ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS
of the
NEW YORK ZOOLOGICAL SOCIETY
VOLUME 34 '3 c>
1949 -s1"
NUMBERS 1-20
Published by the Society
The Zoological Park, New York
December 30, 1949
NEW YORK ZOOLOGICAL SOCIETY
General Office: 30 East Fortieth Street, New York 16, N. Y.
Publication Office: The Zoological Park, New York 60, N. Y.
OFFICERS
Fairfield Osborn, President
Alfred Ely, Vice-president
Laurance S. Rockefeller, Vice-president
Donald T. Carlisle, Vice-president
Harold J. O’Connell, Secretary
Cornelius R. Agnew, Treasurer
SCIENTIFIC STAFF
General 3 C\ , QG Q 1 A ^
John Tee-Van, Executive Secretary
William Bridges, Editor and Curator of Publications
Sam Dunton, Photographer
Zoological Park
Lee S. Crandall, General Curator
Grace Davall, Assistant to General Curator
Brayton Eddy, Curator of Reptiles and Insects
Leonard J. Goss, Veterinarian
Robert M. McClung, Assistant, Mammals and Birds
Aquarium
Christopher W, Coates, Curator and Aquarist
James W. Atz, Assistant Curator
Ross F. Nigrelli, Pathologist
Myron Gordon, Geneticist
C. M. Breder, Jr., Research Associate in Ichthyology
G. M. Smith, Research Associate in Pathology
Homer W. Smith, Research Associate in Physiology
Department of Tropical Research
William Beebe, Director
Jocelyn Crane, Research Zoologist
Henry Fleming, Entomologist
William K. Gregory, Associate John Tee-Van, Associate
Gloria Hollister, Associate Mary VanderPyl, Associate
Scientific Advisory Council
A. Raymond Dochez Caryl P. Haskins
Alfred E. Emerson K. S. Lashley
W. A. Hagan John S. Nicholas
George M. Smith
Editorial Committee
Fairfield Osborn, Chairman
William Beebe Lee S. Crandall
William Bridges Brayton Eddy
Christopher W. Coates Leonard L. Goss
John Tee-Van
CONTENTS
Part 1. May 16, 1949.
PAGE
1. Paradilepis simoni n. sp., a Cestode Parasitic in the Osprey. (Ces-
toda: Dilepididae) . By Robert Rausch. Text-figure 1 1
2. A Contribution to the Study of North American Cestodes of the
Genus Paruterina Fuhrmann, 1906. By Robert Rausch and
Everett L. Schiller. Text-figures 1-12 5
3. Behavioral Interactions in a Herd of Barbary Sheep ( Ammo-
tragus lervia) . By Irwin Katz 9
4. The Pericopidae (Moths) of Kartabo, British Guiana, and Cari-
pito, Venezuela. By Henry Fleming 19
5. Report on a Collection of Phalangids from Rancho Grande, Vene-
zuela. By Clarence and Marie Goodnight. Text-figures 1-4 21
6. Fresh-water Crabs of the Genus Pseudothelphusa from Rancho
Grande, Venezuela. By Jocelyn Crane. Text-figures 1-3 25
Part 2. August 10, 1949.
7. Comparative Biology of Salticid Spiders at Rancho Grande, Vene-
zuela. Part III. Systematics and Behavior in Representative New
Species. By Jocelyn Crane. Text-figures 1-8 31
8. The Swifts of Rancho Grande, North-central Venezuela, with
Special Reference to Migration. By William Beebe. Plate I;
Text-figures 1-3 53
9. Eastern Pacific Expeditions of the New York Zoological Society.
XL. Mollusks from the West Coast of Mexico and Central Amer-
ica. Part VII. By Leo George Hertlein & A. M. Strong. Plate I.. 63
10. Fishes That Rank Themselves Like Soldiers on Parade. By E. W.
Gudger. Plate I ; Text-figures 1 & 2 99
11. Notes on Seasonal Changes in Creatophora cinerea, the Wattled
Starling. By Lee S. Crandall. Plate 1 103
12. Insect Migration at Rancho Grande in North-central Venezuela.
General Account. By William Beebe. Plates I & II ; Text-figure 1 107
Part 3. November 30, 1949.
PAGE
13. The Behavior of Two Captive Specimens of the Lowland Gorilla,
Gorilla gorilla gorilla (Savage & Wyman). By B. F. Riess,
Sherman Ross, S. B. Lyerly & H. G. Birch. Plates I & II;
Text-figures 1 & 2 Ill
14. Migration of Papilionidae at Rancho Grande, North-central
Venezuela. By William Beebe. Plate I; Text-figure 1 119
15. Notes on Ergasilus Parasites from the New Brunswick, New
Jersey, Area, with a Check List of All Species and Hosts East
of the Mississippi River. By Roland F. Smith 127
16. An Analysis of Reproductive Behavior in the Mouth-breeding
Cichlid Fish, Tilapia macrocephala (Bleeker). By Lester R.
Aronson. Plates I-III; Text-figures 1-10 133
Part 4. December 30, 1949.
17. Comparative Biology of Salticid Spiders at Rancho Grande, Ven-
ezuela. Part IV. An Analysis of Display. By Jocelyn Crane.
Plate I; Text-figures 1-9 & 159
18. Differential Effects of Estradiol, Estradiol Benzoate and Preg-
neninolone on Platypoecilus maculatus. By Margaret Cordsen
Tavolga. Plates I-V ; Text-figures 1-5 215
19. Eastern Pacific Expeditions of the New York Zoological Society.
XLI. Mollusks from the West Coast of Mexico and Central
America. By Leo George Hertlein & A. M. Strong. Plate 1 239
20. Tettigellidae and Gyponidae (Homoptera) of Kartabo, Bartica
District, British Guiana. By Z. P. Metcalf. Text-figures 1-8.... 259
Index to Volume 34 2wl
Rausch: Paradilepis simoni, n. sp., a Cestode Parasitic in the Osprey
1
1.
Paradilepis simoni n. sp., a Cestode Parasitic in the Osprey.
(Cestoda: Dilepididae).1
Robert Rausch2.
Department of Veterinary Science, University of Wisconsin, Madison 3
(Text-figure 1).
The knowledge of helminths parasitic in
North American birds is very incomplete,
especially for the region west of the Miss-
issippi. Not only is this true in regard to
parasite-host ecology, but a little work with
almost any group of birds discloses unde-
scribed forms or species unreported from
North America, as well. In fact, anyone wish-
ing to carry out host-parasite studies must
devote considerable time to describing spe-
cies-time which could be more profitably
spent otherwise. The osprey might be men-
tioned to illustrate this situation. Of this
bird the writer has examined but three speci-
mens, one each from Ohio, Wisconsin and
Wyoming. From this small series of birds,
four species of helminths were collected; of
these, two species were undescribed and two
had never been recorded from North Amer-
ica. While it is true that a comparable situa-
tion is not to be expected in every case, it
soon becomes obvious from work with a
given host-group that much remains to be
done before the helminths encountered in
birds can be readily identified.
The cestodes with which this paper is con-
cerned were obtained from the small intes-
tine of an osprey, Pandion haliaetus carolin-
ensis (Gmelin), collected on June 3, 1948,
near Moran, Wyoming. This osprey was one
of 267 birds collected by the writer for hel-
minthological study from the Jackson Hole
region of Wyoming.
A total of more than 75 worms was ob-
tained. Whole-mounts were prepared of
specimens stained with Semichon’s acetic
carmine and Delafield’s haematoxylin. Serial
sections, cut at 15/x, were also studied.
This cestode is named in honor of Mr.
James Simon, Director of the Jackson Hole
Wildlife Park, whose generous cooperation
contributed much to the success of the field
work in Jackson Hole.
1 Contribution of the 1948 Research Program of the New
York Zoological Society at Jackson Hole Wildlife Park.
4 Now with U. S. Public Health Service, Anchorage.
Alaska.
3 Section on Parasitology, B. B. Morgan. Project Leader.
Paradilepis simoni n. sp.
(Text-fig. 1 A-E).
Diagnosis : Strobila from 50 to 90 mm.
long; greatest width, up to 450 p, attained in
terminal gravid segments. External seg-
mentation absent ; strobila very delicate and
translucent in the living worm. Scolex large
and distinct from neck; from 470 to 596 p
in diameter. Suckers from 180 to 220 p in
diameter. Well-developed rostellum slightly
over 100 p long; armed with about 36 hooks
arranged in a double row. Large hooks from
98 to 102 p long ; small hooks 68 to 72 p long.
Hook shape typical for genus. Neck from
250 to 270 p wide, narrowing gradually to
a distance of about 2 mm. posterior to scolex ;
from this point the strobila widens to reach
greatest width at posterior enu. Musculature
consists of two layers; the first layer, of lon-
gitudinal fibers, is from 1 to 3 bundles deep;
directly beneath it is a layer of transverse
fibers. Excretory canals typical in arrange-
ment; the ventral longitudinal canal meas-
ures from 6 to 20 p in diameter; the dorsal
and transverse canals about 3 p in diameter.
Internal segmentation best recognized by the
arrangement of the transverse excretory ca-
nals, which divide the strobila into about 30
“segments” per mm. of length in the mature
region. Genital organs not confined entirely
to space between transverse canals, but over-
lap into adjacent segments. Genital Anlagen
appear about 2 mm. posterior to scolex. Geni-
tal pores unilateral and sinistral; genital
atrium about 16 p deep. Genital canals pass
dorsal to longitudinal excretory canals. Five
spherical to ellipsoidal testes, not all in same
plane, in each segment; testes measure from
26 to 33 p in diameter in mature segments.
