. The ability to turn the head in a direction opposite
to that which the body “faces,” a convenient adaptation for any perching
animal, relieves the necessity of lessening the hold on the perch.

e. Ears.

The membraneous ears are large and almost naked on the inner and
outer surfaces except at the rim where there is a sparse covering of fur
(PI. I, Fig. 1). The inner surface, as E. Geoffroy-St. Hilaire first and
A. Smith later noted, is marked by four transverse ridges which seem to
terminate in an outer vertical ridge (PI. I, Fig. 1). An outstanding galago
pecularity is the ability not only to move the ear as a whole forward, later-
ally or upward, but also to fold the pinna along the ridges somewhat like an
accordion, throwing its outer tip back over the upper transverse ridge. In
addition to the accordion-like folding, the whole pinna may be further
pressed back against the head so that all evidence of large projecting ears
disappears.

The ears of the galago are perhaps its most expressive feature. They
move singly or in unision in the direction of the least unexpected sound.
When looking over a new object or person the ears seem as alert as the nose
and the eyes. In sleep there is a tendency to fold the pinnae along the

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ridges (PI. II, Fig. 2, lower center) and in anger there is also some ten-
dency to corrugate them (PI. II, Fig 2, right). The final flattening of the
folded pinnae against the head occurs when the animal enters an enclosed
space. When seeking food which I have placed in the cupped palm of my
hand, the galago thrusts forward both head and hand, invariably flattening
the ears against the sides of the head. This is an admirable device for the
protection of large and delicate ears from the dense foliage and perhaps also
from the perils of battle.

Tarsius ears, though much smaller relatively, seem to be of a similar
type, according to Ernest Huber (page 27), and the musculature which
moves them is highly developed.

f. Jaivs, Teeth and Tongue.

The round head and short face are characteristic of the several varieties
of Galago senegalensis and demidoff ii. The upper jaws and the snout with
the crescent-shaped nostrils do not project forward to the extent that they
do in Galago crassicaudatus or in the Malagasy lemurs. The lower jaws,
which are also short, bear the usual two pairs of procumbent incisors
flanked by the elongate modified canines that project forward almost hori-
zontally and have been known as the “tooth-comb” of the lemurs. The teeth
of this incisor-canine complex of the lower jaw converge at their tips so
closely that, as M. Russell Stein (1936) points out in the case of the true
lemur, the term “comb” is probably inaccurate. “Scraper” is perhaps a more
accurate term. The* first premolar of the lower jaw has become modified into
a somewhat caniniform structure though less so than in the true lemurs. By
working against the canine of the upper jaw, this tooth acts as a shearing
or a nipping apparatus. The remaining molars and premolars are more ad-
vanced than those of the typical insectivorous lemurs as Le Gros Clarke
(1934) has recorded, since thet third premolar of the upper jaw is more
molarized and the first two molars are quadri-tubercular, having one more
cusp than those of the typical lemurs. The two pair of upper incisors, sepa-
rated by a space in the center, are so small that it has been suggested they
are functionless. Because of the strong roots possessed by these teeth in the
case of the true lemurs, M. Russell Stein questions this idea. Can these
upper teeth function with the lower horizontally placed incisor-canines or
can they be used in any other way? I can confirm the statement made by
Stein that the upper incisors are functional. The galagos in my possession
will frequently bite when handled, and several times the bite has been suffi-
ciently severe to leave a clear imprint of the tooth marks. In each instance
the imprints of the upper canines and the two pair of incisors which lie
between have been evident, as well as the imprint of the pair of lower can-
iniform premolars and the scraper itself.

The question of the use of the incisor-canine complex or scraper is an
interesting one. In an article published in 1939 on “The Feeding and Groom-
ing Habits of the Galago,” I attempted to show that the scraper was used
as a toilet or grooming structure. Since then I have experimented further
to determine whether, as suggested by Stein, this device is ever used for
scraping fruits and biting off leaves. On placing a pear upon the feeding
shelf, I found that the scraper was never used. The galago attacked the
pear with the canines of the upper jaw and after making a hole in the skin
it then inserted its long, thin tongue into the aperture and licked out what
it wanted from the interior. Again, when a tree was placed on the porch
some of the smaller branches were bitten off. In each case the canines and
the first premolars of the lower jaw were used, never the incisors. The gen-
eral conclusion is that the scraper of the lower jaw, in the case of the galago
at least, is essentially a grooming implement though it has not yet been
proved that it may not act upon occasion as a food-scraping device.

* Homologous with P2 of the primitive mammalian dentition.

f P4 superior of the primitive mammal.

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The long, thin tongue, like that of the true lemurs, is used for licking and
lapping. Although of somewhat limited movement, its action is straight
forward over the incisor-canine complex of the lower jaw which acts as a
trough. The tongue is long enough to reach the moist rhinarium over which
it can curve.

It is interesting to note that Tarsius also uses its tongue as a licking
and lapping structure.

The Limbs.

a. The Hand and its Action.

It is well known that the fourth digit of the hand among the Lemuroidea
is the longest of the five. In this characteristic, they differ from all other
primates, including Tarsius, and from Mammalia generally where the third
digit is the longest. The hand digits of Galago s. moholi as a whole are
relatively long. The thumb is short and separated from the index finger b'y
a considerable interval; the index finger is also short and separated from
the third digit by a less marked interval (PI. Ill, Fig. 6). The third and
fourth fingers grow progressively longer with the fifth somewhat shorter.
The last three digits are more closely placed (PL III, Fig. 4).

In 1838 W. Ogilby demonstrated at a meeting of the Zoological Society
of London the structural peculiarities of the hand of a living specimen of
galago, then known as Otolicnus garnetti. This peculiarity consisted in the
partially opposable character of the index finger of the hand. It was shown
that the thumb and the index finger could be used as a unit to grasp one
side of a branch and that the remaining three fingers were used on the other
side, much as the koalas would do. Mr. Ogilby remarked, “The anterior
index finger in all the ‘inferior’ Lemuroidea is weak and powerless and it
has the same tendency to divide with the thumb instead of with the other
fingers in the rest of the galagos as well as in the Nycticebi, the Microcebi,
the Cheirogalei, and Tarsii whilst in the Potto it is reduced almost to a
tubercle.” In watching the action of the anterior digits of the galago, I find
that my observations do not agree with Mr. Ogilby’s in all respects. The
index finger lies almost midway between a large space which separates the
thumb from the third' finger (PI. Ill, Fig. 6) and, in grasping, it is just
as likely to be found with the third, fourth and fifth digits as with the
thumb. Its disposition depends somewhat on the size of the object grasped.
If it is small enough to fit in the space between digits two and three, digit
two is then likely to be found on the side of the thumb (PI. I, Fig. 1) ; if,
however, the object or branch is large, the index finger then lies almost
between the two. In any event the index finger is so small that its effective-
ness as an opposable structure is very limited.

An interesting feature of the digits of the hand is the size of the indi-
vidual phalanges of all but the thumb. In each instance the proximal phalanx
is long and the second one is slightly less so while the third or distal phalanx
is minute and very little larger than the nail. The relative length of the
phalanges is an admirable adaption for grasping limbs and small branches
of trees. Two peculiarities, however, are evident because perhaps of the
elongation of the proximal elements. 1. When the galago places its hand
upon a flat surface, the palm of the hand with its pads rests flat upon the
surface; the digits, however, are not extended straight forward but are
flexed, the pads on the distal ends of the digits alone resting upon the
surface. There seems to be a flexure at the knuckles which permits the rais-
ing of the proximal phalanges away from the ground. Furthermore, the
joints between the first two phalanges instead of pointing directly upward
are bent to the side, even the index finger may be involved (PI. Ill, Fig. 4).
A recent motion picture of Dr. John Fulton’s live Tarsius shows the same
flexed condition of the digits when the hand rests upon a flat surface. 2. A

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second peculiarity is to be noticed when the galago gx-asps a piece of food.
Among the other primates the digits flex over the object at the knuckles
and the palm of the hand is involved. In this instance, however, the proximal
phalanx of each digit (the longest) remains straight on a plane with the
back of the hand, and the gx-asping is done by flexing the two distal phalanges
over the proximal. The thumb acts in opposition and the palm is not covered
by the digits (PI. Ill, Fig. 5).

b. The Foot and its Action.

The enormous hallux of the foot is widely separated from the other four
toes which act as a unit (PI. Ill, Figs. 6, 7 ; PI. IV, Fig. 8). The fourth digit,
in this instance like that of Tarsius, is again the longest of the five, while
the second toe is short and, as in the case of all other lemuroids, beai’s a
flexed claw (PI. IV, Fig. 10). All the other digits of both hands and feet
have nails which are flat with relatively straight edges (PI. Ill, Fig. 4)
(A. Smith, 1836). The great distinction of the foot of the galago as com-
pared with other lemuroids is the extraordinax-y elongation of the navicular
(scaphoid) and the calcaneum first noted by E. GeoflToy-St. Hilaire. The
digits and distal portion of the metatarsals ai’e the only elements which
rest upon the ground. The proximal pox'tion of the metatax-sals ax-e well as
the tax-sals are always off the ground; in fact, as Pocock (1918) has ad-
mix-ably shown, the ventral aspects of this ax-ea of the foot are covered
with fur and are usually raised some distance from the ground (PI. Ill,
Fig. 7; PI. IV, Figs. 8, 9). The total length of the foot is 2 to 21/4 inches,
about half the length of the hind leg. It is this unusual lengthening of the
tarsal segment which makes possible the extx-aordinax-y leaps chax-acteristic
of this fox-m.

c. Volar and Solar Pads.

The well developed pads on the palm and the sole of the galago are
quite px’imitively disposed iix a chax-acteristic manner (PI. Ill, Figs. 6, 7).
There are two px-oximal pads, the thena and the hypo-thena, and four distal
intex-digital pads. In both hand and foot the thena pads tend to fuse some-
what with the first iixterdigital and the hypo-thena with the fourth inter-
digital. The intex-digital pads of the foot are not all of equal size, the
third being much smaller than the others.

Pads also are to be found on the toes aixd fingex-s (PI. Ill, Figs. 4, 6, 7).
Along each digit between the palm or sole, and its tip, a long, thin pad is
found which enlax-ges at the tip. Although this enlax-gement or pad is not
equal in size to that found on the tip of the Tarsius digits, it is, howevex-,
an effective sux-face. The big toe has a broad distal and px-oximal pad. The
first intex-digital which follows is a round, conspicuous pad, protruding be-
yond the confines of the digit aixd the sole (PI. Ill, Fig. 7). Oblique papil-
lax-y ridges are to be seen on the pads. Dr. Charlotte Wolff (1938) has found
“unique scale-like roughness” in two species of galago ( moholi and crassi-
caudatus ) as well as in Loris and in the genus Lemur. She describes them
as more expressed on the sole than on the palm and are found in the center
of the palm and sole aixd on the lower phalanges but not on pads. She also
finds that all the terminal phalanges have tactile corpuscles.

d. The Hind Limb and Its Action, aa. Climbing,
bb. Moistening of Pads. cc. Jumping.

Not only is the hind limb vex-y long, but its strength is also enormous.
The animal can hold on by its hind feet alone (PI. IV, Fig. 9) ; it can lower
itself down into a jar, or suspend itself fx-eely, head downward. By the power
of the muscles of its hind legs, combined with the strength of the muscles
of the foot, it can lift itself once more to its ox-iginal position without the
aid of the anterior limbs. One of the tamer individuals box-n in captivity will

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grasp a finger of my hand by the huge opposable big toes and the other
digits of its hind feet and will let itself down into a paper bag suspended
from my hand. The strength of the grasp of the large toe and the other
digits upon my finger seems out of proportion to the size of the animal.
There seems also to be a particular pressure exerted by the hallux and the
inner surface of the sole of the foot. This is the area where the interdigital
suction pads are especially well developed. The added pressure exerted at
this point flattens out the adhesive pads and makes them more effective
as a clinging device. The use of the pads as an aid to clinging is shown in
the illustration (PI. Ill, Fig. 6). This photograph was taken from the inner
side of a glass door on the outer side of which the galago was clinging. The
pads of the hind feet press against the flat surface of the glass, obliterating
the spaces between the pads and show the way in which the large pads of
the big toe and the first interdigital adhere firmly to the glass surface. This
explains the animal’s ability to climb up and down any perpendicular surface
which offers an edge, such that the thumb and great toe can get a purchase.
The galago always descends head first.

Grasping and climbing are facilitated not only by the elongation of the
fourth digit over that of the third and by the strength and opposability of
the very large, widely abducted hallux, but also by the moistening of the
pads of both hands and feet. The latter action is one characteristic of the
galago which is accomplished in a definite way at frequent intervals through-
out the active period, whether climbing or sitting still. By placing the palm
of one hand under the urethral aperture, the animal collects a drop of urine.
It then rubs the moistened palm of the hand on the sole of the foot of the
same side, repeating the action with the other hand and foot. This auto-
matically moistens the naked surfaces of the pads and renders them more
effective surfaces of adhesion. No one seems to have recorded this action
except E. G. Boulenger, who, in his popular book, “Apes and Monkeys,”
states that the galago has “a strange habit of moistening the palms of the
hands and the soles of the feet at frequent intervals when climbing” (p.
206), although he gives no clue as to the way in which this is accomplished,
nor does Mr. Boulenger record this as a common practice even when the
galago is sitting quietly.

The galago, essentially a hopping and jumping animal, normally perches
in high places (PI. IV, Fig. 8). Although it uses all four feet for climbing
and usually for perching, it does not normally run or walk on all fours. The
structure of the posterior limbs is responsible for the characteristic hopping
and leaping motions. When jumping on a horizontal plane this form can
span as much as six feet. Vertically it can spring upward as much as five
feet, landing on a perch with unerring precision. When jumping downward
diagonally from a higher to a lower perch much greater distances are
covered.

As an example, one of the adults sprang from the balcony rail of a
studio room to the top of an open door in the room below, a distance of
twenty feet. Although the thickness of the door was only two inches, the
landing was made with precision. When, and rarely, the adult galago makes
an imperfect landing, it seems to be due to an insecurity of grasp rather
than inaccuracy of gauging distance. The speed and extent of the jump is
particularly great in this variety and exceeds that of the larger species
( crassicaudatus ) .

The preliminary stages of jumping are expressed by a crouching of
the body. The hind feet are placed with heels together off the ground and
toes out. The spring is made with great speed and the feet are kept in the
same relative position as attained when taking off, though somewhat farther
apart. From a front view of the approaching animal the soles of the feet
can be seen slightly up-raised. The heels are no longer touching, but may

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remain relatively close together. This explains the ability of the galago to
jump even though his legs are bound together above the heel. Donald Carter,
in his field notes of his trip in South Africa (1938), recorded the following:

“Balovale, September 4. — A native brought us four (two males and two
females) galagos alive in a gourd. Upon removing the wad of grass which served
as a plug for the hole, one of the animals jumped out landing on the table.
From there it jumped a good six feet to one of the posts supporting the tent
and without a moment’s hesitation he took another jump to the trunk of a small
tree under which the tent was pitched. This jump must have been eight feet. Up
this tree he scrambled and hid among the boughs. I sent a boy up the tree and
with some difficulty the animal was dislodged and jumped to the ground where
he was pounced upon by about five small boys who were waiting underneath. It
was not until he was retrieved that I noticed that the animal had his two hind
legs securely tied together at the ankles by a piece of grass. Hampered as he
was he easily and gracefully made these two jumps landing both times where
he had planned.”

Before taking off to a new position the eye of the galago always gauges
the distance to be covered quickly and carefully. The jump to a new perch
may involve a diagonal or backward leap. The animal, however, always
manages to face the new perch on landing, though it may mean a 180
degree turn in mid-air.

Occasionally the galago will stand up straight, balanced on its toes with
the hind limbs not Hexed. In such a position, it immediately becomes quite
tall (11 inches) and of a short-waisted appearance because of the relative
length of the hind legs and feet (PI. IV, Fig. 10). The tail, from eight to
ten inches long and somewhat bushy at the end, serves as a balancing struc-
ture when jumping or when standing upright. In the latter action the tail
moves up and down to help maintain balance. It may also serve to protect the
eyes from the light while sleeping, though it is more frequently curled over
the breast at such times. While the tip of the tail continues to be slightly
curled when the animal first awakens, it straightens out as soon as the galago
becomes active.

The Reproductive Organs.

R. I. Pocock in his admirable article, “On the external Characters of
the Lemurs and Tarsius,” (1918) has described in detail the structure of
both male and female reproductive organs of Galago senegalensis, crassi-
caudatus and montieri as well as other forms of the Lemuroidea. The pres-
ence of baculum and spines on the penis seems to be a condition common to
all male Lemuroidea but invisible of course in the living state.

The male in conformity with the lemuroid type (except the genus Lemur)
has an external scrotum covered with fur. The penis is relatively short with a
long, inverted tip. Recently, just before the period of heat of the female,
(December, 1939) I saw the everted tip of the penis. Its shape differed from
those described by Pocock for crassicaudatus, senegalensis and demidoffii.
The everted portion was a relatively narrow cylinder, over an inch in length,
which expanded suddenly into a wide bulb at the tip. As I watched, the bulb
suddenly contracted with startling speed to almost the dimensions of the
rest of the penis. As I was fully six feet away from the cage when I saw
this structure, it was not possible to see whether the bulb after contracting
formed a structure comparable to the “frill” which Pocock described as
present in the case of the tip of the penis of Galago crassicaudatus.

The female Galago s. moholi, as Pocock has shown in the case of galagos
of other species, possesses a long, pendulous clitoris, at the terminus of
which is the urethral aperture. This peculiarity makes it difficult to deter-
mine the sex of the individual as the structure resembles superficially a
penis in shape and length. The inverted tip of the penis and the presence of
the permanent scrotum, however, differentiate the male.

446 Zoologica: New York Zoological Society [XXV:27

Disposition.

The galago is a very quick and nervous animal, which responds to the
slightest unexpected noise or movement. It will crouch in terror and then
spring away from the direction of danger with lightning speed. Like most
lemurs, though easily tamed, it is somewhat truculent and uncertain in its re-
lations to humans. It resents handling. Balancing on its haunches, it will hurl
itself forward, striking out first with both hands like a diminutive prize
fighter, and then grabbing hold of the object. Since the nails are flat, this is a
harmless gesture. But this attack is followed by a quick forward thrust of
the head and a very nasty bite may be inflicted by the sharp canines and the
first premolars of the lower jaw. At times even the scraper is involved.
During the attack the animal utters a querulous chatter. The truculence of
the galagos seems to be directed solely toward humans. I have never seen
them fighting among themselves. Even the young ones are independent and
dislike handling, though they will tolerate a gentle rubbing of a finger along
the jaw or behind the ears. The presence of several people does not disturb
them providing no quick movement or noise is made. Completely fearless of
humans, the twin galagos born in capitivity enjoy jumping on a shoulder
or sitting on the top of someone’s head.

The galago is as inquisitive as a monkey. Unlike that animal, however,
curiosity manifests itself by smelling instead of handling the strange ob-
ject. The galago will investigate the face and wearing apparel of even a
total stranger. Anything new brought into the cage or enclosure is always
smelled carefully.

I have attempted to show in another article (1939) that Galago crassi-
cauclatus, when alone, and dependent upon humans for companionship, was
a relatively affectionate animal which treated me much as it would a fellow
galago. Without doubt the Galago s. moholi would be more demonstrative
to humans were it dependent on them for companionship. It cannot be said,
however, that this group of four display any real affection beyond their
own circle. Their interest in me, I suspect, is due to the food I supply. As a
group they play together in utmost harmony. They will pursue each other,
roll about, and playfully grapple. At times they will hang suspended from
a horizontal stick by their hind feet the while beating at each other with
their free arms like two inverted pugilists. At feeding time it is a question
of each galago for himself, although when the babies were beginning to eat
and for a few months thereafter, their mother would sometimes give up
part of her catch to a hungry young one who had jumped up beside her to
help itself. Food in the possession of other galagos is much more attractive
than any food in a plate. They will spend a great deal of time pursuing
succulent morsels in the hands of one of their companions, even though
similar morsels may be present for them. There seems to be no resentment
on the part of the pursued if its prize is taken from it. The galago imme-
diately proceeds to recapture the diminishing tid-bit.

Behavior of the Adults.

Although the behavior recorded in this section has been observed while
the galagos were in the state of captivity, nevertheless the conditions under
which they were kept have approximated, so far as is possible, their natural
environment. Hence, I am justified in feeling that this general behavior
does not differ fundamentally from that of the wild state.

Feeding.

Observations of this variety of Galago confirm the recorded statements
that while they are insectivorous, they eat other things as well. Live moths,

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grasshoppers and meal worms are great favorites. Milk, a constant in their
diet, has never been refused. Vitamin B and cod-liver oil have been added
occasionally to milk, especially after the birth of young. Sweetened fruit
juices and melted ice cream are relished. Although water has frequently
been placed before them, there is no evidence that they drink it. They will
eat various kinds of thinly sliced raw vegetables and fruits at any time;
more freely in the winter months when insects are difficult to obtain. They
will also eat buttered bread or bread spread with honey in which wheat germ
meal and rice coating flour are mixed. When cut flowers from the garden
are available, such as bergamot and honeysuckle, they will bite off the heads,
pull apart the petals and chew at their bases.

In the case of live food, as Geoffroy (1796) first recorded, the galago
fixes its gaze intently on the insect. It never attempts to jump at it in mid-
air; instead, it leaps up to a perch close beside the insect. Then holding on
with its hind feet, the galago reaches out and grabs the moth with one or
both hands. After putting the live moth in its mouth, it jumps away to a
secure perch somewhat removed from its companions. Sitting up and holding
the insect in one or both hands and closing its eyes to protect them from
the frantic beating of the insect’s wings, the galago begins at the head and
consumes all but the wings. This is the same procedure which Cooke has
recorded in the case of the Tarsius (1939). The wings are discarded by using
the tongue to shove out over the scraper, which thus seems to be passively
used as a trough. I have frequently placed moths in a wide-mouthed jar cov-
ered with a loosely fitting lid. The galagos quickly learned to push off the lid
by using the tip of the nose. Standing up, stretched to full height, one will
peer in and leap to the rim with all four feet bunched together. Then liber-
ating the front legs but still holding on with the hind feet, the galago will
dive in head first, locate and grab the insect with one hand and haul itself
back by the power of its hind legs without loss of balance. Even when there is
plenty of food flying about, the galagos tend to grab it from each other.

Vegetables and other inert food are always smelled carefully before
sampling. If acceptable, the animal takes a small piece directly in the mouth,
jumps away, and holding the morsel in one or both of its hands, bites off
what it wants and drops the rest. Liquids are usually lapped up by the long
tongue; at times, however, the hand is plunged into the fluid and subse-
quently licked dry.

Feeding takes place regularly. In the morning milk is placed in a shallow
container close to the sleeping box. By the end of the day most of this has
disappeared. At about 8 o’clock in the evening various foods and a second
supply of milk are provided and left available throughout the night. The
animals which have been active for some time are now hungry. They rarely
consume a whole piece of vegetable or fruit; their habit is to take a few
bites and drop the rest. In the morning the enclosure is strewn with frag-
ments.

Throughout their active period, fecal elimination is very plentiful; the
more so when they first become active at night. Urinary elimination is also
plentiful and frequent, and usually independent of moistening the foot-pads,
which has already been described. Though they rarely soil their sleeping
box, they seem to have little further concern about the place for elimination.
If handled during the day, they express fear or nervousness by immediate
urinary and fecal elimination.

Grooming.

The galago is an unusually clean animal. Even its hands when soiled by
food or foreign substance are carefully licked clean. General grooming in-
volves the frequent use of the procumbent scraper, the tongue, and the oc-

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casional use of the claw which serves as a scratcher for such spots, other-
wise inaccessible, as behind the ear.

The chief grooming instrument, the procumbent scraper, long called
the “comb,” is used by the galago in vigorous action on his own pelt or on
that of one of his companions. This action is a series of quick thrusts through
the fur deep down to the integument. By this scraping action the animal
can dislodge and remove any dead skin or foreign substance in the pelt
(Lowther, 1939). The tongue completes the process of grooming with a
thorough gentle licking.

Begun on first awakening, before the start of much activity, the groom-
ing is repeated for short periods frequently during the active period. Groom-
ing may be a mutual operation. I have seen two galagos grooming each other
simultaneously; and two young ones may work on one of their parents at
the same time. As for the action of grooming on the part of a galago of
another species ( crassicaudatus ) with no companions other than myself,
I have had personal experience. This animal frequently perched on my arm
or hand and vigorously dug into my integument with its scraper as though
I were a fellow galago. The intensity of the digging and scraping made it
clear that any foreign object in the fur of a galago could be effectively
removed. While the present variety of galago has not bothered to attempt
much grooming of their mistress, the young ones occasionally have repeated
the action of the former galago, particularly after licking off some sweet
morsel adhering to my fingers. At times they have used the scraper perhaps
to get off the last remnants of sweetness.

Calls.

The galago has several notes characteristic of different conditions,
clearly differentiated but difficult to describe. 1. The alarm note; a shrill
sound on a high pitch, which starts somewhat like a chipmunk’s scolding
note, though shriller, but which ends in a whistle. This note quiets the group
into frightened stillness. 2. An automatic cry, the cause of which is un-
known, and which may continue for an hour. It is a piercing noise with
two pitches, high and low. While the animal is making this noise it will
continue its activity, eating and jumping. It does not affect the others. 3.
The low clucking note like the brooding hen, but on a lower register; used
when annoyed. 4. The sex note, used by the male when pursuing the female.
This is a soft questioning sound of two notes. 5. The conversational note;
when separated, both male and female call to each other. This note is softer
than the sex note but has the same two pitches, high and low. 6. A chattering
note used by the female as an expression of annoyance at the attention of the
male. 7. The maternal note; a very gentle, soft, caressing sound, used by the
mother when talking to the young in the nest. 8. The squeak of the young ;
suggestive of young mice. It might possibly be called a squeaky chirp.

Sex Behavior.

The female of this form will accept the male only during periods of
oestrus, which may last as long as five or six days. At this time she has a
colorless discharge which perceptibly excites the male. He constantly smells
her genitalia and just before the act of copulation the male is likely to lick
the female with his long tongue. During the period of sex activity copulation
has been observed to occur three or four times a night and it has also been
seen in the morning when the animals would normally be sleeping. During
the long periods between oestrus, the male continues his interest, although
the female refuses to accept him. As he pursues her around the cage or
enclosure, he utters a soft, plaintive call which I have described as the

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“questioning sex note.” At times the female merely keeps one jump ahead
of the male and seems undisturbed by his attentions. If, however, they are
continued for any considerable time, she finally turns upon him in annoy-
ance with a chattering cry. This usually effectively discourages his atten-
tions. Once, however, I saw the female become so angry, after chattering
at the male without effect, that she turned upon him with such fury that
he took to his heels in alarm. Generally speaking, the adult pair is a friendly
and affectionate couple. They are frequently found perching side by side,
grooming each other, wrestling and at times embracing (rubbing noses).
During the period of pregnancy the female’s disposition became far more
truculent. The continued attentions of the male were repelled with increas-
ing vigor and shortness of temper.

Nesting Habits.

The galago in the wild is in the habit of nesting in the hollow places
among the tree-tops, according to E. Geoffroy-St. Hilaire, Pitman, Short-
ridge, and others.

E. Geoffroy, 1796:

“Ils nichent dans des trous d’arbres ou ils preparent a leurs petits un lit
qu’ils tapissent d’herbes.”

Pitman, 1934:

“I have often come across the leafy beds these animals prepare for expected
offspring at the bottom of hollows in trees.”

Shortridge, 1934:

“Galago moholi granti is strictly nocturnal, sleeping during the day in
hollow trees where it may generally be taken in small family parties.”

In captivity, I have found that these animals show a tendency to tear
up bits of available paper and to carry them into the sleeping box. While
on the porch where a growing apple tree is situated, they bit off the ends
of leafy twigs and carried these to the box up under the eaves of the porch
roof. They would also gather bits of string, wool, heads of flowers and
almost any other small soft object which happened to be loose. Just before
the birth of young ones, the female was particularly active about the con-
struction of a lining for the sleeping box. The galagos obviously prefer a
darkened area in which to sleep during the day. Whenever a box was given
them they used it rather than some secluded corner of the cage or porch.

F. Wood-Jones, p. 116 (1929) states that:

“Nest-building is a habit that is widespi'ead among the Lemurs, and it
occurs both in the species found in Madagascar, and in those living in continental
Africa and in Asia. The curious Aye-Aye “Chiromys” builds an elaborate nest
which has been described by Baron as a structure about two feet in diameter
and entered by a hole in the side. Shaw has described the nest of “Chirogale
milii” as consisting of “leaves and Dry Grass,” and that of “Microcebus smithii”
as resembling a bird’s nest.”

Breeding Season and Period of Oestrus.

Galago Activities.

When received in September, 1937, the female was immature. Ten
months later (July 19, 1938), she experienced the first onset of oestrus. She
did not become pregnant, however. After an interval of five months a second
period of oestrus occurred (December, 1938) which resulted in the birth of
twins four months later (April 14, 1939). Oestrus did not re-occur until

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December 10, 1939, although the young stopped nursing sometime in July,
At intervals of six weeks three additional short periods developed, and a
fourth occurred twelve weeks later. A table of these data follows :

Female matured July 19, 1937
Second oestrus Dec. 15, 1937
Third oestrus Dec. 10, 1939
Fourth oestrus Jan. 22, 1940
Fifth oestrus March 7, 1940
Sixth oestrus May 29, 1940
** Oestrus recurred Dec. 7, 1940.

Length of
oestrus

Result

5-6 days
5-6 days
at least 3 days
at least 3 days

7

3 days

See Appendix, pj

no fertilization
twins (born Apr.
no result
no result
no result
no result
461.

14)

The only conclusions which can be drawn from these limited data are:
1. the period of gestation is four months; 2. a restricted season of sexual
activity is indicated. Whether December normally initiates the sexual season
in the wild state and the first expression of oestrus in July was atypical, or
whether the latter is typical and the shift from July to December is an
adaptation of the animals to their new environment, cannot be determined.
In any event this evidence still indicates the time for the birth of the young
is limited to the period between April and November. 3. It is also evident
that like many other forms with a limited breeding period, in the absence
of fertilization oestrus will reoccur at intervals, thus establishing the galago
as poly-oestrus within the limits of their sexual season.

Comparison with Breeding Habits of Other Lemuroidea.

Zuckerman substantiates the conclusions that the moholi galagos, like
the lemurs, have a restricted breeding season. He states that although the
records are too few to place any definitive reliance on them, they neverthe-
less indicate that the breeding season is betwen April and September, with
the majority of births taking place between April and June. He bases these
conclusions on the record of seven births of Galago s. moholi which have
occurred in the London Gardens between 1856 and 1927.

Donald Carter, mammalogist of The American Museum of Natural His-
tory, who recently returned (1938) from an expedition in South Africa, has
made available to me his field notes and many specimens of Galago s. moholi,
most of which were taken at Balovale, Northern Rhodesia, on the Zambezi
River.

Between August 23 and October 15, Mr. Carter secured more than one
hundred adults of which he brought back seventy-two to the museum. Two
of the 29 females contained embryos, and one he noticed was nursing young.
(He tells me that he did not record whether the remaining females were
nursing) . In addition to the adults 20 young galagos were brought into camp
by the natives. The following is a digest of Mr. Carter’s field notes concern-
ing the young taken at Balovale :

August 23 1 pair shot, female nursing young

September 4 1 female with a single embryo

September 14 1 female with two fully formed embryos

September 18 2 newly born

September 20 1 young

September 24 2 young

September 26 4 young

September 27 3 young

September 28 2 young

October 1 2 young

October 10 2 young

October 15 2 young

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An analysis of these data reveals that:

1. The relatively uniform dimensions of the adults indicate that they
were at least a year old.

2. A comparison of the pelage and measurements of the young in the
museum collection with those taken during the growth of the pair
in my possession shows that the largest were not more than four
or five weeks old.

Zuckerman (1932) states that three births among Galago garnetti oc-
curred, one in May and one in June, 1930, and one in September, 1929, in
the London Gardens. This corresponds with reports of births in the London
Gardens among the true lemurs where only very exceptionally were there
records of births outside the period March to June. Zuckerman states that,
“All the sixty-six births occurring in the London Gardens Zoo were distrib-
uted in the period March to September, sixty-four between March and June,
and two in September.” Major S. S. Flower (1933) presents additional data
which “provide a very good reason for arguing that species belonging to
the genus Lemur have a demarcated breeding season. The following table
showing the months in which 120 lemurs were born in the Giza Zoological
Gardens confirms Zuckerman’s opinion”:

March 26 births Twins Triplets

April 42 births 3 1

May 32 births

June 15 births

July 4 births

October 1 birth

120

From this additional evidence it is conclusively established that the
galago, like the lemur, has a restricted breeding season ranging from April
to October, but that probably the galago south of the equator is more apt
to have young in September and October than the lemur.

Zuckerman (1932) states that, “The. breeding habit of the African
members of the series Lorisiformes appears to be different from that of the
Asiatic species belonging to the same subdivision of the Lemuroidea. Thus,
the few available records regarding births in the genus Galago suggest that
the animals belonging to this genus may have a restricted season, whereas
the Hubrecht data for the slow Loris (Nycticebus coucang) prove that this
animal breeds throughout the year.” According to him, however, its curve
of fertility seems to rise toward the end of the year.

Discussion of Environmental Causes of Oestrous Change.

An interesting article written by F. H. A. Marshall (1937) presents
evidence that the reproductive rhythm may be influenced by extero-ceptive
or other environmental influences, and that the oestrous cycle may be reversed
in animals living in temperate climates after transference across the equator.
He concludes that “among the insectivores, carnivores, rodents and non-
ruminating ungulates with very few exceptions outside the tropical areas
breed in the spring or first half of the year in the Northern Hemisphere.”
Also, “That they react to seasonal changes in a remarkable way is shown
especially by those natural experiments in which animals of various species
that normally breed once annually have crossed the equator and as a conse-
quence have been induced to have two sexual seasons in one year.” Marshall
emphasizes the potency of more or less light as an activating factor. He
further states that “Tropical and subtropical forms, however, such as ante-
lopes (red and blue duiker) which live under comparatively uniform con-
ditions as regards daylight, may have an extended breeding season or breed

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all the year. They do not adjust themselves to the changing conditions of
temperature in regard to sexual activity.”

Following this analogy, could not the differences between the breeding
habits of the lorises and the galagos be accounted for by a study of differ-
ences constant in their normal habitats? Since all these forms are nocturnal
and tropical or subtropical, the value of more or less light as a breeding
factor is not as important as conditions insuring a constant food supply, or
other environmental influences. The Galago senegalensis moholi and Galago
crassicaudatus and their subspecies live in the open savanna country where
there is a prolonged dry season lasting from April through October, during
which their young are born. The lorises, however, live in forested tropical
regions where there are no such extremes of humidity and dryness and
where a restricted breeding season would have no survival value. It would be
interesting to determine whether Galago demidoffii, a small form found in
the rain forest of Equatorial Africa in the very regions in which the duikers,
cited by Marshall, live, would, like them, have continuous or irregular breed-
ing season as a result of more uniform environmental conditions.

Period of Gestation.

As I have reported, the first period of heat, experienced by the young
female under my observation, was not followed by pregnancy. The second
period, however, which began five months later, proved to be a fruitful one.
No peculiar conditions were noted for two months. The female then dis-
played increasing irritability at the attention of the male. She showed pref-
erence for the warmth of the radiator at the corner of the room and gave
up jumping to high places. Her pelt became patchy between the shoulders.
She looked thin and in bad condition. During the third month the female’s
irritation at the attentions of the male was so great that I separated them,
giving each a cage. Paradoxically the female would answer the male’s con-
versational note and seemed to enjoy talking to him from a safe distance.
She showed enormous interest in food, eating very much more than usual,
and seemed particularly fond of buttered bread. Since she refused cod-liver
oil in milk, I smeared it on her fur, so that in licking it off she acquired an
adequate dosage. During the intervals when both were liberated for exer-
cise, the truculence of the female increased. A week before the birth of the
young the female became very active in building a nest in her sleeping box.
Tearing up pieces of newspaper, she carried them into the box. On the nights
of April 12 and IB, 1939, she was unusually active.

Birth and Number of Young.

On the evening of April 14, I opened the doors of both cages to give
the animals their customary exercise. The female was still in her sleeping
box. On returning to the room a half hour later, I found her perched on
the edge of the cage, holding in her mouth a dangling infant. All the while
she was uttering a strange, protesting cry, directed at the male who had
entered her cage and was watching the scene attentively. Obviously dis-
turbed at his presence, the female jumped out of the cage onto a portiere
and from there to the picture molding, her young one still clutched in her
mouth. She finally made a downward leap of fully ten feet onto the bed, and
from there retired to a dark corner of the room where she deposited the
squeaking infant. I found a second young one clinging to the perpendicular
side of the cardboard sleeping box. I cannot say just when the young were
born. The pelage of one seemed still wet. The box was clean and there was
no evidence of the placental membranes, which must have been eaten by
the mother.

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The birth of twins among galagos of this species seems to be a common
occurrence, although single births do occur.

Donald Carter’s field notes indicate that two young are frequently found
in a nest, and Shortridge (1934, page 00) states:

“Twins seem to be of frequent occurrence. In Northern Rodesia — Ndola — -
several females were found carrying twins.”

“A female in a Pretoria Zoo suckled two young ones. Another female gave
birth to two young in October.”

Haagner (1920, page 10) states that two is the usual number at a
birth. There have been other instances, however, of gravid females contain-
ing but a single foetus.

Other galagos such as crassicaudatus and crassicaudatus monteiri are
more likely to be carrying single foetuses (Pitman, 1934, page 159). This
resembles more closely the habit of the Malagasy lemur where, as the statis-
tics show (S.S. Flower, 1933) (vide supra, p. 451), among 120 births there
were but three instances of the birth of twins and but one of triplets.

Post-natal Condition of the Young.

The young, male and female, were about the size of my index finger,
and almost as slender. Their eyes were partly open. The tail was curled in
a loose spiral, and they were sparsely covered with a homogeneous gray
pelt.

The fact that the infant galagos were able to cling to the perpendicular
surface of the box, and actually to stand on all fours shortly after birth, is
of interest.

The young creep about, and after the first day or two of clinging to
the mother with all four feet, they may be found under her, feeding, or in
the nest. While they never cling to the parent as she moves about, frequently
they will climb on her back or creep between her legs when she is crouched
quietly in the cage. (PI. VI, Figs. 15, 17). When in motion, however, they
are either left behind, or she picks each up in her mouth, by the neck or
the back, much as a cat would her kitten, and carries them about one at a
time (PI. V, Figs. 11, 12, 13). The passive infant accommodates itself to
this treatment by drawing up its legs close to the body (PI. V, Fig. 12).

Donald Carter reports that the young Galago s. moholi brought in by
the natives were found in the nests and never found clinging to an active
parent. These observations are at variance with those recorded by Short-
ridge (1934) and Haagner (1920) in the cases of captive animals in the
Zoological Gardens of South Africa. Shortridge states:

“The young cling to the under side of the mother and are carried about in this
manner until more than half grown, the mother being much hampered in her
movements, when the young become larger.”

Haagner states (p. 19) :

“They cling to the mother like the young of the ordinary Ringtail Lemur and
South American Marmosets, and ride on her back or hang underneath her as
opportunity offers.”

These statements by Shortridge and Haagner imply that the mother
jumps about with the young clinging to her ventral surface or her back.
Careful observation of the family under discussion has never disclosed any
such action. When feeding the young are on their backs on the floor of the
cage and the mother remains quietly in one position. To be sure, the young
do cling to the under surface of the mother by their front legs, but the
hind legs are always free and protruding beyond the body of the mother.
They may also be found sitting on her back (PI. VI, Figs. 15, 16) or later
on the father’s back, but this is only when the animals are quiet or sleeping.

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There never is any clinging action (as Haagner has suggested) such as one
finds among the monkeys when the parent is active or jumping about.
Shortridge’s statement about the mother’s being much hampered in her
movements is misleading. She obviously could not move when the young
were nursing, and during activity they were never attached to her in any
way, unless she decided to move them by carrying them in her mouth.

In contrast with these findings, however, F. Wood-Jones5 6 and P. L.
Sclater0 both describe the mother of the genus Lemur as the passive agent,
the young clinging to the ventral surface of the parent, and using its own
tail to help hold itself on to the parent body. The young lemur therefore is
the active agent.

Hill, in 1937, described the birth of twins of the slow loris ( Nycticebus
tardigradus) . He writes that the young are sturdy and that the mother
placed them on the ground at the very beginning. He does not state how
the mother carried them about, but implies that she does the carrying, and
the young are passive.