Usually 4 testes are aporal, and one is poral
of female organs; at times 2 may be poral.
Flask-shaped cirrus sac extends to middle of
mature segments, or beyond, dorsal to testes;
it measures from 100 to 132 p long by 30 to
40 p wide. Cirrus heavily spined. Internal
and external seminal vesicles absent; ductus
ejaculatorius coiled within bulb of cirrus sac.
Text-fig. 1. The morphology of Paradilepis simoni n. sp. The drawings were made in
part with the aid of a projector. A. Ventral view of a typical mature segment. B. Typical
scolex. C. Dorsal view of a section of gravid segments. D. Hooks from rostellum. E. Cross-
section of genital atrium region, showing relation of cirrus to vagina.
1949]
Rausch: Paradilepis simoni, n. sp., a Cestode Parasitic in the Osprey
3
Well-developed vas deferens with numerous
convolutions situated in dorsal part of seg-
ment. Cirrus sac provided with strongly-
developed retractor muscles. Thin-walled va-
gina opens ventral to cirrus sac; it enlarges
gradually, attaining greatest diameter near
place where longitudinal excretory canals are
crossed ; it narrows after this point and runs
medially to join small seminal receptacle dor-
sal to ovary. Ovary rather variable in shape
and position; usually 4-lobed, situated near
middle of segment. Spherical to ellipsoidal
vitelline gland dorsal to posterior part of
ovary; it increases in size toward posterior
end of strobila, attaining a maximum diam-
eter of about 40 p. Uterus develops as two
lateral, spherical sacs situated ventral to
ovary, and connected by a narrow neck. Grav-
id uterus fills entire segment; unlobed and
sac-like when completely gravid. Cirrus sac
and vagina persist into terminal gravid seg-
ments. Eggs, from 27 to 33 p in diameter, are
arranged in 3 to 4 rows across the segments.
Embryonic hooks about 6 p in length.
Host: Pandion lialiaetus carolinensis
(Gmelin) (Osprey).
Habitat: Small intestine.
Locality: Moran, Wyoming.
Type : Three slides of cotype material have
been deposited in the Helminthological Col-
lection of the U. S. National Museum, No.
46403.
Discussion.
As far as could be determined, the genus
Paradilepis Hsii, 1935, has not been previ-
ously recorded from North America. Neither
has the writer discovered any record of ces-
todes parasitic in the osprey.
Cestodes of the genus Paradilepis are typi-
cally parasitic in pelicaniform birds, particu-
larly in cormorants, Phalacrocorax spp. The
genus Paradilepis was established (Hsii,
1935) for cestodes from a Chinese cormor-
ant, with P. duboisi as type. Hsii also assigned
Dilepis scolecina (Rudolphi, 1819) to the
genus Paradilepis. According to Joyeux and
Baer (1935), P. duboisi is identical with P.
scolecina; consequently P. scolecina (Syn. P.
duboisi ) becomes type species. The examina-
tion of the original preparations of Oligor-
chis delachauxi Fuhrmann, 1909, led Joyeux
and Baer (1935) to place it in the genus
Paradilepis. It had been earlier assigned by
the same writers (1930) to the genus Dilepis
Weinland. Further study of their African
material disclosed that they were deaiing
with two species, referred to as Dilepis de-
lachauxi (Fuhrmann, 1909). As a result, a
new name, P. macracantha, was proposed
(Joyeux and Baer, 1935) for Dilepis dela-
chauxi Joyeux and Baer, 1930 nec Fuhrmann,
1909.
Burt (1940) described Paradilepis brevis
from a Ceylon cormorant, apparently without
referring to the work of Joyeux and Baer
(1935). It is possible that P. brevis is iden-
tical with P. scolecina.
Joyeux and Baer (1935) suggested that
Oligorchis longivaginosus Mayhew, 1925,
might also belong to the genus Paradilepis.
This is of particular interest in connection
with the present paper, since O. longivagino-
sus was collected from a white pelican from
Yellowstone Park, Wyoming. Apparently
this species has a single crown of hooks, in-
stead of a double row as seen in Paradilepis ;
external segmentation also seems evident.
The number of species of the genus Para-
dilepis is at present indefinite, and must re-
main so until some of the material is studied
further. Regardless of this situation, P.
simoni is readily differentiated from any
others previously assigned to the genus in
that it possesses 5 testes in each segment,
instead of 4.
Although the presence of 4 testes is con-
sidered a generic character by Joyeux and
Baer (1935, 1936), we do not consider it
justifiable to erect a new genus for P. simoni
on the basis of this character alone. It is
otherwise very similar to the other members
of the genus. Since the previously known
species have been described from pelicani-
form birds, it is not strange that they are
morphologically similar. If, in addition to
P. simoni, cestodes of this genus are recorded
from other host groups, a much better con-
cept of morphological variation within the
genus may be had.
It is possible that P. simoni is an “acci-
dental” parasite of the osprey, and occurs
naturally in cormorants. It would be of in-
terest to examine cormorants from the colony
north of Jackson Hole in order to determine
whether they are parasitized by any species
of Paradilepis. Since all the hosts are pis-
civorous, presumably species of fish might
act as the intermediate hosts of cestodes of
this genus. At present, there is no reason to
doubt that the osprey is the natural host of
P. simoni.
References.
Burt, D. R. R.
1940. New species of cestodes from Chara-
driiformes, Ardeiformes, and Pelicani-
formes in Ceylon. Ceylon Jour. Sci.,
sect. B, 22 1-63.
Hsu, H. F.
1935. Contributions a l’etude des cestodes de
Chine. Rev. Suisse Zool., 42: 477-570.
Joyeux, Ch. and Baer, J. G.
1930. Mission saharienne Augieras-Draper,
1927-1928. Cestodes. Bull. Mus. Nat.
Hist., second ser. 2: 217-223.
1935. Notices helminthologiques. Bull. Soc.
Zool. France, 60: 482-501.
1936. Faune de France 30. Cestodes. Paris,
613 pp.
Mayhew, R. L.
1925. Studies on the avian species of the
cestode family Hymenolepididae. III.
Biol. Monogr., 1 0 : 7-125.
.
.
.
-
.
.
*
....
■
Rausch & Schiller: North American Cestodes of the Genus Paruterina
5
2.
A Contribution to the Study of North American Cestodes
of the Genus Paruterina Fuhrmann, 19061.
Robert Rausch2 and Everett Schiller.
Department of Veterinary Science, University of Wisconsin, Madison 3
(Text-figures 1-12).
Two of the 16 apparently valid species of
Paruterina Fuhrmann, 1906, have been re-
corded from North American birds. Par-
uterina similis (Ransom, 1909) occurs in the
yellow-billed cuckoo, Coccyzus a. americanus
(L.), and P. candelabraria (Goeze, 1782) is
the most frequently encountered cestode
parasitic in owls. The latter occurs in Europe
as well as in North America, and infects sev-
eral species of owls (Wolffhiigel, 1900;
Rausch, 1948). Evidence to the present
would indicate that a high degree of host
specificity has been developed in the cestodes
of this genus.
It is the purpose of this paper to describe
two species of Paruterina, and to include
some remarks concerning the two previously-
recorded Noi’th American species. The un-
described species were collected by one of
us (R. R.) from birds in the Jackson Hole
region of Wyoming. Both were taken from
hosts whose parasites probably have not
been previously studied.
The Wyoming birds parasitized by ces-
todes of the genus Paruterina were a rock
wren, Salpinctes o. obsoletus (Say), and a
green-tailed towhee, Chlorura chlorura (Au-
dubon), which were collected from the same
area, along with numerous birds of other
species. The wren was collected from the
southeast slope of a hill, at an altitude of
about 7,000 feet. Sandstone outcroppings
were numerous here, and rock wrens were
rather commonly observed among them. A
marmot, Marmota flaviventris nosophora
Howell, was the characteristic mammal of
this zone. The towhees were common a few
hundred feet lower, where a sage, Artemesia
tridentata Nutt., was the characteristic
plant. Brewer’s sparrow, Spizella b. breweri
Cassin, was also characteristic of this habi-
tat.
In view of the fact that representatives
of the genus Paruterina have not been often
reported in North America, it seemed un-
1 Contribution of the 1948 Research Program of the New
York Zoological Society at Jackson Hole Wildlife Park.
- Now with U. S. Public Health Service, Anchorage,
Alaska.
3 Parasitology Section, B. B. Morgan, Project Leader.
usual to collect two undescribed species from
so small an area. However, since the mor-
phological differences are quite distinct,
there can be no doubt as to their specific
validity. As pointed out earlier (Rausch,
1948), the helminth parasites of the North
American avifauna are only poorly known.
Paruterina ch lorurae n. sp.
(Text-figs. 1-4).
Diagnosis: Strobila from 35 to 50 mm.
long; greatest width, attained in terminal
gravid segments, about 1 mm. Strobila con-
sists of about 140 segments; margins of
latter not serrate. Segments wider than long
in mature segments, with a gradual increase
in length as segments become older; terminal
gravid segments, in well relaxed strobilae,
considerably longer than wide. Scolex about
550 g in diameter, not set off from neck.
Suckers rather weakly developed, about 180
g in diameter. Rostellum armed with a dou-
ble row of hooks, from 40 to 42 in number;
large hooks 20 g long, and small hooks 16 g
long. Handle of larger hook about same
length as guard and blade; guard of smaller
hook relatively shorter, with blade and guard
of nearly equal length.
Ventral longitudinal excretory canals
measure about 33 g in diameter; dorsal
canals about 10 g in diameter, median to
ventral canals. Transverse canals about 3 g
in diameter. Genital pores irregularly alter-
nate ; genital ducts pass between longitudinal
excretory canals. Musculature well devel-
oped; longitudinal bundles numerous.
Cirrus sac anterior to vagina, from 105
to 119 g long by 23 to 29 g wide. Cirrus sac
does not extend to level of ventral longi-
tudinal excretory canal. Internal and ex-
ternal seminal vesicles absent. Vas deferens
well developed and strongly coiled in area
between poral ventral excretory canal and
ovary. Testes spherical, from 10 to 12 in
number; about 50 g in diameter in mature
segments. Testes lateral and posterior to
female genital organs, not extending anterior
to vagina on poral side, nor antei'ior to ovary
on aporal side.
6
Zoologica: New York Zoological Society
[34: 2
1949]
Rausch & Schiller: North American Cestodes of the Genus Paruterina
.7
Vagina runs directly from genital pore
toward ovary; poral to latter it enlarges to
form a well-developed seminal receptacle.
Ovary slightly lobed, about 50 by 60 y in
mature segments; situated on mid-line near
middle of segment. Vitelline gland spherical
to ellipsoidal, about 20 y in diameter; situ-
ated at posterior margin of segment on mid-
line, directly posterior to ovary. Uterus
appears as a crescent-shaped organ ventral
to ovary; the arms lengthen until the organ
assumes an inverted V-shape. In terminal
gravid segments, arms of uterus become
somewhat sinuous. Parauterine organ de-
velops slowly from anterior margin of early
uterus; it becomes elongate and finally at-
tains anterior margin of segment. Spherical
eggs, observed only in the uterus, measure
from 43 to 50 y in diameter.
Host : Chlorura chlorura (Audubon).
(Green-tailed towhee).
Locality : Near Moran, Wyoming.
Habitat-. Small intestine.
Type : Cotype material has been deposited
in the Helminthological Collection of the
U. S. National Museum, slide number 46421.
Paruterina chlorurae is differentiated
from the other species of the genus by shape,
size and number of rostellar hooks. Differ-
entiation of this species is considered more
fully under the discussion below.
Paruterina morgani n. sp.
(Text-figs. 5-8).
Diagnosis: Strobila about 40 mm. long;
maximum width, attained in gravid seg-
ments, about 500 y. Strobila consists of
about 150 segments; margins of latter
strongly serrate. Mature segments wider
than long ; they increase gradually in length
as they become older, with gravid segments
being slightly longer than wide. Scolex about
250 y wide, distinctly set off from neck;
suckers about 100 y in diameter. Rostellum
armed with a double row of 34 to 36 hooks ;
large hooks measure 66 y long; short hooks
measure 40 y long. Blade of large hook,
slightly longer than handle, curves down-
ward abruptly at end; guard, near middle
of hook, inconspicuous. Blade and handle of
small hook nearly equal in length; guard, at
middle hook, about Yz as long as blade.
Ventral longitudinal excretory canals
about 13 y in diameter; dorsal canals about
4 y. Transverse canals about 3 y in diam-
eter. Genital pores irregularly alternate;
genital ducts pass between longitudinal ex-
cretory canals. Musculature well developed ;
two rows of longitudinal and a single row
of transverse fibers occur in close contact.
Longitudinal muscle fiber bundles not numer-
ous; accurate count not obtained.
Cirrus sac clavate, anterior to vagina; it
extends beyond poral ventral excretory
canal, and measures from 86 to 105 y long
by 16 to 20 y wide. Internal and external
seminal vesicles absent. Vas deferens well
developed and strongly coiled; convolutions
fill greater part of poral half of segment,
from end of cirrus sac to level of mid-line of
ovary. Testes spherical, from 15 to 18 in
number; about 50 y in diameter in mature
segments. Testes lateral and posterior to
female genital organs; not extending an-
terior to vagina on poral side, nor anterior
to ovary aporally.
Vagina runs directly from genital pore,
without convolution, to form a well-devel-
oped seminal receptacle posterior and poral
to ovary. Ovary unlobed, ellipsoidal; about
120 y long by 80 y wide in mature segments;
situated in posterior half of segment, at
mid-line. Vitelline gland ellipsoidal; about 60
y long, situated directly behind ovary some-
what anterior to posterior margin of seg-
ment. Uterus appears as an elongate organ
lying transversely in the posterior part of the
segment, ventral to ovary. It enlarges gradu-
ally, and finally forms an elongate, irregular
sac, situated at posterior margin oi com-
pletely gravid segments. Parauterine organ
grows gradually from anterior margin of
uterus; it does not reach anterior margin of
segment. Eggs spherical, observed only in
uterus; from 36 to 43 y in diameter.
Host: Salpinctes o. obsoletus (Say).
(Rock wren).
Locality: Near Moran, Wyoming.
Habitat: Small intestine.
Type : Cotype material has been deposited
in the Helminthological Collection of the
U. S. National Museum, slide number 46422.
Paruterina morgani is differentiated from
the other members of the genus by size,
shape and number of hooks, as well as by
other, less obvious details. This cestode is
named in honor of Dr. B. B. Morgan, Depart-
ment of Veterinary Science, University of
Wisconsin.
Discussion.
At least 18 species have been assigned
to the genus Paruterina; of these, 2 species,
P. fuhrmanni Baczynska, 1914, and P. meli-
erax (Woodland, 1929) have been trans-
ferred to other genera. Three of the remain-
ing species, P. angustata Fuhrmann, 1906,
P. guineensis Joyeux and Baer, 1928, and
P. southwelli Hilmy, 1936, have unilateral
genital pores, and are immediately separated
by this character from the species described
in the present paper.
Of the North American species, Paru-
terina similis (Ransom, 1909) has been re-
described by Linton (1927). This species
was placed in the genus Paruterina by Jones
(1929). Certain morphological details of
this species have never been completely de-
scribed; Linton (1927, page 50) stated
“There is a short rostellum surmounted by
a double circle of very short hooks. Their
exact number was not satisfactorily made
out, but there appear to be in the neighbor-
hood of 40 . . .” Jones (1929) examined both
Linton’s material, and that of Ransom, but
did not give further details concerning the
hooks of P. similis. We found that P. similis
possesses from 50 to 52 hooks, arranged in
8
Zoologica: New York Zoological Society
[34: 2: 1949]
a double row. The large hooks measure 13 g
long, while the small hooks measure 11 g
long. They are essentially the same in form,
except that the larger hooks have a larger,
more rounded guard (Text-fig. 12). Our ob-
servations were made under oil immersion
on hooks which had been removed from the
scolices, and were lying flat on the slide.
Paruterina chlorurae and P. morgani are
differentiated from P. similis and P. candel-
abraria by hook size, shape and number
(Text-figs. 2, 7, 10, 12). It is of interest to
note that the North American species can
also be separated by differences in the ar-
rangement of the parauterine organ and the
uterus in the fully-gravid segments (Text-
figs. 4, 8, 9, 11). In fact, differences here
are more obvious than are those seen in the
mature segments. It might also be mentioned
here that cestodes of this genus can easily
be recognized as such macroscopically, at
the time they are removed from the intestine
of the host, by the appearance of the gravid
segments.
The remaining 11 species, widely dis-
tributed geographically, are best separated
by hook characters. All of these (P. bovieni
Iliibscher, 1937; P. bucerotina Fuhrmann,
1909; P. cholodkowskii Skrjabin, 1914; P.
daouensis Joyeux, Baer, and Martin, 1936;
P. javanica Htibscher, 1937; P. meggitti
Johri, 1931; P. otidis Baczynska, 1914; P.
parallelipida (Rud. 1809) ; P. purpurata
(Dujardin, 1845) ; P. septotesticulata Moghe
and Inamdar, 1934; P. vesiculigera (Krabbe,
1882) all differ appreciably in hook size,
shape and number.
There are also differences in testes number
in most cases. Where there is overlapping
of this character, hook differences serve to
separate the species involved. Other taxon-
omic details need not be discussed here in
order to separate the species described in
the present paper.
References.
Baczynska, H.
1914. fitudes anatomiques et histologiques
sur quelques nouvelles especes de ces-
todes d’oiseaux. Bull. Soc. Neuch&t.
Sci. Nat., .40:187-239.
Fuhrmann, 0.
1906. Die Tanien der Raubvogel. Centralbl.
Bakt. Parasit. (orig.), -41:212-213.
Johri, L. N.
1931. A new cestode from the grey hornbill
in India. Ann. Mag. Nat. Hist., 8, ser.
10, pp. 239-242.
Jones, M. F.
1929. Tapeworms of the genera Rhabdo-
metra and Paruterina found in the
quail and yellow-billed cuckoo. Proc.
U. S. Nat. Mus., 75, Art. 20, pp. 1-6.
Linton, E.
1927. Notes on cestode parasites of birds.
Proc. U. S. Nat. Mus., 70, Art. 7, pp.
1-73.
Moghe, M. A. and Inamdar, N. B.
1934. Some new species of avian cestodes
from India with a description of
Biuterina intricata (Krabbe, 1882).
Rec. Ind. Mus., 36: 7-16.
Rausch, R.
1948. Observations on cestodes in North
American owls, with the description of
Choanotaenia speotytonis n. sp. (Ces-
toda: Dipylidiinae) . Amer. Midi. Nat.
40 (2) : 462-471.
1949. Paradilepis simoni n. sp., a cestode
parasitic in the osprey. Zoologica, 5-4
(1) : 1-3.
Skrjabin, K. I.
1914. Vogelcestoden aus Russisch Turkestan.
Zool. Jahrb. (syst.) 57:411-492.