Hill, 1937, page 388:

“She will rush toward the baby and pick it up if danger threatens, and
place it down again when the risk is over. She will leave her food to do this. If
the baby is marooned, it squeaks for its mother, and she will try to get to it, if she
is able. If the baby is picked up for observation and replaced, she has not dis-
carded it, but smells it, licks it clean, and allows it to go to sleep again.”

Cuming, in 1838, described the habits of a Tarsius with her young
which seems to have a striking similarity with Galago s. moholi. He states
that the young when born have their eyes open, are covered with a good
pelt of fur, are able to creep around the cage shortly after birth, and the
parent carries the young around, in the mouth, just as in the case of
Galago s. moholi.

H. Cuming, 1838, page 68:

“The young appeared to be rather weak, but a perfect resemblance to its
parent; the eyes were open, and the body covered with hair; it soon gathered
strength, and was constantly sucking betwist its parents legs, and so, well-covered
by its mother, that I seldom could see anything of it but its tail; on the second
day it began to creep about the cage with apparent strength and even climb up to
the top by the rods of which the cage was composed. Upon persons wishing to see
the young one covered over by the mother, we had to disturb her, upon which the
dam would take the young one in its mouth, in the same manner as a cat, and
carry it about for some time; several times I saw her when not disturbed trying
to get out of the cage, with the young one in her mouth as before. It continued to
live and increase in size for three weeks, when unfortunately some person trod
upon the tail of the old one, which was protruded through the cage, a circumstance
which caused its death in a few days; the young one died a few hours after, which
I put into spirits.”

Le Gros Clark (1924), however, does not corroborate this observation
with relation to the Tarsius. He states that:

“I have kept under observation several female Tarsiers and young and have
never been able to confirm this” . . . (carrying of the young by mother in mouth).
“The young cling to the fur of the mother’s abdominal wall, grasping with both
hands and feet, and are not in any way held by the mother. In this way the little
animal is very often wholly concealed from view when the mother is in a resting
position, clinging to a branch” — He further states that : “At birth the Tarsier has
reached a comparatively advanced stage of development. The eyes are open, and
though its movements are uncertain, the animal can cling to a vertical branch,
and scramble about the branches in a hesitating way. When disturbed it assumes
an appearance of alertness.”

5 Wood-Jones, 1929, page 115. “Lemurs do not nurse or handle or carry their young ones, for
it is not the mother that clasps the offspring, but the offspring that grips the mother.”

6 P. L. Sclater, 1885, p. 672. “Young lemur lies nearly transversely across the belly of its mother,
and, passing its long tail around her (mother) back and so on to its own neck, uses it as a prehensile
organ to hold on by.”

1940] Lowther: Study of Galago senegalensis mofroli 455

Nursing and Care of the Young.

The female has two pair of mammary glands. Those of the anterior
are so laterally placed that each lies practically on a line with the inner
border of the arm. The inguinal pair lie closer to the median line. During
the first two days the young clung to the mother by all four feet. I saw one
nursing from a pectoral mamma while its body was attached transversely
under the mother’s arm and around onto her back. Within a few days, how-
ever, they began to stretch out on their backs in the nest, gripping the
mother with their front feet, while their hind feet protruded grotesquely
from under her body. Occasionally they would give a convulsive kick. At
first the young were so small that the mother had no difficulty in covering-
all but their protruding legs with her own body. As they grew larger, how-
ever, she was obliged to lift herself more and more to accommodate them,
causing her front legs to be bowed over them in a ludicrous fashion. Finally
they grew so large that she could accommodate only one at a time.

During the period of nursing, which lasted three and a half months,
the mother displayed an enormous interest in food. Her consumption of milk
was more than twice as great as before pregnancy. And she would now even
tolerate in it cod-liver oil, which perceptibly improved her pelage.

The mother kept the young clean by grooming their fur herself. At
their nursing time, while they were lying on their backs, with hind legs
extended, she would lick clean their genitalia.

During the period of nursing the mother’s urine carried the odor of
sour milk.

Parental Attitudes.

The mother galago displayed great solicitude for her young. When
they were taken out for inspection, she would go to the length of jumping
close to and even brushing my arm in her agitation. When the young were
put down, she promptly gripped one in her teeth by the neck and jumped to
the cage, using the pendant youngster instead of her nose to lift the cage
door. Having recovered one infant, she then proceeded to rescue the second,
in a similar manner. A thorough licking and smelling of the young followed
their return to the nest. During the first three weeks the male was kept in a
separate cage except during the periods of exercise. He seemed restless and
curious about the occupants of the other cage, eating relatively little and
keeping his eyes glued on the activities across the room. When liberated for
exercise the male invariably made his way to the cage and tried to open the
then locked door. The mother, permitted to exercise with the male, seemed
glad of her freedom and was willing to play to some extent. When the young
were three weeks old the galagos were all placed in the same cage without
ill effects.

Perhaps the separation of the parents was an unduly cautious move,
but the circumstances seemed to warrant this action. For the first few weeks
the mother made very soft caressing sounds to her young and they were
frequently heard making small mouse-like squeaks. As the infants grew,
the mother continued and the father assumed solicitous attitudes, and even
up to the age of nine months they obeyed the call of the mother. During
the summer months when the family occupied a screened-in porch, I often
heard and watched the mother give the signal for the hour of retirement
after a night’s activity. Just before sunrise, she would start a soft, con-
versational note and the young as well as the mature male would answer her
call and immediately jump to her side. Within five minutes all four would
have retired into the sleeping box for the day’s rest. The action of the
mother indicates a greater sense of responsibility toward the young than
that of the father. It is she who warns the group of any danger. When a

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cat lurked on the steps leading to the porch, it was the mother who gave the
alarm call ending in a shrill whistle, which warned the family and sent me
on a tour of investigation. In the winter the family was moved to a large
enclosure in the college greenhouse, where a shelf has been made into a
sleeping box by the addition of a sliding door in which an aperture has been
cut. For several minutes the family jumped and climbed and smelled about
this new region. It was the mother who located the small aperture of the
sleeping box and the dark quarters within. And after investigation it was
she who began the conversational call which brought father and children to
her side. The entire family is affectionate. The mother frequently caresses
the young even though they are now more than a year old.

The maternal solicitude continued unabated and even when they were
far too heavy to be carried, (up to 3 months), she still tried to lift them in
her mouth and to get them back into the nest when the young were molested.
After handling, also, she would lick them solicitously.

Development of the Young Galago.

From the beginning, as has already been stated, the young galagos
could cling to surfaces and walk about on all fours. When not nursing, they
were either under the mother, on her back (PI. VI, Figs. 15, 16), or were
curled up in a little nest made for them of soft tissue. In two weeks time
there was a perceptible increase in size. Their pelt was heavier, but still
homogeneous in color. They not only could walk about, but had begun to
take small leaps of four and five inches, clumsily trying to balance on their
hind legs, making efforts to climb on the struts of the cage and even to jump
a few inches. Their bellies were very fat and their tails still curled some-
what. Between the second and third weeks, the young ones began to use
the tongue in licking and grooming each other; they even began to lick me.
When three weeks old they were approximately twice their original weights.
They had minute incisor and canine teeth. They were very playful and would
nip each other’s tails and frolic about.

When four weeks old they could jump from the side of their cage a foot
or more to its floor and back again. They could walk along a horizontal bar
about a half-inch in diameter, sometimes losing their balance, but never
their grip. They would now play with each other like kittens and could
stand on their hind legs although their balance was still uncertain.

At six weeks the fur had become adult in appearance, the end of the
hair assuming a lighter color, giving a frosted look. The chest and the
lateral surfaces of the legs and arms were light orange-yellow. The longer,
darker hairs standing out on the back, which are characteristic of the adult,
were more conspicuous in the young ones, like a soft black fuzz. The dark
markings on the face, however, were not yet as evident as those of the adult.
At this time they began the moistening of the hand and foot-pads. They
made great work of this operation, rubbing with energy the palm of the
hand onto the sole of the foot on the same side. They scrubbed for some
time, and were not completely sure of their balance.

The young galagos, which could jump from two to three feet in a
horizontal position, watched with interest the action of their parents as
they made vertical leaps from a window-sill and portiere to the picture
molding. The first attempt at a vertical jump like those of the parents was
made at the age of seven weeks when one succeeded in reaching the top of
the window trim. One of the parents immediately jumped to a position be-
side it, and licked its fur. By the eighth week the young ones began coming
down a perpendicular surface, gripping the edge in a regular adult fashion.
At this time the young began making the protesting adult growl when
handled. By the end of the second month, their balance was greatly improved
and they could stand upright with considerable confidence.

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From the age of one month the young attempted to eat solid food which
was placed in the cage for the mother. By the end of the second month
they were eating solid food regularly and drinking milk in considerable
quantities.

Although still nursing, they were now so large that they had to be
fed one at a time. By this time, the galagos had been moved into the country,
and now occupied a screened-in porch, with freedom of action during the
entire night. Flying moths were included in the diet of the parents. While
at first the young had little interest in these flying insects, it grew as they
watched the eagerness of the parents. They did not attempt to catch any
themselves, but as the parents caught a moth, the young would try to take
it away. The mother seems fairly willing to give up portions to her offspring.
The samples were evidently pleasant, and by the twelfth week the young
ones were capturing their own moths. They were not yet as quick or as
dexterous as their parents, and even after succeeding in a catch, they fre-
quently were obliged to give up their prize to a hungry parent. Picking
up the discarded wings, the young would chew on the base of these for
what nourishment there might remain. Cooke (1939) notes that adult
Tarsius also chew on the base of moth wings. At three months of age they
had learned to balance themselves on the rim of a jar and to dive in and
extract the fluttering insect without loss of balance. By the end of the third
month, the young had stopped nursing. The only evidence of infancy was
the tendency to take moths from the mother, which she continq d to per-
mit them to do. They were now less rotund, and except for the soft black
fuzz on their backs they looked like their parents in miniature.

Sleep.

During sleep these animals have a tendency to crowd together, one fre-
quently sleeping on top of another. In cold weather it is not unusual to find
one adult completely hidden by a companion. They protect their eyes from
the light by burying the head in the pelt of another galago. If the light is
too intense, they try to crawl under any movable object for protection. At
times, too, the tail is wrapped over the eye to exclude the light.

This variety does not usually sleep with the neck arched and the head
tucked between the hind legs as does Galago crassicaudatus , although on
rare occasions, when one Galago s. moholi has been isolated, I have seen
this method employed. The large membranous ears remain somewhat folded
during sleep.

Acknowledgments.

I give grateful acknowledgment to those who have aided me in the
preparation of this article and whose knowledge of this field has rendered
their criticism and assistance of special value. They include Professor
J. Howard MacGregor of Columbia University, Professor W. K. Gregory of
Columbia University and the American Museum of Natural History, and
Mr. John Tee-Van of the New York Zoological Society who read the manu-
script; Mr. Harry Haven, Dr. James Chapin, Dr. George Pinkley, Dr.
George G. Simpson, Dr. J. Eric Hill and Mr. Donald Carter, all of the Ameri-
can Museum of Natural History, who contributed information of peculiar
value. I warmly acknowledge my indebtedness to Dr. Agnes Townsend of
Barnard College who took many of the photographs which illustrate this
article. I also record my appreciation of the kindness of Mr. A. M. Vida of
the American-South Africa Line who brought back from Africa the original
pair of galagos with which this study is concerned. Thanks are also due
Mr. William Metzler and Mr. John Connolly for their watchful care of the
animals at night while in the college greenhouse.

458

Zoologica: New York Zoological Society

[XXV :27

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1929. Man’s Place Among the Mammals. Arnold, London, pp. 81-125.
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1940]

461

Lowther: Study of Galago senegalensis moholi
Appendix.

The activities outlined here occurred after the manuscript was in type.

On the evening of December 7, 1940, oestrus recurred after an interval
of six months. At this time a new factor was introduced. For some weeks
prior to this date the young male, now twenty months old, had given evidence
of sexual maturity with the result that, at the onset of oestrus, both males
were accepted by the mother. The young female, however, has not yet become
sexually active.

EXPLANATION OF THE PLATES.

Plate I.

Fig. 1. Head of adult male enlarged 2 X. Crescent-shaped rhinarium shown
with deep median cleft. Upper lip bounded by fur. No “snarling” ex-
pression of upper lip though animal is angry. Eyes with vertical pupils.
Lashes visible on upper lid of eye to right. Vibrissae evident lateral,
median to and below the eye, also at side of mouth on left. Note method
of grasping small stick. Thumb and index finger on one side, the three
outer fingers on the other.

Plate II.

Fig. 2. Life size group of family of four in characteristic sleeping position,
taken when young were a year old. Note corrugation of ears of animals
to right and lower center.

Fig. 3. Photograph of family on rafter under porch roof. Upper eyelids of
animal third on right shown extended over bulging eyes. This galago
had been perched on the rim of the flash light reflector and was looking
at the bulb when it was discharged. For a short time the animal was
unable to tolerate even a subdued light.

Plate III.

Fig. 4. Photograph of hands X2. Illustrates method of resting palm on a flat
surface with the flexed digits somewhat bent to one side. Note flat nails
and pads along under surfaces of digits ending in enlargements at tips.
Interdigital pads shown protruding beyond palm of hand to left.

Fig. 5. Phalanges of hand flexed for grasping food. The two distal phalanges
bend over the proximal phalanx of each digit except that of thumb.
Thumb opposed.

Fig. 6. Photograph of under surfaces of hand and foot, taken from the inside
of a glass door on the outerside of which a galago is clinging. The
index finger is shown to lie almost midway between the large space
which separates the thumb and third finger. Illustrates the way in which
the pads of the palm and sole flatten out when applied to a flat surface.

Fig. 7. Photograph of under surfaces of feet taken when animal was in same
position as in Fig. 6. The pads of the soles are not flattened out. Note
fur-covered hinder portion of ventral surface of foot and large size of
the pads of hallux, the interdigitals, thenar and hypothenar.

Plate IV.

Fig. 8. Photograph showing method of grasping branch by feet alone, leaving
hands free for feeding.

Fig. 9. The animal is hanging head down with hands free, grasping screen-trim
by hind feet. Note the widely abducted, powerful hallux.

462

Zoologica: New York Zoological Society

Fig. 10. Adult male and female. Male standing on toes reaching for insect. Note
length of foot and hind limb. Tail serves as balancing structure. Digits
of the hands are somewhat flexed, which is their usual position unless
grasping food or perching.

Plate Y.

Figs. 11, 12, 13. Method of carrying young. (Fig. 11). Mother while on table-
top grasps young in region of shoulder. (Fig. 12). Young lifted from
table, mother poised to jump to floor; feet of young drawn up. (Fig. 13).
After jump, mother depositing young on floor.

Plate VI.

Fig. 14. Photograph of young when two weeks old.

Figs. 15, 16, 17. Different postures when awakened from sleep. Young are about
five weeks old. (Fig. 15). One baby under the mother, trying to get away
from the light while the other lies on top of parent.

(Photographs reproduced as Figs. 1-10 were taken by Agnes Townsend; Figs.
11-17 enlargements of 16 mm. motion picture film taken by author).

LOWTHER.

PLATE I.

FIG. 1 .

A STUDY OF THE ACTIVITIES OF A PAIR OF GALAGO SENEGALENSIS MOHOLI IN CAPTIVITY,
INCLUDING THE BIRTH AND POSTNATAL DEVELOPMENT OF TWINS.

LOWTHER.

PLATE II.

FIG. 3.

A STUDY OF THE ACTIVITIES OF A PAIR OF GALAGO SENEGALENSIS MOHOLI IN CAPTIVITY,
INCLUDING THE BIRTH AND POSTNATAL DEVELOPMENT OF TWINS.

LOWTHER.

PLATE ML

FIG. 6.

FIG. 7.

FIG. 5.

FIG. 4.

A STUDY OF THE ACTIVITIES OF A PAIR OF GALAGO SENEGALENSIS MOHOLI IN CAPTIVITY,
INCLUDING THE BIRTH AND POSTNATAL DEVELOPMENT OF TWINS.

LOWTHER.

PLATE IV.

A STUDY OF THE ACTIVITIES OF A PAIR OF GALAGO SENEGALENSIS MOHOLI IN CAPTIVITY,

as

LOWTHER.

PLATE V.

FIG. 12.

FIG. 13.

A STUDY OF THE ACTIVITIES OF A PAIR OF GALAGO SENEGALENSIS MOHOLI IN CAPTIVITY,
INCLUDING THE BIRTH AND POSTNATAL DEVELOPMENT OF TWINS.

LOWTHER.

PLATE VI.

FIG. 15.

FIG. 14.

FIG. 16.

FIG. 17.

A STUDY OF THE ACTIVITIES OF A PAIR OF GALAGO SENEGALENSIS MOHOLI IN CAPTIVITY,
INCLUDING THE BIRTH AND POSTNATAL DEVELOPMENT OF TWINS.

Ratcliff e: Diets for a Zoological Garden

463

28.

Diets for a Zoological Garden: Some Results During a Test Period

of Five Years.1

Herbert L. Ratcliffe

Advances in animal nutrition have not been applied extensively to feed-
ing problems of zoological gardens. Yet many of the difficulties inherent in
maintaining health and vigor of wild animals in this environment probably
could be solved in terms of information available either for related domes-
ticated species or for man. Presumably adequate substitutes for natural foods
would permit both the exhibition of more nearly normal animals and the
maintenance of breeding colonies of many species. The economic aspects of
either of these results are obvious.

During the first quarter of 1935 new diets were introduced at the
Philadelphia Zoological Garden. For more than ten years before this time
feeding practices had not changed materially. Therefore to aid in planning
new formulae, records of mammals and birds that had been autopsied from
1924 to 1934 were reviewed for definite or presumptive evidences of malnu-
trition. Estimates of deficiencies, obtained by this review, were substantiated
by comparing the values of foods that had been available to these wild
specimens with those necessary to meet the known requirements of the
nearest related species, whether man or domesticated animal.

Thus far, three mixed rations have largely replaced the various com-
binations of foods that had been used. One of these, with supplementary
fruits and vegetables, is fed to the so-called omnivorous mammals and birds.
A second, with mixed timothy and clover hay, provides for the herbivorous
mammals. A third is a complete diet for the greater number of the Carni-
vora. This system includes neither fish-eating mammals and birds, nor spe-
cialized insectivorous animals.

The degree to which requirements have been met by these diets may
be estimated at the present time only by changes in the causes of death,
the death-rate, the birth-rate and the development of the offspring. More
exact standards have been precluded to a large extent by conditions under
which this study was conducted and by the types of animals involved.

Because of the great number of species represented in this collection,
changes in the causes of death may be considered more completely only
when more extensive material has been accumulated. But within the five-
year test period just passed, changes in death rates, birth rates and in the
development of young of several animal groups seem to justify the conclu-
sion that these food mixtures have been reasonably successful. Increasing-
interest in certain wild animals as experimental subjects and requests for
information on feeding them, suggest that some of these observations may
interest others than those immediately concerned with the management of
zoological gardens.

1 From the Penrose Research Laboratory of the Zoological Society of Philadelphia and the
Department of Pathology of the University of Pennsylvania.

464

Zoologica: New York Zoological Society

[XXV :28

At the present time, discussion must be limited to the reactions of cer-
tain species of animals and birds as members of relatively large taxonomic
groups rather than as individuals. These groups, as such, are believed to
have had approximately equal opportunity for breeding before and since
the diets were changed. Exposure to morbid influences also has been equal,
except of course, as modified by diet, or, in some instances, by reduction of
the number of individuals within enclosures.

Omnivorous Animals.

The ration designed for this rather indefinite class of mammals and
birds is made up of 1 part of ground, boiled horse-meat and 9 parts of the
following mixture :

Rolled oats 20 per cent.

Whole wheat meal 20 “ “

Soybean oil meal 20 “ “

Peanut oil meal 10 “ “

Yellow corn meal 10 “ “

Buttermilk (dry) 5 “ “

* Brewer’s yeast (dry) 5 “ “

Alfalfa leaf meal (dehyd.) 5 “ “

Oyster shell flour 2 “ “

Iodized salt 2 “ “

**Cod-liver oil concentrate 1 “ “

These materials are mixed in bulk and used as needed. The ration usu-
ally is prepared the day before it is fed. First the ground meat and the dry
materials are mixed thoroughly, then enough meat broth (water in which
the meat is boiled) is added to form a stiff mash. This is pressed into
shallow pans of convenient size and refrigerated overnight at 44 to 46 de-
grees F. after which it has hardened sufficiently to be cut easily into
pieces appropriate to the size of the animals.

Allowing for natural variation in foodstuffs the approximate composi-
tion of this ration is:

Protein

Fat

Carbohydrate
Calcium . . .
Phosphorus
Iodine

25
5
45

1

0.5

0.017

per cent.

Energy value=3 calories per gram.

(potassium iodide
in “iodized salt.”)

A variety of materials has been included in this ration with the inten-
tion of offering a palatable food of which animals would not tire quickly. The
sources of protein are varied and the mineral and vitamin content have
proved adequate for growth and reproduction except in instances that will
be mentioned later. Obviously the formula may be varied and other sub-
stances added or substituted for those now in use.

Subhuman Primates. Thus far, animals of the following genera have
been available for observation: Gorilla, Pan, Pongo, Hylobates, Cercopi-
thecus, Erythrocebus, Macaca, Theropithecus, Papio, Mandrillus, Saimiri,
Cebus, Lagothrix, Ateles and Lemur. Some of these generic groups contain
relatively large numbers of species of which few have been represented in

* Brewer’s Blended Dried Yeast. 1938 assay claims 30 international units of vitamin Bi and 15
Sherman units of vitamin B2 per gram.

** Assay claims: 3000 U.S.P. units of vitamin A and 400 A.O.A.C. “chick” units of vitamin D
per gram.

1940]

Ratcliff e: Diets for a Zoological Garden

405

the collection. With the exception of rhesus monkeys ( Macaca mulatto),
these animals are kept in heated quarters during cold weather, and have
little exposure to sunlight except in summer.

Prior to 1935, more than half the total diet of these animals consisted
of polished rice, boiled sweet potatoes and bananas. Green foods, citrus
fruits, milk and eggs, were fed in significant quantities only to anthropoid
apes (Corson-White, 1922).

Since 1935, about one-half the energy intake of subhuman primates of
all ages has been supplied by the mixed ration, and, with exceptions to be
noted, the remainder of their diet made up of uncooked fruits and vege-
tables : sweet potatoes, carrots, apples, bananas, citrus fruits and green vege-
tables. In addition to these food stuffs, anthropoid apes also receive from
1 to 4 liters of fresh whole milk each day. The two gorillas now in the
collection also are fed about 200 grams of uncooked ground horse-meat each
day, and an equal quantity of boiled horse liver 3 or 4 times each weex.
Pregnant and lactating females and immature animals of all types are given
from 0.5 to 2 liters of a milk and egg mixture (1 egg per liter) and 5 to
20 cubic centimeters of a preparation containing some 800 units of vitamin
A and 250 units of vitamin D per cubic centimeter.

This diet, mixed ration and fruits and vegetables, is estimated to supply
at least 10 to 12 per cent of protein, 3 to 4 per cent, of fat and 30 to 40
per cent, of carbohydrate. Because of the relatively small amounts of cal-
cium and phosphorus contained in vegetables and fruits the percentages of
these materials in the mixed ration is reduced by at least half in the total
diet.

The known daily energy needs of man and lower animals permitted
the assumption that food requirements per unit of body weight would vary
widely with the several species of subhuman primates in the collection. In
consideration of size and spontaneous activity, allowances of 100 to 200
calories per kilogram of body weight per day did not seem excessive. In
actual practice, however, amount of food taken may vary widely from day
to day, and dominant members of a group must always be overfed so tha:
others will be allowed adequate amounts of food. Also, frequent feedings2
seem to be the most economical method of maintaining this type of animal,
lessening waste and preventing overeating of any one food with subsequent
loss of appetite for that or other substances. To accomplish this best, the
keeper must return to the cages two or three times.

Table I is a summary of breeding and mortality records for each of the
five years preceding and following the change of diet. “Total Stock” listed in
the first column of the table includes all animals because records do not
permit separation of potential breeding stock.

2 Feeding schedule developed with the aid of Mr. M. I. Tomilin.

Monkeys 8:30 A.M. Mixed ration (except on Sunday)

11 :30 A.M. to 12 M. Green leafy vegetables

2:30 P.M. to 3 P.M. Sweet potatoes and carrots

Offer water twice daily.

From 4 to 16 ounces of mixed ration with 2 to 3 times these weights of vegetables will be required.
Amounts are determined by size of the animals.

Chimpanzees and Orangs

8:30 to 9:00 A.M.
9:00 to 9:30 A.M.
11:00 to 11:30 A.M.
1:00 to 1:30 P.M.
1:30 to 2:00 P.M.
3:30 P.M.

4:30 to 5:00 P.M.

Gorillas

Oranges (from October to May because of cost)
Whole milk
Green vegetables
Water

Sweet potatoes and carrots or tomatoes
Whole milk

Mixed ration except on Sundays when bananas
are substituted

Feed according to the schedule for Orangs and Chimpanzees except that from
11 :30 A.M. to 12 M. they receive raw ground horse-meat every day, and from
3 :30 P.M. to 4 P.M., three or four times a week, each receives boiled horse liver.

466

Zoologica: New York Zoological Society

[XXV :28

TABLE I.

Records of Births and Deaths for the Subhuman Primates at the Philadelphia
Zoological Garden During Each of Five Years Before and Five Years After the

Diet Was Changed,

Year

Stock

8

$

Acquisitions

Births

Deaths

Sales

1930

58

48

27

5

51

0

1931

48

39

21

2

19

0

1932

49

42

8

3

15

1

1933

48

38

9

3

11

0

1934

46

41

11

5

18

0

1935

45

40

30

6

17

15

1936

43

46

19

6

8

19

1937

34

53

13

5

7

4

1938

35

59

8

7

14

19

1939

28

48

9

12

5

0

From 1930 to 1935 the average annual birth rate was 1 for 11.6 fe-
males. This increased to 1 for 6.6 females after the diet was changed. Prior
to 1935, breeding was limited to a few species of Cercopithecus monkeys,
mainly Macaca mulatta, and to Orang-utans ( Pongo ) and Chimpanzees
(Pan). All infants that survived breast feeding to the age of 12 to 16
weeks developed rickets.

With the present diet, Macaca irus, M. mulatta, Cercocebus fulginosus,
Cercopithecus sabeus, C. grivet, C. mozambicus, Cebus fatuellus, Papio
papio, and Pongo pygmaeus, have bred about as frequently as may reason-
ably be expected. Except for an occasional death that could not be attributed
to mulnutrition, all ofispring, except those of P. papio, the Guinea baboon,
have developed normally. These baboons, of which there have been five, de-
veloped rickets, however. All of them were born in the autumn, as were
many of the others, and had little exposure to sunlight. They recovered
promptly with appropriate artificial feeding. Possibly the greater vitamin D
requirement of the baboon will prove to be characteristic of the type and
may be related to the rapid growth of the facial bones to form the typical
dog-iike features; for it is in this part of the body, rather than in the
thorax and extremities, that abnormal bone formation first appears.

Also to be noted in Table I is the reduction in the number of males in
proportion to females in the colony. Breeding possibly has been influenced
by this change, but the death rate was not materially affected. Injury by
cage mates has contributed little to mortality rate at any time, although
occasional losses from this cause occurred throughout the period. The
average yearly death rate of 12 per cent, from 1935 to 1940 is to be com-
pared with that of 25 per cent, from 1930 to 1935.

This diet for subhuman primates is based to a considerable degree
upon recognized requirements of man, especially as to vitamins and min-
erals. It is essentially a modification of the diets suggested for these ani-
mals by Corson-White (1931) and by Tinklepaugh (1931, 1933) but is less
complex than either of these. In consideration of the known food habits of
the species of subhuman primates available for study it seemed that their
dietary needs might be met adequately by using a mixed ration composed
of easily available foods for domesticated animals. This has given an
economical and palatable mixture which, combined with accessory foods,
seems to be a reasonably complete diet. An obvious fault possibly is the
low fat content, but thus far there is no indication that this is an important
lack.

Whatever criticisms may be offered to this system of feeding, it seemed
reasonable to assume that the mixed ration with some of the supplements
also would meet the requirements of many other species of mammals and

1940]

Ratcliff e: Diets for a Zoological Garden

467

birds. In deciding upon the relative amounts of mixed food, vegetables
and fruits for these animals, preferences as well as presumed needs, as
indicated by experimental studies of diet, field studies of food habits and
the structure of the intestinal tract, have served as guides.

Other Mammals. The mixed ration that has been described constitutes
from 50 to 60 per cent, of the diet of rodents of the following genera:
Glaucomys, Sciurus, Citellus, Hystrix, Erethizon, Dasyprocta and Neotoma.
Supplementary foods are apples, carrots, sweet potatoes and green vege-
tables. This diet, the mixed ration and vegetables, also forms from 60 to 70
per cent, of the intake of the bears (Ursidae) and raccoons, coatis and
kinkajous (Procyonidae) . Horse-meat and fish supply the remainder of
their intake. The present quarters for these animals do not favor breeding,
but as judged by appearance, activity, growth of fur and postmortem
examinations, the diet seems adequate for growth and maintenance.

Birds. A large number of species of so-called seed- and insect-eating
birds, Passeriformes, many of the parrots, Psittaciformes, all Columbi-
formes, Galliformes, and Anseriformes, are fed one diet, the chief element
being the mixed ration that was developed for the apes and monkeys. This
food makes up about 70 per cent, of their intake. Supplementing it are
green vegetables, sprouted grains and fruits. In view of the known food
habits of many species of Passeriformes, the mixed ration is probably too
low in fat content to be completely satisfactory, and it has proved to be
unsuited for unfeathered young of local species of this order. Further
investigation is necessary to determine whether or not a special diet for
the adults of this order is economically justified. During the present test
period these birds have shown noteworthy improvement in plumage colors,
and lessened morbidity and mortality.

Breeding as a measure of the adequacy of the diet must, however, be
limited to birds of the families Phasianidae and Anatidae, for few others
are exhibited under conditions favorable to this activity. Prior to 1935,
about 70 per cent, of the intake of these two families consisted of a mixture
of equal parts of cracked corn and wheat screenings, supplemented by
green vegetables. Crushed limestone and oyster-shell were supplied ad
libitum to the pheasants.

Phasianidae. The pheasants, junglefowl and larger varieties of quail
are exhibited in pens that provide from 150 to 200 square feet of floor
space. Small types of quail have less space while wild turkeys are exhibited
in a much larger pen, and peafowl are at liberty on the grounds. Visitors
may pass within a few feet of all enclosures. From two to seven pheasants
of one species, with one adult male to the group, occupy each pen. Jungle-
fowl, turkeys and quail are kept in small flocks in which there may be more
than one male.

From 1930 to 1935, breeding in this group was limited chiefly to silver
pheasants ( Gennaeus nycthemerus ) but occasional young were produced
also by Swinhoe’s pheasant ( Gennaeus swinhoii), junglefowl {Gallus gallus ),
the common peafowl (Pavo cristatus) and stubble quail ( Coturnix coturnix) .

Since the present diet was offered, five species of pheasants have pro-
duced fertile eggs regularly, but none has had greater than 15 per cent,
fertility. Compared to this is the fertility rate of more than 50 per cent, for
wild turkeys and three species of quail. But in spite of the relatively un-
favorable conditions under which these birds live, the number of young
produced by this group from 1935 to 1940 was four times greater, in pro-
portion to the breeding stock, than from 1930 to 1935, and the annual
death rate fell from 38 to 24 per cent.

Anatidae. Many of the members of this family are exhibited in one
body on an artificial pond. Others are kept in smaller enclosures along a

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[XXV :28

stream. The mute swan, ( Cygnus olor), is the only species provided with
a separate space where the pair may nest. In all locations visitors may
approach within a short distance of the birds.

From 1930 to 1935, breeding was limited to Canada geese, ( Branta
canadensis) and mallard ducks, (Anas platyrhynchos) , which hatched an
average of twelve young per year. With the present diet these continue to
breed and, in addition, mute swans, ( Cygnus olor), and upland geese
( Choephaga magellanica) have produced young regularly, and blue geese
( Chen coerulescens) and Egyptian geese ( Alopochen aegyptiacus) have
occasionally nested. From 1935 to 1940, an average of 43 young were
hatched each year. Young birds of both the family Anatidae and Phasiani-
dae are fed the mixed ration with about 10 per cent, of finely chopped
green vegetables. Growth has been entirely satisfactory and loss negligible.
This diet for Phasianidae and Anatidae corresponds in all essentials to that
developed by Callenbach, Murphy & Hiller (1932) for ring-necked pheasants.
More recent experimental studies of the nutritional requirements of
chickens, pheasants, turkeys, ducks and geese, (Barnum, 1935), (Baird &
Green, 1935), (Bethke, Record & Kennard, 1936), (Norris, Elmore, Ringrose
& Bump, 1936), (Milby & Henderson, 1937), show that the food supplies all
necessary elements in adequate quantities, especially those essential to
high fertility and good “hatchability.”

Breeding of both Phasianidae and Anatidae probably has been limited
more by small enclosures, crowding by other species and by disturbances
of visitors than by any other factors. Pheasants should have larger pens,
and geese and swans probably should be separated in pairs for the best
results.

Herbivorous Animals.

The composite ration used for herbivorous animals contains beet pulp,
chopped vegetables and a portion of the following dry mixture:

Brewer’s grains (Dry) 400 lbs.

Yellow cornmeal 200 lbs.

Crushed oats 200 lbs.

Soybean oil meal 200 lbs.

Alfalfa meal (Dehyd.) 100 lbs.

"Brewer’s yeast (Dry) 40 lbs.

Oyster-shell flour 40 lbs.

* Iodized salt 10 lbs.

*Codliver oil concentrate 5 lbs.

This mixture also is prepared in bulk for use as needed. The ration is made
up in the late morning and fed in the early afternoon. The beet pulp is wet
with an approximately equal weight of water and soaked for 1 to 2 hours.
An amount of the dry mixture equal to the dry weight of the beet pulp, and
chopped cabbage or carrots in an amount about equal to 10 per cent, of the
total dry weight, are added and the whole well mixed. This forms from 60 to
70 per cent, of the diet of the herbivorous animals. The remainder is ob-
tained from mixed timothy and clover hay. With different species of
herbivorous animals the feeding allowances of the mixed ration vary
between 10 and 40 grams per kilogram of body weight, or 1 to 4 pounds of
the completed mixture per 100 pounds of body weight.

The approximate composition of the completed ration is:

Protein 15 per cent.

Digestible protein 11.5 “ “

Total digestible nutrients 70 “ “

Calcium 1.16 “ “

Phosphorus 0.25 “ “

* See formula for omnivorous animals.

1940]

Ratcliff e: Diets for a Zoological Garden

469

This mixture is a modification of rations that have been developed for
cattle and sheep (Morrison, 1939). In view of the type of hay available, the
protein content is lower than that usually recommended. But the problem
here is simply one of maintaining breeding stock in good condition. Neither
milk production nor rapid gains in weight are considerations. Beet pulp is
used as palatable diluent because concentrates are not tolerated well by
deer in captivity. Vegetables are included also for palatability. The cost
of the mixture compares favorably with others that provide equal amounts
of digestible nutrients with smaller numbers of constituents.

Artiodactyla. The majority of herbivorous animals that have been
available for study are included in the families Cervidae, Bovidae and
Camelidae of the order Artiodactyla. Some species of cervines and bovines
always have bred in the garden. Hence consideration will be given mainly
to these groups.

All of the deer, Cervidae, except delicate tropical species such as the
barking deer, Muntiacus muntjak, all of the Camelidae and the hardier
Bovidae, are kept in open paddocks throughout the year. These are provided
with tight shelters. African antelopes, Bovidae, live in heated quarters
during the cold weather, but have access to outdoor enclosures during the
warmer months. Natural pasturage is scanty or absent in all pens.

Prior to 1935, approximately half the diet of these ruminants was made
up of mixed grains (equal parts by volume of cracked corn, crushed oats and
wheat bran) and the remainder of mixed timothy and clover hay. Rock
salt was the only mineral supplement. African antelopes received small
amounts of fresh vegetables twice weekly.

TABLE II.

Records of Births and Deaths for Three Families of Artiodactyla (Bovidae,
Cervidae and Camelidae) at the Philadelphia Zoological Garden During Each of
Five Years Before and Five Years After the Diet Was Changed.

Year

Stock

Acquisitions

Births

Deaths

Sal,

S

9

1930

70

48

1

20

14

6

1931

77

42

3

19

15

4

1932

74

48

2

17

24

2

1933

67

48

0

21

12

8

1934

59

57

15

16

24

10

1935

60

53

6

20

28

14

1936

53

44

12

14

18

9

1937

53

43

15

16

10

16

1938

54

47

0

18

13

24

1939

41

41

6

20

7

10

Neither the birth rate nor the death rate of these animals has changed
materially since the present diet was offered. From 1930 to 1935, the
average annual birth rate was 1 for 2.6 females and from 1935 to 1940, 1 for
2.3 females. The average annual death rate was approximately 15 per cent,
throughout the period, but would have fallen to 12 per cent, after the change
of diet, except for the sacrifice of 16 animals as excess or as unfit specimens.

From these observations it appears that the present diet offers little
advantage, but the response of the ruminants is not well expressed by these
data. Certain species of Cervidae and Bovidae seemed to be able to live and
reproduce fairly well on the former diet. Examples of these are the Barbary
sheep, Ammotragus lervia, Sika deer, Sika nippon, and possibly the Central
American deer, Odocoileus virginianus nemoralis. On the other hand,
American bison, Bison bison, elk, Alces alces, fallow deer, Dama dama, red
deer, Cervus elaphus, Axis deer, Axis axis, and white-tailed deer, Odocoileus

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Zoologica: Neiv York Zoological Society

[XXV :28

leucurus, also reproduced but young matured slowly, rarely attaining the
stature of wild-bred specimens. Many offspring of all species died shortly
after birth, either because of lack of vigor or failure of the mother to
lactate; and adult elk and white-tailed deer commonly developed progressive
paralysis of the posterior etremities, due possibly to vitamin A deticiency.

The present diet seems adequate for maintenance of all species of
Artiodactyla with which it has been tested. White-tailed deer have not
developed paralysis of the posterior extremities during the five-year test
period. Elk were not available for part of the time. Occasionally, new-
born young, especially of hog deer, Hyelaphus porcinus, are lost shortly
after birth, but in general, offspring mature normally. As one example, two
generations of barking deer, Muntiacus muntjak, have bred while confined
entirely within doors.

More recent studies of the nutritional requirements of domesticated
ruminants, (Thurston, Eckles & Palmer, 1926), (Bechdel and co-workers,
1928), (McElroy & Gross, 1939), (Winegar, Pearson & Schmidt, 1940), indi-
cate that the present diet for wild ruminants may be unnecessarily complex.
Possibly it is, but the use of such a mixed ration seemed to be justified by
the conditions under which the animals are exhibited, and by the fact that
it is fed to non-ruminant herbivores such as the elephant, rhinoceros,
hippopotamus, tapir, kangaroo and hyrax.

Carnivorous Animals.

Mammals and birds of this dietary group receive a ration which con-
tains raw, ground horse-meat (approximately 20 per cent, glandular organs)
60 per cent., ground green vegetables, 10 per cent., and 30 per cent, of the
following mixture :

Soybean oil meal 30 per cent.

Peanut oil meal 25 “ “

Buttermilk (Dry) 25 “ “

Oyster-shell flour 6 “ “

Alfalfa leaf meal (Dehyd.) 4 “ “

*Brewer’s yeast (Dry) 4 “ “

*Cod-liver oil concentrate 4 “ “

"Iodized salt 2 “ “

This is fed to animals of the families Canidae, Mustellidae, Viverridae,
and Didelphiidae, and to birds of the orders Strigiformes and Accipitri-
formes. For these animals daily feeding allowances are 75 to 100 grams
per kilogram of body weight, or 1 to 4 ounces per pound. It also forms part
of the diet of other mammals and birds, but it has not been a successful
diet for adult wild Felidae. On this diet, “husky dogs,” foxes, wolves, skunks
and opossums have bred successfully and offspring have matured without
obvious defects. Judged by breeding, development of young and growth of
fur, the ration seems to be adequate for these animals.