W OLFFHUGEL, K.
1900. Beitrag zur Kenntnis der Vogelhel- ,
minthen. Inaug. Diss. Basel. 204 pp.
Katz: Behavioral Interactions in Barbary Sheep
9
3.
Behavioral Interactions in a Herd of Barbary Sheep
( Ammotragus lervia).1
Irwin Katz.
The University of Buffalo.
Introduction.
Studies of social behavior in animals have
generally been of two types, the naturalistic
field investigation and the laboratory experi-
ment. Field studies of ungulates have been
made by Darling (5) on the red deer, and
Mills (10), Davis (6) and Spencer (13) on
the Rocky Mountain bighorn sheep. The ex-
perimental method has produced an exten-
sive literature on dominance relationships
and aggressive behavior in many species.
Collias (4) has reviewed the work on aggres-
sive behavior among vertebrates up to 1944.
Studies on dominance have been too numer-
ous even to be mentioned briefly in the pres-
ent paper.
Carpenter (2) has pointed out that the
development of a science of comparative so-
cial behavior requires that the results of field
investigations and those of the laboratory
should be systematically co-related. He also
has stated that the standards of scientific
research which apply in the laboratory can
and should be applied in the field. Recogniz-
ing the research potentialities of an inte-
grated approach to animal behavior, Scott
(11) recently combined systematic observa-
tion and experimentation in a study of a
small flock of domestic sheep living under
semi-natural conditions.
The methods and aims of the following
study of a herd of Barbary sheep were sug-
gested in large part by the work of Scott
and the theoretical discussions of Carpenter
(2, 3). Carpenter (2) has listed 11 types of
behavioral interactions found in primate
societies. It was hoped that the first six of
these might be studied in the Barbary sheep.
They are: 1, Interactions among adult males
of organized groups ; 2, among adult females
of organized groups ; 3, between adult males
and adult females; 4, between adult males
and young; 5, between adult females and
1 A report submitted to the New York Zoological Society
on research performed as a Summer Research Fellow of
the Society during July, August and September, 1947.
The writer is indebted to his colleagues. Dr. N. E.
Collias and Dr. B. F. Riess, who made many valuable sug-
gestions and participated in the observation and experi-
mentation from time to time. Special gratitude is due a
third associate, Mr. D. Lehrman, who did most of the work
with the Bristol Recorder, and who shared equally in some
of the experiments. The Summer Research Fellows worked
under the general direction of Professor C. R. Carpenter.
young; and 6, among the young. The data
were to be compared with information on the
domestic sheep and the Rocky Mountain big-
horn. In addition, the investigator sought to
obtain data relevant to the hypothesis that
deprivation is an effective instigator of ag-
gressive behavior.
The Herd.
The Barbary sheep, or aoudad ( Ammotra -
gus lervia ) is very distinct in appearance
from all other wild sheep, its most unique
features being a mane of long hairs over the
fore-quarters, the length of tail and the large
size of the female’s horns. Its color is uni-
form rufous tawny. The habitat of the
Barbary sheep is the arid southern slopes
and foothills of the mountains of North
Africa, extending from near the Atlantic
seaboard to Egypt. Lydekker (8) quotes re-
ports that the animals go about in groups of
four or five and may drink as seldom as once
in four or five days.
The herd at the New York Zoological Park
is descended from stock brought to the Park
during the years 1901-1906. No new stock
has been introduced since then. During the
summer of 1947 the herd consisted of four
rams, four ewes and four lambs. One of the
lambs was a yearling, while the others were
first-season. The sheep lived on an enclosed
field of about two acres. They shared the field
with two elands and a zebra. Human regula-
tion of the activity of the sheep has been kept
at a minimum by the Park authorities. Under
normal circumstances the herd is fed about
one and one-half buckets of grain, which is
spread out on a large, flat rock at about 9 :30
every morning. The elands and the zebra
usually feed from a box some distance away,
although they sometimes wander over to the
rock and feed with the sheep. The grain
ration is supplemented occasionally with hay,
which is placed in one corner of the field.
Vending machines in the Park provide spe-
cial food pellets which visitors may throw
through the fence. During the warm months
the sheep regularly gather at the north fence
in the afternoon to receive these pellets. The
sheep tend generally to avoid the two elands
and the zebra. In recent years a newborn
lamb was killed by the male eland, and
another by the zebra.
10
Zoologica: New York Zoological Society
[34: 3
Methods.
Casual observations were made almost
daily from mid-July until mid-September,
and during all hours of the day, so that a
complete picture of the daily routine could
be obtained. Colored dyes were used to mark
the individual animals until the observer
could recognize them easily by differences in
appearance and behavior. The following ex-
periments were performed repeatedly: (1)
tossing of bread between pairs to ascertain
relationships of dominance-subordination;
(2) placing of daily grain ration in a small
box to study dominance-subordination rela-
tionships in a complex herd situation, as well
as to provide observations on food sharing,
fighting, and related phenomena; (3) fright-
ening of the herd to elicit leadership and
timidity. These experiments were carried out
from July 22 to September 13. In addition,
on two days a Bristol multi-pen recorder was
employed in connection with the feed box
experiment to ascertain the amount of time
each animal actually fed from the box. Mo-
tion pictures were taken of types of behavior
which had been previously recognized and
described.
The study of social relationships was
limited by the fact that the age and par-
entage of adult individuals could not be
established with certainty. Although the
Park maintains complete records of births
and deaths, there is no provision made for
identifying individual members of herds. It
was possible, however, to know three ewe-
lamb relationships on the basis of observed
behavior.
Daily Routine.
The daily pattern of behavior of the herd
was marked by fairly regular periods of
alternating activity and rest. But this was
greatly modified by changes in the weather,
experimentation and marked variations in
the supply of food from visitors. Usually in
the early morning the sheep wandered about
the field. Grazing was desultory, since other
food was available. At about 9:00 A.M. the
sheep generally gathered on an outcropping
of broad, flat rocks situated on a hillock in
the center of the field, and there they rested
until 9:30 A.M., when grain was scattered
on a nearby rock by the keeper. The sheep
ate the grain peacefully, with very little
butting or shoving. By 10:15 A.M. the sheep
were either back at their earlier places on the
rocks or were under a tree, where they re-
mained until after the noon hour.
Shortly afterwards children and adults
would usually begin to gather along the north
fence. The movement of the sheep to the
fence seemed to be associated with the size
of the gathering of people rather than the
hour. On days when very few people visited
the Park the sheep might remain entirely
away from the fence throughout the after-
noon. Once at the fence, the herd remained
there as long as pellets were given to them,
usually until about 5:30 P.M. On hot after-
noons the males made occasional trips to a
nearby water hole. Here they cooled them-
selves by sinking down into the shallow water
and rolling in the mud.
In the evening the sheep wandered and
rested until dark. The lambs played actively
at this time by running and leaping on the
rocks. During late August and September
fighting and attempted breeding occurred
among the males, and most frequently in the
evening. At dusk the herd gathered inside
or near a shed and bedded down for the night.
Sometimes the sheep moved as a group, but
consistent leadership was not apparent. In
general, there was much independent move-
ment among the ewes, rams and lambs.
Matching Tests.
The matching tests were conducted every
few days from July 22 until September 13
to determine dominance-subordination inter-
actions between individual animals. Usually
the tests were made in the afternoon, when
the sheep were gathered at the north fence.
By supplying several willing children with
bread, and placing them along the fence, it
was possible to disperse the sheep so that
all or most of the possible pairings could be
made among the rams, the ewes and the
lambs. The matching of adults and lambs,
or of rams and ewes, was not attempted after
the first day because of practical difficulties.
The matching test was simple. The ex-
perimenter stood at the fence and held a
small piece of bread in his extended hand.
When two sheep, which were not more than
ten feet apart, looked in the direction of the
experimenter, the bread was tossed so that
it landed approximately equidistant between
them. No score was recorded unless both
animals moved toward the bread. The one
which obtained possession of the bread by
causing the other to withdraw was consid-
ered dominant. The behavior elicited in this
situation was clear and unambiguous ; if both
sheep advanced toward the food, one always
threatened or butted and the other always
withdrew. Sometimes the bread landed much
closer to the animal known to be subordinate.
In such a case the subordinate sheep might
obtain the bread, but this was usually fol-
lowed by vigorous butts from the dominant
animal. Often, however, a quick dash by the
dominant sheep caused the other to retreat,
even when the bread lay directly at its feet.
On three occasions the sheep appeared to
be uniformly unmotivated with regard to the
bread. Two of these days were extremely
hot, and the third was marked by a morning
of heavy feeding. At all other times compe-
tition was keen and sustained. The method
of scoring is somewhat defective in that no
scores were recorded for those tests in which
only one animal moved toward the food. The
assumption here is that of “no contest” and
this is, of course, questionable, since the ani-
mal’s lack of a positive overt response to the
food might be due to the presence of the
dominant animal. However, the almost per-
1949]
Katz: Behavioral Interactions in Barbary Sheep
11
feet consistency of the results presented in
Table I, and the agreement between these re-
sults and behavior observed in other situa-
tions strongly suggest that the method is
highly valid.
In the majority of contests, dominance was
decided by a sudden tivisting movement of
the dominant sheep’s head in the direction
of the other sheep. At this “signal” (or sign)
the subordinate sheep stopped advancing.