The hawks and owls, Accipitriformes and Strigiformes, have not had
opportunity for breeding but the drop in mortality rate has been so pro-
nounced as to warrant mention. From 1930 to 1935, the stock of Accipitri-
formes averaged 39.6 birds each year of which about 30 per cent. died. With
almost the same average stock, 40.8 birds, from 1935 to 1940, the annual
death rate has dropped to less than 10 per cent. The average number of
Strigiformes exhibited from 1930 to 1935 was 19.8 birds of which 59 per
cent, died each year. From 1935 to 1940 an average stock of 32.2 birds also
lost less than io per cent. Prior to 1935 these birds had been fed only
horse-meat. The chief causes of death were either non-specific inflammatory
disease of the intestinal tract or simply failure of appetite. These condi-
tions are no longer factors in mortality.

* See diet for omnivorous animals.

1940]

Ratcliff e: Diets for a Zoological Garden

471

This diet for carnivores is one of many combinations that have been
tested. Early in the work, 30 per cent, of the dry mixture for omnivores
was used instead of the present dry material with the result that much of
the grain-meals were undigested and stools were soft. This is in keeping
with the observations of Hodson & Maynard (1938) on mink. In its essen-
tials the present diet corresponds to the suggestions of Kellogg (1939).

On the whole this work has been an attempt to adapt available stand-
ards of nutrition to the needs of a zoological garden. The aim has been
to provide acceptable, adequate and economical diets. Little training is
necessary to induce the animals to take the foods, and costs of feeding have
been reduced approximately 20 per cent. Whether or not the diets are com-
pletely adequate, in that continued breeding in captivity will be possible,
remains to be determined. In all probability, the mixed rations that are
being used could be further simplified without loss. The inclusion of iodized
salt in all mixtures may be questioned, but its need seemed indicated by the
frequent occurrence of hyperplasia of the thyroid in both mammals and
birds that were examined prior to 1935.

It must be emphasized that a five-year test period is a relatively short
time in which to evaluate reactions to these diets. This interval is only
part of the normal life-span of all except the more short-lived specimens.
The responses of the second and third generations must be awaited.

In planning the formulae efforts were directed to simplifying mixtures,
using easily weighed amounts or standard bulk quantities as much as
possible. Also, in actual practice food allowances are not arranged on a
grams per kilogram of body weight per day basis. Instead, easily under-
stood measures of the prepared foods are employed so that neither the keep-
ers nor the workmen who prepare the foods are compelled to do more than
follow a simple routine.

Summary.

A rather elastic classification of mammals and birds according to known
food habits and the general structure of their alimentary tracts, permits the
subdivision of the greater number of species that are exhibited in zoological
gardens into three general groups. Three diets, the make-up of which was
determined by the known requirements of related species, whether man or
domesticated animal, by the preferences of the wild specimens, and by prac-
tical considerations, have been used for five years. Response to these diets,
as judged by death rates, birth rates and development of young, indicate
that advances in human and animal nutrition may be successfully applied
to the feeding problems of zoological gardens.

Literature Cited.

Baird, F. D., & D. J. Green

1935. The comparative vitamin D requirements of growing chickens, tur-
keys and pheasants. Poultry Sci., vol. 14, pp. 70-82.

Barnum, G. L.

1935. The vitamin E content of eggs as related to the diet of the hen
and to hatchability. J. Nutrition, vol. 9, pp. 621-635.

Bechdel, S. I., H. E. Honeywell, R. A. Dutcher & M. H. Knutsen

1928. Synthesis of vitamin B in the rumen of the cow. J. Biol. Chem., vol.
80, pp. 231-238.

Bethke, R. M., P. R. Record & D. C. Kennard

1936. The relation of vitamin G to the hatchability of hen’s eggs. J. Nutri-
tion, vol. 12, pp. 297-307.

472

Zoologica: New York Zoological Society

Callenbacii, E. W., R. R. Murphy & C. A. Hiller

1932. The artificial propagation of ring-necked pheasants. Poultry Sci.,
vol. 11, pp. 150-157.

Corson-White, E. P.

1922. Osteomalacia. Arch. Int. Med., vol. 30, pp. 620-628.

1931. Degenerative bone disease in primates. Report Lab. and Mus. Comp.
Path., Zool. Soc. Philadelphia, pp. 31-35.

Hodson, A. Z., & L. A. Maynard

1938. Digestion and metabolism studies with mink. Am. Fur Breeder, vol.
10, pp. 38-41.

Kellogg, C. E.

1940. Nutrition of fur animals. Yearbook of Agriculture, U. S. Dept, of
Agriculture, 1939, pp. 871-892.

McElroy, L. W., & H. Gross

1939. Report on four members of the vitamin B complex synthesized in the
rumen of the sheep. J. Biol. Chem., vol. 130, pp. 437-438.

Milby, T. T., & E. W. Henderson

1937. The comparative growth rates of turkeys, ducks, geese and pheasants.
Poultry Sci., vol. 16, pp. 155-165.

Morrison, F. B.

1939. Feeds and feeding. 20th ed. Ithaca, N. Y.

Norris, L. C., L. J. Elmore, R. C. Ringrose & G. Bump

1936. The protein requirements of ring-necked pheasant chicks. Poultry
Sci., vol. 15, pp. 454-459.

Thurston, L. M., C. H. Eckles & L. S. Palmer

1926. The role of the antiscorbutic vitamin in the nutrition of calves.
J. Dairy Sci., vol. 9, pp. 37-49.

Tinklepaugh, O. L.

1931. A diet for chimpanzees and monkeys in captivity. J. Mammalogy,
vol. 12, pp. 400-404.

1933. Correction to a diet for chimpanzees and monkeys in captivity.
J. Mammalogy, vol. 14, pp. 68-69.

Winegar, A. H., P. B. Pearson & H. Schmidt

1940. The synthesis of nicotinic acid in the body of the sheep. Science,
vol. 91, pp. 508-509.

Hamilton: Biology of Smoky Shrew

473

29.

The Biology of the Smoky Shrew (Sorex fumeus fumeus Miller) 1

W. J. Hamilton Jr.

Zoology Department, Cornell University
Ithaca, New York.

( Plates I-IV ; Text-figure 1).

The long-tailed shrews (Sorex) are cosmopolitan creatures, represen-
tatives of the genus occurring throughout most of North America, Europe
and Asia. They reach their greatest numbers in temperate and sub-arctic
lands, occupying a variety of habitats but seeming to prefer damp situations.
Often these shrews are the most abundant mammalian representatives of
extensive habitats, but their habits are little known. They have been re-
ligiously catalogued by the systematist, but as with most species, we know
little more than their names, colors and measurements. This surprising
dearth of information on such widespread and ubiquitous creatures is not
surprising, for their diminutive size and retiring habits make them un-
worthy subjects for the attention of most naturalists. Nevertheless, these
little mammals hold a peculiar fascination for some, for almost any planned
study on these creatures will provide new information on their habits. The
long-tailed shrews, if for no other reason than their very abundance, must
play an important role in the habitat which they occupy.

The rate of growth, food habits, reproduction and behavior of Sorex has
been little studied in North America. The best accounts, which are all too
fragmentary, have been written by European students, the subject of their
attentions being largely confined to the common shrew ( Sorex araneus) .

In 1927 I commenced a study of the small forest mammals of New York.
Special efforts were made to secure life history data on the long-tailed shrews
(Sorex). As time and opportunity have permitted, attention has been di-
rected to solving the life history of the smoky shrew (Sorex fumeus). My
repeated efforts to secure live gravid females, so that the young might be
raised and described, have failed, but considerable data have been accumu-
lated during the past ten years of study. Most of my studies have been made
about Ithaca, in central western New York. The summers of 1937 and 1938
were spent on the Edmund Niles Huyck Preserve at Rensselaerville, Albany
County, in eastern New York where large numbers of shrews were collected
and population studies completed. Five hundred and sixteen shrews have
been collected during the course of the study, and these have provided
sufficient fresh material for rather complete studies on the various phases
of the life of the smoky shrew.

Specimens were secured in small snap-back mouse traps baited with
walnut meat, while a few shrews were captured alive in Sherman metal
boxtraps.

1 This study has been aided by a grant from the Elizabeth Thompson Science Fund of Harvard
University which the writer gratefully acknowledges.

474 Zoologica: New York Zoological Society [XXV:29

Description.

The smoky shrew is a relatively large Sorex, heavier and more power-
fully built but with a shorter tail, than Sorex dispar which occupies part of
its range. It is much larger than the common masked shrew, Sorex cinereus
cinereus, which is occasionally found in the same runways.

The winter pelage differs decidely from that of the summer pelage. In
winter the upperparts are deep mouse gray or slaty; the underparts are
very much lighter, almost silvery. Some specimens have a butty surfusion in
the post-abdominal region. In summer pelage the upperparts are pale grizzled
brown, the hair tips of the underpart being paler and gray at the base. The
tail at all seasons is bicolored, straw yellow below and brown above. I have
seen no aibinistic specimens but old specimens occasionally have the hair
tipped with white, or there may be a small patch of white fur on the thigh.

Summer specimens may be confused with Sorex cinereus (Townsend,
1935), but their gi’eater size and darker color serve to distinguish them
from their lesser kin.

Measurements. Three conventional measurements are made of mam-
mals when they are caught and before they are made into scientific speci-
mens, viz., total length (nose to end of last tail vertebrae) ; tail (base of tail
to end of last tail vertebrae) ; hind foot (heel to end of longest claw). These
measurements, unfortunately, vary with the collector and his methods, the
condition of the specimen whether fresh, bloated, in rigor mortis, etc., and
are largely a matter of personal equation. All the measurements listed below
were made by myself, from freshly caught specimens taken in New York
State.

Adult males. Twenty-six specimens average: total length, 116.5 mm.;
tail, 41 mm.; hind foot, 13 mm.

Adult females. Twenty-two specimens average: total length, 116.3 mm.;
tail, 43 mm.; hind foot, 12.6 mm.

Immature males. Seventeen specimens measure: total length, 113.5
mm.; tail, 41.4 mm.; hind foot, 12.7 mm.

Immature females. Fourteen individuals measure: total length, 113.1
mm.; tail, 41.2 mm.; hind foot, 12 mm.

Thus the measurements of 48 adults of both sexes average : total length,
116.4 mm.; tail, 41.9 mm.; hind foot, 12.8 mm. Thirty-one immature speci-
mens average: total length, 112.8 mm.; tail, 41.3 mm.; hind foot 12.6 mm.

It is thus evident that the conventional external measurements provide
little data on relative age, at least with this species. It is my belief that this
is true of many small mammals. This belief is based on measurements of
many hundreds of small mammals of various species in northeastern United
States. Weights, external appearance (pelage, hairy tail tufts or bare tail
tips, scarring of tail and feet) and above all, weight give a good index to
the age of this shrew, and combined, are sufficient in themselves to give a
ready clue to the approximate age of the animal. Tooth color and wear are
infallible, and should always be resorted to in order to determine the age
of the individual.

Weight. The smoky shrew has a variable weight, depending upon the
amount of food it has recently ingested and also on its condition. Sufficient
numbers, however, taken throughout the year, show a remarkable difference
between the immature and mature animals, and the two may readily be
separated on this factor alone.

Recently weaned shrews, about one month old, weigh 4 grams. They
gain weight slowly and throughout the first 7 to 10 months of their lives
weigh from 4 to 6.8 grams. Immature individuals of both sexes collected
from June to November weigh from 5.5 grams to 6.8 grams, the average of
97 specimens of both sexes being 6.19 grams. There is no sexual variation
in the weight of immature shrews.

Hamilton: Biology of Smoky Shrew

475

1940

During March, the shrew population (those which have been born the
previous year) gain weight rapidly and continue to become heavier through-
out the spring and summer until they die. Twenty-nine adult males collected
from March to August average 8.94 grams, the extremes ranging from 6.4
grams to 11 grams. Twenty-four females collected during the same period
average 7.44 grams, the extremes for these, with embryos removed in gravid
individuals, being 6.7 grams to 10 grams. It is very unusual to take these
adult shrews after August, and it will be shown later that they die of old
age at this season.

Molt. It has already been shown that the summer pelage varies markedly
from that of the winter, but the change by no means occurs at the same time
each year, and it may occupy several months in any one region.

Jackson (1928) states that in the southern part of the range of Sorex
fnmeus the spring molt may begin as early as the middle of April, though
most specimens at that time are still in full winter pelage. He records indi-
viduals showing molt from April 19 in the south to July 18 in the north.
Jackson observed that the fall molt occupies the period during October and
November, depending in part upon the latitude.

In New York it is usual to find these shrews in full winter pelage from
late September until early May, although the majority of museum specimens
are in summer pelage. This merely indicates that collectors are most active
during the summer months. The progress of the molt is a rather insidious
affair, sometimes well marked and again the hairs being shed generally over
the entire body so that little demarcation can be seen between the gray
pelage of winter and the brown hairs of summer. The earliest record of
molt is seen in an April 22 specimen, which has a large patch of summer fur
on the belly and chin.

Usually by mid-May the summer pelage has been obtained, although it
may be long delayed. An adult male taken on May 6, 1938 has a few hairs of
the winter pelage remaining irregularly over the back, the inguinal region
and particularly the rump. On June 21, 1936, I collected an adult female
which had attained the complete summer pelage except for a small patch
of winter fur about the base of the tail.

The summer pelage is retained until late September or early October.
Three immature shrews collected in the last week of September, 1935 and
1936, have a few patches of summer fur, usually about the head and base of
the tail. Fifty-seven immature shrews collected from September 27 to
October 9 were all in winter pelage except one, which had a few hairs of the
summer pelage about the ears. Males tend to molt earlier in the spring and
fall than females. Breeding is well under way and the first litter produced
before the females have completely shed their winter coat. This is con-
trary to the findings of Green (1930) who trapped a gravid female on April
18, 1928, at Ricketts, Pennsylvania, in which the molt was beginning on the
abdomen.

A smoky shrew is seldom collected which shows any sharp demarcation
between the winter and summer pelage. The molt appears to commence on
the belly, and extends over the shoulders and middle of the back, the old
fur persisting longest about the ears and rump.

Inasmuch as shrews in any given locality exhibit all stages of molt
over a rather long period, particularly during the spring, it might be as-
sumed that the change of fur in the individual occupies several weeks.
Wilcke (1938) observed that a captive Sorex araneus shed its long gray
shaggy winter coat during the first two days of captivity, assuming the
dark shiny summer coat in rapid time.

On May 6, 1938, I collected a large male at Ithaca, New York, which
showed a prominent molt over the back and rump. Long winter hairs stood
out irregularly from the new short brown fur of summer. During the two
days this animal remained alive, all the winter fur was lost.

476 Zoologica: New York Zoological Society [XXV :29

Habitat.

The smoky shrew is a northern species, reaching its abundance in the
cool forested regions of Pennsylvania, New York and New England. In the
beech-hemlock-striped maple woods with a ground cover of loose leaf mold
and black friable soil, it is often found in considerable numbers, and, unlike
its congener, Sorex cinereus, appears to be largely restricted to such a
habitat. In New York it is often abundant in birch-hemlock woods, but is
only poorly represented in the thin-soiled, rocky, dry, maple woods. While
the shrew perhaps favors damp woods, and the vicinity of streams, it is often
taken in areas far removed from water. This species seems particularly fond
of the moss-covered boulders which lie along the wooded south slopes of
valleys, where the sun does not penetrate far in the summer.

In the deep leaf mold of northern forests, one can often push a finger
through the surface layer to find a cavernous void beneath the inch or two of
soil which covers so completely the world beneath. By tearing away this
superficial layer, the tortuous galleries and broad runways of many small
forest mammals are exposed. It is in this retreat, made by the larger
Blarina, red-backed mice, hairy-tailed moles, and kindi'ed forms, that the
smaller Sorex are often found in profusion, a highway of mammal travel
scarcely visualized by the inexperienced observer. During the hot days of
August, 1938, I made several records of temperature in these retreats. In
the open woods and the borders, my thermometer read 89° F. Within the
woods, where little direct sun penetrated, the temperature was 81° F. In the
covered galleries of the small mammals, 6 to 10 inches below the level of the
forest floor, the temperature read 69° F. Conversely, in the winter, when the
forest floor was free of snow, the galleries of these animals, hidden below
the forest litter, had a temperature ranging from 20° F. to 31° F. while the
exposed forest floor had a temperature of 8° F.

The clumps of yew ( Taxus canadensis ) which grow from the deep leaf
mold are favorite retreats for these small shrews and many have been
captured in such places. The runways of moles, the tunnels canopied by
rotting logs and the leaf-filled depressions which obscure so well the laby-
rinth of tunnels, all yield their Sorex.

The smoky shrew population is by no means evenly distributed through-
out the forest. While there is no evidence of a communal or gregarious
nature, they prefer certain localities to others and here they may often be
found in some numbers. A typical habitat is shown in Plate I, Fig. 1.

Activity.

In general, shrews are active at all hours, trapping records to date not
indicating that these animals have any particular time in which to feed or
move about. The three common shrews of eastern United States, Blarina
brevicauda, Sorex fumeus and S. cinereus, are active at all hours of the
day and night. It appears possible that the presence or absence of suitable
cover may limit their movements to certain periods, but my trapping records
do not indicate this. By far the greater number have been taken at night,
but this is accounted for by the collecting methods employed. In the course
of the investigation, traps have been generally set in the afternoon and not
attended until the following morning, so that no positive record has been
obtained for the majority of specimens collected.

In order to determine at what hour catches were made, large numbers
of traps were set at various times of the year and visited at 6-hour inter-
vals. The results appear in Table I.

1940]

Hamilton: Biology of Smoky Shretv

Table I.

477

Activity of Sorex fumeus as determined by time of catch.

June 12-14, 1932
150 traps set
5-6:30 AM June 12

April 14-15, 1939
130 traps set
1-2:30 PM April 14

June 24-27, 1937
200 traps set
6 PM June 24

Oct. 6-7, 1938
140 traps set
3 PM Oct. 3

6 AM-12M.
2

3

12M-6 PM.
4

2

6 PM-12M.

3

5

12M-6 AM.
2

3

5 3 8 6

3

5 15

From the table above it can be seen that 27 shrews were taken from
6 A.M. to 6 P.M. while 33 shrews were trapped from 6 P.M. to 6 A.M. Al-
though the number is admittedly small, it shows that these shrews are active
at all hours. Inasmuch as these shrews live in the deep leaf mold and forest
litter, they are not particularly susceptible to predation at any particular
time. Their diminutive eyes probably hardly distinguish light from darkness.

Associates.

The forest floor which serves these little shrews, provides a home for
many other species. Within the same habitat many other small mammals
are to be found, perhaps competing with these mammal sprites for nest
sites and food. These associates vary in their numbers both from year to
year and in the various habitats which one selects to study. The relative
population of these to other small species may best be illustrated by noting
all mammals taken on a trap line over a unit of time in similar habitat.
The following examples will serve.

Maple-beech-hemlock woods, good leaf mold. Ithaca New York, June 28-
29, 1939. Three hundred trap nights. Sorex fumeus, 9; Blarina brevicauda,
27 ; Napaeozapus insignis, 10; Peromyscus leucopus noveboracensis, 9. Sorex
fumeus represents 16.4% of catch.

Beech-hemlock woods with deep leaf mold. Rensselaerville, New York.
August 6-14. 1938. Seven hundred and fifty trap nights. Sorex fumeus, 18;
Blarina brevicauda, 60; Peromyscus (principally P. leucopus novebora-
censis), 66; Synaptomys cooperi, 6; Zapus hudsonius, 2; Microtus pennsyl-
vanicus, 1. Sorex fumeus represents 11.8% of the catch.

Beech-hemlock woods with deep leaf mold. Ithaca, New York. Oct. 4-7,
1939. 800 trap nights. Sorex fumeus, 27 ; Blarina brevicauda, 24 ; Peromys-
cus leucopus noveboracensis, 26; Cleth?-ionomys gapperi, 7; Pity my s pine-
torum scalopsoides , 2; Microtus pennsylvanicus, 2; Parascalops, 1. Sorex
fumeus represents 20.2% of the catch.

In the records above, it is seen that Sorex fumeus comprises 11.8 to
20.2% of the small mammal population in ideal shrew habitat for certain
regions in New York. If these totals be averaged, it is found that, in the
three areas discussed, Sorex fumeus comprises 18.3% of the small mammal
population. In the northern part of the state, as the Adirondack region,
and much of northern New England, certain species become more abundant,
such as Clethrionomys and Sorex cinereus, and the fumeus population in
these regions would comprise a smaller percentage of the catch.

478

[XXV :29

Zoologica: Neiv York Zoological Society
Burrows and Nests.

The weak feet of these small shrews make digging a difficult procedure
and as a consequence they utilize and occupy burrows made by larger
mammals. Deep in the leaf mold are prominent subterranean galleries,
some the diameter of a baseball, and others whose bore scarcely exceeds
the diameter of a penny. In these burrows, from an inch to nearly a foot
below the leaves and humus ceiling, a host of small mammals move, feed,
breed and die. These burrows are made chiefly by the powerful hairy-tailed
mole, the short-tailed shrews and certain small rodents, notably the red-
back mice, jumping mice and lemming mice. Chipmunks often tunnel for
some distance below the soil, and the loose texture of the rotting logs and
stumps make progress easy beneath these decaying objects. The tiny Sorex
utilize these burrows, and are often caught in such places. They occasionally
occupy the punky moss-covered logs which lie half hidden in the trash and
rotted leaf mold of the forest.

As evidence that little burrowing is done by these diminutive creatures,
I placed smoky shrews in sizeable aquaria partly filled with friable loam
pressed to a consistency found in the normal habitat of these animals.
Little effort was made to dig, and when the animals did try to displace
dirt with their fore paws, the efforts were weak and ineffective. It seems
probable that they make no burrows but utilize entirely the runways of
other species.

The nests are placed at various places in these tunnels, a favorite
site being beneath a stump or rotten log. I have found one nest composed
almost entirely of the hair of a cottontail rabbit, situated beneath a
stump from which a shrew ran when the nest was exposed. Other nests
which I have examined have been made of shredded leaves, usually situated
from 4 to 9 inches below the sui'face. These nests are roughly spherical in
shape and approximate a baseball in size. They are more compact and some-
what smaller than the bulkier nests of Blarina. One nest which I presume
had been made by a smoky shrew (a specimen was trapped in a runway
leading from the nest) was situated in a punky log which crumbled apart
easily. The nest was placed at one end of the log near its base and com-
municated with several burrows leading into the soil beneath. Another
was found at the bottom of a large mass of rotten logs which had been piled
by a woodcutter, and allowed to disintegrate. This nest was placed directly
on the ground, between two logs which formed part of the foundation of
the pile.

None of the nests contained remains of food, but piles of scats were
usually found within a few inches of the nest. Captive shrews are certainly
sanitary creatures, often reserving a corner of the aquaria farthest removed
from the nest in which to deposit their feces. All the nests which I have
examined have been free of parasites.

Voice.

The long-tailed shrews utter diverse little notes, all with probably some
meaning. When alarmed, or approached by a larger mouse or shrew, the
smoky shrew utters a high pitched grating note, not unlike that of the
smaller bats. If greatly disturbed, it will throw itself on its back, and with
spread and waving legs, repeatedly utter these squeaking staccato notes.
When foraging for food these little shrews utter an almost indiscernible
twitter. I have twice seen these little forest mammals rooting through the
leaf mold or appearing on the forest litter about rotted logs, the twitching
nose and vibrissae held aloft, and this faint, almost inaudible twittering
was kept up continually.

1940] Hamilton: Biology of Smoky Shrew 479

Senses.

In spite of the diminutive eye (Plate I, Fig. 2) sight appears to be
of some importance. In very subdued light where captives were housed,
the shrews experienced little difficulty in avoiding obstacles and actually
turned from their trails to explore new objects which were placed in the
pens while the animals were exercising. In the thick carpet of leav'es and
the dark tunnels beneath the forest litter, there is little need of good eye-
sight and it is probable that these organs are degenerating.

The sense of smell does not appear to be particularly well developed.
Little opportunity was had to test this sense, but when odorous food was
placed in the boxes of hungry shrews, the animals often appeared to
experience difficulty in locating the food items. Indeed, on several occasions
they came within a few inches of small pieces of liverwurst, which were
shallowly buried, but did not find the food until they had hunted for some
time.

Hearing is reasonably acute. Gentle squeaking will alarm the shrews,
while loud and prolonged clapping of the hands will cause the shrews no
disturbance. Blossom (1932) refers to the apparent immunity of Sorex
cinereus to loud sounds. Sleep is often so sound that snapping the fingers
or loud whistling fails to disturb the shrews immediately.

The tactile sense appears to be well developed, and the long facial
vibrissae may well enable the shrew to intercept disturbances which the
other senses fail to detect. Indeed, there is some reason to believe this,
else why do these little creatures forever hold their long cartilaginous snouts
in the air, the mobile tip and associated vibrissae moving so rapidly that
it is all the observer can manage to follow these extraordinarily rapid
motions?

Food.

The smoky shrew is admirably adapted for gleaning the forest floor.
Its long snout, with stout inner cartilaginous support, enables it to push
through the leaf mold and loose litter, and its strong hooked and pro-
truding incisors act as efficient tweezers to gather the minute life which
swarms in this habitat. Elsewhere (Hamilton, 1930) I have reported on
the stomach contents of 31 S. fumeus which were taken from February to
December. All of these contained insect remains, these being, in the order
of their abundance, undetermined larvae, Coleoptera, Diptera and Hymenop-
tera. In addition, centipedes, sowbugs, earthworms, salamanders and vege-
table matter were found.

I now have the analyses of 168 stomachs to report. These were taken
in New Yoi’k during every month, but the greater majority were secured in
the spring and the fall. Specimens taken while a deep layer of snow
covlered the ground contained fragmentary remains of insects, principally
small golden pupal cases and fragments of beetles which were presumably
dormant. One contained what appeared to be the meat and skin of a beech-
nut. A dozen shrews secured in March while the ground was well frozen
contained quantities of insect fragments, including dipterous remains and
the legs of centipedes. During May the shrews feed chiefly upon insects,
particularly beetles and their larvae, and eat a few spiders. One shrew had
the toes and flesh of a small salamander, while another had parts of a snail,
probably Polygyra. Still another had eaten several small feathers, possibly
secured from a dead bird, for it is improbable that these delicate little
mammals could successfully overcome a sleeping bird, even though it also
be of diminutive size. I have seen j uncos sleeping among the Taxus clumps
when examining my shrew traps at night.

480 Zoologica: New York Zoological Society [XXV :29

During the summer months the principal food of the shrews appears
to be insects, but the stomachs of at least two taken during July contained
a mass of flesh and fur. Again it seems improbable that the animal was
not a victim of the shrew, but rather met its death in some other manner.
None of the trap victims had been touched, however, upon the occasion
when these two were taken. Snails, small earthworms and a quantity of
vegetable matter have been recognized in autumn specimens but the food
appears to differ little from that of the summer.

Table II.

Food habits of Sorex fuvieus, determined from examination of 168
stomach analyses made throughout the year. The figure below the month
indicates the number examined. Other figures denote the percentage fre-
quency of occurence of the different food items.

Food

Jan.

Feb.

Mar.

Apr.

May

June

July

Augr.

Sept.

Oct.

Nov.

Dec.

Total

4

5

14

27

20

10

19

8

17

21

ii

6

168

Insects

75

80

85.7

77.7

70

93.7

94.7

100.

88.2

52.4

81.8

66.6

80.0

Earthworms

7.1

14.4

15

6.2

10.5

12.5

9.5

23.3

10.1

Vegetable

50

20

14.1

14.4

25.0

18.7

5.3

50

18.2

50

14.9

Centipedes

25

21.4

18.5

5.0

18.7

15.8

5.9

9.5

18.2

16.6

13.1

Snails

20

11.1

10.0

10.5

17.6

23.8

9.1

10.1

Salamanders

3.7

5.0

5.3

1.8

Mammals

3.7

10.0

10.5

3.0

Sowbugs

11.1

6.2

21.1

9.1

5.3

Spiders

20

14.4

io.o

6.2

12.5

9.1

5.9

Birds

5.0

.6

Food Requirements. On several occasions I have had captive smoky
shrews for variable periods and have observed their feeding activities. These
shrews are not the prodigious feeders that many naturalists would have us
believe; they never eat food equivalent to their own weight daily, at least in
captivity. Nevertheless, Dixon (1924) has demonstrated that the little gray
shrew ( Notiosorex ) will eat 75% of its own weight each day. Blossom
(1932) observed that a captive female Sorex cinereus, weighing 3.6 grams,
ate an average daily weight of 11.7 grams, that is, an average of approxi-
mately 3.3 times its own weight every twenty-four hours. Wahlstrom (1928)
states that the food requirements of Sorex vulgaris are enormous and the
ability to fast, even when the animal is in good health, is very low. On the
other hand, Rorig (1905) maintained that a small amount of food was
quite enough to maintain a captiv'e Sorex araneus in good health. The
shrew ate an average of 7 grams of mealworms, grubs and flesh, and was
maintained for nearly three months. Inasmuch as an adult shrew of this
species weighs about 12 grams, it may be seen that slightly more than half
of its weight in food each day is sufficient to keep the animal in good
health.

It may be that conditions under which captive shrews are kept deter-
mine their ability to survive on a moderate amount of food. Moreover, a
varied assortment of natural foods appear to keep the captives more con-
tented than a restricted diet of mealworms or cockroaches. My captive
shrews had an assortment of small snails, mouse flesh, small earthworms,
beetles, centipedes, sowbugs and siftings from the forest floor, including
many spiders'. All seemed to do well on this variable diet, and maintained
themselves many days on an amount not exceeding half of their weight.
Still, this quantity is an incredible amount for such a diminutive animal to
consume. Much of the food is bulky and indigestible, for the chitinous
remains of the prey pass with the feces in a state which frequently makes
identification of the parts quite possible. The shrews eat at all hours, the
only stay to their hunger being the necessity for short naps and the actual
search for food, which seldom appears to entail any great labor.

1940]

Hamilton: Biology of Smoky Shrew

481

Feeding Habits in Captivity. The most important sense utilized in
searching for food appears to be that of touch. While hearing is rather
acute, sight and smell seem to play a minor role, and the ever moving snout
and long vibrissae probably are of major importance in apprising these
shrews of their food. They utilize the burrows of many small forest rodents
and the larger shrews, and within the confines of these tunnels the small
prey has little chance to avoid capture. Moreover, the leaf mold may be
riddled with their temporary tunnels and it is suspected, from the actions of
captive specimens, that much of their prey is secured by the sensitive nose.
Small prey, like flies, tiny isopods and centipedes, soil worms and the lesser
beetles are merely bitten and swallowed. The larger food items, as earth-
worms, snails, salamanders or newborn mice are held down with the fore
feet while pieces are bitten off. Wireworms, mealworms, the larger beetles
and kindred prey are first paralyzed by being bitten in the head. Small
salamanders ( Plethodon cinereus ) are held firmly to the ground and quickly
destroyed by being bitten in the head or back, severing the spinal cord.
The feet, head and viscera of these amphibians are first eaten, the remainder
often being neglected for other food.

It appears improbable that these small shrews are capable of overcoming
deer mice and the other small rodents which inhabit their domain. A small
Peromyscus, weighing 14 grams, was liberated in a large aquarium with
an adult fumeus weighing 10 grams. The shrew made several abortive
efforts to capture the mouse, and once closed in, grasping its active prey by
the flank. The mouse immediately broke this hold, and the discouraged
shrew thereafter paid no heed to its intended victim.

Sorex fumeus probably obtains sufficient water from its food, for cap-
tives drink only a limited quantity, dipping the snout into the pan of water
and tilting back the head like a chick. Blossom (1932) gave no water to
his captive S. cinereus over nearly a three month period. Wahlstrom (1928)
found his captive Sorex vulgaris never drank; indeed they seemed to fear
the water, and one shrew upon falling into its drinking receptacle avoided
it thereafter. On the contrary, captive specimens of Sorex araneus observed
by Wilcke (1938) drank very freely, in fact after every meal.

Reproduction.

Male Reproductive Organs. The following description is based on
the genital organs of a sexually mature male (Plate II, Fig. 3). The testes
are oval and measure 4 by 8 mm. but occasionally are larger. The caput
epididymis is often diffused and ill defined and usually partly embedded in
a mass of yellow fatty tissue. The cauda epididymis is large, and in breed-
ing animals the convoluted tubules may be distinctly seen with the aid of a
low powered binocular microscope. If the tubules are visible through the
walls of the epidydimis, they invariably contain sperm. This serves to de-
termine whether the males are in a breeding condition. The proximal end
of the vas deferens commences as a narrow tube, gradually swelling until
its distal two-thirds is reached. Here a prominent constriction occurs, the
distal part being enormously swollen into a pouch which gradually tapers
until its junction with the urethra is reached. The distal swelling has a
diameter 4 or 5 times as great as the tubal portion of the vas deferens. A
similar condition prevails with Blarina, but is never so well marked.
According to Brambell (1935) these swellings are composed of alveolar
pouches which store sperm. During the spring, one can slit the distal
pouches and make a smear which usually contains numerous sperm. Two
large paired organs, with numerous villi, somewhat larger than the testes,
and lying dorsad to the vasa deferentia and the bladder, have been desig-
nated the prostate glands by Arnback-Christie-Linde (1907). These never-
theless appear to be the seminal vesicles, for they appear essentially the

482

Zoologica: New York Zoological Society

[XXV :29

same as these latter structures in other small mammals. Their true homo-
logies cannot be determined until a careful developmental study is made of
these and associated structures in several insectivores. The large bean-
shaped Cowper’s glands, about two-thirds the size of the testes, lie at the
base of the tail, and dorsally give one the impression of anal glands. The
penis is a large organ for so small an animal. When retracted it is folded
on itself so that the tip lies over the most proximal portion. When extended,
it measures 16 to 18 mm. There is a suggestion of diffuse prepucial glands
but no ducts are visible. The penis is without a baculum.

Weber (1898) implies that the testes in the Soricidae leave the pelvic
cavity seasonally or Voluntarily. I have never found the testes situated
other than in a cremaster sac outside the pelvic cavity and lying near the
dorsal tip of the pubis. Even in very young shrews, the testes may be
teased into view with the forceps by grasping the sac and drawing the
testes out. In immature shrews, the caput epididymis is more prominent
than the testis, both in size and in the lighter color. Apparently the
enlargement occurs within the sac, without change of position of the testes
during their enlargement.

The increase in size of the reproductive organs at the onset of the
breeding season must occur with great rapidity. Specimens secured in
March have either very minute testes whose associated structures are barely
visible macroscopically, or else the organs are much enlarged. From exten-
sive field observation and dissection of more than a hundred males taken in
the late winter or early spring, I should judge that the enlargement of the
testes, seminal vesicles and Cowper’s glands does not occupy more than 3 or
4 days.

The Female Reproductive Organs. The female reproductive organs of
Sorex araneus have been well described by Brambell (1935). These organs
in Sorex fumeus (Plate II, Fig. 4) do not differ markedly from the former
species. Inasmuch as these organs are often of minute size, and their parts
not easily determined, it seems desirable to give a brief description of them
so that American investigators may recognize the different structures.

The ovaries of immature and anoestrous adults are minute, oval yellow-
ish bodies, measuring no more than 1 mm. in length. The much larger
ovaries of breeding animals appear like small mulberries, with corpora
lutea and follicles projecting above the surface. The corpora lutea, num-
bering 2 to 5 for each ovary, often occupy the entire ovary. A prominent
periovarian sac completely invests the ovary. The oviduct is well marked
from the uterine cornu, its ovarian part having a much greater diameter
than the uterine end. The entire oviduct is bent in the form of an S.
Brambell (1935) states that the Fallopian tube passes diagonally through
the uterine wall and projects slightly into the uterine lumen. When stretched
taut, the uterine horns join at right angles to the elongated vagina, but
when in situ they are coiled and thus occupy little space. The vagina is
unique in being bent back upon itself, the distal portion lying dorsad to
the proximal portion, so that the junction of the vagina with the uterine
horns lies near the site of the bladder. This is an adaption for the excessive
length of the vagina which, if it lay in position similar to other small mam-
mals, would reach nearly to the kidneys. The lower part of the vagina
merges into the urogenital sinus, where the urethra opens. The clitoris is
not well marked.

The three pairs of teats are inguinal, their glands occupying much of
the post abdominal and inguinal region in lactating individuals. The first
pair of glands are by far the largest, occupying a large area in front of the
thighs and extending almost to the sacral vertebrae. In immature shrews
and anoestrous females they are small and inconspicuous.

The Breeding Season. The season of reproduction normally extends
from early April into August.

1940]

Hamilton: Biology of Smoky Shrew

483

Males. Males secured in January and February are all immature,
weighing from 4.2 to 6 grams. The testes of these animals are minute and
contain no sperm. In order to determine what individuals were fecund, the
testes were removed, freed from all extraneous tissue and weighed. Micro-
scopic examination of the epididymis was made to determine the presence
or absence of sperm. It was soon evident that size of testes and macro-
scopic examinations of the epididymis were sufficient to determine fecundity.
When the combined weight of the testes exceeded 75 milligrams, they in-
variably contained mature and abundant sperm. Those individuals which
were not fecund had small testes, seldom exceeding 2 mm. and together
not exceeding 10-15 milligrams. Sexually mature individuals possessed
testes measuring 4X6 mm. to 4.5 X 8 mm. and the pair weighed from
105 to 145 milligrams.

The change in size of the testes and the increase in weight of the
shrew at the commencement of the breeding season is accomplished with
great rapidity. Several specimens taken in mid-March of successive years
weigh only 5 or 6 grams and have minute testes. During the last week of
March a number have been secured weighing 8-11 grams and containing
enlarged descended testes which have numerous sperm. On the same dates
a few have been collected which still retain their small obscure testes and
give no indication that they will, in a few days, become fecund. I have
taken no immature shrews from early April until early June. All males
secured during this period have had enlarged testes, with swollen vas
deferens and seminal vesicules (Plate II, Fig. 3). Adult males continue
to maintain their fecundity through August, and the only adult captured
in early September likewise had enlarged testes and abundant sperm.

Females. The females, like the males, are all immature during Janu-
ary and February, and show little sexual activity during March. Four
females collected on March 29 weigh 6.7 — 10 grams, and all but one show
no sign of breeding. This individual has a swollen uterus and indicates
sexual activity, but there are no corpora lutea nor any sperm in the vagina.
Nine females collected from April 12-14 are, judged by weight, mature
animals but none were pregnant. Shrews taken on April 22 contain em-
bryos measuring from 1-7 mm. Individuals taken on May 1 contain em-
bryos but are not nursing. On May 5 I took 2 females, one of which con-
tained five 5 mm. embryos and another which was actively nursing. The
uterine horns show only faint placental scars, so parturition probably
occurred prior to May 1.

Mating may follow partus as it does with numerous cricetine rodents,
for a large female taken on May 16 was actively nursing and contained
eight 2 mm. embryos. Two other females taken on the same date were
nursing but were not gravid. In late May a number of lactating females
have been examined, but none of these were pregnant. Twenty-four adult
females taken during June were all in breeding condition. Of these, 4
were pregnant and the remainder were nursing individuals. Thirteen adult
July females were all nursing, and 2 were pregnant. During August, adult
shrews become increasingly scarce, only one of the 4 adult females which
were examined during the second week of this month being in a breeding
condition. No breeding occurs during September. An interesting instance
of October breeding is given by Green (1930) who took a recently nursing
shrew on October 11 at Ricketts, Pa. This is a most unusual circumstance
and is difficult to explain, inasmuch as I have examined 68 females during
this month, and all have been immature animals.*

From these data it may be seen that males become fecund earlier in
the spring than the females, and individuals usually continue in a breeding
condition after the females have all died or at least ceased to breed. It is
thus concluded that the female controls the breeding season, a condition

* Of 32 females taken at Ithaca, N. Y„ from Oct. 3-10, 1940, one was a recently lactating adult.

484 Zoologica: New York Zoological Society [XXV :29

similar to that of Blarina, Peromyscus and several other small mammals
which I have studied.

Duration of Pregnancy. Little data are at hand regarding the gesta-
tion length of shrews. This is, in large measure, due to the difficulty of
keeping these animals in captivity a sufficient time to breed and thus
observe the gestation period. Blarina has a probable gestation period of
21 days (Hamilton, 1929). According to Brambell (1935), the probable
gestation of Sorex araneus is between 13 and 19 days, with a similar lacta-
tion period.

If sufficient numbers of adult females are examined prior to the incep-
tion of the breeding season, and suitable samples obtained at several day
intervals, it is possible to estimate the probable length of the gestation.
Prior to April 15, no pregnant shrews have been secured, although more
than 20 individuals have been examined from March 15 to April 15. On
the latter date, vaginal smears from one individual indicated the shrew
was in oestrous. On April 22 and 23, shrews which I examined contained
embryos having a crown-rump measurement from 1 to 7 mm. On April 26,
29 and 30 and on May 1, several shrews contained embryos measuring
from 3 to 10 mm. The earliest records of lactating shrews occur on May
5, when 2 nursing specimens were secured. One had faint placental scars
while the other had produced young very recently. Females taken on May
3, 5 and 9 were all actively lactating.

If the earliest pregnancies occur on or about April 15 (based on the
7 mm. embryos secured on April 23) and the earliest parturition date on
or about May 4, this would place the gestation length at approximately 3
weeks. It appears likely that the gestation of this shrew is no longer than
3 weeks duration, and probably somewhat shorter.

Litter Size. Based on counts of embryos and placental scars in 42
females, the litter size averages 5.5 embryos per female. Absorption of the
embryos occasionally occurs, particularly in the late litters so that an
embryo count is not a positive criterion of the young actually produced.
Nevertheless, the six mammae suggest that 5 or 6 young are the usual
number born (Plate III, Fig. 5). The largest embryo count was 8, the
smallest 2. Middleton (1931) has shown that the litter size of Sorex
araneus becomes smaller in July, at the close of the breeding season. One
might suppose that as the females became senile and approached death
during mid- to late summer those continuing to breed would have smaller
litters. Such diminution is not well marked. In April the litter size, based
on embryo counts, was 5.8. During May it averaged 6.2; for June, 5.5, and
for July, 4. A July 12 female contained 2 embryos. The single pregnant
August female contained 6 embryos, one of which was being resorbed,
while a lactating shrew taken on August 12 had 4 placental scars.

Dermal Glands.

Side Glands. All soricids possess side glands, small oval areas of
agglutinized hairs on the side of the body midway between the fore and
hind limbs. These glandular areas are composed of enlarged sweat glands
lying in the deeper part of the corium of the skin and of large sebaceous
glands above the sweat glands in the upper portion of the corium (Eadie,
1938).

Arnback-Christie-Linde (1907) found side glands restricted to the
male in Sorex vulgaris and Crossopus fodiens but present in both sexes in
Crocidura. Both sexes of Sorex fumeus possess these glands, although
they are better developed in the male than in the female. The glands may
not be evident in immature females, whereas in immature males the
glandular areas are always marked on the flesh side of the skin.

A noticeable enlargement of the side glands occurs in the late winter,

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Hamilton: Biology of Smoky Shrew

485

coincident with the enlargement of the reproductive organs. The glands
then become highly vascularized (Plate III, Fig. 6) and are evident as
large reddish patches on the skin surface. Johnsen (1914) believes the
scent glands, at least in the male, play an important part in the sex life of
the animal. He has observed that in male shrews, the growth of the sweat
glands and preparation of the secretory stages parallel the ripening of the
testes. This condition certainly obtains for Sorex fumeus. Throughout
the breeding season, the glands remain prominent in both sexes, although
non-breeding adult females do not exhibit pronounced glands.

The function of these glands is problematical, but it seems possible
they act in some manner to attract the sexes during the breeding season.
Their relation to reproduction is evident, for only at this season are the
glands well developed.

On June 12, 1932, I captured a female smoky shrew in a tin trap. The
animal had been dead a short time. As I was standing near the trap, a
shrew appeared in the runway, ran about the trap several times and
appeared to be much excited. Its side glands were very evident, the hairs
about these structures being laid back so that the semi-glabrous glandular
area could readily be seen. No odor was visible until I knelt down and
smelt closely of the burrow it had occupied, when the peculiar and char-
acteristic smell of this shrew was quite evident.

Life Span.

It has long been apparent that the long-tailed shrews caught at periods
other than the spring and summer months (late March to early Septem-
ber) were all immature individuals. A critical examination of these
specimens suggests that the smoky shrew does not become mature until
late winter and almost the entire adult shrew population dies during the
late summer.

Many observers have remarked on the supposed autumnal mortality of
Sorex, but Adams (1910) was the first to suggest the true cause. He could
trap no adult males nor females after December, and concluded that the
autumnal epidemic is due to nothing other than old age, this being reached
in the case of Sorex araneus and S. minutus in thirteen or fourteen months.
Middleton (1931), apparently unaware of Adams’ study, reached a similar
conclusion, viz., that all adult shrews ( Sorex araneus), following the short
but prolific breeding season, gradually die out, so that none are left by the
following winter. Recently Brambell (1935), from a study of 1,064 trapped
shrews ( Sorex araneus), likewise concludes that the females live over a
single winter only and die at the close of their first breeding season, the
year following that in which they were born, and that the young males do
not breed in the season in which they were born, and do not live over a
second winter.

Montague (1922) questions the abundant testimony in support of this
short life span, for a specimen of Sorex araneus grantii which he collected
in June appeared to be undoubtedly above normal age, the teeth being almost
completely worn away, while the coat is grizzled.

Certain distinguishing characters readily separate the immatures from
the adults of Sorex fumeus. Shrews weighing 7 grams or more may be con-
sidered adult. Those with glabrous tails, the tip more or less rounded and
blunt, with few or no hairs and often with numerous scars, may be con-
sidered adult. Prominent tooth wear is characteristic of adult shrews
(Plate IV, Fig. 8).

Perhaps the most distinctive feature of young shrews is the general
hairy condition of the tail, particularly at its tip. In young shrews, this
member retains its hairy condition until April, when the shrews are eight to
eleven months old (Plate III, Fig. 7).

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[XXV :29

During the year in which they are born, none of these young shrews
become sexually mature. I have carefully examined the reproductive organs
of more than 300 fresh immature specimens, and all indicate a quiescent
stage, in which the testes are minute, seldom exceeding a millimeter in
length, and the ovaries microscopically showing no indication of sexual
activity. Moreover, I have collected only one adult shrew later than the first
of October. It is probable that a few adult females live into the fall, for
Green (1930) secured a nursing Sorex fumeus in northern Pennsylvania
on October 11, 1927. Brambell (1935) states that immature females of
Sorex araneus may exhibit signs of precocious sexual activity in the autumn
of the season in which they are born. Even though follicular growth and
vaginal cornification occur there is no evidence that these immature shrews
become functionally mature. Such animals gradually pass into the typical
winter condition.

Text-figure 1.

Graph demonstrating the relation of weight to age. A pronounced accelera-
tion in weight is evident in March, when both sexes, which have overwintered as
immature individuals, become sexually mature. From April to September, adults
may be recognized by the increasingly worn condition of the teeth and the
glabrous condition of the tail. The pencil of hairs is usually lost in May or June,
but a few may be retained into August. It may be seen that the adult shrew
population dies during August. A few may survive into the fall, but such is
most unusual. The immature shrews lose weight in mid-winter, but their tran-
sition in March from immaturity to adults is remarkably rapid.

Examination of Text-figure 1 will show the marked disparity between
the weights and tail condition of immature and adult shrews. We may thus
conclude that, with Sorex fumeus, all of the adults, after completing their
reproductive duties, die of old age when they are fourteen to seventeen
months old.

Populations.

Our small native mammals are, for the most part, cyclic in nature, their
populations varying from year to year. Some seasons they may exist in
considerable numbers, while in other years only a few may be trapped.

During July and August, 1938, I trapped various habitats on the Huyck
Preserve at RensselaerVille, Albany County, New York, to determine popu-
lation levels of small mammals. In the more favorable habitats of beech
and hemlock, with ground cover of Taxus and broad leaved herbaceous
species, deep leaf mold and litter, I established several quadrats measuring

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Hamilton: Biology of Smoky Shrew

487

100 by 100 feet (.23 acre). Within these areas 300 traps were placed in the
most promising runways, particularly in subterranean burrows and trails
beneath the leaf mold. These traps were visited at dusk and dawn for a
three-day period, or until the catch had been so reduced it was assumed the
major part of the population had been removed. On one of these quadrats
65 small mammals were taken in three nights, 12 of which were Sorex
fnmeus. On another plot in similar habitat a quadrat yielded 68 small
mammals, 10 of them being Sorex fumeus. In an open, sparsely wooded
second growth stand of beech, maple and hemlock, with thick carpet of
leaves and moss, 41 small mammals were collected on a quadrat. Six of
these were smoky shrews. Thus on three quadrats of nearly a quarter acre
each, there were 12, 10 and 6 smoky shrews collected, or an assumed average
population of between 25 to 50 Sorex fumeus per acre. This was in the
very best shrew habitat. One well wooded ravine, with a permanent cold
stream and with black loamy slopes covering loose rock, appeared to be
particularly attractive to these shrews. Yellow birch, mountain and striped
maple, hemlock, witch hobble, with ground cover of liverworts, mosses and
forest perennials, provided a cool retreat, and the black friable soil sup-
ported a particularly rich invertebrate fauna. In 1937 I trapped 70 fumeus
during a three-week period in an area scarcely exceeding 1.5 acres. This
intensive collecting had its effects, for a brief trapping period in 1939 indi-
cated the population had been much reduced.

These high populations are exceptional, and are only met with in the
most suitable habitats during years when shrews are numerous. In 1936
several quadrats in a beech woods near Ithaca, New York, on which I col-
lected produced an estimated population of 9 and 14 fumeus per acre
respectively.

While these shrews exhibit definite variable populations, and give every
indication of being cyclic, I am not certain how long the cycle runs its course.
About Cleveland, Ohio, Bole (1939 ) finds the smoky shrew very susceptible
to drought, and believes they are affected by cycles as well. During the
1937-38 Sorex peak their numbers were estimated at 58 per acre, one of
the highest ever discovered for any species by Bole. He remarks that in
habitats other than climax forests this shrew is distinctly uncommon, the
average being less than 1 per acre. Wherever I have found Sorex fumeus,
the population has never been this low, although it is probable in some
of the areas which I have trapped in western New York, the numbers ot
smoky shrews would not exceed 5-6 per acre.

The autumn population is composed almost entirely of young shrews,
and as no breeding occurs from September until April, there is naturally
some attrition during this season. Some years it may be rather high.
Collecting was conducted on Connecticut Hill, near Ithaca, New York, in
beech-hemlock woods with deep leaf mold. Traps were grouped in lots of
5-10 about stumps, beneath fallen logs, in leaf mold runways and kindred
places. In 1,000 trap nights 155 mammals were collected, 17 being Sorex
fumeus. During the first week of May, 1939, a similar trap line, in compar-
able habitat and under similar conditions was attended with markedly less
success. Forty-six mammals were taken, of which 4 were Sorex fumeus. The
population increased decidedly during the summer. From October 3 to 8,
1939, a thousand trap nights under situations comparable to the previous
year produced 101 mammals, but of these 30 were smoky shrews. During
the first week of May (May 1-5), 1940, the same number of traps produced
28 specimens and 6 of these were fumeus.

Thus we may conclude that, in these two years, scarcely more than 20
to 25% of the shrew population survived the winter. The winter mortality
does not appear to be so great with the smaller rodents and Blarina, but even
with these the loss may be 60% of the entire population.

488

[XXV :29

Zoologica: New York Zoological Society
Enemies.

Every small abundant mammal has a host of enemies ; the smoky
shrew has proved to be no exception.

The larger predators, such as the fox and bobcat, are known to capture
and eat Sorex. Indeed the only item in the stomach of a large Vermont
bobcat which I captured contained the entire body of a Sorex cinereus, and
it is unlikely that they would shun fumeus. Hawks and more particularly
owls, are known to prey upon them. Fresh pellets from a nesting pair of
long-eared owls collected during June, 1926, near Ithaca, New York, con-
tained the skulls of this species. Weasels capture considerable numbers of
the long-tailed shrews (Dearborn, 1932; Hamilton, 1933). All of these
larger predators probably make serious inroads on the shrew population
but there is a more deadly and ubiquitous predator which possibly levies a
fearful toll, and may be, in part, responsible for drastic reductions in the
Sorex population. I refer to the ever-present Blarina, which may weigh
two to four times as much as its smaller relative. Its greater size and
formidable dentition are more than a match for the largest and strongest
fumeus. On May 18, 1938, I caught a 20 gram Blarina whose stomach
contents consisted entirely of the fur, flesh and toes of a smoky shrew. No
Sorex had been captured in nearby traps, so it is good presumptive evidence
that Blarina had overpowered and devoured its lesser kin. It is not without
reason to suspect that this happens not infrequently in areas occupied by
both species.

Parasites. The following parasites have been determined by the staff
of the Bureau of Animal Industry, U. S. Department of Agriculture.

One seldom encounters a shrew without its quota of ectoparasites.
Fleas are seldom found, and are never as abundant as they are on the larger
Blarina. The only fleas which have been recovered from the smoky shrew
are Ctenophthalmus pseudagyrtes and Doratopsylla blarinae, both commonly
found on Microtus, Peromyscus and Blarina. Mites of the genus Myobia,
Haemogamasus and Protomyobia (probably P. clarapedi) are often
abundant, and immature individuals may exceed a hundred to a shrew. The
small gamasid species burrow into the skin, particularly about the ears and
vent, and these are the parasites which cause inflammation and pruritus.
The anal region is often extensively scarred and pitted by the action of
the mites.

Encloparasites apparently are restricted to a few nematodes. One,
Porrocaecum sp., is found coiled in a sheath among the muscles, the external
Avails of the stomach and attached to the viscera generally. It is a sizeable
roundworm, measuring 40 mm. or so in length, and there may be 10 or 12
when the infestation is heavy. It is thought that the adults occur in birds of
prey but no experiments have been conducted to demonstrate this. Similar
roundworms occur with great frequency in Blarina.

Summary.

A life history study of the smoky shrew, Sorex fumeus, in New York
has been made over the past ten years to secure data on the biology of this
little known species. Five hundred and sixteen shrews have been examined
in the flesh and a number of captive animals have been studied, providing
much new information on their biology.

It is shown that external measurements are of little Value in deter-
mining age classes, but weight is of decided value, readily separating im-
mature from mature individuals.

The smoky shrew undergoes a biannual molt. The change from the

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Hamilton: Biology of Smoky Shrew

489

gray winter pelage to the brown summer pelage occurs from late April
until late June, and the fall molt occurs from mid-September to early
October.

The smoky shrew is a northern species, attaining its greatest abundance
in the cool forested regions of Pennsylvania, New York and New England.
It is most numerous in the hardwood and coniferous forests which have a
deep leaf mold.

This species is active at all hours. It is found in company with several
other small mammals, whose runways it occupies. These associates are
Blarina brevicauda, Sorex cinereus, Peromyscus leucopus and P. manicula-
tus, Clethrionomys gapperi, Synaptomys cooperi, N apaeozapus insignis,
Parascalops breweri and several other species. In the habitats of this shrew
where collecting has been most extensive, Sorex fumeus represents 18.3%
of the small mammal population.

Little burrowing is done by the smoky shrew; it appears to occupy the
runways of other small mammals, building its nests beneath stumps, rotted
logs and in the galleries of Blarina and various rodents.

Sorex fumeus has diverse little notes, from high pitched grating, bat-
like squeaks to indiscernible twitters. Sight and smell are not well devel-
oped, hearing is reasonably acute, but reliance is placed on the acute tactile
sense.

On the basis of 168 stomach analyses, made from shrews taken through-
out the year, it is concluded that insects are by far the most important
food, being found in 80% of the stomachs. Vegetable matter was found
in 14.9%; snails, 10.1%; spiders, 5.9%; sowbugs, 5.3%; mammals,
3.0%; salamanders, 1.8% and birds, .6%. Captive shrews maintain them-
selves well on an assortment of natural foods equal to half their weight
daily. Captive individuals seldom drink.

The gross anatomy of the reproductive organs are described, attention
being directed to the rapid increase in size of these organs at the onset of
the breeding season. There are six teats, post-abdominal and inguinal in
position. The mammary glands are very large, the first pair nearly en-
circling the body. The breeding season commences in late March and lasts
through August, although only a small number of females breed in the
latter month. Inasmuch as males become fecund earlier in the spring and
remain so later in the summer than the females, it appears probable that
the female controls the length of the breeding season. The duration of
pregnancy is assumed to be somewhat shorter than three weeks. The litter
size averages 5.5.

Dermal side glands are found in both sexes, but are more prominent in
the males. They must bear some close relation to the reproductive period,
for only at such a time do they enlarge and secrete the odorous material
common to shrews.

The life span of the smoky shrew is short, for all the adults die of old
age when they are 14 to 17 months old. This conclusion is based on the
fact that no shrews breed in the year in which they are born, but pass their
first winter as immature individuals. Weight, degree of hairs on the tail
and tooth wear combined are infallible criteria of age, and serve readily
to separate the adults from the immature shrews. Following the breeding
season, the adult population of shrews die, usually in August.

The smoky shrew is probably a cyclic species, for its numbers fluctuate
in different years. The shrew population varies from 5-50 individuals per
acre during the late summer, but is much reduced during the winter, only
20-25% of the population surviving.

These shrews have many enemies, including predatory mammals and
birds. It is pointed out that Blarina may be one of the most important

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[XXV :29

enemies. Ectoparasites include fleas ( Ctenophthalmus pseudagyrtes and
Doratopsylla blarinae), mites ( Myobia , Haemogamsus and Protomyobia) ,
while a nematode, Porrocaecum, is often abundant.

Literature Cited.

Adams, Lionel E.

1910. A hypothesis as to the cause of the autumnal epidemic of the common
and the lesser shrew. Memoirs of the Manchester Literary and
Philosophical Society, Vol. 54, No. 10: 1-13.

Arnback-Christie-Linde, A.

1907. Der bau der Soriciden und ihre beziehungen zu andeim saugetieren.
Morph. Jahrbuch, Vol. 36: 463-514.

Blossom, Philip M.

1932. A pair of long-tailed shrews ( Sorex cinereus cinereus) in captivity.
Jour. Mammalogy, Vol. 13, No. 2: 136-143.

Bole, B. P., Jr.

1939. The quadrat method of studying small mammal populations. Cleveland
Museum of Natural History. Scient. Publ. 5, No. 4: 15-77.

Brambell, F. W. Rogers

1935. Reproduction in the common shrew ( Sorex araneus Linnaeus) I. The
oestrous cycle of the female. Philos. Trans. Royal Soc. London. Series
B. Vol. 226, No. 518: 1-49.

Dearborn, Ned

1932. Foods of some predatory fur-bearing animals of Michigan. Univ.
Michigan School of Forestry and Conservation, Bull. 1 : 1-52.

Dixon, Joseph

1924. Notes on the life history of the gray shrew. Jour. Mammalogy, Vol. 5,
No. 1: 1-6.

Eadie, W. Robert

1938. The dermal glands of shrews. Jour. Mammalogy, Vol. 19, No. 2:
171-174.

Green, Morris M.

1930. A contribution to the mammalogy of the North Mountain region of
Pennsylvania. Ardmore. Privately Publ. pp. 1-19.

Hamilton, W. J., Jr.

1929. Breeding habits of the short-tailed shrew, Blarina brevicauda. Jour.
Mammalogy, Vol. 10, No. 2: 125-134.

1930. The food of the Soricidae. Jour. Mammalogy, Vol. 11, No. 1: 26-39.

1933. The weasels of New York; their natural history and economic status.
Amer. Midi. Naturalist, Vol. 14, No. 4: 289-344.

Jackson, Hartley H. T.

1928. A taxonomic review of the American long-tailed shrews. North Amer-
ican Fauna No. 51: 1-238.

Johnsen, Sigurd

1914. Uber die seitendriisen der Soriciden. Vorl. Mitt., Anat. Anz. Vol. 46,
No. 12: 139-141.

Montagu, Ivor G. S.

1922. On a further collection of mammals from Inner Hebrides. Proc. Zool.
Soc. London, Vol. 43: 929-941.

1940]

Hamilton: Biology of Smoky Shrew

491

Middleton, A. D.

1931. A contribution to the biology of the common shrew, Sorex araneus.
Proc. Zool. Soc. London, I: 133-143.

Rorig, G.

1905. Uber den nahrungsverbrauch einen spitzmaus. Arb. K. Gesuntheits-
amte. Biol. Abt. 4(1903)1: 121-122.

Townsend, M. T.

1935. Studies on some small mammals of central New York. Roosevelt Wild
Life Annals. Vol. 4, No. 1: 1-120.

Wahlstrom, A.

1928. Beitrage zur biologie von Sorex vulgaris L. Zeitschr. fur Saugetier-
kunde. Vol. 3: 284-294.

Weber, Max

1899. Studien uber Saugethiere. Zweiter Theile. Jena. p. 77.

Wilcke, Gerhard

1938. Freilands-und gefangenschaftsbeobachtungen an Sorex araneus L.
Zeits. fur Sdugetierkunde, Vol. 12, No. 3: 332-335.

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Zoologica: Neiv York Zoological Society

EXPLANATION OF THE PLATES.

Plate I.

Fig. 1. Habitat of the smoky shrew. Beneath half -rotted logs and in the deep
leaf mold of beech, yellow birch and hemlock woods, these small shrews
are often numerous. In the area shown, twenty smoky shrews have been
taken in a two-year period, in addition to scores of deer mice, red-backed
mice, woodland jumping mice, short-tailed shrews and an occasional hairy-
tailed mole. Ithaca, New York.

Fig. 2. Head of a freshly killed smoky shrew. The prominent ear, small eyes
and long snout with prominent tactile hairs are characteristic of this
species.

Plate II.

Fig. 3. Reproductive organs of an adult male Sorex fumeus. A, testes; B, caput
epididymis; C, cauda epididymis; D, vas deferens; E, distal swelling of
vas deferens; F, seminal vesicle; G, Cowper’s gland; H, bladder; I, penis.

Fig. 4. Female reproductive tract of the smoky shrew. The uterus shows a
condition about two weeks following partus. Four placental scars may
distinctly be seen. A, suspensory ligament; B, ovarian capsule enclosing
the ovary; C, proximal region of fallopian tube; D, uterine cornu; E,
uterine vessels; F, bladder; G, diffuse prepucial glands; H, vulva.

Plate III.

Fig. 5. Mastology of nursing smoky shrew. The teats are always limited to
three pairs in the position shown.

Fig. 6. Side glands viewed from the flesh side of an adult male Sorex fumeus
collected on April 19, 1936. A strip of skin has been removed from the
back and sides. The highly vascularized glandular area is prominent
against the pale hide.

Fig. 7. Tails of immature and mature shrews. The upper figure is that of an
immature shrew taken in October. Note the well-furred tip. The lower
figure is that of an adult male collected in April. The absence of hairs
on the tail tip and the eventual loss of hair over the entire tail, in addition
to the scarring, is an indication of maturity.

Plate IV.

Fig. 8. Tooth wear in Sorex fumeus. A, an immature specimen collected on June
11 at a probable age of one month; B, immature shrew taken in November;
C, sub-adult shrew collected in March, showing the prominent wear re-
sulting in almost total loss of the brown tipped hooks of the first incisors ;
0, characteristic tooth row of mature shrew, as indicated in July and
August specimens. The teeth of these old shrews have all lost their brown
tips.

HAMILTON.

PLATE I.

FIG. 1.

FIG. 2.

THE BIOLOGY OF THE SMOKY SHREW (SOREX FUMEUS MILLER)

HAMILTON

PLATE II.

THE BIOLOGY OF THE SMOKY SHREW (SOREX FUMEUS MILLER).

HAMILTON

PLATE III.

V

FIG. 6.

THE BIOLOGY OF THE SMOKY SHREW (SOREX FUMEUS MILLER)

HAMILTON.

PLATE IV.

FIG. 8.

THE BIOLOGY OF THE SMOKY SHREW (SOREX FUMEUS MILLER).

Shlaifer & Breder: Behavior of Small Tarpon

493

Social and Respiratory Behavior of Small Tarpon.
Arthur Shlaifer & C. M. Breder, Jr.

New York Aquarium
(Plates I & II; Text-figure 1).

Introduction.

As part of the program covering a study of the life history and habits
of the tarpon, Tarpon atlanticus (Cuvier & Valenciennes), a number of small
specimens were brought to the laboratories of the New York Aquarium.
These were shipped from the field station of that institution on Palmetto
Key, Florida, for laboratory study. They had been collected on Sanibel Island
by Mr. Marshall Bishop (Breder 1939 a and b).

A consideration of the social attitudes and respiratory behavior of these
specimens forms the basis of the present report. These studies have been
reinforced by others made in outdoor pools, both in New Jersey and at the
Florida field station.

Aside from the actual details which this study works out, the suitability
of small tarpon for laboratory work early became apparent. In addition to a
most remarkable hardihood, the peculiar specializations of this species lends
itself, to a remarkable degree, to a variety of purposes. Some of these will
become apparent on a consideration of the body of the report.

For the chemical analysis in Table X by the laboratory of the Bureau of
Water Supply of New York City under the direction of Dr. F. E. Hale, v/e
wish to express our thanks. We also wish to thank Mr. R. S. Mathews of the
New York Aquarium for assistance during the course of the work.

Materials and Methods.

In order to obtain some measure of the attitude of one fish to its fellows,
two methods of approach were employed. One consisted of estimating the
locomotor behavior of the fishes and the other of estimating the extent of
imitation in the specialized respiratory behavior of this species.

The first method, that established by Schuett (1934) and Escobar,
Minahan & Shaw (1936) and subsequently used by others, consists of esti-
mating the amount of locomotor activity by noting the travel in a given
period of the fish under observation by means of projected cubes from
rulings on the glass walls of aquaria.

In these experiments rectangular metal framed aquaria of 50 liters
capacity, measuring 60 x 30 x 27 cm. and containing 48 liters, were used.
The four sides, which were of transparent glass, were ruled in 8 cm.
squares and two mirrors were used to observe movements at the ends. Ob-
servation periods of 15 minutes were employed, the distance covered being
computed from the number of projected 8 cm. cubes the fish passed through

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in that amount of time. The centimeters traversed were obtained by multi-
plying by 8 the number of cubes traversed.

Each fish was observed for 15 minutes of each hour of a three-hour
experimental period. The tarpon used in each day’s experiments were
selected from a group of ten specimens of the same size, about 10 cm. from
the tip of the snout to the base of the tail. In general, experimental tarpon
were alternated every day between the various experimental conditions of a
particular series so as to distribute evenly any possible daily variations. No
observations were made until at least 21 hours had elapsed after a tarpon
had been placed in an experimental arrangement. The tarpon were fed
chopped herring three times a week at the close of the day’s experiments. In
general, new water was used at the start of each set of experiments and
was not changed during the course of their running. When such water was
not oxygenated artifically the oxygen content was 2.0 to 2.7 cc. per liter.
When oxygenated it ran from 5.6 — 5.8 cc. per liter.

Day by day laboratory results for all experiments were checked against
the daily barometric pressure readings of the U. S. Weather Bureau, whose
New York City station is located one block from the New York Aquarium.
No correlations were obtained.

Tarpon respire atmospheric oxygen and for that reason rise to the
surface to gulp air at more or less definite intervals. Since this feature
might be useful in estimating metabolic activity, such behavior was noted
along with the locomotor activity. A marked tendency to rise in groups was
evident and this association of fishes in their respiratory movements was
likewise studied. Since it is possible to make such studies in the field, ob-
servations on the rising of tarpon not confined in small aquaria were also
taken for comparison. These field observations on rising tarpon were made
in part in a private garden pool of fresh water in northern New Jersey and
in part in a pool at the New York Aquarium’s Florida field station. The New
Jersey pool, of irregular shape, has a surface of 15 square meters and is
about 60 cm. deep in its deepest part, and has a volume of 4,300 liters. The
water is standing, additions being made only for evaporation, and is for
most part crystal clear. The Florida pool is nearly rectangular in outline and
has a surface of 115 square meters and is 1 meter deep in its deepest part.
The water is standing, supplied only by rain except on the highest spring
tides wThen sea water enters it through a narrow channel. The color is a
deep tan. The volume of water is 70,000 liters.

Results op Observations on Aquarium Material.

Data obtained in the laboratory, together with their statistical analysis,
are given in Tables I to V. Considering only the locomotor portions of these
tables, it is evident that under the conditions of these experiments there is
little or no significant difference in the locomotor activity between solitary
or homotypically grouped tarpon, excepting only some indications of mini-
mal activity in groups of two. This is in contrast to the marked differences
found in other species where large differentials have been obtained. Hetero-
typic groups of tarpon and goldfish indicated a considerable increase in the
locomotor activity of the tarpon in association with the normally more active
goldfish.

Tables I to V also give the data on aquarium observations of the
respiratory efforts. Like the locomotor-activity figures, these show little if
any significant relationship between rises for air and various degrees of
homotypic grouping. Heterotypic grouping with goldfish shows a general
increase in this type of activity paralleling the increased locomotor index.

There appears to be an imitative factor in this rising to the surface for
air, as the number of nearly coincidental rises in a group shows a well

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495

marked mathematical significance. Furthermore, it will be shown that
models properly manipulated will induce such rises.

Effect of Isolation, Grouping and Tonicity on
Locomotor and Respiratory Activity.

Research on the goldfish, a non-schooling form, by Schuett (1934),
Escobar, Minahan & Shaw (1936), Breder & Nigrelli (1938) and Shlaifer
(1938) has established the fact that goldfishes are significantly more
active when isolated than they are when grouped. Tarpon are, like the
goldfish, non-schooling forms; however, they are definitely aggregating
forms, more so than are goldfish. Accordingly, it was deemed appropriate
to observe the behavior of young tarpon in isolation and in groups as
regards locomotor activity. Even in well oxygenated water, tarpon rise to
the surface periodically to gulp air. They possess alveolar tissue in their
lung-like swim-bladder (Babcock, 1936). This pattern of respiratory be-
havior was employed as an indirect check on metabolic rate and the air-gulp
rate (respiratory activity) was noted. Since tarpon live in both fresh and
salt water, experiments were performed in both media to determine whether
this difference in the chemical environmnt of the tarpon was reflected in any
way in locomotor or respiratory activity.

The statistical analysis of the data in Table I indicates, in general,
that there is no significant difference in locomotor and respiratory activity
between a tarpon in isolation and in groups either in fresh or salt water
and that locomotor and respiratory activity are, in general, the same in both
media. There is, however, some indication of minimum locomotor activity in
the group of two in fresh water.

Effect of Heterotypic Grouping.

When individual goldfishes were placed in various heterotypic groups
of fishes (Shlaifer, 1940), the decreased activity that obtained in homotypic
groups disappeared. It was thought desirable to investigate the effect of
heterotypic grouping on individual tarpon, although it must be remembered
that a tarpon in a group of four has the same rate of locomotor activity as
does one in isolation. Tarpon were observed in isolation, in homotypic
groups of two and of four and in heterotypic groups containing in one case
the experimental tarpon and three orange-colored common goldfishes 10 cm.
in length and in the other the experimental tarpon and three golden shiners,
Notemigonus crysoleucas (Mitchill), colored olive-green and also 10 cm. in
length. In the experiments performed on the goldfish heterotypic group and
its control homotypic group, the medium was 90% fresh water and 10% sea
water and it was not oxygenated, having an average oxygen content of 2.40
cc. per liter. In the experiments performed on the shiner heterotypic group
and its control homotypic group, the water was completely fresh, was oxy-
genated with compressed air and contained 5.62 cc. of oxygen per liter.

The data in Table II again indicate minimum locomotor and respiratory
activity in the homotypic group of two tarpon ; especially evident, however,
is the greatly increased rate of locomotor and respiratory activity in in-
dividual tarpon when in the heterotypic situations. At the same time, the
data in Table II and Table III indicate that goldfishes and shiners are
much more active than are tarpon. Though goldfishes differ markedly in
color and to some extent in form from tarpon, shiners do not show nearly
such great differences. It is probably, then, merely the disturbing effect of
greater activity by the goldfishes and shiners that is responsible for in-
creased activity in the young tarpon. At no time were goldfishes or shiners
observed to chase the tarpon and thus directly affect activity. The data also
show a distinctly lower rate of respiratory rises when the tarpon are in the
highly oxygenated water used in the shiner experiment and its control.

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Effect of Visual Contact with the Same and Other Species.

Shlaifer (1939, 1940) demonstrated that the activity of an isolated gold-
fish was significantly decreased when it was in contact through vision alone
with others of its kind as well as when it was actually grouped with them.
This line of evidence helped to establish the fact that the group effect was
integrated on a visual basis. When goldfishes were grouped with various
other species or were in visual contact alone with them, no diminution in
activity obtained (Shlaifer, 1940). Since goldfishes and shiners do pro-
duce an increase in the activity of individual tarpon grouped with them, it
was deemed desirable to pursue the analysis further by eliminating all fac-
tors but vision. Table III indicates the results.

Reference to Table III indicates that visual contact alone with goldfishes
will also increase significantly the rate of locomotor and respiratory activity
of isolated tarpon. In one case, IV vs. IVa, the increase in rises is not sig-
nificant but from the trend probably would be so with more data. Cross-
comparisons between the first parts of Table II and Table III show no
significant differences for isolated fishes and those in a group of four in
visual contact or grouped together. Though the volume available to the
experimental tarpon in the visual contact experiments is only one-half that
in Table II, there is no apparent decrease in activity. Amount of available
swimming space undoubtedly affects activity in these forms but not between
these limits. It will be noted that the activity of the experimental tarpon
in actual grouping with three goldfishes is significantly higher than the
activity of a tarpon in visual contact alone with these goldfishes. Thus, it may
be concluded that while visual contact with goldfishes is enough to increase
significantly the activity of isolated tarpon, it is not as effective as when the
tarpon is actually grouped with them directly ; presumably this is due to the
greater chances for disturbing stimuli in the latter case. Additional data
in Table III reveal that the goldfishes used in the experiments listed for
this table had the same rate of activity as did those in Table II, although the
volume available was reduced by half. The goldfishes were, however, signi-
ficantly more active than the tarpon.

“Imitation” in Respiratory Behavior.

A very interesting pattern of behavior found in the tarpon is the rising
to the surface to gulp air by more than one tarpon at a time in a group of
two or more. One tarpon will rise to the surface and its rise may be fol-
lowed immediately by that of one or more other tarpon. This imitative rise
does not occur constantly, but apparently, as further data will attempt to
show, only when a physiological threshhold has been reached. That such
imitative rises are truly imitative and not merely coincidental may be
determined by the laws of chance. An imitative rise, or at least what we
consider to be one, occurs within a second after the initial rise. The minimum
time between the rises of any given tarpon that are not imitative is 300
seconds; in other words, tarpon when most active in water not too well
oxygenated will rise only once every five minutes. If they are less active
and the water is well aerated, the interval is even greater. The data in
Table IV were obtained during the running of other experiments.

The data in Table IV indicate that in a group of two, 36.8% of the
time a rise by one tarpon will induce a rise by the other. In a group of
four, 69.3% of the time a rise by one tarpon will induce a rise in one or
more other tarpon in the group. If imitative rises do not occur at random
but only when the animal is physiologically ready for an air gulp, then the
results are in accordance with expectations. We find in the group of four
that the percentage of induction of rises decreases as the number of fishes
rising at one time increases. Presumably, the more tarpon, the less the
chance that they will all be in a physiologically receptive state for the in-
duction of imitative rises at the same time. However, comparing the
group of two with the group of four it is seen that the larger the group the

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Shlaifer & Breder: Behavior of Small Tarpon

497

greater the chance that the rise of one fish will induce an imitative rise in
others. Under “additional data” in Table IV may be found evidence for the
visual nature of the imitative rise. We find many cases in which the rise
of a tarpon on one side of a transparent glass partition in an aquarium will
produce imitative rises by tarpon on the other side.

Effect of High Oxygen Content.

Tarpon in nature may be found in salt water that is highly oxygenated
and in fresh or brackish water with very low oxygen content. This ex-
periment was designed to investigate the effect of high oxygen content on
the locomotor and respiratory activity of tarpon in the laboratory. In pre-
vious experiments the water used was not artifically oxygenated and on the
average contained 2. 0-2. 7 cc. of dissolved oxygen per liter. In this experi-
ment compressed air was constantly bubbled through the water and main-
tained an average oxygen content in the aquaria of 5.81 cc. per liter.

The data in Table V, part A, again indicate no effect of grouping on
activity. There is again some indication of minimum activity in the group
of two but no significance statistically. When the data for locomotor and
respiratory activity are compared with comparable data in Tables I and II,
it is seen that while the locomotor activity is the same at the higher oxygen
content, the rate of surface rises is significantly less. Thus when the
oxygen content of the medium is higher, tarpon apparently utilize more
dissolved oxygen and less atmospheric oxygen.

Effective Limits of the Induction of Imitative Rises.

Observations were taken upon individual experimental tarpon in a group
of four in order to determine the mean and extreme ranges for non-imita-
tive and imitative rises by the experimental animals and for rises by the
non-experimental fishes which did not induce rises in the experimental.
Table V, part B, lists the results.

The mean time for an imitative rise by a tarpon, following its previous
rise, imitative or non-imitative, is sufficiently close to the mean time for a
non-imitative rise to indicate the importance of a physiological respiratory
threshhold being reached before a rise by one tarpon will induce an imitative
one in another. It should be remembered that we consider a rise by a
tarpon imitative only if it occurs within a second after a rise by another
tarpon in the group. This is not a purely arbitrary assumption, for if the
fish does not rise within one second it usually will not rise for a minute or
more. The ranges from lowest to highest are great but most data fall close
to the mean. It is not easy to explain these extreme ranges. Probably
long periods between rises indicate greater oxygen uptake at the last gulp
and very short periods indicate very low oxygen uptake. It will be noted
from the column titled “mean activity. per minute” that at the extreme
ranges in the last column the activity per minute (obtained by dividing
the centimeters by the number of minutes) does not differ markedly from
these means and hence would eliminate the possibility that very low or very
high rates of activity were responsible.

Induction of Imitative Rises by a Tarpon Model.

While it is known that the rising of a tarpon to the surface to breathe
air may induce an imitative rise in others in a group, it was thought de-
sirable to determine whether such imitative rises could be induced by ob-
jects other than the fish themselves. Accordingly, the following experiment
was performed :

Four tarpon were placed in water 90% fresh and 10% salt. The average
temperature was 21 degrees C. and the average oxygen content 2.69 cc. per
liter. Attempts were made to induce rises in the tarpon by periodically

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raising to the surface in a typical tarpon respiratory rise a carved wooden
model. This model was 10 cm. long, painted silver, and was a reasonably
accurate though rough duplication of a tarpon in general body form. It
had no fins, eyes nor mouth and was manipulated by the observer by means
of two strings tied at its opposite ends.

Many attempts were unsuccessful, as is to be expected if a tarpon does
not respond imitatively even to another of its kind unless it has reached a
respiratory threshhold. However, over a period of 12 hours of observation
there were 47 successful inductions of rises in one or more tarpon. The
average time for such imitative rises by a tarpon following its last rise
was 4.35 minutes. The lowest elapsed time was 1.50 minutes and the highest
was 12:00 minutes.

Results of Observations on Pool Material.

Data obtained in the field, together with analysis, are given in Tables
VI to X. Under the conditions, estimates of locomotor activity were im-
possible, resulting in the confinement of these studies to the respiratory
efforts of the tarpon. These fish were mostly of larger size than the aquarium
material, the measurements being given in the tables.

“ Imitation ” in Respiratory Behavior.

The figures obtained on “imitative” rises in the two pools studied are
in general agreement with those observed in small aquaria, differing only
in the lesser extent of the imitation, Table IX. Since these pools were much
larger and the fish able to retreat much further from each other and since
the other centrifugal influences of the outdoors were present, this is exactly
what should be expfected. In accordance with this it is evident that there is
a tendency for more fish to rise in groups in closer confinement or larger
groups. This, again, would be expected, in that a greater opportunity of
seeing a fish rise spontaneously is directly proportional to the number
present in a given body of water.

Effect of Temperature on Respiratory Behavior.

In the laboratory the temperature differences were not great enough to
give any measurable difference in the number of rises for respiratory pur-
poses relative to thermal differences in the environment. With the large
temperature differentials found in the outdoor pools, it was possible to es-
tablish a relationship between temperature and the amount of aerial respira-
tory activity encountered. Table VIII gives the rises per fish per hour
tabulated according to the temperature sea.e in steps of 5 degrees Centi-
grade. Text-figure 1 illustrates some of the correlations between tempera-
ture and various elements in the respiratory rises of these practically wild
fish. Table VIII and Graph “A” in this figure show a consistent increase in
the number of rises per hour with an increasing temperature, from the
lowest temperature to which the fish could be safely subjected without en-
dangering their lives, 16° to 20° C., to a range of 31° to 35° C. Above this,
36° to 40°, which is close to their upper thermal limits, the respiratory
rises actually fell off. At this temperature range there was, however, a
marked difference in their activity. At the ranges lower than this one they
swam about actively, which activity no doubt contributed to their respira-
tory needs and consistently added to their basic metabolic rate at any given
temperature. At this higher level, however, they rested quietly in one place,
evidently uncomfortable and making no more effort than was necessary to
rise periodically to the surface. This is interpreted to mean that the in-
hibiting influence on their activity of this high temperature actually re-
duced their oxygen demand below that at the next lower temperature range
where they were normally active.