Sometimes a token butt was delivered, but
seldom was a more forceful attack necessary
to effect retreat. Counter attacks by subordi-
nates occurred rarely and were never suc-
cessful. Among the rams, the ewes and the
lambs straight line dominance orders were
revealed on the first day and remained al-
most stable during the 53-day period of test-
ing. Only two instances of reversals occurred
during a total of 272 matching tests. Al-
though interactions between rams and ewes,
and between adults and lambs, were not for-
mally tested, it was apparent that all rams
were dominant over all ewes, and all adults
over all lambs.
The results of the matching tests are pre-
sented in Table I. The dominance order is as
follows: Ram 1> Ram 2> Ram 3> Ram 4>
Ewe 1> Ewe 2> Ewe 3> Ewe 4> Lamb 1>
Lamb 2> Lamb 3> Lamb 4>. The attempt
was to test at least twice a week every pos-
sible combination of individuals within each
of the three subgroups. But this could not
always be done because of the difficulty of
bringing certain of the sheep together. For
example, matchings between lambs were
often disrupted by the sudden approach of
one or more adults.
Feed Box Experiments.
The feed box experiments were intended
to furnish information on social behavior in
a competitive group situation. The matching
tests had indicated the existence of a clear,
stable relationship between any two animals
which were made to compete for a small
food object while in relative isolation from
the other members of the herd. But it could
not be assumed that these relationships
would hold in all types of competitive situa-
tions, especially in those where more than
two animals are interactive. Maslow (9)
found that stable dominance-submission re-
lationships which were established between
monkeys by the method of paired matching
tests broke down when three or more indi-
viduals were placed together.
The food incentive box was heavy and
made of wood, typical of those used in the
Park for the feeding of large animals. Its
sides were about two feet long and about
one and one-half feet high. The box was
modified so that the interior sides measured
15" by 12", with the depth remaining un-
altered. The interior was large enough to
hold more than a bucket of grain without
spilling by the feeding animals. The size of
the opening was such that two adults could
not feed simultaneously without frequent
contact, while simultaneous feeding by three
Table I.
Results of the Matching Tests.
Rams*
Number
of matchings
1-2
25
1-3
20
1-4
22
2-3
22
2-4
21
3-4
12
4-2f
1
2-lf
1
Eives*
1-2
22
1-3
18
1-4
15
2-3
15
2-4
13
3-4
9
Lambs*
1-2
7
1-3
6
1-4
5
2-3
12
2-4
10
3-4
14
Groups
Total
Rams
124
Ewes
94
Lambs
54
* Numbers indicate sheep according to position in domi-
nance order. Number of dominant animal precedes that
of subordinate,
t Reversal.
adults would result in almost constant con-
tacts. It was hoped that food sharing, and
the conditions surrounding this behavior
might result as well as competition for food.
The feed box experiment was conducted
12 times. On mornings when the experiments
were performed, the experimenter moved the
elands and the zebra from the field to adja-
cent pastures. At about 9:30 or 10:00 the
box was placed on the rock where grain nor-
mally was scattered by the keeper. Then the
experimenter emptied one bucket of grain
into the box and withdrew behind a gate
about 20 yards away. The sheep were ob-
served by means of binoculars and their be-
havior was recorded immediately in a note
book. Usually at the end of an hour it was
necessary to place more grain in the box.
The general pattern of social interaction
at the feed box was similar throughout the
entire series of experiments. During the se-
ries of group tests an order of dominance
was formed which conformed closely to that
observed during the matching tests. Rams
1 and 2 always dominated the other animals
at the box during the first 15 or 20 minutes
of feeding. The other sheep milled around
the feed box but were not permitted to eat.
The two dominant rams ate alternately.
Whenever Ram 1 raised his head to chew or
rest, Ram 2 ate from the box. As Ram 1 again
lowered his head, Ram 2 usually withdrew
his head. A high degree of orderliness usu-
ally characterized the feeding of these two
sheep. Often Ram 2 did not withdraw until
12
Zoologica: New York Zoological Society
[34: 3
ha was threatened or mildly butted by Ram
1. In the main, Ram 1 butted and shoved
Ram 2, and the latter in turn kept the other
sheep from the box. Upon being forced from
the box by the more dominant animal, Ram
2 might circle the box and butt all the sheep
in his path.2 After the first few minutes the
other animals ceased to crowd around the
box. Some of them moved to a nearby tree
and others formed a wide circle about the
feeding place.
When he had completed his first feeding,
Ram 1 left the feeding area. Ram 2 would
either leave at the same time, or continue
feeding. Then Ram 3, or Rams 3 and 4 to-
gether, moved up to the box, and interactions
very similar to the previous ones were ex-
hibited. During the first 30 or 40 minutes
the box was controlled constantly by a pair
of rams. But after the initial feeding of Rams
1 and 2 the pairings shifted frequently due
to the movements to and from the box of
dominant rams. From time to time ewes and
lambs attempted to feed, usually with little
success. The subordinate ram of a pair did
most of the butting and chasing of the other
members of the herd.
During the second hour the rams spent
less time at the box, and when there mani-
fested increasing tolerance toward the lambs
and ewes. The order of feeding among the
ewes was also determined mainly by domi-
nance status, while a lamb’s ability to feed
depended on the tolerance of its own ewe.
Often Ewe 4 and Lamb 1 (the yearling) ob-
tained little or no food during an entire ex-
periment. Being of low dominance status,
the ewe was excluded, and the lamb likewise
because it lacked high dominance maternal
protection. The experiment usually ended
shortly after the noon hour, when the sheep
began to move toward the north fence for
pellets offered by visitors.
Dominance. In Table II are presented the
butts and threats given and received by each
sheep during the series of 12 feed box ex-
periments.3 Except for two instances, threats
were always directed by dominant animals
against subordinate ones. The butt more
frequently was directed by a subordinate
sheep against a dominant one. Nineteen
butts, of a total of 198, fall in this category.
The data on rams in Table II indicate that
the dominance-subordination relationships
among these animals were somewhat less
rigid and involved more behavioral inter-
action among individuals than in the match-
ing tests. However, it must not be assumed
that relationships at the feed box were less
stable. Stability cannot be inferred from the
ratio of butts given and received. Nor would
2 These attacks by Ram 2 against subordinate sheep
appeared to be clear instances of displaced aggression,
and will be discussed later on in this paper.
3 A threat is defined as an aggressive movement or pat-
tern of movements which one sheep directs at another,
but which does not end in physical contact. The typical
threat consisted of a sudden lowering of the head and
slight movement toward the other animal. But sometimes
a mere lowering and twisting of the head composed the
pattern. The object of a threat usually withdrew imme-
diately or modified his behavior in some observable way.
a mere tabulation of instances of food shar-
ing and food hoarding provide a valid basis
for inferring dominance status. For example,
Rams 1 and 2 usually ate together with little
overt indication of dominance-subordination.
Often they fed alternately for three or four
minutes without observable conflict. But
upon closer examination it could be seen that
Ram 1 'permitted, Ram 2 to eat with him, and
even to shove him occasionally. Over-vigor-
ous shoving or persistent crowding on the
part of Ram 2 always elicited a sharp attack
from Ram 1. Exchanges of butts might be
equal in number but they always ended with
Ram 1 in control of the box.
Wide variations in the “social distance”
between different rams are apparent. Reci-
procity of aggression was relatively high
between Rams 1 and 2 and between Rams 2
and 4, but Ram 3 never aggressed against
Ram 1 or Ram 2, and was almost never ag-
gressed against by Ram 4.4 The meaning of
these differences in “social distance” will
become clearer at a later point in the dis-
cussion. Among the sheep included in Table
II, frequencies of butts and threats decrease
in almost perfect rank order. The data on
ewes and lambs has not been analyzed in
detail because of the low frequency of ag-
gressions. Lambs 2, 3 and 4 are not included
in the Table only because their scores on
both items were zero.
Feeding time. Scores for feeding time
were computed from the Bristol recordings
of an experiment on August 11 according to
the method described in the first footnote to
Table III. The time score for each sheep is
the number of 10-second periods during
which the animal had its head in the box for
two seconds or longer. In Table III individual
scores are given for each of nine consecutive
periods. The periods are 16.6 minutes in
length. Individual totals for the whole experi-
ment indicate a lack of positive relationship
between dominance status and feeding time,
although the differences between rams and
ewes, and adults and lambs, are on the whole
substantial. The lack of relation between
feeding time and dominance status may be
due to wide individual differences both in
rate of food intake and nutritional needs.
For most of the sheep there is a single
period during which much more feeding oc-
curred than during other periods. This would
seem to justify a comparison of the periods
in which individuals made their highest
scores. Such scores have been indicated in
the Table by a small circle. The circles follow
a line which gradually descends from left
to right, indicating that time-of-maximum-
feeding is closely related to the dominance
order. The near-zero scores of Ewe 4 and
Lamb 1 have already been discussed.
Coordinate-feeding. An important type of
social interaction among the sheep was that
of coordinate-feeding, or food sharing. The
concept and the unit of measurement em-
ployed are described in the first footnote to
•* See footnote to Table II.
1949]
Katz : Behavioral Interactions in Barbary Sheep
13
Table II.
Total Butts and Threats for 12 Feed Box Experiments*.