1940]

Shlaifer & Breder: Behavior of Small Tarpon

499

Comparison of the respiratory behavior of small tarpon according to tem-
perature in outdoor pools.

The solid line in this chart indicates the mean value as taken from
Table VIII while the two dotted lines indicate the maximum and minimum
values obtained.

Practical considerations made it impossible to take oxygen readings of
the waters of these two pools at the time the observations were made. How-
ever, since both were supporting other fishes which are not able to respire
air at the same time, they were clearly not near an anaerobic condition.
Aquarium experiments in which oxygen content was measured showed that
at a content of 2.5 cc. per liter, tarpon rose on the average of 11 times per
hour in the 21° to 25° range. This is higher than any reading in the field,
where we have good reason to suppose that the oxygen content was at all
times considerably above that figure. In similar aquarium experiments at
an oxygen content of 5.7 cc. per liter, the same fish rose on an average of
2.8 times per hour. This value is between the mean and minimum of the
field observations, and there is reason to suppose that the field conditions
more closely approached the latter figure than the former. These two values
are indicated on Text-figure 1 by circled dots. It is noteworthy in this
connection that the thermally lower parts of this chart were taken in New
Jersey and the higher in Florida. Considering the differences in the dis-
solved salts in these two waters (see Table X), it is rather remarkable that
the entire line is so consistent, tending to confirm the aquarium observa-
tions with regard to the apparent indifference that these fishes show to the
tonicity of their environment.

The field observations were each made for stated periods and tabulated
by minutes. Consequently there were always some whole minutes which
were represented on the note sheet by blanks. If a similar graph, “B” in
Text-figure 1, is plotted showing these minutes without rises, expressed as

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percentage of an hour, against a temperature scale, a line is obtained ap-
proximately the inverse of that of “A.” It is, however, clearly not its mirror
image and has the advantage of showing certain features not evident in
the former. It, among other things, takes into account the manner of the
grouping of the rises and smooths out a simple count of the number in an
hour, since it is a measure of arbitrary units (minutes during which the
tarpon were “holding their breath.” If the imitative efforts of the tarpon
modified this need for air seriously, this line should not so closely agree
with the first. For example, fish rising a minute or more apart, as com-
pared with all working within one minute in approximate synchronization,
would result in vastly different lines. It is notable in this connection that
while the number of rises per hour in “A” shows greater consistency
(spread between minimum and maximum) in the highest temperature range,
in “B” there is a reverse effect, this range showing the greatest spread, in-
dicating that while the fish are probably doing little more than answering
physiological needs they are at the same time paying less attention to their
fellows.

These data may be handled in still a third way. If the greatest
elapsed time between rises in minutes and seconds be plotted against a
temperature scale, we obtain a line showing still other tendencies. For prac-
tical reasons the figures on this graph are limited to observations concern-
ing five fish. This figure “C” in Text-figure 1 has rather different character-
istics and actually the solid line of means is less valuable for consideration
than the dotted maximum and minimum, for reasons given below. The line
showing the maximum time between rises recorded is a measure of
how long the fish can “hold their breath” and actually gives an estimate of
the physiological needs of the fish apart from its socialized imitative be-
havior. The minimum line on the other hand gives an estimation of how
soon a fish will follow a fellow to the surface after already having filled his
"lung.” In other words, this line is more of an index of social attitude than
of respiratory need. Thus the mean becomes a time of averages between
physiological demand and social influence. It is notable in this connection
that the greatest spread between “physiological need” and “sociability” is
registered in the lowest temperature range measured. This is interpreted
to mean that while tarpon can hold their breath a long time at this tem-
perature, they are also at liberty to b$ fully social. The line of “maximums”
drops sharply with increased temperature, as should be expected from what
we have already seen in “A” and “B.” The line of “minimums” remains
relatively closer to a horizontal line, indicating that their attitude toward
their fellows is much less influenced by temperature than is their need for
oxygen. It is noteworthy that the time of their greatest sociability is also
at the range of their greatest respiratory activity, 31° to 35° C.

A considerable amount of differential behavior was noted in these field
observations which is not amenable to statistical treatment.

For example, the pool at the Palmetto Key laboratory showed marked
variations in the physical environment from time to time. None of the
figures in the calculations show any significant differences associated with
these changes, although the observed behavior of the tarpon was distinctly
different.

At times when the surface of the water was clear and mirror-like, the
fishes generally took their air with a resounding gulp and considerable
agitation of the water. The surface disturbance so made is shown in Plate
I, Figure 1. The center of the major circle is where the head protruded and
the lesser one, to the right, was caused by the tail. This is typical of the
rises at such times.

At other times the surface was covered with a scum of green algae
that at times formed into a complete blanket. During the processes of
formation the rises were much less evident and usually made without any
tail splash. Plate I, Figure 2, shows the conditions during the formation of

1940]

Shlaifer & Breder: Behavior of Small Tarpon

501

such an algal scum. A fish had just risen to the left of center. This repre-
sents about the maximum disturbance under such conditions.

Later when the scum completely covered the pool, the disturbance in-
cident to rises was even less. The algal blanket was such that it entrained
and held the exhaled air, which floated on the surface for some minutes as
bright green bubbles. This condition is shown in Plate I, Figure 3. The
various bubbles are almost entirely from tarpon exhalations. As the fish
normally emit a bubble from each gill opening, these were frequently seen
as paired bubbles. The two largest to the left of center are such a pair, from
the most recent rise.

While some of these features are referable to the physical condition of
the water, the impression was gained that the tarpon disliked coming in
contact with the green scum any more than was necessary, avoiding the
usual tarpon “roll” of clear water.

Discussion.

Since differences in the locomotor behavior of fishes may be used as
one kind of measure of their mental state or psychic attitude toward their
fellows, the recent development of a simple method of estimating these
differences has stimulated work along such lines. Up to the present all
species used for this type of study have shown distinct differences in the
amount of locomotion in association with various groupings, homotypic or
heterotypic. The species used have been Carassius auratus, Schuett (1934),
Escobar, Minahan & Shaw (1936), Shlaifer (1938, 1939, 1940) Breder &
Nigrelli (1938); Gambusia affinis, Escobar, Minahan & Shaw (1936). All
considered the extent of travel per unit of time. In addition, Breder &
Nigrelli (1938) considered the configurational pattern in three dimensional
space and found similar differentiable behavior.

With this as a background, it came somewhat as a surprise to find that
the tarpon, a markedly aggregating form, gave no mathematically significant
figures from its locomotor behavior when alone or in aggregations. If such
is present, it is extremely feeble as compared with the species previously
studied, as all studies have been based on a reasonably comparable amount
of data. It thus appears that this method may or may not be useful merely
on a basis of whether it “works.” Evidently fishes are no more uniform
from group to group in this regard than they are in others.

This, obviously, is not to say that tarpon are indifferent to one an-
other, for even a most superficial examination will show their strong aggre-
gating tendency. Very possibly the difficulty lies in their relative inactivity
as compared, for example, to the goldfish or shiners. When in the presence of
the latter they show a marked increase in travel. That this relative in-
activity is not merely a condition imposed by close confinement is evident
when it is considered that in the garden pool of clear water their behavior
was closely similar to that of the small aquaria; i.e., at nearly all times they
were quietly aggregated in a relatively compact group, spaced about as they
were in the aquarium.

That the metabolic rate is not vastly different under various conditions
of grouping is shown by the similarity of their rising to breathe under
different groupings; it rises notably only in the presence of goldfish, which
no doubt is a result of their greater muscular exertions incident to their
increased activity in swimming. Thus we are obliged to leave this portion
of the study with the conclusion that the method of estimating social
attitude from a basis of locomotor activity is not useful in estimating the
social attitude of tarpon, on a purely pragmatic basis.

Since tarpon will rise to breathe air for purely physiological reasons
and will also imitate each other in this item of behavior, there are thus
two elements involved. While the full separation of them would be exceed-

502 Zoologica: New York Zoological Society [XXV :30

ingly difficult, it has been shown in the tables that there is a high degree
of imitation involved.

Very low oxygen concentrations induce more rises per hour, as would
be expected, but grouping and changes in volume seem to have little effect.
An unknown variable that must be considered in this connection is the
amount of air exchanged on any given gulp. Judging from the figures,
however, this would seem to have scant bearing on the social part of this
behavior. The work of Shlaifer (1939) on the oxygen consumption of the
goldfish, in which he showed that the rate varied with the grouping, could
only be used in the present species in a properly arranged metabolic chamber,
since both aerial and aquatic respiration are involved. As evidenced by the
social factors to be seen in the rising behavior, such a study should be ex-
pected to give similar results.

In all these observations a rise has been considered imitative only if
made a second or less after the initial rise. Those of a larger time interval
have been considered independent. The question arises as to how long such
behavior can be considered imitative, or, in other words, what of the memory
image and inhibiting influences that give delayed responses? It would seem
that first the tarpon must be in a physiologically receptive state, i.e., have
not recently gulped air, for two gulps in rapid succession have never been
noted. Thus, if a tarpon has partly used up its air supply it may respond
with more or less immediateness on the sight of an individual rising to
breathe in response to a purely physiological need.

Summary.

1. The locomotor behavior of tarpon is not a useful index in estimating
their homotypic group behavior, nor is the extent of their rising to the
surface to breathe.

2. There is a marked tendency toward imitation in tarpon rising to breathe,
which is of social rather than physiological significance. More or less
crude models properly manipulated will induce rises to breathe.

3. Observations in the field, checked against those in small aquaria, agree as
to respiratory behavior and indicate the validity of the use of small
aquaria for such purposes.

4. Goldfish and shiners, which are more active, induce in tarpon a marked
increase in both locomotor and respiratory activity, as do goldfish with
which tarpon are in contact through vision alone.

5. Environmental changes including barometric, chemical (salt and fresh
water) and capacity of aquaria, do not significantly change the locomotor
or respiratory behavior of tarpon, but a large reduction in dissolved
oxygen increases respiratory activity. Large temperature changes also
result in the alteration of the respiratory activity.

Table I.

Effect of Grouping on the Locomotor and Respiratory Activity of Small Tarpon.

I. In Isolation II. In Group of Two IV. In Group of Four

Exp. Mean Mean No. Cm. per Mean Mean No. Cm. per Mean Mean No. Cm. per

Cond. Activity3 of Rises4 Rise Activity3 of Rises4 Rise Activity3 of Rises4 Rise

1940]

Shlaifer & Breder: Behavior of Small Tarpon

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Table II.

Effect of Isolation, Homotypic and Heterotypic Grouping on the Locomotor and Respiratory Activity of Small Tarpon1.

Part A — Goldfish.

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Degree of Imitation in Respiratory Activity1.

506

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1 All observations taken on homotypic groups of tarpon.

2 Data based on observations taken during the running of A of Table I.

3 Data based on observations taken during the running of B of Table I.

4 Data based on observations taken during the running of the experiments of Table IIA.

Table V.

Effect of Various Factors on Small Tarpon.

1940]

Shlaifer & Breder: Behavior of Small Tarpon

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508 Zoologica: New York Zoological Society

Table VI.

Respiratory Activity in a Fresh-Water Pool in New Jersey1.
(One-half Hour Observation Periods).

No. of rises of all fish to surface in groups of:

No.

Date

Time

°c

Fish

1

2

3

4

5

6

7

8

9 Total

6/82

1:28

24

5

3

6

6

0

0

— 15

6/82

2:04

24

5

6

10

6

4

0

—

—

—

— 26

6/93

11:00

24

5

1

0

3

0

0

—

—

—

— 4

6/93

11:45

24

5

5

0

0

0

0

—

—

—

— 5

6/94

3:40

23

5

8

4

3

0

0

—

—

—

— 15

6/91

4:16

23

5

3

2

3

0

0

—

—

—

— 8

7/145

11:21

21

5

13

0

0

0

0

—

—

—

— 13

7/145

11:51

21

5

10

0

0

0

0

—

—

—

— 10

7/20°

2:35

26

9

11

4

0

0

0

0

0

0

0 15

7/206

3:05

26

9

16

4

3

0

0

0

0

0

0 23

8/257

11:00

18

5

0

2

0

0

0

—

—

—

— 2

Rises per
fish hour 1

Fish rising in
2 3 4

Calculations.

Minutes without rises
in percent8

groups by percent of total :

5 6 7 8 9 All Per fish

6.0

20

40

40

00

00

—

—

—

—

73

14.6

10.4

24

37

24

15

00

—

—

—

—

57

11.4

1.6

25

00

75

00

00

—

—

—

—

93

18.6

2.0

100

00

00

00

00

—

—

—

—

83

16.6 ‘

6.0

54

26

20

00

00

—

—

—

—

73

14.6

3.2

38

25

37

00

00

—

—

—

—

83

16.6

5.2

100

00

00

00

00

—

—

—

—

70

14.0

4.0

100

00

00

00

00

—

—

—

—

70

14.0

3.3

73

27

00

00

00

00

00

00

00

63

7.0

5.1

70

17

13

00

00

00

00

00

00

53

5.9

0.8

00

100

00

00

00

—

—

—

—

98

19.6

l Fish

used in

this experiment

: 9.5,

12.5,

11.25,

10.25,

10.75

cm. in

standard

length

, placed in

pool June 7. Added July 19: 11.1, 10.0, 11.9 cm. in standard length. This pool, recently made of
concrete, is fed from a pipeline running from a private home. Water lillies and some Lebistes
reticulatus, together with an occasional volunteer Rana catesbiana, make up the remaining fauna.

2 Fish remaining in deepest part of pool in a small aggregation. Partly cloudly with a
slight haze ; light s-w breeze. Rain threatening.

3 Fair with light easterly breeze. Rain had fallen from 7:00 to 10:00 a.m.

4 Thunder with rain threatening. Hazy and overcast. A small trickle of fresh water entering
pool. Rain began at 4:35 p.m.

5 Sun bright, water crystal clear. Fish active, swimming in a larger group than before, over
an area of about 4 ft. in diameter.

6 Humid and partly overcast; hazy. Light drizzle at 2:40 p.m. Fish in n-w corner, rising
very quietly. Fish near surface for most part.

7 Cloudy, chilly. Weather has been unseasonally cold for some time.

8 Based on initial data, not shown in first part of table.

509

1940] Shlaifer & Breder: Behavior of Small Tarpon

Table VII.

Respiratory Activity in a Brackish Water Pool in Florida1.
(One Hour Observation Periods).

Date

Time

°C

No. fish Number of rises of all

12 3

fish in groups of:

4 5 Total

6/232

9:27

33.5

5

31

22

15

8

5

81

6/253

9:42

30.5

5

29

8

6

0

0

43

6/264

9:52

33.0

5

25

16

12

0

0

53

6/265

4:52

36.5

5

25

16

12

0

0

53

6/296

9:34

34.0

5

50

16

3

0

5

74

6/297

3:31

39.0

5

34

6

3

4

0

47

6/308

11:58

37.0

5

20

18

12

4

0

54

7/1 3

9:25

33.5

5

13

12

9

4

5

43

7/210

2.27

36.0

4

30

6

0

0

—

36

7/411

11:52

34.0

3

67

Calculations

0

0

— — 67

Minutes without

Rises per

Fish

rising in

groups

by percentage of total:

rises

in percent13

fish hour

1

2

3

4

5

All

Per fish

16.2

38

28

18

10

06

43

8.6

8.6

67

19

14

00

00

50

10.0

10.6

47

30

23

00

00

46

9.2

10.6

47

30

23

00

00

51

10.2

14.8

68

22

04

00

06

31

6.2

9.4

72

13

06

09

00

50

10.0

10.8

37

33

22

08

00

53

10.6

8.6

30

28

21

09

12

66

13.2

Mean12

11.2

52

25

16

04

03

49

9.8

9.0

83

17

00

00

60

15.0

22.3

100

00

00

—

—

30

10.0

1 This pool, actually an old alligator hole, which had been artificially enlarged two years
previous, is connected with the sea by a cut channel and receives sea water only at the highest
spring tides. This channel is choked with fallen palmetto leaves, effectively preventing the passage
of even small fish. The pool is usually sufficiently fresh for the island fauna to use for drinking
purposes. These include various birds, coons and gopher tortoises. See Table X for analysis of this
water. Gambusia, Mollienesia, Cyprinodon and both fiddler and blue crabs make up the remainder
of the aquatic fauna.

2 Clear and sunny. Water turbid ; reddish colored. The fish used in this experiment measured
48.89, 31.16, 33.02, 30.48, 29.21 cm. in standard length. The first two were removed in the order
listed and had been placed in pool March 26, 1939, not having been disturbed since then.

3 Sky overcast, distant thunder. Pool covered with scum of algae, forming bright green bubbles
wherever tarpon rolled. Fishes rising very quietly.

4 Rain most of previous night. Green scum on pool much reduced. Water much clearer.

5 Clear, shadows of palmettos reaching pool. Algal scum forming again. Fish scattered out
all over pool. Formerly in deepest part.

6 Light clouds passing over sun. Scum no longer green ; grayish-brown. A little rain previous
night. Tarpon rolling all over pool.

I Clear. Surface of pool cleaner. Tarpon making vigorous splashes.

8 Clear. Surface still cleaner. Fish roll well but lazily.

9 Clear. Scum as day before.

10 Clear. Scum completely gone from pool. Day before fish had been caught and measured. One
died. Rain in the morning. Fish rising in all parts of pool.

II Clear. Spring tides. Day before, after the observation period, salt water entered and as
this observation was being made water was again running in. Some slight scum on water. Another
fish died. Fish feeding vigorously all around edge. Rain previous night. Pool overflowing banks
in places. Fish widely scattered.

12 The preceding footnotes indicate why, with the changed conditions, the last two periods have
been omitted from some of the calculations.

13 Based on initial data, not shown in first part of table.

510

[XXV :30

Zoological Neiv York Zoological Society

Table VIII.

Temperature and Rises in Pools.

Calculations Based on Tables VI and VII.

Behavior Temperature in 5° intervals centigrade

16-20 21-25 26-30 31-35 36-40

0.8 6.0 3.3 16.2 10.6

10.4 5.1 10.6 9.4

1.6 8.6 14.8 10.8

Rises per fish hour 2.0 8.6 9.0

6.0 22.3

3.2

5.2

4.0

Mean

0.8

4.8

5.7

14.5

10.0

Maximum

0.8

10.4

9.6

22.3

10.8

Minimum

0.8

1.6

3.3

8.6

9.0

19.6

14.6

7.0

8.6

10.2

11.4

5.9

9.2

6.2

18.6

10.0

10.0

10.6

16.6

13.2

15.0

% minutes without rises,

14.6

10.0

per fish

16.6

14.0

14.0

Mean

19.6

15.0

7.6

10.2

10.5

Maximum

19.6

18.6

10.0

13.2

15.0

Minimum

19.6

14.0

5.9

8.6

6.2

6:46

3:10

2:61

4:31

4:57

3:16

2:40

4:18

Greatest unit of

19:18

6:03

3:43

3:23

time between rises

8:33

6:05

3:22

5:14

in minutes and seconds1

5:08

2:57

5:02

8:18

2:05

3:41

7:26

5:09

U:20

5:07

5:34

U :01

3:16

2:U

Mean of all

8:07

4:09

3:43

4:04

Maximum

19:18

6:05

5:34

5:14

Minimum

4:57

3:10

2:05

3:23

Mean (5 fish only)

8:07

3:13

3:04

4:01

Maximum

19:18

3:16

5:34

5:14

Minimum

4:57

3:10

2:05

3:41

1 In this tabulation, the hour periods of the brackish water pool have been divided in two and
read as recorded. All entries refer to groups of five fish except those in italics. From left to right
these groups are of 9, 3 and 4 fish respectively.

1940]

511

Shlaifer & Breder: Behavior of Small Tarpon

Table IX.

Comparison of “Imitation” in Various-sized Groups and in
Various-sized Volumes of Water.

Aquaria

N. J. Pool

Florida Pool

Number of fishes in groups

2

4

5 9

3

4

5

Total time of observation
(in hours)

36

36

4% 1

1

1

8

No. fish in group rising'
1

63

No.

31

of rising fish in percent
58 72 100

83

52

2

37

29

16 22

00

17

25

3

—

25

24 06

00

00

16

4

—

15

02 00

—

00

04

5

—

—

00 00

—

—

03

6

— ■

—

— 00

—

—

—

7

—

—

— 00

—

—

— •

8

—

—

— 00

—

— ■

■ — -

9

—

—

— 00

—

—

—

Area in sq. meters
Volume H2O in liters
% of rises alone (weighted means)
% of group rises (weighted means)

Aquai

0.196

48

47

53

Table X.

•ia N. J. Pool

15
4,300
61
39

Florida Pool
115
70,000
60
40

Chemical Analysis of Pools in New Jersey and Florida.

Date (1940)

Florida

June 30 July 4

New Jersey
Aug. 28

Albuminoid ammonia (p.p.m. Nitrogen)

14.000

11.000

.200

Free Ammonia (p.p.m. Nitrogen)

21.000

15.000

.220

Nitrate (p.p.m. Nitrogen)

.25

.25

.25

Chlorine (p.p.m.)

17,400.

18,250.

3.6

Hardness (p.p.m. Calcium carbonate)

8,100

8,100

92

Alkalinity (p.p.m. Calcium carbonate)

106

116

78

POi (p.p.m.)

1.2

1.7

. —

SOi (p.p.m.)

1900

2010

16

Iron (p.p.m.)

.05

.10

.30

Turbidity

1

2

1

Color (p.p.m. Platinum)

220

200

10

Total solids (p.p.m.)

55430

46770

252

512

Zoologica: Neiv York Zoological Society
Bibliography.

Babcock, L. L.

1936. The Tarpon. 4th Ed.:l-75. Privately printed, Buffalo, N. Y.

Breder, C. M., Jr.

1939a. On the Trail of the Tarpon. Bull. N. Y. Zool. Soc., 42(4) :98-110.

1939b. The Tiniest of Tarpon Now at the Aquarium. Bull. N. Y. Zool. Soc.,
42(5) :154-155.

Breder, C. M., Jr., & Nigrelli, R. F.

1938. The Significance of Differential Locomotor Activity as an Index of the
Mass Physiology of Fishes. Zoologica, 23(1) :l-29.

Escobar, R. A., Minahan, R. P., & Shaw, R. J.

1936. Motility Factor in Mass Physiology: Locomotor Activity of Fishes
under Conditions of Isolation, Homotypic Grouping and Heterotypic
Grouping. Physiol. Zool., 9(l):66-78.

Schuett, V. F.

1934. Studies in Mass Physiology: The Activity of Goldfish under Different
Conditions of Aggregation. Ecology, 15:258-262.

Shlaifer, A.

1938. Studies in Mass Physiology: Effect of Numbers upon the Oxygen Con-
sumption and Locomotor Activity of Carassius auratus. Physiol. Zool.,
11 (4) :408-424.

1939. An Analysis of the Effect of Numbers upon the Oxygen Consumption
of Carassius auratus. Physiol. Zool., 12(4) :381-392.

1940. The Locomotor Activity of the Goldfish, Carassius auratus, L., under
various conditions of Homotypic and Heterotypic Grouping. Ecology,
21 (4) :488-500.

EXPLANATION OF THE PLATES.

Plate I.

Respiratory behavior of tarpon under varying conditions. See text for

explanation.

Fig. 1. A typical rise through a clean water surface. July 4, 1940.

Fig. 2. A typical rise through an algae-covered surface to left of center. June
26, 1940.

Fig. 3. Two sets of paired respiratory bubbles left entrained on algae-covered
surface in lower left. June 25, 1940.

Plate II.

Fig. 4. Two tarpon rises at less than a second’s interval. The first is represented
by the rings of large diameter. Bubbles may be seen at its center from
the fish which has already descended. The second, left of center, has not
as yet started the Newton’s ring formation. Drops are still falling from
the splash formed as the fish broke the surface.

SH LAI FER a BREDER.

PLATE I.

FIG. 1.

FIG. 2.

FIG. 3.

SOCIAL AND RESPIRATORY BEHAVIOR OF SMALL TARPON.

SHLAIFER a BREDER.

PLATE II.

SOCIAL AND RESPIRATORY BEHAVIOR OF SMALL TARPON.

Candela, Wiener & Goss: Blood Group Factors

518

3L

New Observations on the Blood Group Factors in Simiidi and

Cercopithecidae* f

P. B. Candela, A. S. Wiener & L. J. Goss.

(From the Department of Anatomy of the New York Medical
College, the Serological Laboratory of the Office of the Chief Medical
Examiner of the City of New York, and the New York Zoological
Society) .

This paper will serve to describe some new findings with regard to the
blood group factors in a lowland gorilla (G. gorilla), and in monkeys belong-
ing to two species of macaque (M. mulatta and M. irus) .

Since these findings are a direct outgrowth of the discovery that cer-
tain conclusions previously reported by one of us (Candela, In press) were
incorrect, it may be profitable to recount the successive stages in the inves-
tigation in a more or less chronological order.

It has been known for some time that the blood group factors A, B and
0, whose presence or absence determine the four blood groups in man, are
to be found also in the blood of the anthropoid apes. This fact was first
suggested by the finding of group A in one chimpanzee, whose blood was
examined by v. Dungern & Hirszfeld (1910-11), and established by the
systematic study of Landsteiner & Miller (1925), which revealed that the
blood of anthropoids contained also the agglutinogen B, as well as A.

Since that time, the continuation of such investigations of the bloods
of Simiidi and Hylobatidae has made it apparent that although all four
groups are represented among the apes, each of the genera lacks at least
one of the groups1. Thus, the chimpanzee, on the strength of about 100
individuals, has only groups 0 and A, while the orang-utan and the gibbon
have only groups A, B and AB. Until recently, the gorilla, represented by
only four individuals, was reported to have only group A2.

This distribution of the blood group factors seemed to imply that the B
factor was not present in either of the two African anthropoids. Since the
gorilla and the chimpanzee are generally conceded to resemble man more
closely than do the Asiatic apes, the apparent lack of the B factor was
anomolous, and stimulated various theories postulating its independent
origin in the Asiatic apes, and in African man.

* Aided by a grant from the Committee on Scientific Research of the American Medical Associa-
tion.

t A more extended report, including details of technique, will be published elsewhere. (Wiener,
Candela & Goss, In preparation).

1 This observation applies also to certain of the races of man, the most outstanding examples
being the American Indians, the Polynesians and the Australian natives.

2 Actually, as will be pointed out in a later paper (Wiener, Candela & Goss, In preparation),
the results quoted on those four gorillas are entirely unreliable. The results reported by Landsteiner
(1928) have been misquoted, since this investigator stated that he could not assign a definite group
to the single gorilla which he examined. The results reported by Weinert on the remaining three
gorillas have been shown by Dahr (1938) to be incorrect, since they are based on an invalid technique.

514

Zoologica: Neiv York Zoological Society

[XXV :31

The importance of these questions to anthropology, and to the problem
of human evolution, prompted one of us (C.) to attempt to determine the
blood groups of the gorillas in the zoological parks of the United States, by
means of tests upon their urines. Such tests are dependent upon the well-
known fact that in man, at least, the group-specific substances responsible
for the individual blood differences are present in water-soluble form in
almost all the tissues and body secretions3.

The results of the tests on the urines of seven gorillas, three chim-
panzees and one orang-utan have been detailed in a previous publication
(Candela, 1940). It is sufficient to state that two mountain gorillas (G.
berengei) were found to belong to group A, three lowland gorillas (G.
gorilla) to group B, while two other lowland gorillas appeared from the
reactions to belong to group AB, and were so reported. Actually, the
diagnoses upon these last two were erroneous, for reasons which will
presently appear (v.i.).

The dependability of the results obtained by testing ape urine was
demonstrated by control tests on twenty human beings, whose groups were
determined by independent tests on urine and blood, with complete corre-
spondence. In addition, the blood of the orang-utan, and that of one of the
chimpanzees were obtained and tested, and here also the results of the urine
tests were confirmed by the reactions of the blood.

Another reason for accepting the urine reactions at their face value lay
in the fact that those urines which gave A or B reactions, did so at the same
dilution levels found to be suitable for testing human urines. As for the
AB reactions, in which the samples removed both the oc and the f3 agglu-
tinins from the test sera, at least one of these reactions, namely that
obtained with the urine of the orang-utan, was confirmed by tests upon his
blood. It therefore seemed plausible that the two other AB reactions,
obtained from the urines of the two gorillas, Janet and Susie, respectively,
were equally valid.

Tests upon the Blood of the Gorilla Janet.

An opportunity to check the accuracy of the urine tests on one of these
two gorillas came with the transfer of Janet from the New York Zoo-
logical Society to the Laboratory of Physiology, The School of Medicine,
Yale University, where certain studies were carried out under general
anaesthesia.

Blood samples obtained at that time were examined by us, and the
unexpected finding was made that the gorilla did not belong to group AB ;
in fact, it did not correspond exactly to any of the four human groups'1,
although it resembled most closely blood of group B.

The serum contained cc agglutinins of moderate strength5, but no /3
agglutinins, and therefore corresponded with human group B serum. The
erythrocytes were tested with anti-A and anti-B testing fluids, prepared
from immune rabbit sera. No agglutination at all was obtained with the
anti-A serum, and only very weak agglutination with the undiluted anti-B
serum, although the latter gave distinct reactions with human B cells in dilu-
tions as high as 1 to 32. Evidently, therefore, the gorilla blood contained
an agglutinogen related to, but not identical with the human agglutinogen B.
Thus, it is no longer accurate to state that the group factors in the blood of
anthropoid apes are in all cases indistinguishable from those found in
human blood.

Since the reactions of the gorilla’s blood in anti-B serum were weak,

3 This statement applies only to “secretors.” In “non-sec retors,” who constitute about 20% of
white populations, the group-specific substances are absent from the secretions, or present only in
traces.

4 In this respect, our results correspond with those of Landsteiner (footnote 2).

5 These were accompanied by relatively weak anti-human species agglutinins.

1940]

Candela, Wiener & Goss: Blood Group Factors

515

they might have been open to question, in the absence of additional sup-
porting evidence. Accordingly, advantage was taken of the death of the
gorilla to study the group-specific substances in its organs. Aqueous ex-
tracts of the submaxillary gland were found to inhibit human A (anti-B)
serum in relatively high dilutions, but to have no effect on group B (anti-A)
sera. This confirmed the grouping of the gorilla’s blood as “B-like, oc

The discrepancy between the results obtained from the tests upon the
b'.ood, and those previously obtained on the urine of this gorilla, necessitated
the re-testing of the urine, not only of this gorilla, but also that of Susie,
of the Cincinnati Zoological Society, who likewise had been thought to
belong to group AB.

An interesting phenomenon was observed, in that while the undiluted or
slightly diluted urines gave AB reactions, when they were diluted 1:6,
they both gave B reactions6, inhibiting the (3 agglutinin but not the oc
(Table 1). Since the inhibition of the a: (anti-A) agglutinin was thus
much weaker than that of the anti-B, the former cannot with any degree
of certainty be attributed to the presence of group-specific substance A7.

Table 1.

Results of the re-testing of specimens of urine from
Janet and Susie, and three other apes.

Name

Species

Urine

Residual Agglutinins In:

Dilution

Group A serum

Group B serum

Janet

G. gorilla

1:3

—

—

Susie

G. gorilla

1:3

—

—

M ’bongo

G. berengei

1:3

+ + + +

—

Gargantua

G. gorilla

1:3

—

+ + + +

Mike

P. pygmaeus

1:3

—

—

Janet

v.s.

1:6

H — 1 — 1 — b

Susie

u

1:6

—

+ + + +

M ’bongo

u

1:6

+ + + +

H — 1 — b

Gargantua

u

1:6

+ + +

+ + + +

Mike

<<

1:6

+ +

+ +

Accordingly, it must be pointed out that the inaccuracy in reporting the
groups of Janet and Susie as AB carries no implication as to the correctness
of the conclusions arrived at with regard to the remaining nine apes in the
series. This is particularly true since eight of these reacted as either group
A or group B. With regard to the orang-utan, whose urine gave an AB
reaction, his blood also gave an AB reaction, as has been already mentioned.

In summaiy, it may be said that, (1) The urines of the gorillas Janet
and Susie had some adsorptive power for a as well as /?, exceeding even
the specific reactions of the urines of the remaining nine apes in the series
and of the twenty human controls, but when tested at suitable dilution
levels, they gave only distinct B reactions. (2) The blood cells obtained
from Janet gave very weak reactions in anti-B serum, and none in anti-A
serum, while the extract of the submaxillary gland gave a distinct B
reaction. The serum of this gorilla contained oc agglutinins but no (3
agglutinins. (3) Based on these new findings, it appears that of the seven
gorillas tested to date, all five of the lowland species (G. gorilla ) belong or
are related to group B, while both of the mountain gorillas (G. berengei )
belong or are related to group A.

6 But at this dilution, the urines of the other apes did not afford clear reactions, nor did the
human urines (Table 1).

7 The phenomenon of the non-specific adsorption of both the oc and the /3 agglutinins, with the
production of false “AB” reactions, has been noted by one of us to occur also in the blood-grouping
of human bones (Candela, In press).

516 Zoologica: New York Zoological Society [XXV :31

Tests Upon the Secretions and Blood of Old-World Monkeys.

The interesting results obtained from the examination of the secretions,
blood and tissues of the gorilla Janet suggested the performance of similar
investigations upon monkeys.

The available information on the subject of the blood groups of monkeys
is based on the examination of the blood alone. The data concerning old-
world monkeys can be summarized as follows: (1) The erythrocytes of the
Cercopithecidae are not agglutinated by anti-A or anti-B sera. (2) The
sera of certain of these monkeys nevertheless contain anti-A or anti-B
agglutinins. In the rhesus monkey (M. mulatto), anti-A agglutinins but
not anti-B are regularly present in the serum ; on the other hand, in
Cercopithecus, anti-B is said to be present, but not anti-A. In baboons and
other monkeys, one or the other agglutinin may be present, depending on
the individual serum (Landsteiner, 1936; Thomsen & Kemp, 1930; Buch-
binder, 1933).

From these observations, it has been concluded that the reciprocal
or complementary arrangement of agglutinogens and agglutinins found in
man and the anthropoid apes, does not hold true for monkeys. In man, and
in the Simiidi and Hylobatidae, when the A factor is present alone in the
blood cells, the serum contains agglutinin f3, (or anti-B) ; when only the B
factor is present, the serum contains oc (anti-A) ; the absence of both
factors from the blood cells is accompanied by the presence of both agglu-
tinins in the serum; finally, when both factors A and B are present in the
blood cells, neither iso-agglutinin is found in the serum.

The finding of a definite B factor in the salivary gland of the gorilla
Janet, accompanied by distinct anti-A agglutinins in the serum, in the
face of the extremely weak B reaction of the blood cells, suggested a new
interpretation of the so-called blood group formula. It seemed possible that
the lack of reciprocity between the reactions of the blood cells and those of
the serum in monkeys might be reconciled by the results obtained from the
examination of the tissues and secretions.

The first opportunity to test the validity of this idea arose when the
death of a Java macaque (M. irus) made available for examination the
sublingual gland, together with a block of tongue tissue8. The aqueous ex-
tract prepared from this material was found to inhibit, in high dilutions,
human group A serum, but not to affect human group B serum, thus estab-
lishing the presence of group-specific substance B in the tissue extract.

Subsequently, the submaxillary gland and a sample of blood were obtained
from a rhesus monkey (M. mulatto). The blood serum was found to contain
a moderately strong oc agglutinin, but the blood cells were not agglu-
tinated by either anti-A or anti-B test sera. The extract prepared from the
submaxillary gland inhibited group A serum, but not group B serum, thus
yielding a definite B reaction, as anticipated from the presence of the oc
agglutinin but not the /? agglutinin in the serum of the monkey.

In order to determine the presence of group-specific factors in the
natural secretions of the macaques, specimens of urine from fifteen monkeys,
identified only by numbers, were tested by one of us (C.). Four different
kinds of reactions were obtained. In the first four samples, the tests all
resulted in group B reactions. Of the remaining eleven, seven gave group A
reactions. Three inhibited the A and B test sera equally; these could not be
definitely interpreted, nor could a definite conclusion be reached upon the
blood group of the monkey whose urine gave only a slight trace of inhibition
of the oc (anti-A) agglutinin.

The key list obtained after the completion of the tests disclosed the
unexpected fact that, although all fifteen specimens of urine derived from
macaques, the first four, which had given B reactions, had come from

8 Except as otherwise indicated, all the material used in this study was personally collected and
prepared by one of the authors (G.).

1940]

Candela, Wiener & Goss: Blood Grou-p Factors

517

rhesus monkeys (M. mulatta) , while the remaining eleven, which had
yielded seven group A reactions, had been obtained from Java macaques
(M. irus) .

Samples of saliva, obtained from the same fifteen monkeys by means of
cotton swabs applied to the floor of the mouth, were tested independently by
another of us (W.). These afforded confirmation of the validity of the four
B and seven A reactions, and revealed that three of the doubtful individuals
belonged to group AB. The fourth, which had yielded an extremely weak A
reaction, gave an equally weak A reaction with the saliva. The decision as
to group is much easier to make in tests upon the saliva, since this secretion
gives specific reactions which are much stronger than those of the urine.
For this reason, the failure of the last mentioned specimen (Table 2, number
11), to give a distinct reaction even in tests upon the saliva, suggested the
possibility of group 0.

The final step in the clarification of the blood group characteristics of
this series of monkeys consisted in the testing of their bloods. The results
of these tests were in harmony with previous findings obtained with the
blood of old-world monkeys, in that the cells could not be agglutinated, while
the sera contained group-specific agglutinins.

However, the identification of A-like and B-like factors in the tissues
and secretions now made it evident that a reciprocal relation exists in
monkeys between group-specific substance in the secretions, and the agglu-
tinins in the serum, just as it does in man and the anthropoid apes between
the agglutinogens in the blood cells and the agglutinins in the serum.
Every monkey whose tissues or secretions contained the B factor alone,
had the a , or anti-A, agglutinin in its blood serum; those in which the A
factor alone was found, possessed the /?, or anti-B agglutinin in their sera;
those three which had both the A and the B factors in their secretions,
lacked both iso-agglutinins from their sera, while the one monkey whose
secretions suggested group O, had both agglutinins <x and f3 in its serum.

Familial Relationships.

After the completion of the blood group studies upon the monkeys, the
family relationships of sixteen of them were clarified by reference to the
records of the New York Zoological Society. These sixteen monkeys fall
into three families, and their mutual relationships, indicated in Table 2 by
the letters x, y and z, may be summarized as follows:

Family x : Java macaques. Father: monkey number 00; mother: monkey
number 7; children: monkeys numbers 6 and 15. In this family, the father
gave a group B reaction, while the mother and two children reacted as group
AB.

Family y: Rhesus monkeys. Father: monkey number 2; mother: monkey
number 1; children: monkeys numbers 3 and 4. All members of this family
gave group B reactions.

Family z: Java macaques. Father: monkey number 5; mother: not
available; children: almost certain, number 8; certain: monkeys numbers
9, 10, 11, 12, 13, and 14. All members of this family gave A reactions, with
the exception of number 11, which gave reactions corresponding to group O.

The laws governing the inheritance of the blood group factors in man
are, in essence, two: (J.) Neither the A nor the B factor appears in an
offspring unless it is present in one or the other parent. (2) A parent
belonging to group O cannot have a child of group AB, and a parent
belonging to group AB cannot have a child of group O.

Although the numbers of monkeys herein reported are too small to
permit definite conclusions, it can be said that in these three families of
macaques, at least, no exceptions were found to the laws which govern the
transmission of blood group factors in humans.

Table 2.

Results of tests upon the secretions, tissues and bloods of a gorilla and seventeen macaques.

518

Zoologica: New York Zoological Society

[XXV :31

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Candela, Wiener & Goss: Blood Group Factors

519

Summary of Results.

1. A gorilla whose blood group had previously been diagnosed as AB
from tests upon the urine, was found actually to belong or be related to
group B, as a result of tests upon the blood and tissues. The apparent dis-
crepancy was due to the reactivity of the urine for anti-A as well as anti-B
sera, at dilution levels suitable for other specimens; in higher dilutions, the
urine gave distinct B reactions. The reactions obtained by tests upon the
blood serum and upon an extract of the gorilla’s submaxillary gland closely
paralleled those of a human group B individual. However, the erythrocytes
were only feebly agglutinated by anti-B sera, indicated that the agglutinogen
present in the blood was similar to, but not identical with the human B
agglutinogen.

Of a total of seven gorillas tested, the revised results reveal that all five
of the lowland gorillas (G. gorilla ) belong or are l'elated to group B, while
both of the mountain gorillas (G. berengei) belong or are related to group A.