Threats
Butts
Ram
#1
Ram
#2
Recipients
Ram
#3
Ram
#4
Ewes &
Lambs Total
Ram
#1
Ram
#2
Recipients
Ram
#3
Ram
#4
Ewes &
Lambs Total
Ram #1
X
21
1
2
21
45
X
42
5
5
15
67
Ram #2
0
X
4
2
16
22
9
X
2
13
16
40
Ram #3
0
1
X
8
11
20
0
0
X
14
19
33
Ram #4
0
0
1
X
18
19
0
8
2
X
15
25
Ewe #1
0
0
0
0
5
5
0
0
0
0
17
17
Ewe #2
0
0
0
0
6
6
0
0
0
0
12
12
Ewe #3
0
0
0
0
0
0
0
0
0
0
8
8
Ewe #4
0
0
0
0
0
0
0
0
0
0
3
3
Lamb #1
0
0
0
0
0
0
0
0
0
0
4
4
* Intermittent fighting was observed between Ram 2
and Ram 4. and between Ram 3 and Ram 4, during the
second week in September. Butts exchanged during these
fights have not been included in the table because on the
two days that these fights occurred the experimenter was
occupied with taking motion pictures. However, only one
Table IV. The unit of measurement is, of
course, arbitrary, but it has the merits of
being easily computed from the raw data
and of yielding quantitative relationships
which are in close agreement with observa-
tions made during 12 experiments. Table IV
contains data on coordinate-feeding among
the rams. Both absolute frequencies and co-
ordinate-feeding ratios indicate that among
those pairings in which the possibility of co-
ordinate-feeding existed relatively often its
frequency varied widely. The ratios are high
for Rams 1 and 2 and Rams 2 and 4, and low
for Rams 2 and 3 and Rams 3 and 4. Parallel
differences among pairs have already been
noted with regard to butts and threats. Ag-
gressive interactions and food sharing vary
together. But although they are associated,
it would be wrong to assume that one is a
primary cause of the other. Rather, both are
directly related to the amount of social dis-
tance between individuals. Social distance is
psychological rather than spatial, and is de-
fight might be said to have ended in favor of Ram 4. This
fight was between Ram 4 and Ram 2. Since these fights
were associated with sexual excitement in Ram 4 the table
as presented is representative of social relations before the
onset of rutting behavior. Fighting and rutting behavior
will be discussed in a later section.
termined by the willingness-for-contact, or
tolerance, of the dominant sheep with regard
to the subordinate. Lack of unilateral or bi-
lateral aggression between two sheep may be
an indication of very low tolerance on the
part of the dominant animal. Thus, on the
day for which coordinate-feeding data is pre-
sented, Rams 3 and 4 were the only rams to-
gether at the box for a longer time than
Rams 1 and 2 were present togther, yet the
only aggression between the former individ-
uals was a single threat by Ram 3, and the
coordinate-feeding ratio was 4/25. In con-
trast, Ram 1 threatened or butted Ram 2 six-
teen times and received three butts from the
latter, while the coordinate-feeding ratio
was 16/21.
Quantitative data on coordinate-feeding
among the ewes was not obtained because
during the experiment on August 11 one or
more rams were almost always present at the
box. The recorded observations of all 12 ex-
periments show that when no ram was pres-
Table III.
Feeding Time Scores of Sheep in Experiment Lasting Two and One-half Hours.*
Periods!
Ram
#1
Ram
#2
Ram
#3
Ram
#4
Ewe
#1
Ewe
#2
Ewe
#3
Ewe Lamb
#4 #1
Lamb
#2
Lamb
#3
Lamb
#4
I
87°
82°
2
3
1
5
0
3
0
0
0
0
II
27
18
54°
8
6
3
0
0
0
0
0
0
III
0
13
28
18
26°
15
14
3
0
9
5
8
IV
10
26
6
39°
7
0
11
0
0
18
0
0
V
0
0
15
12
0
7
7
0
0
11
0
0
V I
9
9
28
25
14
22°
13
0
0
21
4
0
VII
12
0
5
15
12
0
35°
0
0
36°
0
11
VIII
0
28
1
28
0
16
0
0
0
8
11°
0
IX
0
43
19
19
15
7
3
0
2
1
6
14°
Totals
145
219
158
167
81
75
83
6
2
104
26
33
* This was the only occasion upon which a Bristol re-
cording was made for an entire experiment. The experi-
ment was conducted on August 11, and was the sixth in
the series of 12. Time scores were computed from the
recorded data by counting for each sheep the number of
10-second periods during which the animal had its head
in the box for two seconds or longer. This method of
scoring, while less accurate than the very laborious pro-
cedure of counting actual time in seconds, does not intro-
duce a serious bias, in the opinion of the experimenter.
t The total time of two and one-half hours was divided
into nine periods of 16.6 minutes each.
° Maximum score for a single period.
14
[34: 3
Zoologica: New York Zoological Society
Table IV.
Incidence of Coordinate-feeding Between Rams During Experiment Lasting Two
and One-half Hours.*
Pairings
Frequency of
coord-feeding
Highest frequency
possiblef
Coord-feeding
ratio
Rams 1 and 2
16
21
16/21
Rams 1 and 3
1
5
1/5
Rams 1 and 4
1
4
1/4
Rams 2 and 3
3
14
3/14
Rams 2 and 4
8
11
8/11
Rams 3 and 4
4
25
4/25
* Coordinate-feeding between two rams is said to occur
when both of these rams obtain feeding scores of 3 or
more during a 100-second period, while the other two rams
obtain scores of zero (c.f. first footnote to Table III for
unit of scoring) . The coordinate-feeding data were obtained
from the Bristol recordings and synchronized field notes
of the feed box experiment on August 11.
t The highest frequency possible is the total number of
100-second periods during which a given ram could have
engaged in coordinate-feeding with the dominant ram by
virtue of the fact that the dominant ram was (a) the only
other ram at the box, or (b) accompanied at the box by a
ram subordinate to the ram in question.
ent Ewes 1 and 2 dominated other animals at
the box. Food sharing was common among
Ewes 1, 2 and 3. The lambs almost never had
exclusive possession of the box, and when at
the box they showed no overt aggression.5 6
The amount of time that they fed depended
on the tolerance of the lambs by the ewes.
Hence, differences in the ability of individ-
ual lambs to feed are related to differences
in ewe-lamb relationships. Field observa-
tions furnished ample evidence of the follow-
ing mother-young relationships: Ewe 3 and
Lamb 2, Ewe 1 and Lamb 3, and Ewe 2 and
Lamb 4.G The yearling was not associated
with a ewe in any observable way, either at
the box or in the field.
There is reason to believe that the very
high feeding score of Lamb 2 was due to an
especially close and permissive relationship
with its mother. Lamb 2 usually stood very
close to its own ewe at the box, and fed when-
ever she did. Ewe 3 never butted her lamb,
although she did not tolerate other lambs.
Lambs 3 and 4, on the other hand, did not
stay close to their ewes, and often were not
at the box when their ewes were feeding. It
will be seen in Table III that the feeding
scores of Ewe 3 and Lamb 2 are very similar
from period to period. The average difference
in scores for the same period is only 2.8. Anal-
ysis of the Bristol recordings reveals that, of
a total of 20 100-second periods during which
Ewe 3 fed, her lamb also fed during 17 peri-
ods. In contrast with this, Ewe 1 and Ewe 2
fed about as frequently as Ewe 3, but Ewe 1
shared with her lamb only twice, and Ewe 2
never shared with hers.
The lack of agreement between the domi-
nance status of ewes and their lambs may
have been noted. Dominance order among the
first-season lambs shows no relation to dom-
inance order among the mothers. Of course
the number of sheep is much too small to war-
rant generalizing, but there are two possible
correlates of dominance order among first-
season lambs that might be mentioned. First,
5 The lambs often shoved each other, but never threatened
or butted when at the box.
6 Sucking and following were the principal behavioral
indications of mother-young relationships during July,
August, and September.
it is possible that dominance among these
lambs is related to order of birth, so that
lambs born in February tend to be dominant
over those born in March or later. There is no
way of checking this hypothesis in the pre-
sent study, since the birth dates of individu-
als are not known. Stewart and Scott (14)
have found that age is favorable to domin-
ance in a herd of goats.
A second hypothesis is that the amount of
social distance between a ewe and her lamb
will have a direct bearing on the dominance
status of the lamb. In the case of Ewe 3 and
Lamb 2 extreme social closeness is associated
with dominance of this lamb over two other
lambs born in the same season. On the other
hand, the relationship between Ewe 2 and
Lamb 4 (the lamb of lowest dominance sta-
tus) was the weakest of the three mother-
young relationships. Ewe 2 was the least will-
ing to share food with her lamb or to be
sucked. Further credence is given to this sug-
gestion by Scott’s (11) observations of two
orphan lambs which were placed with a flock
of domestic sheep. He noted that, “Both or-
phans appeared to show less fighting than the
other sheep . . . and the ram was not aggres-
sive toward other males even in the breeding
season.” The hypothesis could be tested in
a large herd by testing dominance inter-
actions among the ewes and among the lambs
over a period of time beginning shortly after
the birth of the lambs, and making frequent
observations of each ewe with her lamb in
isolation from the other sheep.
Leadership
Recent studies indicate that leadership
may be a behavioral characteristic quite un-
related to dominance status maintained by
fighting. The reports of Darling (5) on red
deer, Mills (10) and Davis (6) on Rocky
Mountain bighorn sheep, and Scott (11) on
domestic sheep, all mention that the usual
leader in a herd is an old female. Lack of cor-
relation between leadership and dominance
interactions has been noted by Allee et al (1)
in a flock of ducks, and Stewart and Scott
(14) in a herd of goats.
In the present study, clear instances of
1949]
Katz: Behavioral Interactions in Barbary Sheep
15
leadership occurred only when the sheep were
in a conflict situation involving both a source
of attraction and a source of danger in close
proximity to each other. The source of attrac-
tion was the feed box; danger usually was
represented by the presence of a strange per-
son, such as the experimenter. The experi-
menter discovered that if he stood a few
yards behind the box after filling it, the
sheep would flock into the center of the field
and not advance to feed for several minutes.
Finally, Ewe 1 slowly moved forward about
ten yards and then halted. Her lamb imme-
diately ran to her. Then the other ewes and
lambs, and finally the rams, moved up to her
advanced position. If the experimenter with-
drew further from the box the process of ad-
vancing and halting under the leadership of
Ewe 1 might be repeated several times, until
the herd finally reached the box. The pattern
of advance might vary from day to day, but
Ewe 1 always led the others. Sometimes she
advanced 15 or 20 yards in front of the herd
before they followed her. Sometimes the en-
tire herd moved in single file, with Ewe 1 in
the lead and the rams bringing up the rear.
The rams rarely came up to the box until Ewe
1 had begun to eat.
Leadership was observed 15 times, always
in connection with the feed box experiments.
Once the rams dashed ahead of Ewe 1 when
she was about five yards from the box, and
on two occasions Ewe 2 took the lead after
Ewe 1 had led most of the way. But at all
other times Ewe 1 moved in advance of the
others. There is ample evidence that the
sheep were following Ewe 1, rather than just
moving toward the feed box. Seven times
Ewe 1 did not take the most direct path to the
box, but turned and walked at right angles
to it for several yards. When she did this, the
other sheep continued to follow her just as
as though she were approaching the box.
When no person was in the field, the sheep
moved toward the box more or less independ-
ently of each other, but the rams still tended
to stay behind the ewes.
On general grounds it would be expected
that boldness and leadership were related.
In the present study there were no opportu-
nities for observing differences in boldness
among the ewes. However, the rams did ap-
pear to be more timid than the ewes. A lone
ram rarely stayed at the box for more than
a few seconds. In all likelihood, when he
raised his head from the box and saw that
the others had left he would quickly run to
where they were. On the other hand, a single
ewe might continue feeding alone indefi-
nitely. When the zebra, which had been placed
in an adjacent enclosure, suddenly galloped
toward the fence, the rams were the first to
run from the box and the last to return. This
was also true when the experimenter inten-
tionally frightened the herd. The zebra
frightened the sheep away from the box
about eight times, and each time Ewe 1 led
them back. Thus there is considerable evi-
dence that the role of Ewe 1 as leader was
not due merely to a greater familiarity with
humans.
In the field studies cited above, leadership
usually was an important factor in the
normal moving about of the sheep and deer.
But among the Barbary sheep, instances of
leadership were quite l'are outside of the spe-
cial conflict situations described. In wander-
ing and grazing the herd often was scattered
widely over most of the field. The rams, ewes
and lambs often formed into separate and
dispersed sub-groups, yet no consistent lead-
ership was apparent in any one of the sub-
groupings. What little leading and follow-
ing there was occurred between lambs and
ewes, and between rams and ewes with the
onset of sexual activity.
Ewes and lambs. During approximately 80
hours of observation from mid-July to mid-
September, each first-season lamb attempted
to suck its ewe about 20 or 30 times. The
usual duration of sucking was only a few sec-
onds, and often the attempt consisted of a
single, brief thrust at the udder. Generally
the ewe was passive while the attempt was
made. Lamb 4, however, often was rejected
by its ewe, even though it tried to suck less
often than the others. For a while, in fact,
its maternal origin was not clear. The lamb
alternated between following Ewe 2 and Ewe
4, and twice tried to suck from Ewe 4. Re-
peated observations confirmed its relation-
ship with Ewe 2. Sucking was accompanied
by a certain amount of following of ewes by
their lambs. When the herd rested a lamb
often lay beside its ewe. The yearling asso-
ciated wth the other lambs and joined in the
general movements of the herd, but did not
favor a particular ewe.
Rams and ewes. As the rams began to
manifest sexual interest in the ewes they
gradually spent more time in the company
of the ewes and tended to follow them closely
during early morning and evening. Before
August 15 relatively little following occurred.
Perhaps the following of the ewes by the
rams in the conflict situations is, in part, the
result of conditioning which develops during
the rutting season.
Fighting.
The Barbary sheep would seem to fall
about half way between the domestic sheep
and the bighorn with regard to amount of
fighting. Scott (11) mentioned pushing and
shoving among domestic sheep competing for
food in winter, and some butting between
rams following the same ewe in heat. In con-
trast, Mills (10) stated that fights between
big-horn rams in rut might result in “bleed-
ing noses, splintered horn tips, limping, and
skull fractures.” No complete comparison can
be made with the bighorn because the Bar-
bary sheep were not observed during the
height of rutting. There were no observable
injuries, and it is probable that the fighting
witnessed was far less serious than that as-
sociated with breeding.
Two main types of fighting occurred
16
Zoologica: New York Zoological Society
[34: 3
among the Barbary sheep. One type con-
sisted of a series of head-on charges, usually
between rams. The sheep walked away 10 or
15 yards, turned, and walked rapidly toward
each other, gradually picking up speed and
breaking into a run shortly before they col-
lided. Just before impact their heads were
lowered and turned slightly to opposite sides.
They attempted to meet squarely with their
noses crossed. At the beginning of the
charge, the sheep got in step and then tried
to keep in step until they struck. If one got
out of step they broke off the charge and
walked away to charge again. Spencer (13)
described bouts between bighorn rams which
were very similar in detail. He termed such
fights “playful” because one ram would not
attack if the other was off balance or not pre-
pared. A fight of this type between Ram 1
and Ram 2 continued intermittently for 25
minutes on August 25. Before that time
fights had never lasted more than five min-
utes.
The second type of fighting consisted of
close butting, and locking and twisting of
horns. Usually the sheep stood head to head,
facing in the same or in opposite directions,
and engaged their horn tips. Each tried to
twist the head of the other by pulling down-
ward and away. Also, attempts were made
to hook the belly or the flank. Fighting of
this sort might continue for several minutes.
It sometimes started at the feed box as a kind
of maneuvering for position.
Until mid-August fighting was almost as
common among the ewes as among the rams.
The usual fight between ewes consisted of a
brief exchange of butts, perhaps with lock-
ing of horns and twisting. Charges were
very rare. Ewes did not fight with rams.
From mid-August until the termination
of the study fighting among the rams in-
creased in frequency, duration and vigor,
and was connected with sexual excitement.
Rams 2 and 4 were the first to show in-
creased pugnacity and sexual behavior.
These rams fought with each other and
with the other two rams. Sometimes the
penis of a fighting ram emerged briefly from
its sheath. By September 10 all four rams
had reached a high level of sexual arousal
and aggressiveness. Most of the fighting
took place in the early morning and in the
evening, with relatively little aggressive in-
teraction at the feed box or during the
matching tests. Even when Ram 4 was be-
ginning rut and displayed strong aggres-
sion toward Rams 2 and 3, he remained
rather submissive at the feed box. During
the final two days at the box, Ram 4 started
fights with Rams 2 and 3. He fought with
each male about three times, the average
duration of fighting being about two or three
minutes. But only once, in a contest with
Ram 2, did it appear that he had achieved
temporary dominance over his opponent.7
And only once did Ram 4 achieve dominance
~ The butts exchanged have not been included in the data
because the experimenter was engaged in taking motion
pictures, and so could not take notes.
in a paired matching test. His opponent in
this test was Ram 2. Possibly in the two
experimental situations Ram 4 was inhibited
by previous experiences of defeat and sub-
ordination. Seward (12) with rats, and
Ginsburg and Allee (7) with mice, have
shown that an animal could be conditioned
to defeat much more readily than to victory.
Although they did not study the factor of
physical environment, it seems reasonable
to expect that aggression would be most
strongly inhibited in the place where sub-
ordination had been experienced most fre-
quently.
Play fighting was frequent among the
lambs. Often in the evening they scrambled
about, pushing and butting each other, in
order to gain a position on top of the rocky
hillock in the center of the field. This ac-
tivity was similar to the children’s game,
“king-of-the-hill,” and has been reported by
Darling (5) as occurring among red deer
fawns. Sometimes fighting in lambs had a
more serious appearance. Two lambs might
butt head-on forcefully and in rapid suc-
cession until both seemed quite exhausted.
There was one instance of a lamb fighting
with an adult. Late in August Ewe 4 butted
Lamb 3 and the lamb immediately butted
back. A short fight ensued, ending with the
lamb’s retreating and then attacking Lamb 4.
Sexual Behavior of the Rams.
The first witnessed attempt to mount a
female was by Ram 2 on August 10. The
ram reared on his hind legs and his penis
emerged about three inches from its sheath
for a few seconds. The ewe ran off. The
ram then tried to mount another ram.
Toward the end of August attempted mount-
ings by rams of both rams and ewes were
common. This behavior was not observed to
occur in the ewes or lambs. Female urine
had an excitatory effect upon the rams. The
ram sniffed the urines, then curled the up-
per lip, extended the neck, and tilted the
nose in the air. This pattern, according to
Spencer (13) is found in bighorns, as well
as in other ungulates. Rams sometimes lay
on their backs and sucked their penises for
short periods. Ejaculation of semen was not
observed to occur. Often in the evening, sex-
ual activity and fighting occupied the rams
continuously until dark. At no time during
observation was a ewe receptive. A ram did
not persist in attempting to mount the same
ewe. Two tries were usually enough to dis-
courage him. There was no chasing about
the field. A ewe had only to walk or run a
few yards to get rid of a ram.
Displaced Aggression.