2. Examination of the blood of one rhesus monkey ( M . mulatto) con-
firmed the well-known fact that the cells of this primate are not agglutinated
by anti-A and anti-B immune testing fluids, while the blood serum contains
a (anti-A) agglutinin, but no /? (anti-B) agglutinin.

3. The examination of the secretions or the tissues of five rhesus
monkeys (including the one mentioned above) elicited the significant fact
that a B-like factor, which had been thought to be entirely absent, is
actually present, being demonstrable, not in the erythrocytes, but in the
tissues and secretions.

4. The Java macaque ( M . irus ) was found to differ in its serology from
rhesus, who belong exclusively to group B, in that reactions were obtained
corresponding to groups A and AB, as well as group B. In seven of the
twelve Java macaques examined the secretions gave a group A reaction,
while the serum contained the /? or anti-B agglutinin. The eighth individual
was found to possess a B-like factor in his tissues; blood from this individ-
ual was not available for testing. Of the remaining four, three reacted as
AB, in that their secretions inhibited both the <x and the f3 agglutinins,
while their sera did not contain either anti-A or anti-B agglutinins. The
twelfth Java macaque gave very faint group A reactions from his urine and
saliva, but was found to possess both the anti-A and anti-B agglutinins in
his blood serum. Although these findings would seem to suggest a diagnosis
of group 0, further study is necessary for a definite decision.

The finding of an A-like factor in seven Java macaques is believed to be
the first demonstration of the presence of the A factor in any monkey.

5. A correlation of the results of the blood group tests with the
familial relationships of the three families of macaques discloses no excep-
tions to the rules which govern the heredity of the blood group factors in
man.

Conclusions.

The results of our investigations upon the blood group characteristics of
a lowland gorilla, five rhesus monkeys and twelve Java macaques permit
certain conclusions to be drawn. These may have new implications for the
general problem of the blood groups, and for the questions of the serological
interrelationships of the primates.

The findings with respect to the secretions, tissues and blood of the
gorilla provide an exception to the rule which holds for the other anthropoid
apes, whose bloods gave reactions indistinguishable from those of man.

It is believed that our results have established for the first time the
presence, in the tissues and secretions, of both A-like and B-like factors in
the two varities of macaques which were studied. These group-specific

520

Zoologica: New York Zoological Society

[XXV :31

factors are in each case accompanied by complementary agglutinins in the
serum. Thus, a reciprocal arrangement of agglutinogen and agglutinin
previously found only in man and the great apes, is now shown to hold true
also in monkeys.

The apparent lack heretofore of such reciprocity was due to the fact that
blood group investigations on monkeys have been confined to tests upon their
bloods. The reactions obtainable from the secretions and tissues should con-
stitute a valuable implement for future studies in the field of primate
blood groups.

There is a suggestion in our results that certain blood group formulae
may be distinctive for some of the species of anthropoids and monkeys. The
large numbers of studies previously made on the rhesus monkey, re-
interpreted in the light of our findings, justify the belief that all members
of this species are related to group B. In the instance of the Java macaque,
however, even the few herein reported suffice to show that they possess both
the A and the B factors. Among gorillas, results from only seven individuals
remain available at the present writing, since the four previously reported
must now be discounted. These are insufficient to permit deductions; still
it may be significant that all five of the lowland gorillas are now known to be
similar to group B, and both individuals of the mountain species to group A.

It is significant that the only species of primates in which group O
has been found are man and the chimpanzee9-10. This suggests the possibility
that the 0 property may have arisen by mutation from either the A or the B
factor (probably the former), rather than the reverse mechanism, accepted
by some writers, that A and B represent mutations from an original back-
ground of group O.

Acknowledgements.

The authors wish to express their gratitude to Dr. John F. Fulton,
of the Laboratory of Physiology, of the School of Medicine, Yale University,
for his generous co-operation in supplying the blood samples and the
salivary glands of the gorilla Janet. We are indebted also to Dr. Karl Land-
steiner of the Rockefeller Institute for Medical Research for the blood sample
and the salivary gland of a rhesus monkey.

References.

Buchbinder, L.

1933. The blood grouping of Macacus rhesus. J. Immunol., 25, 33.

Candela, P. B.

1940. New data on the serology of the anthropoid apes. Amer. Jour. Phys.
Anthrop., 27, 209.

In press. New data on the serology of the anthropoid apes. Addendum.

In press. Reliability of blood group tests on human bones. Amer. J. Phys.
Anthrop., 27, no. 3.

Dahr, P.

1938. Uber blutgruppen bei menschaffen. Ztschr. f. Rassenphysiol., 10, 78.

v. Dungern, E. & Hirszfeld, L.

1910-11. Ztschr. f. Immunitats., 8, 526.

9 The O property is not common even in the chimpanzee, since of the one hundred individuals
tested, only twelve have been reported to be group O.

10 Although one of the Java macaques which we tested (number 11) appears to resemble a group
O individual, some features of the results indicate caution in the acceptance of such a diagnosis.
Further study of this monkey may reveal it to be a non-secretor, with atypical agglutinins (see
footnote 3 ) .

1940]

Candela, Wiener & Goss: Blood Group Factors

521

Landsteiner, K.

1928. Sur les proprietes serologiques du sang des anthropoides. Comptes
rend. Soc. de Biol., 99, 658.

1936. The specificity of serological reactions. C. C. Thomas, Springfield, 111.

Landsteiner, K. & Miller, P.

1925. Serological studies on the blood of primates. II., The blood groups in
anthropoid apes. J. Exper. Med., 43, 853.

Thomsen, 0. & Kemp, T.

1930. Ztschr. f. Immunitdts., 67, 251.

Wiener, A. S., Candela, P. B. & Goss, L. J.

In preparation. A new approach to the problem of the blood group factors A
and B in the Order of Primates.

Goss: Muscle Dystrophy in Tree Kangaroos

523

32.

Muscle Dystrophy in Tree Kangaroos Associated with Feeding of
Cod Liver Oil and Its Response to Alpha-Tocopherol.

Leonard J. Goss.

It is the purpose of this paper to report muscle dystrophy observed in
tree kangaroos that conforms to that first produced artificially by Goettsch
& Pappenheimer in Guinea pigs and rabbits maintained on a diet deficient
in vitamin E. Similar lesions have been observed in rats by Evans & Lip-
schutz, who thought it to be of nervous origin. Olcott however definitely
associated it with disease of skeletal muscles. A similar condition has also
been seen in lambs and is known as “stiff lamb” disease.

The deleterious effect of oxidation on Vitamin E has long been known.
It has also been shown that Vitamin E is affected by oxidation, when in
association with such oxidizing agents as rancid oil. Madsen, McCay &
Maynard showed that cotton seed oil containing a concentrate of Vitamins
A and D in place of cod liver oil afforded a high degree of protection against
muscular dystrophy in Herbivora and that one of the causative factors of
the disease was in the saponifiable fraction of cod liver oil. McCay, Paul &
Maynard prevented muscle lesions in Guinea pigs by using hydrogenerated
cod liver oil in place of natural cod liver oil. In dealing with encephalomalacia
of chicks, Pappenheimer & Goettsch showed that hydrogenation greatly
stabilizes fats that ordinarily readily become rancid. With these facts in
mind we wish to report the occurrence of muscle dystrophy in tree kan-
garoos, its response to Alpha-Tocopherol treatment and its occurrence when
oils were added to the diet.

In June, 1938, four tree kangaroos were received at the New York
Zoological Park, indirectly from Australia. From June, 1938, until April
20, 1939, these animals were each maintained in the Park on an average
daily ration of 2 small raw potatoes, 1 raw apple, % head of lettuce, % slice
whole wheat bread, 1 medium-sized raw carrot and a small quantity of hay.
The hay varied from alfalfa to mixed timothy and clover. On June 20, 1939,
a fish oil mixture containing 3,400 units of Vitamin A and 660 units of
Vitamin D per gram was added to the ration at the rate of 1 teaspoonful
per animal per day. This oil mixture had been stored in a refrigerator since
February, 1939, in a 29-gallon metal drum which was opened periodically
to withdraw oil through a petcock.

On June 10, 1939, kangaroo No. 1 showed symptoms of stiffness, at
which time the fish oil mixture was omitted from the balance of the usual
ration. For a period of six weeks thereafter she was taken outdoors each
day, weather permitting, and allowed to nibble on fresh green grass. Im-
provement was gradual and after six weeks of such treatment the symp-
toms of muscular stiffness had disappeared.

July 20, 1939, animal No. 2 developed symptoms of stiffness which be-
came progressively worse and resulted in death in 2 days.

On September 22, 1939, animal No. 3 became stiff and died in 7 days.

524

Zoologica: New York Zoological Society [XXV :32 :1940]

The fish oil was removed from the diet when symptoms first appeared. All
muscles, including the tongue, were severely affected.

On October 27, 1939, animal No. 4 became stiff and died in 6 days. All
the dead animals showed extensive gross and microscopic lesions of muscle
dystrophy, characterized by edema, opaqueness, a yellowish white coloration
and in the later stages a gritty appearance. All voluntary muscles, includ-
ing the tongue, were affected. The microscopic changes consisted of necrosis
of the fibres, loss of fibrillar structure and calcium impregnation associated
with varying stages of inflammatory reaction and myocytic regeneration.

Animal No. 1, which became stiff on June 10, 1939, was never put back
on the fish oil diet but rather maintained on her original diet as outlined
above. On January 19, 1940, she again developed typical symptoms of muscle
dystrophy, this time without having had any oil. On January 20 the symp-
toms had become more marked and 25 milligrams of Alpha-Tocopherol were
given by mouth. Three days later the symptoms had disappeared completely.
This animal died July 3, 1940, of actinomycosis without having had a re-
lapse since the administration of Alpha-Tocopherol. The second attack of
muscular degeneration in the first animal affected is difficult to explain in
the absence of oil in the diet. We must, however, assume that the regular
ration bordered on deficiency in Vitamin E. This would account both for
the occurrence of muscle degeneration in the animals when the oil was
added to the diet and for the second attack in the one animal.

The results obtained when a fish oil mixture was included in the diet
indicate that the oil itself has some action in producing muscular degenera-
tion or that it inhibits the action of Vitamin E in preventing the disease.

References.

Goettsch, Marianne & Pappenheimer, A. M.

1931. Nutritional Muscular Dystrophy in the Guinea Pig and Rabbit. J.
Expr. Med., 54, 145.

1934. Nutritional Nyopathy in Ducklings. Ibid., 59, 35.

Lipschutz, Daniel.

1936. Degeneration of the nerve tracts in young rats lacking Vitamin E.
Rev. Neurol., 65:221.

Madsen, L. L., McCay, C. M. & Maynard, L. A.

1933. Synthetic diets for Herbivora with special reference to the toxicity of
cod liver oil. Proc. Soc. Exper. Biol. & Med., 30, 1434.

Madsen, L. L.

1936. The comparative effects of cod liver oil, cod liver oil concentrate, lard
and cottonseed oil in a synthetic diet on the development of nutritional
muscular dystrophy. J. Nutrition, 11:471.

McCay, C. M., Paul, Henry & Maynard, L. A.

1937. The removal by hydrogenation of the properties of cod liver oil which
are harmful to Herbivora, read before the meeting of the Amer.
Chemical Society in Rochester in September.

Olcott, H. S.

1938. Paralysis in the young of Vitamin E deficient female rats. Proceedings
of the Amer. Soc. of Biol. Chemists, 119 LXXIV (June) 1937. J. Nu-
trition, 15, 221 (March 1938).

Pappenheimer, A. M. & Goettsch, Marianne.

1934. Protection afforded by certain vegetable oils against nutritional en-
cephalomalacia in chicks. Proc. Soc. Exper. Biol. & Med., 31:777.

Nigrelli: Mortality Statistics, 1939

525

33.

Mortality Statistics for Specimens in the New York
Aquarium, 1939-

Ross F. Nigrelli

Pathologist
( Plates I-III ) .

Introduction.

Approximately 1,600 specimens which died in the New York Aquarium
during 1939 were examined to determine the cause of death. These are listed
in Taoies I to IV. A large number of specimens, especially fishes, which are
being collected more or less continually for food and which also become
part of the exhibition, are not included in this report. Such forms represent
about 30% of the total population as given in the annual census. Also not
included are those specimens which died immediately after their arrival;
when large collections are made, from 20% to 30% usually survive long
enough to become part of the Aquarium’s exhibited population. However,
in smaller, more selected collections, as much as 90% or more may survive.
It should be remembered that aquatic species of animals, especially fishes,
are difficult to handle and that those factors which tend to weaken them,
e.g., netting, transportation, change of water, also tend to lower their re-
sistance to infective agents.

It is now pretty well established that host specificity among fishes and
their parasites is the exception rather than the rule. One thing is certain —
that under aquarium conditions such a relationship, if it exists at all, soon
ceases to be, and may even take on an epidemic form, as in several cases re-
ported here. In addition, it is known that infective agents tend to become
more virulent in their effects when established on “abnormal” hosts.

Insofar as aquatic animals are concerned, the most virulent of the
parasitic diseases are usually caused by external parasites, either protozoan
or metazoan (flukes and copepods) living for the most part on the skin, eyes
and gills.

It is very difficult in many instances to separate the primary from the
secondary causes of death. For example, the primary cause which lowers the
resistance of a given specimen may be a physical (such as fighting), chemical
(changes in the water), or physiological (humoral or nutritional) one. If
any of these adverse factors is sustained long enough, death follows; infec-
tive agents in such an instance may be but a secondary cause which only
speeds up the demise.

The histo-pathology of the lesions produced by the various diseases has
been, or will be, reported from time to time and therefore is not included
in this report.

The writer wishes to express his thanks to Mr. James Atz for his assis-
tance in the preparation of this paper.

526 Zoologica: New York Zoological Society [XXV :33

Causes of Death of Marine Fishes. Table I.

The great majority of the marine fishes were kept in a closed circulat-
ing system; some of the local species, however, were maintained in water
pumped from the adjoining bay. This is the principal reason why more
than half of the deaths of marine fishes resulted from infectious and para-
sitic diseases (Table V). The largest number of these was due to Oodinium
ocellatum. This protozoan parasite was first observed in the New York
Aquarium in 1935 (Nigrelli, 1936). At that time the fishes harboring it were
indigenous to Sand Hook Bay. During 1939 the infection appeared on fishes
collected from Florida, especially in those forms belonging to the families
Lutianidae and Haemulidae. It is interesting to note that Oodinium ocel-
latum was first described from West Indian fishes kept in the London Aquar-
ium. We have had an opportunity to study the organism from fishes of both
regions and have come to the conclusion that the parasite is the same.

Among Sandy Hook Bay fishes a new protozoan parasite made its ap-
pearance in 1939. The life-history studies of this Trichodina were reported
by Padnos (1939a and b). The parasite is found in large numbers on the
gills of the common puffer and to a lesser extent on other species. The in-
cidence of infection in puffers from Sandy Hook Bay is very high and
studies indicate that these fish may be a potential source of infection of this
and other parasitic diseases.

In previous years, Epibdella melleni, a monogenetic trematode, was the
major cause of deaths of marine fishes, especially those belonging to the
spiny-rayed group (Nigrelli & Breder, 1934). In 1939 most of the fishes
infected belonged to the families Lutianidae, Haemulidae and Chaetodon-
tidae. The latter includes some of the handsomest of all West Indian fishes,
but is most susceptible to this trematode. Various techniques, however,
are employed to keep this parasite under control, as well as other mono-
genetic trematodes of less importance, such as Microcotyle and Diplectanus.
The life-history of Microcotyle and Diplectanus is being studied by Paul
(1939).

One of the most interesting, but less devastating, of infectious diseases
is lymphocystis (Nigrelli & Smith, 1939). This condition has been at-
tributed to a virus causing tremendous hypertrophy of cells of connective
tissue origin. The condition is usually limited to such cells in the skin and
fins, but in more severe instances hypertrophied cells have been found in the
spleen, gastro-intestinal tract and ovaries. Insofar as could be determined,
no cellular disease of this type has been reported for man.

The largest number of mortalities attributed to parasitic and infectious
diseases in marine fishes occurred during the months of July and August
(Table VI), soon after the arrival of a large collection from Florida. As
was stated in the Annual Report of the Zoological Society (Nigrelli, 1940a),
the rate of mortality is far greater in recently arrived specimens.

One disease about which we as yet know nothing, although it was
sporadic in occurrence, involves a peculiar ulceration of the skin around the
mouth and the fins. Plate I, Figs. 1 & 2, shows a queen triggerfish with
this affliction. The condition is first made evident by sores around the
mouth, which continue to spread, resulting eventually in a sloughing off of
tissues in this region; later the fins become involved, there being an in-
tense inflammation of the skin at their bases. Biopsied and autopsied mate-
rial failed to reveal the presence of a causative agent. The internal organs
were normal to all appearances, but in most cases the optic nerves were
inflamed and the optic muscles completely atrophied.

About 20% of the deaths of marine fishes resulted from non-infectious
diseases, with a variety of causes and occuring throughout the year. Most of
these diseases affected fishes which had been on exhibition for more than
one year, and are of the type, with rare exceptions, that normally would be
expected among any animal population of similar proportions and under

1940]

Nigrelli: Mortality Statistics, 1939

527

conditions of confinement. Some of these may be of parasitic origin, al-
though no causative agents have been recovered; others may be entirely
humoral or nutritional. The above remarks also hold for those non-infectious
deaths among the other two groups of fishes.

Fishes are susceptible to relatively slight adverse changes in the phy-
sical, chemical and thermal conditions of their environment — some more
so than others, but all to a certain degree. Therefore it is not surprising
to find that a number of specimens succumbed as a result of such changes.
The Elasmobranchs are particularly sensitive to changes in specific gravity,
for example.

Causes of Deaths of Temperate Fresh-water Fishes. Table II.

A lower mortality rate occurred among the temperate fresh-water fishes.
This may be attributed for the most part to the fact that these fishes were
kept in an open circulation for most of the year. Any infection on one speci-
men was usually limited to that individual or perhaps its tank-mates. The
present closed circulation was put into operation in October, 1939. However,
the increase in mortality during the last two months of the year resulted
from the influx of a large collection from Illinois, rather than the effects
of the closed circulation (Table VI).

The largest number of deaths among this group was caused by Ichthyo-
phthirius multijiliis , a well known and at times very virulent protozoan
parasite. More than three-quarters of the deaths reported from this cause,
however, occurred in one species of catfish. These fish were from one to two
inches long and were confined in a single tank.

Saprolegnia only attacks fresh-water fishes which have been injured
either mechanically or by parasites. Handling thus presents a constant
hazard, since spores of this fungus are omnipresent.

Among helminth infections, some losses were attributed to worms be-
longing to the monogenetic trematode group referred to as Gyrodactyloides.
These are skin and gill parasites, and a large number of species have been
reported from fresh-water fishes. They constitute the most dreaded parasite
in aquarium and hatchery management, often causing the loss of great
numbers of specimens.

The disease referred to as Myxomata is very interesting and in all
probability of parasitic origin. The general effects of any of the external
parasites is to produce an irritation of the skin which results in an ex-
cessive production of mucus, the most important protective mechanism of
fishes. Sometimes a translucent, soft growth remains, covering the whole
fish or parts of it. This growth is microscopically seen to be composed of
stellate-shaped connective tissue cells. One difference between this type of
tumor and those of similar nature reported for man and other mammals is
that the myxomata in the latter are richly supplied with capillaries.

The number of deaths recorded under Temperature is almost entirely
accounted for by losses incurred among the trouts during the summer
months. Mortalities recorded under Chemical Poisoning refer to a single
species. A tank of muskellunge on open circulation was treated for a slight
infection of I chthy ophthirius with a protein — silver salt, a compound con-
tinually used for such purposes. A chemical interaction between this chemical
and the chlorine in the water at that time resulted in the formation of
silver chloride, a highly toxic substance to fishes.

Causes of Death of Fresh-water Tropical Fishes. Table III.

These comprise some of the smallest of fishes, and it is consequently
very difficult to perform adequate autopsy. For this reason, many deaths
resulting from undetermined causes are listed here.

528

Zoologica: New York Zoological Society

[XXV :33

It is known among ecologists, aquarists and tropical fish fanciers that
fishes living in small standing bodies of water alter the chemical content of
their environment, apparently not only making it more suitable for them-
selves, but sometimes rendering it completely unsuitable for even rather
closely related species. This has been called water conditioning. It is a
very important factor in the maintenance of tropical fish, and a very large
portion of the 23% of deaths recorded under Acclimatization resulted from
these small fishes being unable to adjust themselves satisfactorily to a new
aquatic environment. Just what this conditioning is has so far evaded
successful analysis. Evidence indicates that some specific animal proteins
may be involved.

Because these fishes are kept in completely separated tanks, the
incidence of infectious disease is practically negligible.

The largest number of deaths among reptiles was attributed to a general
enteritis. Just what was producing this condition in the Aquarium speci-
mens has not been determined. Since no parasitic infection was evident, it
is altogether possible that the condition may be nutritional.

The most interesting of reptilian diseases so far studied is the one
developed by large alligators, previously reported as fatty degeneration
(Nigrelli, 1940a). Large deposits of encapsulated fatty substance may be
found just beneath the skin ; between layers of muscles (Plate III, Fig. 7) and
possibly extending into muscular tissues ; between the muscles and the
peritoneum, lying just outside the latter; and also inside the peritoneal
cavity. There are indications that organs may be involved, too. Animals
suffering from this disease may lie in the pools for weeks, refusing all
food. Just before death they may have one or more severe hemorrhages
through the snout, mouth or anus.

Although the great majority of invertebrates on exhibition were not
autopsied, it is definitely known that these creatures are extremely sensitive
to changes in the chemical composition of the water about them, Table V.
Thus it is believed that the small sample autopsied does not at all exaggerate
the importance of this factor as a cause of death.

Causes of Death of Reptiles. Table IV.

Causes of Death of Invertebrates. Table IV.

Table I.

Causes of Mortality in Marine Fishes.

No.

Disease

Oodinium ocellatum Brown
32. 6%!

Fish

Deaths

Holocentridae

Holocentrus ascensionis (Osbeck),
Squirrelfish

5

Cheilodipteridae

Apogon maculatus (Poey), Spotted
Cardinal Fish

5

Serranidae

Centropristus striatus (L.), Sea Bass
Epinephelus adscensionis (Osheck),
Rock Hind

Epinephelus morio (C. & V.), Red
Grouper

Petrometopon cnientatus (Lac.),
Graysby

9

2

4

6

1

1 Percentage of total number of recorded deaths.

1940] Nigrelli: Mortality Statistics, 1939 529

Rypticus saponaceus (B. & S.), Soapfish 1

Lutianidae

Lutianus analis (C. & V.), Muttonfish 2

Lutianus apodus (Walbaum),

Schoolmaster 7

Lutianus griseus (L.), Gray Snapper 4

Lutianus jocu (B. & S.), Dog Snapper 2

Lutianus synagris (L.), Spot Snapper 9

Ocyurus chrysurus (Bloch), Yellowtail 17

Haemulidae

Anistoremus virginicus (L.), Porkfish 40

Haemulon albium C. & V., Margate Fish 1
Haemulon flavolineatum (Desmarest),
Yellow Grunt 15

Haemulon macrostomum Gthr., Gray
Grunt 5

Haemulon plumieri (Lac.), White Grunt 8
Haemulon sciurus (Shaw), Blue-striped
Grunt 18

Haemulon sp.. Iron Grunt 1

Sciaenidae

Eques acuminatus (B. & S.), Cubbyu 4

Menticirrhus saxatilis (B. & S.),

Northern Kingfish 6

Pomacentridae

Abudefduf saxatilis (L.), Sergeant-major 2
Dascyllus aruanus (L.), Jesuite 8

Scaridae

Scarus caeruleus (Bloch), Blue
Parrotfish 10

Chaetodontidae

Angelichthys isabelita Jordan & Ritter,

Blue Angelfish 4

Pomacanthus arcuatus (L.), Black
Angelfish 23

Pomacanthus paru (Bloch), French
Angelfish 12

Acanthyridae

Acanthurus caeruleus B. & S., Blue Tang 6

Acanthurus hepatus (L.), Doctorfish 2

Balistidae

Balistes vetula L., Queen Triggerfish 4

Ostraciidae

Lactophrys trigonus (L.), Common
Trunkfish 9

Tetraodontidae

Spheroides maculatus (B. & S.), Puffer 23

Diodontidae

Chilomycterus schoepfii (Walbaum),

Spiny Boxfish 17

Didon hystrix L., Porcupine Fish 2

Triglidae

Prionotus carolinus (L.), Carolina

Sea Robin 8

Prionotus sp., Blue-winged Sea Robin 2

530

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[XXV :33

Trichodina sp.
10.3%

Epibdella melleni MacCallum
17.9%

Carangidae

Caranx crysos (Mitch.), Hard-tailed Jack 9

Serranidae

Centropristus striatus (L.), Sea Bass 5

Roccus lineatus (Bloch), Striped Bass 3

Sparidae

Ste?iotomus chrysops (L.), Porgy 2

Sciaenidae

Menticirrhus saxatilis (B. & S.)

Northern Kingfish 7

Tetraodontidae

Spheroides maculatus (B. & S.), Puffer 43

Diodontidae

Chilomycterus schoepfii (Walbaum),

Spiny Boxfish 12

Triglidae

Prionotus carolinus (L.), Carolina
Sea Robin 6

Prionotus evolans (L.), Striped Sea
Robin 8

Holocentridae

Holocentrus ascensionis (Osbeck),
Squirrelfish 2

Carangidae

Vomer setapinnis (Mitch.), Moonfish 4

Serranidae

Cephalopholis fulvus (L.), Coney 1

Dermatolepis marmoratus Osburn &

Mobray, Marbled Grouper 1

Epmephelus adscensionis (Osbeck),

Rock Hind 1

Mycteroperca interstitialis (Poey),

Princess Rockfish 2

Lobotidae

Lobotes surinamensis (Bloch),

Tripletail 1

Lutianidae

Lutianus analis (C. & V.), Muttonfish 5

Lutianus apodus (Walbaum),

Schoolmaster 1

Lutianus griseus (L.), Gray Snapper 1

Lutianus jocu (B. & S.), Dog Snapper 1

Lutianus synagris (L.) , Spot Snapper 5

Ocyurus chrysurus (Bloch), Yellowtail 1

Haemulidae

Anistoremus virginicus (L.), Porkfish 26

Haemulon flavolineatum (Demarest), 2

Yellow Grunt

Haemulon macrostomum Gthr., Gray
Grunt 10

Haemulon plumieri (Lac.), White Grunt 1

Haemulon sciurus (Shaw), Blue-striped
Grunt 3

1940]

Nigrelli: Mortality Statistics, 1939

531

Sparidae

■ Salema rhomb oidalis (L.), Salema 1

Sciaenidae

Pogonias c?'omis (L.), Sea Drum 1

Labridae

Bodianus rufa (L.), Spanish Hog; fish 5

Julis pavo (C. & V.), Swallow-tailed
Wrasse 3

Lachnolaimus maximus (Walbaum),

Hogfish 16

Trechocopus scrofa (C. & V.), Pink
Wrasse 3

Ephippidae

Chaetodipterus faber (Broussonet) ,

Spadefish 5

Chaetodontidae

Angelichthys ciliaris (L.), Queen
Angelfish 4

Angelichthys isabelita Jordan & Ritter,

Blue Angelfish 29

Angelichthys townsendi Nichols &

Mowbray, Townsend’s Angelfish 1

Chaetodon capistratus L., Four-eyed
Butterflyfish 5

Chaetodon collaris Bloch, Lattice-work
Fish 3

Chaetodon ocellatus Bloch, Common
Butterflyfish 3

Heniochus acuminatus (L.), Long-finned
Butterflyfish 1

Platax sp., Sea Bat 1

Pomacanthus arcuatus (L.), Black
Angelfish 7

Acanthyridae

Acanthurus hepatus (L.), Doctorfish 3

Balistidae

Balistes carolinensis Gmelin, Common
Triggerfish 2

Balistes vetula L., Queen Triggerfish 1

Canthidermis sabaco Poey, Ocean
Triggerfish 4

Microcotyle sp.
.76%

Chaetodotidae

Chaetodon capistratus L., Four-eyed
Butterflyfish 2

Chaetodon collaris Bloch, Lattice-work
Fish 3

Chaetodon pictus Forskal, Striped
Butterflyfish 1

Holocanthus tricolor (Bloch), Rock
Beauty 1

Diplectanus sp.
.54%

Pomacentridae

Abudefduf saxatilis (L.), Sergeant-
major 1

Tetraodontidae

Tetraodon fluviatilis (Buch.), Fresh-
water Puffer 4

532

Zoologica: New York Zoological Society

[XXV :33

Lymphocystis

.43%

Pomacentridae

Amphiprion percula (Lac.) , Clownfish 1

Chaetodontidae

Angelichthys isabelita Jordan & Ritter,
Blue Angelfish 2

Monacanthidae

Ceratacanthus schoepfii (Walbaum)

Orange Filefish 1

Ulceration of skin and fins
.54%

Kyphosidae

Kyphosus sectatrix (L.), Bermuda Chub 2
Ephippidae

Chaetodipterus faber (Broussonet) ,
Spadefish 1

Balistidae

Balistes vetula L., Queen Triggerfish 2

Trematode enteritis
1.30%

Holocentridae

Holocenty'us ascensionis (Osbeck),
Squirrelfish 2

Carangidae

Caranx hippos (L.), Common Jack 1

Serranidae

Centropristus striatus (L.), Sea Bass 1

Epinephelus morio (C. & V.), Red
Grouper 1

Mycteroperca bonaci (Poey), Black
Grouper 2

Roccus lineatus (Bloch), Striped Bass 1

Lutianidae

Ocyurus chrysurus (Bloch) , Yellowtail 2

Scorpaenidae

Scorpaena madurensis (C. & V.) , Madeira
Scorpionfish 2

Cestode enteritis
.65%

Galeidae

Carcharhinus milberti Muller & Henle,
Ground Shark 2

Acipenseridae

Acipenser oxyrhynchus Mitch., Common
Sturgeon 2

Carangidae

Caranx crysos (Mitch.) , Hard-tailed Jack 1
Triglidae

Prionotus carolinus ( L. ) , Carolina

Sea Robin 1

Protozoan enteritis
.11%

Diodontidae

Chilomycterus schoepfii (Walbaum),

Spiny Boxfish 1

Gastritis

2.49%

Ginglymostomidae

Ginglymostoma cirratum (Bonnaterre),
Nurse Shark 1

1

1940]

Nigrelli: Mortality Statistics, 1939

533

Acipenseridae

Acipenser oxyrhynchus Mitch., Common
Sturgeon 4

Serranidae

Centropristus striatus (L.), Sea Bass 2

Roccus lineatus (Bloch), Striped Bass 1

Sciaenidae

Bairdella chrysura (Lac.), Silver
Perch 4

Leiostomus xanthurus Lac., Spot 4

Micropogon undulatus (L.), Croaker 1

Sciaenops ocellatus (L.), Channel Bass 6

Biliary cirrhosis
.22%

Siluridae

Galeichthys felis (L.), Sea Catfish 1

Ephippidae

Chaetodipterus faber (Broussonet) ,
Spadefish 1

Fatty degeneration of liver
.22%

Serranidae

Centropristus striatus (L.), Sea Bass 2

Hepatitis, nematode infection
.11%

Gadidae

XJrophycis chuss (Walbaum), Squirrel
Ling 1

Renal calculi
.11%

Serranidae

Mycteroperca bonaci (Poey), Black
Grouper 1

Kidney degeneration
.54%

Serranidae

Mycteroperca bonaci (Poey), Black
Grouper 1

Ephippidae

Chaetodipterus faber (Broussonet),
Spadefish 1

Batrachoididae

Opsanus tau (L.), Toadfish 3

Ovarian degeneration
.87%

Carchariidae

Carcharias littoralis (Mitch.), Sand
Shark 1

Aetobatidae

Rhinoptera quadriloba (Le S.), Cow-nosed
Ray 1

Acipenseridae

Acipenser oxyrhynchus Mitch., Common
Sturgeon 3

Serranidae

Mycteroperca bonaci (Poey), Black
Grouper 3

Fatty degeneration, general
.87%

Muraenidae

Gymnothorax funebris Ranzani, Green
Moray 1

534

Zoological New York Zoological Society

[XXV :33

Fatty degeneration, general

Serranidae

(continued)

Epinephelus striatus (Bloch), Nassau
Grouper

Mycteroperca bonaci (Poey), Black

3

Grouper

2

Lutianidae

Lutianus griseus (L.), Gray Snapper

2

Swim-bladder disorder

Gadidae

2.27%

Gadus callarias L., Cod

21

Gas aneurisym

Sparidae

.54%

Stenotomus chrysops (L.), Porgy

1

Sciaenidae

Menticirrhus saxatilis (B. & S.) ,
Northern Kingfish

1

Diodontidae

Chilomycte?~us schoepfii (Walbaum),
Spiny Boxfish

2

Triglidae

Prionotus carolinus (L.), Carolina Sea
Robin

1

Malnutrition

Syngnathidae

4.65%

Hippocampus hudsonius ssp. De Kay,
Seahorse

16

Hippocampus sp., Long-nosed Seahorse

7

Scaridae

Pseudoscarus guacamaia (Cuv.), Rain-
bow Parrotfish

Scarus caeruleus (Bloch), Blue Parrot-

8

fish

9

Monacanthidae

Ceratacanthus schoepfii (Walbaum),
Orange Filefish

3

Trauma of eyes

Ephippidae

.43%

Chaetodipterus faber (Broussonet) ,
Spadefish

2

Acanthyridae

Acanthurus hepatus (L.), Doctorfish

2

Exophthalmos

Serranidae

.32%

Centropristus striatus (L.), Sea Bass

3

Tumor of thyroid

Cyprinodontidae

1.08%

Cyprinodon variegatus Lac., Sheepshead
Minnow

10

Senility?

Siluridae

.97%

Galeichthys felis (L.), Sea Catfish

2

Muraenidae

Gymnothorax funebris Ranzani, Green
Moray

2

Carangidae

Trachinotus glaucus (Bloch), Palometa

1

1940]

Nigrelli: Mortality Statistics, 1939

535

Labridae

Halichaeres radiatus (L.), Pudding-wife 1
Thalassoma bifasciatus (Bloch),

Bluehead 1

Cichlidae

Tilapia mossambica Peters 1

Diodontidae

Diodon hystrix L., Porcupine Fish 1

Fractured skull

Galeidae

.32%

Mustelus canis (Mitch.), Smooth
Dogfish

2

Sciaenidae

Cy noscion regalis (B. & S.), Weakfish

1

Eaten by tankmates '

Pomacentridae

.22%

Pomacentrus albofasciatus Schl., Striped
Pomacentrus

2

Jumped from tank

Mugilidae

.43%

Mugil cephalus L., Mullet

1

Ostraeiidae

Ostracion comutus (L.), East Indian
Cowfish

1

Scorpaenidae

Pterois volitans (Bennett), Zebra Fish

2

Fighting

Muraenidae

.54%

Gymnothorax funebris Ranzani, Green
Moray

1

Gymnothorax moringa (Cuv.), Spotted
Moray

1

Pomacentridae

Eupomacentrus leucostictus (Muller &
Troschel), Beau Gregory

3

Handling

Syngnathidae

1.41%

Hippocampus h. hudsonius De Kay,
Northern Seahorse

11

Ephippidae

Cliaetodipterus faber (Broussonet) ,
Spadefish

2

Destroyed

Holocentridae

.54%

Holocentrus ascensionis (Osbeck),
Squirrelfish

5

Water chemistry

Ginglymostomidae

11.3%

Ginglymostoma cirratum (Bonnaterre) ,
Nurse Shark

2

Galeidae

Carcharhinus milberti Muller & Henle,
Ground Shark 1

Mustelus canis (Mitch.), Smooth Dogfish 8

Carchariidae

Carcharias littoralis (Mitch.), Sand
Shark 1

536

Zoologica: New York Zoological Society

[XXV :33

Water chemistry
(continued)

Unknown

4.44%

Rajidae

Raja spp., Rays 10

Dasyatidae

Dasyatis centrura (Mitch.), Stingray 5

Aetobatidae

Rhhioptera quadriloba (Le S.), Cow-
nosed Ray 10

Muraenidae

Gymnothorax funebris Ranzani, Green
Moray 1

Syngnathidae

Hippocampus h. hudsonius De Kay,
Northern Seahorse 30

Carangidae

Seriola zonata (Mitch.), Rudderfish 14

Serranidae

Centropristus striatus (L.), Sea Bass 20

Roccus lineatus (Bloch), Striped Bass 2

Megalopidae

Tarpon atlanticus (C. &V.), Tarpon 1

Anguillidae

Anguilla rostrata (Le S.), Common Eel 1
Mugilidae

Mugil cephalus L., Mullet 3

Pleuronectidae

Paralichthys dentatus (L.), Fluke 3

Pseudopleuronectes americanus
(Walbaum), Winter Flounder 8

Soleidae

Achirus fasciatus Lac., Hog Choker 3

Cheilodipteridae

Apogon maculatus (Poey), Spotted
Cardinal Fish 3

Sciaenidae

Eques acuminatus (B. & S.), Cubbyu 4

Menticirrhus saxatilis (B. & S.),

Northern Kingfish 4

Pomacentridae

Amphiprion melanopus Blkr., Anemone
Fish 3

Ostraciidae

Lactophrys trigonus (L.), Common
Trunkfish 1

Tetraodontidae

Tetraodon sp., Pacific Puffer 1

Scorpaenidae

Scorpaena madurensis (C. & V.),

Madeira Scorpion Fish 2

Echeneididae

Echeneis naucrates L., Sharksucker 4

1940] Nigrelli: Mortality Statistics, 1939 537

Table II.

Causes of Mortality in Temperate Fresh-water Fishes.

No.

Disease Fish Deaths

Saprolegnia sp.

Catostomidae

G.66%1

Ictiobus bubalis (Raf.), Buffalo Fish
Moxostoma aureoleum (Le S.), Redhorse

7

Sucker

1

Siluridae

Ameiurus nebulosus (Le S.), Common
Bullhead

5

Ictalurus punctatus (Raf.), Channel Cat

1

Ichthyophthirius multifiliis

Salmonidae

Fouquet

Salmo irideus Gibbons, Rainbow Trout

1

45.3%

Cyprinidae

Campostoma anomalum (Raf.), Stone-
roller Minnow

2

Emcymba buccata Cope, Silver-mouthed
Minnow

2

Hybognathus sp.

Pimephales promelas Raf., Fat-headed

4

Minnow

Semotilus atromaculatus (Mitch.),

1

Horned Chub

3

Siluridae

Ictalurus punctatus (Raf.), Channel Cat

80

Centrarchidae

Ambloplites rupestris (Raf.), Rock
Bass

Micropterus dolomieu Lac., Small-

1

mouthed Bass

1

Cyclochaeta domerqui

Centrarchidae

Wallengren

Apomotis cyanellus (Raf.), Green

.47%

Sunfish

1

Chilodon sp.

Centrarchidae

1.43%

Helioperca macrochira (Raf.), Blue-
nosed Sunfish

Huro salmoides (Lac.), Large-mouthed

1

Bass

1

Micropterus dolomieu Lac., Small-
mouthed Bass

1

Protozoan, mixed infection

Siluridae

1.43%

Ameiurus nebulosus (Le S.), Common
Bullhead

1

Centrarchidae

Micropterus dolomieu Lac., Small-
mouthed Bass

2

Myxobolus sp.

Centrarchidae

.47%

Chaenobryttus gulosus (Cuv.),
Warmouth

1

1 Percentage of total number of recorded deaths.

538

Zoologica: New York Zoological Society

[XXV :33

Gyrodactyloides (sp.?)
3.81%

Serranidae

Morone interrupta Gill, Yellow Bass 8

Argulus sp.
.47%

Lepisosteidae

Lepisosteus osseus (L.), Long-nosed Gar 1

Cestode Enteritis
.47%

Esocidae

Esox estor Le S., Pike 1

Gastritis

.47%

Amiidae 1

Amia calva L., Bowfin

Malnutrition

.95%

Catostomidae

Catostomus commersonii (Lac.), Common
sucker 1

Hypentelium nigricans (Le S.), Hog
Sucker 1

Edema

2.38%

Cyprinidae

Carassius auratus (L.), Goldfish 2

Scardinius erythrophthalmus (L.),

Pearl Roach 2

Centrarchidae

Micropterus dolomieu Lac., Small-
mouthed Bass 1

Ovarian degeneration
2.38%

Acipenseridae

Scaphirhynchus platorhynchus (Raf.),
Shovel-nosed Sturgeon 1

Salmonidae

Salvelinus fontinalis (Mitch.), Brook
Trout 1

Salvelinus fontinalis (Mitch.), Albino
Brook Trout 1

Centrarchidae

Micropterus dolomieu Lac., Small-
mouthed Bass 1

Percidae

Perea flavescens (Mitch.), Yellow Perch 1

Myxomata?

6.19%

Salmonidae

Salmo fario (Turton), Brown Trout 3

Salmo irideus Gibbons, Rainbow Trout 2

Centrarchidae

Helioperca macrochira (Raf.), Blue-
nosed Sunfish 2

Huro salmoides (Lac.), Large-mouthed
Bass 2

Micropterus dolomieu Lac., Small-
mouthed Bass 4

Epithelioma

.47%

Salmonidae

Salvelinus fontinalis (Mitch.), Albino
Brook Trout 1

Scoliosis

.47%

Acipenseridae

Scaphirhynchus platorhynchus (Raf.),
Shovel-nosed Sturgeon 1

1940]

Nigrelli: Mortality Statistics, 1939

539

Lordosis

Acipenseridae

.47%

Acipenser fulvescens Raf., Lake Sturgeon

1

Senility?

Amiidae

1.43%

Amia calva L., Bowfin

1

Centrarchidae

Helioperca macrochira (Raf.), Blue-nosed
Sunfish

Lepomis auritus (L.), Red-breasted

1

Sunfish

1

Chemical poisoning

Esocidae

5.24%

Esox masquinongy Mitch., Muskellunge

11

Destroyed

Cyprinidae

.47%

Chrosomus erythrogaster Raf., Red-
bellied Dace

1

Water temperature

Salmonidae

13.3%

Salmo irideus Gibbons, Rainbow Trout
Salvelinus fontinalis (Mitch.), Brook

5

Trout

23

Unknown

Salmonidae

5.24%

Salmo salar sebago Girard, Land-locked
Salmon

2

Cyprinidae

Cyprinus carpio L., Carp
Rhinichthys atratulus (Hermann),

1

Black-nosed Dace

1

Centrarchidae

Eupomotis gibbosus (L.), Pumpkin Seed

4

Pomoxis sparoides (Lac.), Calico Bass

2

Percidae

Perea flavescens (Mitch.), Yellow Perch

1

Table III

Causes of Mortality

in Tropical Fresh-water Fishes.

No.

Disease

Fish Deaths

Saprolegnia sp.

Gymnotidae

t.81%1

Electrophorus electricus (L.), Electric
Eel

4

Symbranchidae

Symbranchus marmoratus Bloch

1

Ichthyophthirius multifiliis

Characinidae

Fouquet

1.81%

Cheirodon piaba Liitken

5

Stregeid

Toxotidae

.36%

Toxotes jaculator (Pallas), Archerfish

1

Ovarian cysts

Cyprinodontidae

.36%

Rivulus harti Boulenger

1

i Percentage of total number of recorded deaths.

540

Zoologica: New York Zoological Society

[XXV :33

Failure to deliver young

2.53%

Poeciliidae

Mollienisia sphenops (C. & V.), Mollie 2

Xiphophorus hellerii Heckel, Swordtail 2

Swordtail X Platy Hybrid 3

Egg bound
3.25%

Characinidae

Copeina arnoldi Regan, Splashing
Salmlet 1

Epicyrtus microlepis Reinhardt, Glass
Characin 1

Hyphessobrycon bifasciatus Ellis, Yellow
Tetra 1

Gymnotidae

Eigenmannia virescens (Val.), Glass
Gymnotid 1

Gymnotus carapo L., Banded Gymnotid 1

Cyprinidae

Barbus conchonius Ham-Buch., Rosy
Barb 1

Barbus cummingti Gthr. 1

Cichlidae

Aequidens moronii Steindachner 1

Geophagus surinamensis (Bloch) 1

Hepatic degeneration
.36%

Anabantidae

Trichogaster trichopterus (Pallas),

Hairfin Gourami 1

Edema

2.17%

Cyprinidae

Barbus conchonius Ham.-Buch., Rosy
Barb 1

Siluridae

Acanthodoras cataphractus (L.) 1

Anabantidae

Betta splendens Regan, Siamese Fighting
Fish 1

Macropodus cupanus (C. & V.) 1

Macropodus opercularis (L.), Forked-
tail Paradise Fish 1

Cichlidae

Aequidens latifrons (Steindachner) 1

Malnutrition

1.44%

Osteoglossidae

Osteoglossum bicirrhosum Vandelli,
Arowana 1

Poeciliidae

Mollienisia velifera Regan, Sailfin Mollie 1
Cichlidae

Pterophyllum scalar e (C. & V.), Angel-
fish 2

Chemical poisoning, paint
.72%

Gymnotidae

Sternopygus macrurus (B. & S.) 2

Jumped from tank
1.08%

Polypteridae

Calamoichthys calabaricus (Smith),

Reedfish 1

<M <N CO

1940]

Nigrelli: Mortality Statistics, 1939

541

Characinidae

Exodon paradoxus Muller & Troschel 1

Siluridae

Clarias sp. 1

Senility?

7.58%

Polyp teridae

Calamoichthys calabaricus (Smith),

Reedfish 1

Notopteridae

Xenomystus nigri Gthr., African Knife-
fish 1

Characinidae

Hyphessobrycon flammeus Myers, Flame
Tetra 1

Hyphessobrycon heterorhabdus (Ulrey) 1
Hyphessobrycon serpae Durbin 1

Moenkausia pittieri Eigenmann 1

Mylossoma duriventris (Cuv.) 1

Nannaethiops unitaeniatus Gthr. 1

Pristella riddlei (Meek) 1

Gymnotidae

Gymnotus coatesi LaMonte, Tiger
Knifefish 1

Cyprinidae

Barbus lineatus Duncker 2

Esomus malayensis E. Ahl, Malayen
Flying Barb 1

Rasbora trilineata Steindachner, Three-
lined Rasbora 1

Aspridinidae

Bunocephalus bicolor Steindachner 1

Siluridae

Acanthodoras sp. 1

Poeciliidae

Belonesox belizanus Kner, Pike-killie 1

Anabantidae

Ctenops vittatus (C. & V.), Croaking
Gourami 1

Ophiocephalidae

Channa asiatica (L.), Snake-head 1

Nandidae

Badis badis (Ham.-Buch.) 1

Cichlidae

Etroplus suratensis (Bloch) 1

Missing

2.53%

Cyprinidae

Rasbora trilineata Steindachner, Three-
lined Rasbora 1

Cyprinodontidae

Aphyosemion calliurum (Boulenger),
Lyretail 1

Poeciliidae

Belonesox belizanus Kner, Pike-killie 2

542

Zoologica: Neiv York Zoological Society

[XXV :33

Missing

(continued)

Hemirhamphidae

Dermogenys pusillus van Hasselt, Fresh-
water Halfbeak 1

Anabantidae

Helostoma temmincki C. & V., Kissing
Gourami 1

Cichlidae

Crenicichla lepidota Heckel 1

Destroyed

1.81%

Siluridae

Acanthodoras sp. 1

Loricariidae

Loricaria sp. 4

Water temperature
3.25%

Characinidae

Creatochanes melanurus (Bloch) 1

Cyprinodontidae

Fundulus chrysotus Holbrook 1

Fundulus ocellaris Jordan & Gilbert 1

Oryzias latipes (Schl.) , Medaka 1

Cichlidae

Cichlosoma severum (Heckel), Deep-
bodied cichlid 3

Herichthys cyanoguttatus Baird and
Girard 2

“Spoiled” water
4.70%

Dasyatidae

Potamotrygon hystrix (Muller &

Troschel), Fresh-water Stingray 2

Characinidae

N annaethiops unitaeniatus Gthr. 2

Cyprinidae

Barbus semifasciolatus Gthr., Half-
banded Barb 2

Cyprinodontidae

Rivulus harti Boulenger 4

Poeciliidae

Micropoecilia branneri Eigenmann 2

Cichlidae

Symphysodon discus (Heckel), Disc
Cichlid 1

Fighting

8.30%

Characinidae

Copeina amoldi Regan, Splashing
salmlet 1

Creatochanes melanurus (Bloch) 1

Leporinus affinis Gthr. 1

Leporinus fasciatus (Bloch), Black-
ringed Groundfish 1

Serrasalmus scapularis Gthr. 1

Gymnotidae

Gymnotus carapo L., Banded Gymnotid 1

Cyprinodontidae

Chriopeops goodei (Jordan) 1

1940]

Nigrelli: Mortality Statistics, 1939

543

Epiplatys chaperi (Sauvage), Chaper’s
Panchax 3

Panchax lineatus C. & V., Striped
Panchax 3

Anabantidae

Betta splendens Regan, Siamese Fighting
Fish 3

Macropodus opercularis (L.), Fork-tailed
Paradise Fish 3

Cichlidae

Cichlasoma cutteri Fowler 1

Cichlasoma festivim (Heckel), Painted
Cichlid 1

Hemichromis bimaculatus Gill, Ruby
Jewel Fish 1

Hemichromis fasciatus Peters 1

Acclimatization

23.1%

Osteoglossidae

Osteoglossum bicirrhosum Vandelli 2

Mormyridae

Morviyrus Jcannume Forskal 1

Characinidae

Anoptichthys jordani Hubbs & Innes,

Blind Characin 20

Carnegiella strigata (Gthr.), Mottled
Hatchetfish 7

Chalceus macrolepidotus Cuv. 1

Chilodus punctatus Muller & Troschel,
Head-standing Fish 1

Copeina arnoldi Regan, Splashing salmlet 3
Corynopoma riisei Gill 4

Gasteropelecus sternicla (L.), Hatchet-
fish 3

Hemigrammus rhodostomus E. Ahl, Red-
mouthed Tetra 1

Pterodiscus laevis, Eigenmann, Flying
Characin 2

Cyprinidae

Rasbora pauciperf orata Weber & de
Beaufort 3

Callichthyidae

Corydoras sp. 1

Cyprinodontidae

Aphyosemion calliurum (Boulenger),
Lyretail 2

Atherinidae

Telmatherina ladigesi E. Ahl 8

Cichlidae

Cichlasoma cutteri Fowler 1

Gobiidae

Boleophthalmus viridus Ham.-Buch. 4

Unknown

32.8%

Mormyridae

Mormyrus Jcannume Forskal 1

544

Unknown

(continued)

Zoologica: New York Zoological Society [XXV:33

Pantodontidae

Pantodon buchholzi Peters, Fresh-water
Butterflyfish 1

Characinidae

Anostomus plicatus Eigenmann 1

Carnegiella strigata (Gthr.), Mottled
Hatchetfish 1

Chalcinus elongatus Gthr. 1

Chalcinus rotundatus (Schomburk) I

Epicyrtus microlepis Reinhardt, Glass
Characin 2

Hemoidus gracilis Gthr. 1

Hemiodus quadrimaculatus Pellegrin 1

Hyphessobrycon innesi Hubbs, Neon
Tetra 2

Myloplus schomburgkii (Jardine) 2

Poecilobrycon eques (Steindachner) 1

Poecilobrycon trifasciatus (Steindachner),
Three-lined Pencilfish 2

Poecilobrycon unifasciatus (Steindach-
ner), One-lined Pencilfish 1

Prochilodus insignis Schomburgk, Flag-
tail 1

Cyprinidae

Barbus conchonius Ham.-Buch., Rosy
Barb 2

Barbus oligolepsis (Bleeker) 1

Barbus phutunio (Ham.-Buch.) 1

Barbus sp. 5

Danio malabaricus (Jerdon), Giant Danio 1
Notropis metallicus Jordan & Meek 3

Rasbora heteromorpha Duncker, Red
Rasbora 1

Rasbora maculata Duncker 2

Cobitidae

Botia sp. 1

Siluridae

Centromochlus aidopy gius Kner 1

Micro glanis parahybae (Steindachner) 1

Callichthyidae

Corydoras arcuatus Ahl 1

Corydoras leopardus Myers

Co?'ydoras melanisteus Regan 1

Loricariidae

Loricaria parva Boulenger 1

Plecostomus sp. 1

Cyprinodontidae

Chriopeops goodei (Jordan) 1

Panchax lineatus C. & V., Striped
Panchax 1

Poeciliidae

Belonesox belizanus Kner, Pike-killie 1

Limia nigrofasciata Regan 1

Limia vittata (Guichenot) 1

Mollienisia latipinna Le S., Mudpusser 2

Phalloceros caudomaculatus (Hensel) 4

Platypoecilus maculatus Gthr., Platy 10

Quintana atrizona Hubbs 1

1940]

Nigrelli: Mortality Statistics, 1939

545

Hemirhamphidae

Dermogenys pusillus van Hasselt, Fresh-

water Halfbeak 1

Anabantidae

Colisa lalia (Ham.-Buch.) , Dwarf
Gourami 2

Ctenops vittatus (C. & V.), Croaking
Gourami 4

Macropodus opercularis (L.), Fork-
tailed Paradise Fish 3

Nandidae

Badis badis '(Ham.-Buch.) 1

Monocirrhus polyacanthus Heckel,

Leaffish 4

Polyceyitrus schomburgkii Muller and
Troschel, Casarab 1

Centrarchidae

Elassoma evergladei Jordan, Pigmy
sunfish 2

Cichlidae

Apistogramma pertense, (Haseman),

Dwarf Cichlid 1

Apistogramma sp. 1

Etroplus maculatus (Bloch), Orange
Chromide 2

Geophagus jurupari Heckel, Demon Fish 1
Geophagus surinamensis (Bloch) 1

Tilapia zillii Gervais 1

Toxotidae

Toxotes jaculator (Pallas), Archerfish 1

Table IV.

Causes of Deaths in Specimens Other Than Fishes.

No.

Disease AMPHIBIANS Deaths

Fungus

Cryptobranchus alleganiensis (Daudin),
Hellbender

2

Necturus maculosus (Raf.), Mudpuppy

5

Proteus anguinus, Olm

2

Red-leg disease

Proteus hydrophilus ( Sanarelli )

Pipa surinamensis, Surinam Toad

1

Myxoma

Necturus maculosus (Raf.), Mudpuppy

2

REPTILES

Fungus

e.1%1

Alligator mississipiensis (Daudin),
American Alligator

5

Chelys fimbriata, Matamata

1

Liver abcess from nematodes

Natrix s. sipedon (L.), Common Water

1.0%

Snake

1

Liver degeneration

Guatemalan Turtle

1

1.0%

1 Percentage of total number of recorded deaths.

546

Zoological New York Zoological Society

[XXV :33

Fatty degeneration
7.1%

Alligator mississipiensis (Daudin),
American Alligator 7

Malnutrition

6.1%

Caiman jacuare (Daudin) 1

Caiman sclerops (Schneider) 4

Gecko sp. 1

Enteritis

56.5%

Caretta caretta (L.), Loggerhead Turtle 1
Chrysemys picta (Schneider), Painted
Turtle 23

Clemmys guttata (Schneider), Spotted
Turtle 6

Clemmys insculpta (Le Conte), Wood
Turtle 6

Graptemys p. pseudo geographica
(Gray), Mississippi Map Turtle 1

Podochemis expansa, Amazon River
Turtle 1

Pseudemys scripta troostii (Holbrook),
Cumberland Turtle 8

Pseudemys scripta ssp., Yellow-bellied
Terrapin 1

Terrapene Carolina (L.), Box turtle 7

Trionyx sp. 1

Chinese River Turtle 1

Floating trouble?
1.0%

Caretta caretta (L.), Loggerhead

Turtle 1

Fighting

2.0%

Caiman sclerops (Schneider) 1

Caretta caretta (L.), Loggerhead

Turtle 1

Drowned

7.1%

Alligator mississipiensis (Daudin),
American Alligator 6

Trionyx ferox (Schneider), Southern
Soft-shelled Turtle 1

Unknown

12.1%

Caiman niger Spix 1

Caretta caretta (L.), Logger-head Turtle 1

Caretta kempii (Garman), Kemp’s Turtle 1

Chelys fimbriata, Matamata 1

himys blandingii (Holbrook), Blanding’s
Turtle 1

Graptemys geographica (Le S.), Map
Turtle' 1

Malaclemys centrata (Latreille),
Diamond-back Terrapin 1

Macrochelys temminckii (Troost),

Alligator Snapping Turtle 1

Podochemis expansa, Amazon River
Turtle 1

Terrapene Carolina (L.), Box Turtle 1

Trionyx sp. 1

East Indian Mud Turtle 1

Tuberculosis?

BIRDS

Spheniscus demersus (L.), Blackfoot
Penguin 2

Fractured leg,
Gangrene

Spheniscus humboldti Meyen, Hum-
boldt’s Penguin 1

1940] Nigrelli: Mortality Statistics, 1939 547

Malnutrition

MAMMAL

Trichechus inunguis Natter er, Amazon
Manatee 1

INVERTEBRATES

Larval trematodes on gills

Scyllarides latus, Flat Lobster

1

Fighting

Octopus sp.

3

Water chemistry

Fasciolaria gigantea Kiener, Giant Band
Shell 1

Fasciolaria tulipa L., Tulip Band Shell 1

Limulus polyphemus (L.), Horseshoe Crab 10
Calappa flammea (Herbst), Queen Crab 2
Libinia dubia Milne-Edwards, Spider
Crab 7

Menippe mercenaria (Say), Stone Crab 2
Panulirus argus (Latreille), Spiny
Lobster 1

Petrochirus bahamensis, Sea-soldier 1

Squilla empusa, Mantis Shrimp 1

Oreaster reticulatus, Giant Starfish 10

Sargartia leucolena, White Anemone 35

Unknown

Cambarus sp., Fresh-water Crayfish
Homarus americanus Milne-Edwards,
Northern Lobster
Papaloapan Fresh-water Crab

5

2

1

Table V.

Summary of Mortality Percentage by Causes.

k

k.

e

e

00

e

k.

Causes

Marine

s £

-s X

eo

£ 8

Fresh- w
tropical

jO

2

a.

g

Reptiles

Birds

e

g

§

%

k.

41

Infectious disease

64.8

60.5

3.9

83.3

1.0

67

0

1.2

Non-infectious disease

19.5

21.0

32.2

16.7

86.9

33

100

3.6

Water conditions 1

11.3

13.3

31.1

0

0

0

0

85.5

Unknown

4.4

5.2

32.8

0

12.1

0

0

9.7

1 Includes density, temperature, pH. and chemical composition.

548

Zoologica: New York Zoological Society [XXV:33

Table VI.

Summary of Mortality Percentage by Months.

©

.P

©

©

© -S

s g

« a.

el

©

f©

50 P,

g ?

’©

eo

P

.©

r°

a

£

CO

P,

CO

V.

CO

'©

§

s

e

CP

a

►o

©

©

§

kb

kb

s

January

2.7

0

7.6

3.4

8.3

3.0

0

0

2.4

February

4.6

2.9

5.4

4.5

0

5.1

33.3

0

1.2

March

4.3

0

12.3

5.2

0

11.1

0

0

0

April

5.1

1.0

8.7

5.2

8.3

6.1

0

0

3.6

May

7.2

6.7

15.5

8.8

8.3

21.2

33.3

0

1.2

June

8.4

3.3

5.8

7.1

0

5.1

0

0

1.2

July

35.5

16.2

16.6

28.9

0

12.1

0

0

7.2

August

12.4

3.3

6.6

9.9

0

6.1

0

100

30.1

September

6.7

2.9

7.6

6.2

16.7

10.1

0

0

9.6

October

5.6

4.3

4.3

5.2

16.7

3.0

0

0

13.2

November

4.1

25.7

5.4

7.6

0

5.1

0

0

18.1

December

3.5

33.8

4.3

8.1

41.7

12.1

33.3

0

12.1

Table VII.

Summary of Mortality Percentage of Fishes by Sex.

§

kb

Male

27.9

14.8

23.5

25.1

Female

37.2

39.0

35.0

37.0

Immature

5.9

40.5

9.4

11.8

Undetermined

29.0

5.7

32.1

26.1

Table VIII.

Longevity of Specimens

CP

©

©

©

5

% *

§

k©

O ©

'p

05

53 g

©

©>

v. ©

“I

©

Unknown

—

—

One month or less

, . 55.5

27.6

Two to twelve months . . .

. . 39.3

60.0

More than one year

. . 5.2

12.4

which Died During 1939.

©

eo

CP

-P

V)

eo

©

©

©

P

.©

r°

eo

©

*©

S

P

r©

©

-P

« CO

*©

-5

§

K

©

1

©

P

. I

.P

©

©5

©

05

-©

©

CO

©

<M

05

CO

p

©

00

©

® p.

©

P

©

©

©

©

©

©

£b

$

6.9

1.3

—

—

—

—

—

22.0

44.8

0

9.1

33.3

100.

12.0

50.2

44.5

100.

87.9

0

0

88.0

20.9

9.4

0

3.0

66.7

0

0

1940]

Nigrelli: Mortality Statistics, 1939

549

Table IX.

Fishes Over Two Years Old which Died During 1939.1

Marine
Ten Years

Gymnothorax funebris Ranzani, Green Moray
Seven Years

Tilapia mossambica Peters
Six Years

Ginglymostoma cirratum (Bonnaterre) ,
Nurse Shark
Five Years

Trachinotus glaucus (Bloch), Palmoneta
Sciaenops ocellatus (Linnaeus), Channel
Bass

Mycteroperca bonaci (Poey), Black Grouper
Four Years

Acipenser oxyrhynchus Mitchill, Common
Sturgeon

Galeichthys felis (Linnaeus), Sea Catfish
Gymnothorax moringa (Cuvier), Spotted
Moray

Seriola zonata (Mitchill), Rudder-fish
Pogonias cromis (Linnaeus), Sea Drum
Halichaeres radiatus (Linnaeus), Pudding-
wife

Thalassoma bifasciatus (Bloch), Blue-head
Three Years

Tarpon atlanticus (Cuvier & Valenciennes),
Tarpon

Roccus lineatus (Bloch), Striped Bass
Scatophagus argus (Bleeker), Scat
Two Years

Epinephelus striatus (Bloch), Nassau
Grouper

Rypticus saponaceus (Bloch & Schneider),
Soapfish

Dascyllus aruanus (Linnaeus), Jesuite
Chaetodipterus faber (Broussonet) , Spade-
fish

Balistes vetula Linnaeus, Queen Triggerfish
Diodon hystrix Linnaeus, Porcupine Fish

Temperate Fresh-water
Six Years

Lepisosteus osseus (Linnaeus), Long-nosed
Gar

Amia calva Linnaeus, Bowfin
Five Years

Acipenser fulvescens Rafinesque, Lake
Sturgeon

Huro salmoides (Lacepede), Large-mouthed
Bass

Morone interrupta Gill, Yellow Bass
Four Years

Scavhirhynchus platorhynchus (Rafinesque),
Shovel-nosed Sturgeon
Esox masguinongy Mitchill, Muskellunge

Ambloplites rupestris (Rafinesque), Rock
Bass

Lepomis auritus (Linnaeus), Red-breasted
Sunfish

Helioperca macrochira (Rafinesque), Blue-
nosed Sunfish
Three Years

Scardinius erythrophthalmus (Linnaeus),
Pearl Roach

Ameiurus nebulosus (Le Sueur), Common
Bullhead

Perea flavescens (Mitchill), Yellow Perch
Two Years

Salvelinus fontinalis (Mitchill), Albino
Brook Trout

Pomoxis sparoides (Lacepede), Calico Bass

Tropical Fresh-water
Six and One-half Years

Channa asiatica (Linnaeus), Snake-head
Five Years

Hyphessobrycon heterorhabdus (Ulrey)
Hyphessobrycon serpae Durbin
Pristella riddlei ( Meek )

Serrasalmus scapularis Gunther
Four Years

Moenkhausia pittieri Eigenmann
Gvmnotus coatesi LaMonte, Tiger Knifefish
Ctenops vittatus (Cuvier & Valenciennes),
Croaking Gourami
Aequi'tens moronii Steindachner
Three Years

Calamoichthys calabaricus (Smith), Reedfish
Esomus malayensis E. Ahl, Malayan Flying
Barb

Rasbora tHlineata Steindachner, Three-lined
Rasbora

Etroplus suratensis ( Bloch )

Two and One-half Years

Xenomystus nigri Gunther, African Knife-
fish

Barbus lineatus Duncker
Two Years

Mylossoma duriventris (Cuvier)

Cheirodon piaba Liitken

Rasbora heteromorpha Duncker, Red Rasbora
Rasbora maculata Duncker
Bunocephalus bicolor Steindachner
Acanthodoras sp.

Centromochlus aulopygius Kner
Microglanis parahybae (Steindachner)

Badis badis (Hamilton-Buchanan)
Cichlasoma festivum (Heckel), Painted
Cichlid

H erichthys cyanoguttatus Baird & Girard
Symbranchus marmoratus Bloch

l Includes only oldest specimen of each species.

550

Zoologica: New York Zoological Society

[XXV :33

Parasites

Parasite

I. Protozoa

A. Marine species

1. Protrichomonas sp.

2. Oodinium ocellatum Brown

3. Trichodina sp.

B. Fresh-water species

4. Cyclochaeta domerqui
Wallengren

5. Chilodon sp.

6. Ichthyophthirius mul-
tifiliis Fouquet

7. Henneguya ameiurensis-
Nigrelli & Smith

8. Myxobolus sp.

Table X.

and Hosts Observed in 1939.1 2
Host

Chilomycterus schoepfii
( Walbaum)

Marine fishes
Marine fishes

Fresh-water fishes

Fresh-water fishes
Fresh-water fishes

Ameiurus nebulosus (Le Sueur)

Chaenobryttus gulosus (Cuvier)

Site of Infection

Intestine

Gills

Gills

Skin & gills

Skin & gills
Skin & gills

Skin

Gills

II. Trematoda, monogenetic

A. Marine species

9. Epibdella melleni
MacCallum

10. Microcotyle sp.

11. Diplectanus sp.

12. Gyrodactyloides (Spp?)

B. Fresh-water species

13. Gyrodactyloides (Spp?)

Marine fishes

Skin & eyes

Marine fishes Gills

Marine fishes Gills

Fundulus heteroclitus (Linnaeus) Skin & gills

Fresh-water fishes

Skin & gills

III. Trematoda, digenetic adults
A. Marine species

14 . Lintonium vibex (Linton) Spheroides maculatus

(Bloch & Schneider)

15. Bianium plicitum (Linton ) Spheroides 'tnacv.latus

(Bloch & Schneider)

16. Probliotrema sp. Amphotistius sabinus (Le Sueur)

17. Tubulovesicula madurensis3 Scorpaena madurensis

Nigrelli

18. Hemiuridae (Spp?)

19. Podocotyle atzi Nigrelli3

20. Allocreadiidae (Spp?)

21. N ematobothrium sp.

22. Sterrhurus branchialis
Stunkard & Nigrelli

(Cuvier & Valenciennes)
Marine fishes
Scorpaena madurensis
(Cuvier & Valenciennes)
Marine fishes
Sarda sarda (Bloch)
Trichiurus lepturus Linnaeus

Pharynx

Intestine

Body cavity
Stomach

Stomach

Intestine

Intestine

Gills

Gills

B. Fresh -water species

23 . Allacanthochasmus varius
Vancleave

24. Allacanthochasmus artus
Vancleave & Mueller

25. Neochasmus umbellus
Vancleave & Mueller

26. Bucephalus pusillis
(Stafford)

27. Allocreadium sp.

28. Crepidostomum sp.

29. Amphistome (Sp?)

M or one interrupta Gill

M or one interrupta Gill

M or one interrupta Gill

Stizostedion vitreum (Mitchill)

Eupomotis gibbosus (Linnaeus)
Helioperca macrochira
(Rafinesque)

Trichechus manatus latirostrus
(Harlan)

Intestine

Intestine

Intestine

Intestine

Intestine

Intestine

Caecum

1 Including species and specimens not on exhibition at the N. Y. Aquarium.

2 See Nigrelli & Smith, 1940

8 See Nigrelli, 1940b.

1940]

Nigrelli: Mortality Statistics, 1939

551

IV. Trematoda, digenetic larvae

A. Marine fishes

30. Cryptocotyle lingua Tautoga onitis (Linnaeus)

(Creplin)

31. Metacercaria (Sp?) Scyllarides latus

B. Fresh-water species

32. Neascus vancleavi Fresh-water fishes

(Agersbord)

33. Clinostomum complanatum Fresh-water fishes
(Rudolphi)

V. Cestoda, adults

A. Marine species

34. Calliobothrium verticil- Mustelus canis (Mitchill)
latum (Rudolphi)

35. Crossobothrium laciniatum Carcharias littoralis (Mitchill)
Linton

B. Fresh-water species

36. Proteocephalus sp.

37. Proteocephalus pinguis

LaRue

38. Corallobothrium sp.

39. Bothriocephalus cuspidar
tus Cooper

VI. Cestoda, larvae
A. Marine species

40. Otobothrium crenacolle
Linton

Huro salmoides (Lacepede)

Esox estor Le Sueur

Ictalurus punctatus (Rafinesque)
Perea flavescens (Mitchill)

Poronotus tricanthus (Peck)

41. Tetrarhynchid forms Marine fishes

42. Tetrarhynchid forms Groupers

B. Fresh-water species

43 . Ligula intestinalis (Goeze) Hyborhynchus notatus

VII. Nematoda

A. Fresh-water species

44. Camallanus sp.

45. Spinitectis sp.

VIII. Hirudinea

A. Marine species

46. Branchellion ravenelli*
(Girard)

47. Pontobdella muricata
(Linnaeus)

B. Fresh-water species

48. Placobdella parasitica
(Say)

(Rafinesque)

Fresh-water fishes
Micropterus dolomieu Lacepede

Dasyatis hastatus (De Kay)
Sphyrna zygaena (Linnaeus)

Fresh-water turtles

IX. Copepoda

A. Marine species

49 .Spliyrion lumpi (Krpyer) Sebastes marinus (Linnaeus)
50. Chondrancanthopsis no- Sebastes marinus (Linnaeus)
dosus (Muller)

51 .Penella instructa Wilson Xiphias gladius Linnaeus

52 . Ergasilidae (Spp?) Marine fishes

53 . Pandarus bicolor Leach Mustelus canis (Mitchill)

54 . Lernaeenicus radiatus Menidia menidia (Linnaeus)
(Le Sueur)

Encysted on
skin & fins
Encysted on
gills

Organs

Skin

Intestine

Intestine

Intestine

Intestine

Intestine

Intestine

Encysted around
vertebrae

Intestine
Encysted in
body cavity

Intestine

Intestine

Intestine

Skin

Skin

Skin

Body

Gills

Body

Gills

Skin

Body

■i See Meyer, 1939.

552

Zoological New York Zoological Society

B. Fresh-water species

55. Lernae sp.

56. Argulus spp.

57. Ergasilus sp.

Carassius auratus (Linnaeus) Body

Fresh-water fishes Skin

Fresh-water fishes Gills

References.

Meyer, M. C.

1939. Demonstration of a species of marine Piscicolidae
Jour. Parasitol. 25: 22.

from

Florida.

Nigrelli, R. F.

1936. The morphology, cytology and life-history of Oodinium ocellatum
Brown, a dinotiagellate parasite on marine fishes. Zoologica, 21 (12) :
129-164.

1940a. Health of the specimens. (In Report of the Director of the Aquarium)
Forty-fourth Annual Report of the New York Zoological Society for
the Year 1939 : 63-66.

1940b. Two new species of trematodes from the deep sea scorpion fish,
Scorpaena madurensis Cuv. & Val. Zoologica, 25 (17) : 263-268.

Nigrelli, R. F. & C. M. Breder, Jr.

1934. The susceptibility and immunity of certain marine fishes to Epibdella
melleni, a monogenetic trematode. Jour. Parasitol., 20 (5) : 259-269.

Nigrelli, R. F. & G. M. Smith

1939. Studies on lymphocystis in the orange filefish, Ceratacanthus
schoepfii (Walbaum), from Sandy Hook Bay, N. J. Zoologica, 24 (9) :
255-264.

1940. A papillary cystic disease affecting the barbels of Ameiurus nebul-
osus (Le Sueur), caused by the myxosporidian Henneguya ameiu-
rensis sp. nov. Zoologica, 25 (8) : 89-96.

Padnos, Morton

1939a. Endosphaera englemanni, endoparasitic in Trichodina infecting the
puffer, Spheroides maculatus (Linnaeus). Anat. Rec., 75 (4, suppl.) :
156.

1939b. The morphology and life history of Trichodina infecting the puffer,
Sphaeroides maculatus (Linnaeus). Anat. Rec., 75 (4, suppl.): 157.

Paul, Allard A.

1939. Life history study of a monogenetic trematode. Anat. Rec., 75
(4, suppl.) : 156.

EXPLANATION OF THE PLATES.

All photographs by S. C. Dunton of the New York Aquarium.

Figs. 1 and 2. Queen triggerfish, showing peculiar ulceration of the skin
around the mouth and fins. The causative agent is unknown.

Fig. 3. Edematous condition in pearl roach. A general anasarca was present
and the kidneys were completely degenerated.

Fig. 4. Viscera of small-mouthed bass, showing ovarian degeneration. This
condition results from the failure to spawn and to resorb the eggs.

Fig. 5. Portion of intestine of pike with heavy infestation of Proteocephalus
pinguis. This infection was acquired in the wild state, since the fish
lived but two months in captivity. The entire intestine was practically
filled with these cestodes. Note the severe hemorrhage produced.

Fig. 6. Intestine of blue-nosed sunfish with hemorrhagic islands caused by
intestinal flukes. Not included in Table II.

Fig. 7. Transverse section of alligator tail showing fatty degeneration. Large
fat masses have pushed aside the muscles. Within these masses may
be seen numbers of dark areas, consisting of hard, granular, yellowish
material.

NIGRELLI .

PLATE I.

FIG. I.

FIG. 2.

FIG. 3.

MORTALITY STATISTICS FOR ANIMALS IN THE NEW YORK AQUARIUM, 1939.

NIGRELLI.

PLATE II.

FIG. 4.

FIG.. 5.

MORTALITY STATISTICS FOR ANIMALS IN THE NEW YORK AQUARIUM, 1939.

NIGRELLI

PLATE III.

FIG. 6.

FIG. 7.

MORTALITY STATISTICS FOR ANIMALS IN THE NEW YORK AQUARIUM, 1939.

Cox, et al.: Electrophorus electricus

553

34.

A Comparison of Some Electrical and Anatomical Characteristics
of the Electric Eel, Electrophorus el'ectricus (Linnaeus).

R. T. Cox

Department of Physics, New York University, and. Biological Laboratory,
Long Island Biological Association;

W. A. Rosenblith1

Biological Laboratory, Long Island Biological Association;

Janice A. Cutler

Washington Square College, New York University ;

R. S. Mathews & C. W. Coates

New York Aquarium.

(Text-figures 1-7).

In a previous paper2 it has been shown that the major and minor
electrical discharges of the electric eel, Electrophorus electricus (Linnaeus),
are produced respectively by the large organs and the organs of Sachs. It
has also been shown3 that the current produced in a conductor joining two
points along an electric organ could be approximately accounted for by
ascribing to the segment of the organ between the two points a constant
electromotive force and a variable internal resistance and considering it
shunted by a constant leakage resistance which may be attributed to the
adjacent tissue. Except during the discharge the internal resistance of the
segment is assumed to be so much higher than the leakage resistance that
the potential difference is negligible between different points along the
organ. The discharge is attributed to a transient drop in the internal re-
sistance, allowing the electromotive force to produce a current.

Let R denote the internal resistance of a segment of the organ at the
peak of the discharge, when the resistance is at its minimum. Let E denote
the electromotive force of the segment and let r denote the leakage resistance,
of the adjacent tissue. Let R' denote the resistance of a conductor con-
nected between electrodes in contact with the fish at the ends of the segment
and let V denote the potential difference between these electrodes at the peak
of the discharge. Then these quantities may be shown to be related by an
equation, which may for the present purpose conveniently be written in the
form

E _ R , r + R
V ~ R7 + ~T~

1 Now at Department of Physics, University of California at Los Angeles.

2 Coates, C. W., R. T. Cox, & L. P. Granath. The Electric Discharge of the Electric Eel,
Electrophorus electricus (Linnaeus). Zoologica , Vol. XXII (Part 1), No. 1, April 5, 1937.

3 Cox, R. T., & C. W. Coates. Electrical Characteristics of the Electric Tissue of the Electric
Eel, Electrophorus electricus (Linnaeus). Zoologica, Vol. XXIII (Part 2), No. 8, July 14, 1938.

(1)

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[XXV :34

If R' is varied in this equation, V varies with it in such a way that for
every change in 1/R' there is a proportional change in 1/V. Consequently
the graph of 1/V plotted against 1/R' will be a straight line. In Text-fig. 1
are plotted measurements of peak voltages obtained with various external
resistances joined to electrodes 10 cm. apart on the large organ of an electric
eel 103 cm. long. The sets of points indicated by different signs show sets of
observations made at several different times. It will be seen that the points
fall near to a straight line. (It should perhaps be pointed out that this does
not prove the hypothesis of a variable internal resistance and constant elec-
tromotive force. This must be based upon other evidence discussed in the
previous paper. The equation just given would still be valid if the internal
resistance were constant and the electromotive force variable) .

From the equation (1), when 1/R' is zero, then

V=

E r
r + R

(2)

This corresponds to the case in which there is no external conductor
between the electrodes. If the leakage resistance r is much greater than the
internal resistance R, the observed peak voltage V will be nearly equal to the
electromotive force E.

If, in the equation (1),1/V is set equal to zero, then

Since negative values of the resistances are impossible, this case cor-
responds to no actual observation. But the line determined by the actual
observations may nevertheless be extended, as in Text-fig. 1, to meet the
axis of 1/R' and the intercept may be noted to determine 1/r + 1/R. If the
leakage resistance r is otherwise known, this last result makes it possible to
determine R, the internal resistance of the segment of the electric organ.
With both these resistances known, the intercept on the axis of 1/R' may be
used to determine the electromotive force E.

With the electric eel already mentioned the leakage resistance r was
measured between electrodes 10 cm. apart along the portion of the body
containing the electric organs. The measurement was made by observing
the current passing through the body of the eel between the electrodes when
a measured potential difference was applied to them. The peak voltages
produced between the same electrodes in the same positions on the fish with
various resistances connecting them were measured in both the major and
minor discharge by means of a cathode-ray oscillograph. The electromotive
force and the internal resistance of each segment 10 cm. long down the
length of the organs were determined as already explained and as illustrated
by Text-fig. 1.

The results of these observations and computations are shown in Table I
for the major discharge and Table II for the minor discharge. The distances
recorded in the first column of each table are measured from the anterior
end of the anal fin, which is a convenient point of reference almost at the
anterior end of the large organ.

TABLE I.

Major Discharge.

Positions of

Electrodes, cm.

r, ohms

R, ohms

E, volts

0, 10

750

175

160

10, 20

765

210

150

20, 30

725

210

130

30, 40

675

180

90

40, 50

600

215

60

50, 60

550

325

30

60, 70

625

605

22

&

60, 80

1215

850

32

1940]

Cox, et al.: Electrophorus electricus

555

TABLE II.

Minor Discharge.

Positions of
Electrodes, cm.

r, ohms

R, ohms

E, volts

30, 40

675

475

4.8

40, 50

600

365

7.7

50, 60

550

365

12

60, 70

625

475

20

70, 80

925

780

22

Text-fig. 1.

Major discharge. Electrodes at 10 cm. and 20 cm. from anterior end of anal
fin. Reciprocal peak voltage vs. reciprocal external resistance.

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[XXV :34

It is hard to estimate the accuracy of these results. As regards the
leakage resistance r, it is not so much the accuracy of the measurement itself
which is in question as the accuracy of the assumption that all the current
caused to flow in the adjacent tissue by the discharge of a segment of the
organ can be treated as following a single path. As was pointed out in the
paper already referred to3 and as must indeed be rather obvious, this can be
only a crude approximation. The error in this assumption will impair the
accuracy of the determination of the internal resistance R, which is subject
also to the errors of the extrapolation in the graphical method illustrated in
Text-fig. 1. In what follows we are principally interested in the electromotive
force E. Its values are computed from the values of the resistances and the
measured values of the peak voltage by the equation

r+R

E— V

(4)

in which V is the peak voltage observed with no external conductor between
the electrodes. It is seen in Table I that for about 50 cm. along the fish from

Text-fig. 2.

Major discharge. Electromotive force per cm. vs. distance along organs from
anterior end of anal fin.

1940]

Cox, et al.: Electrophorus electricus

557

the anterior end of the large electric organ the measured values of r are
much larger than the computed values of R in the major discharge. It seems
probable that here the electromotive force actually is, as it is represented by
the equation, not much greater than the peak voltage observed with no
external conductor. In this case the ambiguities involved in the values of
the resistances will not be very important in the determination of the elec-
tromotive force. We think it likely that the first five values of the electromo-
tive force in Table I are not in error by more than about 10 per cent. The
other values of Table I and those of Table II we consider rather more
doubtful.

Text-fig. 3.

Minor discharge. Electromotive force per cm. vs. distance along organs from
anterior end of anal fin.

In Text-fig. 2 the electromotive force per cm. of the major discharge,
as reckoned from the data of Table I, is plotted as ordinate against the dis-
tance along the fish as abscissa. For plotting the first point on the left the
average value of the electromotive force per cm. between electrodes at 0
and 10 cm. is taken as the value at 5 cm., and similarly for the other plotted
points. Text-fig. 3 is a similar graph for the minor discharge plotted from
the data of Table II.

After the measurements of the voltage and resistance were made, the
fish was killed and its body transsected at a number of places. At these
places the areas of the cross sections of the electric organs were measured.
In Text-fig. 4 the black dots show the cross-sectional area of the large organ
at various points along the fish, measured as in the other data from the
anterior end of the anal fin. The white dots in the same figure show the sum

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[XXV :34

of the cross-sectional areas of the large organ and the organ of Hunter. For
comparison the curve from Text-fig. 2, showing the electromotive force per
cm. of the major discharge at points along the organ, has also been drawn in
the figure, to the scale of volts per cm. indicated by the bracket. The fall of
the plotted points along the curve is very striking. It is plain that there is a
rather close proportionality between the electromotive force per cm. of the
major discharge and the cross-sectional area either of the large organ albne
or of the large organ and Hunter’s organ together. Because of the small
cross-section of Hunter’s organ the ratio of area to electromotive force per
cm. is not much different whether Hunter’s organ be included or left out. In
either case, for each 0.5 sq. cm. of cross-sectional area at any point there is
found at the same point an electromotive force of approximately 1 volt per
cm. along the axis of the organs. By varying the scale, either set of points
can be brought closer to the curve than the other set. The best fit of the
points which show the cross-section of both organs seems to be a little closer

Text-fig. 4.

Curve: major discharge, electromotive force per cm. vs. distance along or-
gans. Black dots: Large organ, cross-sectional area vs. distance along organs.
White dots: Large organ and Hunter’s organ, cross-sectional area vs. distance
along organs.

1940]

Cox, et al.: Electrophorus electricus

559

than the best fit of the points showing the cross-section of the large organ
alone, but there is not much to choose between the two.

The significance of this proportionality between the electromotive force
per cm. and the cross-sectional area is not yet clear. If, in the discharge, the
current were the same through every cross-section of the organ, then the
observed relation would mean that equal quantities of power were generated
in equal volumes of the electric tissue, since power is the product of electro-
motive force and current. But the current can scarcely be continuous along
the organ when the fish discharges under water, since the whole skin is
conducting.

Text-fig. 5.

Curve: minor discharge, electromotive force per cm. vs. distance along or-
gans. Dots: Sachs’ organ, cross-sectional area vs. distance along organs.

In Text-fig. 5, the plotted points show the cross-sectional area of the
organs of Sachs and the curve shows the electromotive force per cm. of the
minor discharge. The scales of area and electromotive force per cm. are not
the same as in Text-fig. 4, but their ratio is the same. By choosing another
value for this ratio, the points could be brought nearer to the curve, but the
proportionality between the electromotive force per cm. and the cross-sec-
tional area is by no means as clear as in the other case. On the other hand,
the measurements are less dependable.

Transverse segments of the body of the fish were also divided sagittally,
and the exposed surface of electric tissue was stained with methylene blue.
The number of electroplaxes per cm. in the direction of the length of the fish,
which is also the direction of the electric polarity, was counted under the
microscope in each of the organs at various points along them. The septa
of the electric tissue are tough but the matter they inclose is watery and

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[XXV :34

the tissue is therefore easily misshapen after being cut. This distortion of
the cut surface could not be entirely prevented but was limited by mechanical
means. The probable error of the count is estimated at less than 10 per cent
along the greater part of the large organ and Hunter’s organ, and somewhat
more elsewhere.

In Text-fig. 6, the black dots show the number of electroplaxes per cm.
at various points along the large organ, and the white dots show the number
per cm. along the organ of Hunter. Except at one place there is no difference
between them too great to be a probable error of measurement. The curve
showing the electromotive force per cm. of the major discharge is also drawn
here for comparison. Again the plotted points fall rather close to the curve,
and here the proportionality between the two plotted quantities has a clear
enough meaning, which is simply that the electromotive force of one electro-
plax is at least roughly the same all along the organ. From the data plotted
in Text-fig. 6 it is about 0.14 volt.

Dr. Kenneth S. Cole, using the needle technique, has found a change
in potential during activity in the squid axon of 0.16 volt, and in Nitella of
0.1 volt. (Personal communication).

Text-fig. 6.

Curve: major discharge, electromotive force per cm. vs. distance along organs.
Black dots: Large organ, number of electroplaxes per cm. vs. distance along or-
gans. White dots: Hunter’s organ, number of electroplaxes per cm. vs. distance
along organs.

1940]

Cox, et al.: Electrophorus electricus

561

Text-fig. 7.

Curve: minor discharge, electromotive force per cm. vs. distance along organ.
Dots: Sachs’ organ, number of electroplaxes per cm. vs. distance along organs.

In the first of the papers referred to above2, the opinion was tentatively
offered that Hunter’s organ produced the irregular discharge to which the
name intermediate was given, the major and minor discharges being more
positively ascribed to the large organ and Sachs’ organ. But it was observed
afterward and mentioned in the second paper3 that the power of the inter-
mediate discharge seemed too great to be generated in Hunter’s organ alone.

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Zoologica: New York Zoological Society

It was concluded therefore that Hunter’s organ produced no separate dis-
charge though it might possibly produce an irregularity in the discharge of
the large organ. The similarity in the structure of adjacent parts of Hunt-
er’s organ and the large organ now leads us to question whether the large
organ ever discharges alone, without Hunter’s organ, or whether, on the
contrary, Hunter’s organ is functionally a part of the large organ, only
separated from it by connective tissue and the remnant of the lateralis imus.

In contrast to Hunter’s organ, which is clearly separated from the large
organ but closely similar to it in structure, the organ of Sachs is hardly
separated at all from the large organ but over much of its length is clearly
different in the size of the electroplaxes. At the two ends of Sachs’ organ,
however, the electroplaxes are about as many to the centimeter as in the
adjacent parts of the large organ, and appear, at least with the crude stain-
ing and under the low magnification employed in these observations, to be
practically identical with them. Especially toward the posterior end the
organs become indistinguishable, the septum between them being no more
marked than the septa within each organ. Toward the anterior end the two
organs remain distinguishable by the sizes of the electroplaxes, at least until
a place is reached at which the cross-section of Sachs’ organ is very small
indeed, containing only a few electroplaxes in parallel.

As shown by the points in Text-fig. 7, the number of electroplaxes per
cm. in Sachs’ organ has by far its highest value at the anterior end, where
the electromotive force per cm. is low, according to the measurements
previously described. This is a surprising finding, as it would imply that the
single electroplax of Sachs’ organ in this part has a much lower electromotive
force than the neighboring and similar electroplax of the large organ. This
seems so unlikely as to require further evidence before it can be accepted.

Preliminary measurements leading to the work here described were
made in 1937 at the Museu Goeldi of Para, Brazil, by the New York
Aquarium-New York University Expedition to study the electric eel. It is a
pleasure to acknowledge again the generous grant to this expedition made by
the Doctor Simon Baruch Foundation and to express again our thanks to
Dr. Godofredo Hagmann of the Museu Goeldi and to many others who aided
the research at that time. The present data were obtained at the Biological
Laboratory of the Long Island Biological Association at Cold Spring Harbor.
We are very grateful to Dr. Eric Ponder and others at the Laboratory for
the help and encouragement we received from them.

Index to Parts 1-4

563

INDEX.

Names in bold face indicate neio genera, species or varieties; numbers in
bold face indicate illustrations; numbers in parentheses indicate the paper
containing the Plate numbers immediately following.

A

Abylopsis ietragona, 304
Acanthephyra, 133
acutifrons, 146, 146
anomala, 185
brachylelsonis, 147
brevirostris, 148, 149, 154
cucullala, 150
curiirostris, 143, 144
eximimia, 147, 148
gracilipes, 149, 150, 151
haeckelii, 140, 141
indica, 150
pulchra, 148
purpurea, 134, 141
sibogae, 150
sp. ?, 151

styloroslrala, 144, 145
lenuipes, 150
Acanlhodoras sp. 549
Acharax, 375
Acila, 388

(Truncacila) caslrensis, 388
Acinonyx jubalus, 276
Acipenser fulvescens, 549
oxyrhynchus, 549

Aclinothoe californica, 217, 217, 218
undala var. anguicoma, 217
Adrana, 389

arcuafa 407, 408
crenifera 407, 408
elongala, 407, 409, (25) Plate II
exoptata, 407, 409, (25) Plate II
janeiroen$is, 408
newcombi, 410
penascoensis, 407, 410
scaphoides, 411
sowerbyana, 407, 410
suprema, 407, 409, 411
taylori, 407, 412
ionosiana, 407, 412, (25) Plate II
Aegina citrea, 313
Aeginura beebei, 309, 310-312
grimaldii, 289, 309, 313
Aequidens raoronii, 549
Agapornis sp. 273
Agelaius ruficapillus frontalis, 273
Agkislrodon blomhoffi, 270
mokasen cupreus, 270
piscivorous, 50, 270
Aidemosyne modesla modesta, 274
Ailuropoda melanoleuca, 261
Aix sponsa, 275
Ajaia ajaja, 272
Aloes aloes, 469

Alicia beebei, 211
Allacanlhochasmus arlus, 550
varius, 550

Alligator mississipiensis, 34
Allocreadium sp., 550
Alopochen aegyptiacus, 468
Amazona barbadensis barbadensis, 272
festiva festiva, 273
ochrocephala oratrix, 272
panamensis, 272
Amblopliles rupestris, 549
Amblyrhamphus holosericeus, 273
Ameiurus melas, 92

nebulosus, 89, 91, 92, 353, 355, 549, 550,
(8) Plate I- VIII
Amia calva, 549

Amnotragus lervia, 277, 278, 469
Amphicondrius granulosus, 337
Amphiodia sculptilis, 337
urtica, 338
vicina, 338, 338
Amphipholis geminata, 336
platydisca, 337
squamata, 337
Amphiura arcystata, 336
Ampholistius sabinus, 550
Anas crecca carolinensis, 272
plaiyrhynchos, 275, 468
platyrhynchos, 275
rubripes tristis, 275
Anguilla japonica, 264
Anodorhynchus leari, 273
Anoptichlhys jordani, 113, (18) Plates I-III
Anthracoceros malabaricus malabaricus, 272
Anthracothorax violicauda violicauda, 273
Anlhropoides paradisea, 274
Apharyngoslrigea, 323
bilobata, 323 (20) Plate I
brasiliana, 324
cornu, 325
egretti, 324
flexilis, 325
garciai, 325
ibis, 325
repens, 325
(?) simplex, 325
Arbacia incisa, 348
Area adamsii, 423
Arctocephalus australis, 277
Ardea cinerea, 325
goliath, 272

herodias herodias, 323, 325
nigra, 326
purpurea, 325
Ardeola ibis ibis, 325

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[XXV

Argidia alonia, 84
azania, 84
hilaris, 84
Argulus spp. 552
Argusianus argus argus, 274
Arius aqua-dulce, 223
Arizona elegans elegans, 270
Asleroidea, 332
Asirapia rolhschildi, 259
siephaniae slephaniae, 272
Astropeclen armaius, 332
californicus, 332
Asiropyga pulvinaia, 348
Ateles cucullatus, 276
Afolla wyvillei, 315
Auripasser Iuteus, 274
Axis axis, 469

B

Badis badis, 549

Bagre marinus 221, 222, 224, 225, 229, 230,
232, 234, 236, (13) Plates I-V
Balistes vetula, 549
Barbus linealus, 549
Basiliscus vilfalus, 34
Benlheocaris exuens, 145
Bianium plicitum, 550
Biiis gabonica, 271
Blarina brevicauda, 476, 477
Boa canina, 271
Boiga dendrophila, 270
Bothriocephalus cuspidatus, 551
Brachyplernus benghalensis benghalensis, 272
Branchellion ravenelli, 551
Branla canadensis, 468
Brotogeris jugularis jugularis, 273
Bubo virginianus virgininanus, 272
Bucephalus pusillis, 550
Bunocephalus bicolor, 549
Buleo jamaicensis borealis, 273
Bycanisles cristalus, 272
sharpii duboisi, 273

C

Calamoichlhys calabaricus, 549
Callinecles sapidus, 227
Calliobolhrium verlicillatum, 551
Callilhrix penicillata, 277
Calcycopsis bigelowi, 290
birulai, 290
borchgrevinki, 290
chuni, 290
geometrica, 290
nemalophora, 290
papillata, 290
prolifera, 303
simplex, 290
simulans, 290, 293
lypa, 290
valdiviae, 290
Camallanus sp., 551
Camelus dromedarius, 278
Campostoma, 2, 6
anomalum, 6, 11
pullum, 7, 9

Cancellaria cumingiana, 371
Cancroma cochlearia, 324

Canis latrans, 278
Carassius auralus, 501, 552
Carcharias littoralis, 551
Cardium magnum, 371

Ca restricta, 86

Carpodacus mexicanus fronialis, 273
Casmerodius albus egrella, 272, 273
Calhaemasia famelica, 327
hians, 326

nyclicoracis, 323, 325, (20) Plate I
reticulata, 326
spectabilis, 327

Catostomus commersonnii commersonnii, 10
Cebus capucina, 276
fatuellus, 466

Centromochlus aulopygius, 549
Cerastes cerastes, 270, 271
Cerococebus fulginosus, 466
torqualus, 276
Cercopithecidae, 513
Cercopithecus grivet, 466
mona cambelli, 276
mozambicus, 466
sabaeus, 276, 466
Cervus axis, 276

canadensis canadensis, 276
duvauceli, 278
eldi, 277
elephas, 469
hortulorum, 277
unicolor, 277
Chaenobryltus gulosus
Chaetodiplerus faber, 549
Channa asiatica, 549
Cheirodon piaba, 549
Chelydra serpentina serpentina, 47
Chiarella centripetalis, 294, 295
Chilodon sp., 550
Chilomyclerus schoeptii, 550
Chlorophonia frontalis frontalis, 272
Chlorostilbon caribaeus caribaeus, 273
Choephaga magellanica, 468
Choeropsis liberiensis, 277
Chondrancanthopsis nodosus, 551
Chromatonema erythrogonon, 297, 298
heriwigi, 297
rubrum, 297

Chrysolophus pictus, 274
Cichlasoma festivum, 549
uropthalmus, 116
ericymba, 116
Ciconia alba, 326

ciconia ciconia, 273
Circus cyaneus hudsonius, 273
macrurus, 325
Cirrhites aprinus, 58
graphidopterus, 58
Cirrhilichthys aureus, 61
corallicola, 58, 60
Cirrhitus, 54
betaurus, 55

rivulatus, 54, 56, 64, (5) Plate I
Clethrionomys gapperi, 477
Clinostomum campanulatum, 323, 327, 551
Clinostomus elongatus, 7
Clypeaster europacificus, 349
ochrus, 350

1940]

Index to Parts 1-4

565

speciosus, 350

Coliuspasser albonotatus, 272
Colobonema typicum, 302
Coluber constrictor constrictor, 36
Columba livia, 272

Constrictor constrictor constrictor, 270
Corallobofhrium sp., 551
Corlurnix corlurnix, 467

Corvus brachyrhynchos brachyrhynchos, 274
Coslelloleda, 398
Crepidostomum sp., 550
Crinifer piscator, 273
Crossobothrium laciniatum, 551
Crossoplilon mantchuricum, 273
Crossophus fodiens, 484
Crotalidae, 49
Crotalus atrox atrox, 270
cerastes, 271

horridus atricaudalus, 270
viridis viridis, 46, 270
Cryptocotyle lingua, 551
Ctenophthalmus pseudagyrtis, 488, 490
Ctenops viilatus, 549
Ctenossaura acanthura, 271
Cunina proboscidea, 306
prolifera, 303, 306
Cunoctona lanzerotae, 309
myosura, 311
nausothoe, 309
Cyanerpes cyaneus, 275
Cyclochaela domerqui, 550
Cygnus olor, 273, 468
Cyprinodontidae, 9
Cyrilla, 389
adamsi, 420
munita, 419

(neopleurodon) subdolus, 420

D

Dama dama, 277, 469
Dascyllus aruanus, 549
Dasyatis hastatus, 551
Dasypus novemcinctus iexanus, 278
Dendroaspis augusliceps, 270
viridis, 270

Dendrocygna autumnalis aulumnalis, 275
discolor, 275

Dendrolagus ursinus, 523
Didelphis virginiana, 276
Diodon hystrix, 549
Diopederma danianum, 343
Diplectanus sp., 550
Dissoura episcopus microcelis, 274
Distoma cornu, 324
Dislomum simplex, 324
Dithadama ? domariia, 87
Doratopsylla blarinae, 488, 490
Dromiceius novaehollandiae novaehollandiae,
275

Drosophila athabascae, 27
azteca, 27
lancefieldi, 26
melanogaster, 27
miranda, 27
pseudoobscura, 23
Drymarchon corais couperi, 270

E

Echinometra vanbrunti, 349
Elaphe obsoleta obsoleta, 37
quadrivittata quadriviltata, 270
vulpina, 38

Electrophorus electricus, 249, 553, (14) Plate I
Ennucula, 382
Ephyrina, 173
bifida, 174
hoskynii, 173
Epibdella melleni, 526, 550
Epicrales cenchris, 271
Epinephelus striatus, 549
Ergasilidae (Spp?), 552
Ergasilus sp., 552
Erithizon dorsalum dorsatum, 277
Erythrocebus patas, 278
Esomus malayensis, 549
(Esox), estor, 551
Esoxlucius, 2

masquinongy, 549
Etroplus suratensis, 549
Eucidaris thouarsii, 347
Euglyphis marbodia, 85
Eumedusa similis, 290
Eunectes murinus, 270
Euphraclus sexcinctus, 277
Euplectes franciscana franciscana, 273
orix orix, 275

Eupomotis gibbosus, 353, 354, 550, (23) Plates

I, II

Eutiara mayeri, 284
Exoglossum, 4, 8

maxillingua, 6, 7, 11

F

Felis cacomitli, 278
cougar, 276
leo, 278
pardalis, 276
Florida caerulea, 272
Forbesella (Chologaster) agassizii, 115
Fundulus bermudae, 97, 98, 99, 100
heteroclitis, 550

G

Galago alleni, 436
crassicaudalus, 436
montieri, 440
demidoffii, 436

senegalensis moholi, 433, (27) Plates I-VI
Galeichlhys felis, 221, 222, 225, 229, 230, 232,
234, 236, 549, (13) Plates I-V
milberti, 235
Gallus gallus, 467
gallus, 273
Gambusia afinis, 501
afinis, 108

holbrooki, 97, 99, 106, 107, 109
Gennaeus leucomelanos melanotus, 274
nycthemerus, 467
swinhoii, 467

Ginglymostoma cirralum, 549
Gonostoma elongatum, 127, 136, 175
Gorilla berengei, 519
gorilla, 513

Grus canadensis tabida, 275

566

Zoologica: Neiv York Zoological Society

[XXV

Guttera edouardi edouardi, 272
Gymnoderus foetidus, 274
Gymnoihorax funebris, 549
moringa, 549
Gymnoiis coatesi, 549

H

Halichaeres radialus, 549
Halicreas minimum, 302
Hapale pygmaea, 276
Harengula pensacolae, 22
Helioperca macrochira, 549, 550
Heloderma horridum, 271
suspeclum, 271
Hemitragus jemlahicus, 276
Hemixus flavala flavala, 275
Henneguya acerinae, 92

ameiurensis, 89, 90, 91, 92, 552, (8) Plates
I- VIII
exilis, 93
gurleyi, 92

Herichlhys cyanogutlatus, 549
Herodias iimorensis, 325
Holoslomum repens, 324
Huro salmoides, 549, 551
Hybognalhus regius, 10
Hyborhynchus nolatus, 551
Hyelaphus porcinus, 470
Hylesia detica, 85
Hymenodora glacialis, 176, 179
gracilis, 175, 176

Hyphessobrycon helerorhabdus, 549
serpae, 549

I

Ichthyophlhirius mullifiliis, 527, 550
Iclalurus punctalus, 93, 551
Iguana iguana rhinolapha, 270
Ilyoplax pusillus, 66

J

Jupiteria, 402

K

Kakaloe galerila galerita, 274
roseicapilla roseicapilla, 274
sanguinea goffini, 274
Kaladesmia, 413

Kilfocincla malabarica malabarica, 274
Koiropolamus koiroplamus kenlae, 277

L

Lagothrix humboldtii, 277
ingumalus, 277
Lama vicugna, 278

Lampropeltis geiulus floridana, 40, 270
geiulus, 271
splendida, 271
subsp., 270

Larus argenlatus smithsonianus, 273
atridilla, 274

novaehollandiae novaehollandiae, 274
Lebisles, 98

reticulatus, 99, 109, 110, 111
Lecilhaster lindbergi, 264
Leda agapea, 394
elecla, 412
Jorlis, 402

inornala, 404
melcalli, 408
pontonia, 403
sowerbyana, 411
laylori, 411

Lepisosleus osseus, 549

Lepomis aurilis, 329, 364, 549 (21) Plate I
cyanellus, 364

Leplochela bermudensis, 131, 132
Leploplilus crumenifer, 327
Leptotila cassini cassini, 274
Lernae sp., 552
Lernaeenicus radialus, 551
Leuckarliara gardineri, 284
zacae, 284, 285, 286
Leucosomus corporalis, 6, 7, 11, 363
Leucotmenis kaieiura, 87
Ligula inteslinalis, 551
Linckia colombiae, 334
Linlonium vibex, 550
Liriope letraphylla, 302
Lophortyx californica californica, 274
Lophura rufa, 273
Lovenia cordiformis, 352
Lucaya bigelowi, 189, 190
Luidia asthenosoma, 332
bellonae, 332
ioliolala, 333
ludwigi, 333

Lutra canadensis vaga, 277
Lyfechinus anamesus, 348
pictus, 348

M

Macaca irus, 276, 466, 513
mulatta, 276, 466, 515
nemestrina, 276
Macaria barticaria, 86
Macrophthalmus depressus, 66
dilatatus, 66
japonicus, 66

Macropus giganfeus giganleus, 277
Malletia, 389, 420
elongata, 422
faba, 423
gigantea, 421
inequalis, 423
magellanica, 424
(Malletia) chilensis, 421
(Minormallelia) arciformis, 423
(Minormallelia) benthina, 423
(Neilo) goniura, 422
patagonica, 424
peruviana, 424
truncata, 424
Medusa mollicina, 304
pilearis, 304
Megaceryle alcyon, 326
Megalaima virens marshallorum, 274
Megayoldia, 414

Meleagris gallopava silvestris, 272
Melopsitlacus undulatus, 274
Melozone leucolis, 272
Menidia menidia, 551
Meningodora mollis, 165
Meoma grandis, 352
Mephitis nigra, 276

1940]

Index to Parts 1-4

567

Microcotyle sp., 550
Microglanis parahybae, 549
Microgonia hildonia, 87
Micropierus dolomieu, 551
Microfus pennsylvanicus, 477
Minormalletia, 422
Moenkhausia pilfieri, 549
Moira clotho, 350

Mollienisia sphenops, 98, 99, 101-105
ssp., 104

Monolepis albicans, 67
inermis, 66, 67, 70, 81
orientalis, 67

Morone interrupfa, 549, 550
Mugil curema, 237
Munia atricapilla atricapilla, 274
Munliacus munljak, 469
Murex recurviroslris, 217
Musophaga violacea violacea, 274
Muslelus canis, 551
Myadesles unicolor, 275
Mycteroperca bonaci, 549
Mylossoma durivenlris, 549
Myxobolus cf. linearis, 92
sp., 550

N

Naja hannah, 27
naja, 270
nivea, 270

Napaeozopus insignis, 477
Narcine brasiliensis, 431
Nasua nasua, 276
Natrix cyclopion floridana, 42
erylhrogasler Iransversa, 43
sepiemvitfaia, 44
sipedon picliventris, 45
sipedon, 9

Neascus vancleavi, 551
Neila, 422
Nemanthidae, 212
Nemanlhus, 212

caliiornicus, 214, 215, 216
nilidus, 213, 214, 216
Nemafobothrium sp., 550
Neochasmus umbellus, 550
Neopleurodon, 419
Neolurris crockeri, 287, 287-289
Nidorellia armala, 333
Nocomis, 2, 6, 7
biguttatus, 6, 361
micropogon, 6, 361
Nofemigonus crysoleucas, 495
Noliopsar curaeus, 274
Notophoyx novaeholliandiae, 325
Noloslomus, 152
beebei, 168
compsus, 156, 157
distirus, 166, 167
fragilis, 165
gibbosus, 170
iaponicus, 168
longiroslris, 172
marplocheles, 158, 159, 160
miccylus, 161 162, 163
mollis, 164, 165
murrayi, 168
paientissimus, 172

perlafus, 170, 171
robuslus, 168, 169
vescus, 153, 154
weslergreni, 171, 172
Notropis cornuius, 1, 361, (1) Plafes I-IV
chryocephalus, 2, 5, 10
cornuius, 1, 5, 362
frontalis, 2, 4
hudsonius hudsonius, 10
pilsbryi, 361
x rubellus, 7, 361, 362
Nucolopsis, 383
Nucula, 379
abrupta, 422
agujana, 379, 384
cahuifensis, 381
chrysocoma, 380, 385
colombiana, 379, 385
crenulata, 380
darwini, 411
dorbignyi, 408
(Ennucula) cardara, 379, 383
(Ennucula ?) linki, 379, 383
(Ennucula) tenuis, 379, 382
grayi, 379, 385
hilli, 384

iphigenia, 380, 386
lanceolata, 411
miliaris, 418

(Nucolopsis) schencki, 384, (25) Plale I
(Nucula) declivis, 380, (25) Plale I
(Nucula) exigua, 380, 381
obliqua, 382
panamina, 379, 386
payiensis, 380, 381, 386
pigafetiae, 380, 386
pisum, 379, 387
proxima, 427
savalieri, 379, 387
schencki, 380
semiornala, 380, 387
suprastriata, 380
taeniolata, 380, 387
tanneri, 379, 388
Nuculana, 378, 390

acapulcensis, 394, (25) Plate I

acrita, 391, 402

agapea, 391 402

bicostata, 391, 394, 402

cestrota, 399

cordyla, 390, 401

(Cosfelloleda) cosiellala, 390, 398 (25)

Plale II

(Cosfelloleda) marella, 390, 399, (25) Plate II

crispa, 391, 403

cuneata, 391, 403

elaborata, 401

excavafa, 391, 404

flexuosa, 399

hamafa, 390

hindsii, 391, 404

lobula, 392, 404

loshka, 390, 404

lucasana, 392, 405

lyrafa, 395

media, 394

ornata, 391, 405

568

Zoologica: New York Zoological Society

[XXV

oxia, 391, 405
peruviana, 390, 405
(Polilolida) polila, 391, 394
ponlonia, 391, 406
pyriformis, 394
rhylida, 392, 406

(Saccella) acula, 390, 392, (25) Plate I
(Saccella) callimene, 391, 393, (25) Plate I
(Saccella) eburnea, 391, 395, (25) Plate II
(Saccella) elenensis, 391, 393, 394, (25) Plate
I

(Saccella) gibbosa, 392, 395, (25) Plate II
(Saccella) impar, 391, 396, (25) Plate II
(Saccella) laeviradius, 391, 396, (25) Plate II
(Saccella) laphria, 391, 397
(Spinula) calcar, 391
(Spinula) calcarella, 391
(Theslyleda) cordyla, 400
(Thestyleda) hamala, 400, (25) Plate III
Nuculidae, 378
Nuculanidae, 389

Nyclicorax nyclicorax hoactli, 323, 325
Nymphicus hollandicus, 274

O

Ocypode, 65

albicans, 66, 69, 70, 71, 74-80
gaudichaudii, 66, 67, 68, 75, 77-80
occidentalis, 66, 69, 70, 74-80
Odocoileus virginianus leucurus, 470
nemoralis, 469
osceola, 276

Oodinium ocellatum, 536, 550
Opheodrys aeslivus, 35
Ophiactis savignyi, 339
simplex, 340
Ophiocoma aelhiops, 341
alexandri, 341

Ophioderma panamense, 341
teres, 342
variegalum, 343
Ophiolepis fulva, 345
grisea, 346
variegaia, 344

Ophiomyxa panamensis, 336
Ophionereis annulala, 340
nuda, 340

Ophiophragmus, 338
Ophiopleris papillosa, 341
Ophiothela gracilis, 340
Ophiothrix spiculala, 340
Ophiozona pacifica, 344
Ophisaurus apodus, 35
Ophiura lulkenii, 343
Opislhonema oglinum, 227
Oplophorus, 132, 184
foliaceus, 185
gracilirostris, 188
grimaldii, 187, 188
spinicauda, 184, 185
Oreasler occidentalis, 333
Orthoyoldia, 414
Oslinops viridis, 272
Olhilia aculeata, 334, (22) Plate I
Otobolhrium crenacolle, 551
Ololicnus garnetli, 442
Ovalipes ocellatus, 227

P

Padda oryzivora, 275
Pandarus bicolor, 551
Pandea conica, 284
rubra, 287
Papio anubis, 276
papio, 466
Paracalops, 477
breweri, 489
Paradisaea gulielmi, 273
Paradoxurus hemaphrodifus, 278
Parapandalus, 192
richardi, 192
zurstrasseni, 194
Paroaria nigrigenis, 274
Parolia lawesi lawesi, 257, 273
sefilala, 257
wahnesi, 257, 257, 259
Parapasiphae, 126

macrodactyla, 126, 128, 129
sp. a., 126, 129
sp. b., 126, 130
sulcalifrons, 126
Pasiphaea alcocki, 123
amplidens, 126
corteziana, 123
hoplocerca, 124, 125
liocerca, 122, 123
longispina, 123, 126
orientals, 126
ralhbunae, 123
scoliae, 123, 126
Paliria miniata, 333
Pavo crislatus, 275, 467
var., 274

Pegantha clara, 305, 306
martagon, 308
smaragdina, 305
triloba, 308

Pegasia dodecagona, 304
Pelagia noctiluca, 315
Pelecypoda, 373
Penella instructa, 551
Perea llavescens, 549, 551
Periphylla hyacinthina, 314
Peromyscus, 22

leucopus noveboracensis, 477, 489
maniculatus, 489
Petrasmus, 377

Phalacrocorax aurilis aurilis, 327

Pharia pyramidala, 334

Pharomachrus mocinno mocinno, 273

Phasianus colchicus versicolor, 274

Phassus guianensis, 88

Phastolomis milchelli, 277

Phataria unifascialis, 335

Philortyx fascialus, 274

Pholostomias guernei, 127

Physetocaris microphthalma, 196, 197, 198

Pionites melanocephala melanocephala, 274

Pitta sordia cucullala, 274

Pituophis melanoleucus melanoleucus, 39, 271
sayi sayi, 270

Pitymys pinetorum scalopsoides, 477
Placobdella parasitica, 551
Plagiobrissus grandis, 351

pacificus, 351, (22) Plate I

1940]

Index to Parts 1-4

569

Plagusia depressa luberculata, 66, 70, 72
Pleclropferus gambensis niger, 273
Plesionika irtarfia, 190, 191
Plethodon cinereus, 481
Pleurodon, 418
adamsii, 418
cinclus, 418
dobergensis, 418
gunleri, 418
hedleyi, 418
microdus, 418
ovalis, 419
woodi, 418
zinndorfi, 418

Ploceus vilellinus vilellinus, 275
Pocillopora, 342
Poecilidae, 97
Podocotyle almon, 265

alzi, 263, 265, 266, 550 (17) Plale I
Pogonias cromis, 549
Politoleda, 397
Polycolpa forskalii, 303
Polyorchis penicillala, 296
Polyxenia cyanosfylis, 304
Pomoxia sparoides, 549
Pongo pygmaeus, 466
Pontobdella muricala, 551
Poronofus Iricanthus, 551

Porphyrio poliocephalus poliochephalus, 272

Polos flavus, 278

Prepiella deicoluria, 88

Prionotus scifulus, 227

Pristella riddlei, 549

Procavia capensis, 277

Probliolrema sp., 550

Proteocehphalus pinguis, 551 (33) Plate V
sp., 551

Protrichomonas sp., 550
Pseudbarydia elipha, 83
Pseudemys floridans ssp., 48
Pseudocirrhiles, 53
pinos, 61, 62

Psittacula eupatria nipalensis, 274
Psiltacus erithacus erilhacus, 273
Ptilinopus jambu, 273
Plyas mucosus, 270
Pucrasia macrolopha macrolopha, 272
Python curtus, 270
molurus molurus, 270
regius, 271
reticulatus, 270

R

Ramphostos discolorus, 274
piscivorous brevicarinatus, 272
swainsonii, 273
Ramplocelus carbo carbo, 275
Rasbora maculata, 549
heleromorpha, 549
trilineata, 549

Rhinicylhys atratulus alrafulus, 9, 10, 14 (1)
Plates II-IV
cataracfae, 7
Rhinocheilus lecontei, 41
Rhodeus amarus, 353
Rhynchotus rufescens rufescens, 275
Rhynocheios jubatus, 273

Rhyticeros subruficollis, 272
Roccus lineatus, 549
saxatilis, 237
Rubigula dispar, 274
Rypticus saponaceus, 549

S

Saccella, 392

Sagittarius serpeniarius, 272
Saiga tatarica, 277
Salvelinus fontinalis, 549
Sarcophilus harrissii, 276
Sarda sarda, 550
Scaphirhyncus platorhyncus, 549
Scardinius erythrophlhalmus, 549
Scatophagus argus, 549
Schizoderma diplax, 343
Scleoporous undulatus iasciatus, 34
Sciaenops ocellafus, 549
Scleraslerias alexandri, 335
crassia var., 335
heteropaes, 335
var., 335

Scopimera globosa, 66

Scorpaena madurensis, 263, 264, 265, 550, (17)

Plate 1

Scyllarides lalus, 551
Sebastes marinus, 551
Semotilus, 2, 6, 7

atromaculatus atromaculatus, 6, 7, 9, 11, 14
(1) Plale I

Seriola zonata, 549
Serranus cabrilla, 237
Serrasalmus scapularis, 549
Sigsbeia lineata, 336
Sika nippon, 468
Simiidi, 513
Solemya, 374

(Acharax) agassizii, 375
(Acharax) dalli, 375
(Acharax) johnsoni, 376
(Acharax) macrodactyla, 376
(Acharax) patagonica, 376
australis, 374
borealis, 377
mediterranea, 374
parkinsonii, 374
(Pelrasma) panamensis, 377
(Petrasma) valvulus, 378
(Solemya) lomitensis, 374
tokunagai, 376
yessoensis, 376
Solemyarina velesiana, 374
Solenella gigantea, 421
Solenodon paradoxus, 276
Solmissus albescens, 308
incisa, 308
marshalli, 308
Sorex araneus, 473
granti, 473
cinereus, 474
dispar, 474

fumeus fumeus, 473 (29) Plates I-IV
minutus, 485
vulgaria, 480

Sparus macrocephalus, 264
Spheroides maculalus, 550

570

Zoologica: New York Zoological Society

[XXV

Sphyrion lumpi, 551
Sphyrna zygaena, 551
Spinilectis sp.,

Spinula, 401

Spinus Iristis Irislis, 274
Stephanoproproides, 326
Slernotherus odoratus, 47
Sterrhurus branchialis, 550
Slizosledion vilreum, 550
Slomolophus agaricus, 316
meleagris, 316
Stomotoca divisa, 283
plerophylla, 283

Streptopelia decaocto decaoclo, 272
Slrongylocenlrolus purpuratus, 349
Sloreria dekayi, 41
Slylocidaris dubius, 347
Suricala suricatta, 277
Sylvacapra grimmi, 277
Symbranchus marmoratus, 549
Synaplomys cooperi, 477
Syslellaspis, 179
braueri, 180
debilis, 181, 195
densispina, 180
echinurus, 180

T

Tachysurus barbus, 227
Taeniopygia castanolis castanotis, 275
Tangara arlhus, 272
cyanoptera, 273
cyaneicollis hannahiae, 272
flava cayana, 273
gullala gullata, 275
heinei, 272
icterocephala, 274
larvata centralis, 273
mexicana mexicana, 273
violacea violacea, 273
xanthogasler exsul, 275
Tarpon atlanticus, 493, 549 (30) Plates I, II
Tautoga onitis, 551
Taxidea taxus taxus, 276
Taxus canadensis, 476
Terrapene Carolina Carolina, 4
Testudo tormieri, 48
Thalassoma bifasciatus, 549
Thamnophis sirtalis sirtalis, 46
Theristicus caudatus, 274
Thestyleda, 400
Thraupis cana cana, 274
Tiliapia mossambica, 549
Tindaria, 389, 425
amabilis, 426
atossa, 426
compressa, 426
mexicana, 427
panamensis, 426
salaria, 427
smirna, 427
thea, 427

(Tindariopsis) sulculata, 425
virens, 428
Tindariopsis, 425
Tolype loisa, 85

Taxopneusles roseus, 349
Trachinotus glaucus, 549
Tragelaphus angasi, 276
Tragopan caboli, 273
Trichechus manalus latirostrus, 550
Trichiurus lepturus, 550
Trichodina sp., 550

Trichoglossus haematod moluccanus, 274
Tridacna gigas, 373
Trimeresurus flavomaculaius, 270
Truncacila, 388

Tubulovesicula anguillae, 264, 265
angusticauda, 265
calilornica, 264, 265
lindbergi, 265

madurensis, 263, 264, 265, 550 (11) Plate I
muraenesocis, 264, 265
pseudorhombi, 264, 265
spari, 264, 265
Tupinambis teguixin, 271
Turdus merula merula, 272
olivaceus olivaceus, 274
Tyto alba pralincola, 272

U

Uca minax, 66
pugilator, 66
pugnax, 66

Uraeginlhus bengalus bengalus, 273
Urobrachya axillaris axillaris, 273
Ursus arctos arctos, 277

V

Varanus sp., 271
Venadicadia cetiona, 86
Vipera russelii, 270

X

Xenomyslus nigri, 549
Xiphias gladius, 551

Y

Yoldia, 389, 412
emersoni, 414
(Katadesmia) vincula, 413
(Megayoldia) martyria, 414
(Orthoyoldia) panamensis, 415
subaequilateralis, 421
thracoaeformis, 416, 421
(Yoldiella) cecinella, 416
(Yoldiella) chilenica, 416
(Yoldiella) dicella, 416
(Yoldiella) granula, 416
(Yoldiella) indolens, 417
(Yoldiella) infrequens, 417
(Yoldiella) leonilda, 417
(Yoldiella) mantana, 417
Yoldiella, 415

Z

Zalophus californias, 277
Zapus hudsonius, 477
Zonurus giganleus, 270
Zygocanna costata, 301
pleuronota, 301
vagans, 299, 300
Zygocannula, 301

NEW YORK ZOOLOGICAL SOCIETY

General Office: 630 Fifth Avenue, New York City

OFFICERS

President, Fairfield Osborn
First Vice-President, Alfred Ely

Chairman, Executive Committee & Second Vice-President , Laurance S. Rockefeller

Treasurer, Cornelius R. Agnew

General Director, Zoological Park and Aquarium, Allyn R. Jennings
Assistant General Director, Harry Sweeny, Jr.

SCIENTIFIC STAFF

Zoological Park

Raymond L. Ditmars, Curator of Reptiles and Insects
Lee S. Crandall, Curator of Birds
Claude W. Leister, Curator of Mammals and Educational Activities
Leonard J. Goss, Curator of Health
William Bridges, Editor and Curator of Publications

Aquarium

Charles M. Breder, Jr., Director
Christopher W. Coates, Aquarist
Ross F. Nigrelli, Pathologist
G. M. Smith, Research Associate in Pathology
Homer W. Smith, Research Associate in Physiology

Department of Tropical Research

William Beebe, Director
John Tee-Van, General Associate
Gloria Hollister, Research Associate
Jocelyn Crane, Technical Associate

Editorial Committee

Fairfield Osborn, Chairman

Allyn R. Jennings Charles M. Breder, Jr.

William Beebe William Bridges

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