One of the most striking behavior pat-
terns observed in the course of this study
was that in which a sheep responded to a
butt, threat or attack from a dominant sheep
by delivering in kind to the nearest sub-
ordinate. Such “displaced aggression” might
continue chain-wise through three or four
1949]
Katz: Behavioral Interactions in Barbary Sheep
17
individuals, each successive one being lower
in dominance. Thus a ram might butt a sub-
ordinate ram away from a piece of bread,
the latter might butt a ewe nearby, and the
ewe in turn a lamb. At the feed box Ram 2
often responded to a sharp butt from Ram 1
by circling the box and butting all the sheep
in his path. The examples of displaced ag-
gression are too numerous to be listed. The
pattern appeared in rams, ewes and lambs
with great frequency. There seemed to be
only two factors determining which sheep
was to receive a displaced attack, physical
proximity and lower dominance status. No
special relationships between individuals
were apparent, other than the usual ones
of dominance-subordination. Winslow (15 >
found displaced aggression in cats made to
compete for food.
Summary and Conclusions.
1. This study represents an attempt to
anaiyze social behavior and group organi-
zation in a small herd of Barbary sheep.
2. Observations and experiments were
made on the herd of four rams, four ewes,
and four lambs at the New York Zoological
Park during the summer of 1947, under con-
ditions with a minimum of human care and
interference.
3. In two types of tests it was found that
stable relationships of dominance-subordina-
tion existed between all individuals, and that
the dominance order of all eight sheep was
one of straight descent through rams, ewes
and lambs.
4. When grain was placed in a small feed
box it was found that individual differences
in total feeding time at the box were not
related to the dominance order. These in-
dividual variations probably were due to
different rates of food intake and differences
in nutritional needs.
5. In the feed box experiments it was found
that there was an order of time-of-maxi-
mum-feeding which was very similar to the
order of dominance.
6. Differences were found in the “social
distance” between any two rams when at
the feed box. These differences were re-
flected in the amount of food sharing that
occurred and in amount and reciprocity of
aggression. Food sharing and aggressive in-
teraction were positively related, and both
appeared to be manifestations of the domi-
nant animal’s tolerance, or willingness-for-
contact with regard to the subordinate.
7. There were individual differences in
amount of food sharing and amount of suck-
ing among three ewe-lamb pairs. The domi-
nance status of the lamb seemed to be asso-
ciated with the social distance between the
lamb and its mother. But there appeared
to be no relation between the dominance
status of the ewe and the dominance status
of her lamb.
8. Consistent leadership appeared only in
conflict situations characterized by a locus
of attraction and a locus of danger in close
proximity to each other. In conflict situa-
tions a ewe always led, and with only two
exceptions it was always the same ewe. The
rams were more timid than the ewes in
strange and potentially “dangerous” situa-
tions.
9. Fighting occurred between rams, ewes
and lambs. With one exception, there were
no fights between ewes and rams, or between
adults and lambs. Ewes fought less than
rams, while among lambs play-fighting often
was observed.
10. As sexual activity appeared in the
rams, fighting became more frequent and
vigorous.
11. Dominance relationships between rams
remained stable throughout the study and
from the time of the first appearance of sex-
ual interest until the study terminated five
weeks later.
12. Sexual activity in the rams consisted
of attempted mounting of ewes and rams
and sometimes of incomplete masturbation.
Ewes were not receptive up to September
14, when the study ended.
13. Instances of displaced aggression were
very numerous. The recipient was usually
the nearest subordinate animal.
Bibliography.
1. Allee, W. C., Allee, M. N., and
Castles, E. E. Concerning leadership
in a flight of white Pekin ducks (Ab-
stract). Bull. Ecol. Soc. of America,
1946, 27, 15-16.
2. Carpenter, C. R. Societies of monkeys
and apes. Biol. Symp., 1942, 8, 177-
204.
3. Carpenter, C. R. Concepts and prob-
lems of primate sociometry. Socio-
metry, 1945, 8, 56-61.
4. Collias, N. E. Aggressive behavior
among vertebrate animals. Physiol.
Zool., 1944, 17, 84-123.
5. Darling, F. F. A herd of red deer: a
study in animal behavior. Oxford
University Press, Humphrey Milford,
London, 1937.
6. Davis, W. B. Summer activity of moun-
tain sheep on Mt. Washburn, Yellow-
stone National Park. J. Mammal., 19,
88-94, 1938.
7. Ginsburg, B. and Allee, W. C. Some
effects of conditioning on social dom-
inance and subordination in inbred
strains of mice. Physiol. Zool., 1942,
15, 485-506.
8. Lydekker, R. Wild oxen, sheep and
goats of all lands. Rowland Ward,
London, 1898.
9. Maslow, A. H. The role of dominance
in the social and sexual behavior of
infrahuman primates: IV. The de-
termination of hierarchies in pairs
and in a group. J. Genet. Psychol.,
1936, 49, 161-198.
18
Zoologica: New York Zoological Society
[34: 3: 1949]
10. Mills, H. B. A preliminary study of
the bighorn of Yellowstone National
Park. J. Mammal., 1937, 13, 205-212.
11. Scott, J. P. Social behavior, organiza-
tion and leadership in a small flock
of domestic sheep. Comp. Psychol.
Monogr., 1945, 18.
12. Seward, J. P. Aggressive behavior in
the rat: II. An attempt to establish
a dominance hierarchy. J. Comp.
Psychol., 1945, 30, 213-224.
13. Spencer, C. C. Notes on the life history
of Rocky Mountain bighorn sheep in
the Tarryall Mountains of Colorado.
J. Mammal., 1943, 24, 1-11.
14. Stewart, J. C. and Scott, J. P. Lack
of correlation between leadership and
dominance relationships in a herd of
goats. J. Comp. Psychol., 1944, 37,
297-314.
15. Winslow, C. N. Social behavior in cats,
1. J. Comp. Psychol., 1944, 37, 297-
314.
Fleming : Pericopidae of British Guiana and Venezuela
19
4.
The Pericopidae (Moths) of Kartabo, British Guiana, and Caripito,
Venezuela.1
Henry Fleming.
Entomologist, Department of Tropical Research,
New York Zoological Society.
[This contribution is the result of various
expeditions of the Department of Tropical Re-
search of the New York Zoological Society to
British Guiana and to Venezuela, all under the
direction of Dr. William Beebe. The Guiana
expeditions were made during the years 1917,
1919, 1920, 1921 and 1924. The expeditions were
arranged so that each month of the year is
represented in the collections. The Venezuelan
expedition, in 1942, during which field work was
carried on from February 19 to September 2,
was sponsored by grants from the Committee
for Inter-American Artistic and Intellectual
Relations and from four trustees of the Zoologi-
cal Society, George C. Clark, Childs Frick,
Laurance S. Rockefeller and the late Herbert
L. Satterlee, and by invaluable assistance from
the Standard Oil Companies of New Jersey and
Venezuela.]
A total of eight species of Pericopidae
were collected at Kartabo and four at Cari-
pito. One species from British Guiana and
two species from Caripito are new locality
records for their respective countries. One
species common to both British Guiana and
Venezuela is represented by a new race in
Venezuela.
Eucyane bicolora (Sulzer).
Sulzer, Gesch. Ins., t. 22, f. 6 (Expl. Tab.)
(1776) (Noctua).
Three specimens taken at Kartabo, two on
October 11 and one on December 2. The
species has been reported from the Guianas,
South Brazil and Peru.
Eucyane temperata Walker.
Walker, List. Lep. Ins. Brit. Mus., 7, p.
1656 (1856).
One specimen taken at Caripito on July 11.
The species has been reported from South
Brazil, Guianas and Colombia, so this is a
new record for Venezuela.
Pericopis c atilina catilina (Cramer),
new status.
Cramer, Pap. Exot., 1, t. 79. f. E. F. (1775)
(Attacus) .
In my opinion Dysschema brotes (Druce)
Ann. Mag. Nat. Hist., 15 s. 6, p. 48 1895
( Anthomyza ) is only the male form and thus
a synonym of Pericopis catilina catilina. The
males in collections are usually named brotes
1 Contribution No. 827, Department of Tropical Research,
New York Zoological Society.
and the females catilina. The male specimens
match Druce’s description better since the
males are usually blackish-brown rather than
the cinnamon brown typical of the females.
Furthermore, the character given in the lit-
erature to separate the genus Dysschema
from Pericopis is not valid. This character,
the length of the pinnae of the antennae
double the width of the shaft, is a male sexual
character typical of many of the species of
Pericopis. I am not synonomizing the genus
Dysschema since the genotype tiresias is not
available, but the present generic character
does not justify the genus. Thus, on the basis
of this character the male specimens of
catilina are assignable to Dysschema, hence
brotes, and the females to Pericopis, hence
catilina.
One male captured at Kartabo on May 24.
Recorded from Brazil, Guianas and Colombia.
Pericopis catilina angustilineata, new
sub-species.
Length of
Specimen Sex Date forewing Type
42487 male March 15 35 mm. Holotype
42488 female March 16 38 mm. Allotype
4239 male March 11 38 mm. Paratype
42489 male April 15 35 mm. Paratype
42490 male March 15 37 mm. Paratype
42491 male June 5 35 mm. Paratype
42492 female April 8 38 mm. Paratype
Head as in c. catilina. In the male the
length of the pinnae of the antennae is twice
the width of the antennal shaft, while in the
female the pinnae are barely as wide as the
antennal shaft.
Ground color of both the fore and hind-
wings blackish-brown to cinnamon brown
with bands of greenish-yellow.
The forewing with two semi-hyaline
greenish-yellow bands, one median and the
other apical as in c. catilina with brown or
brownish-black veins crossing the bands. The
bands differ from those of c. catilina in being
much reduced; little more or not more than
half the width of the bands of c. catilina. If
one assumes the nomenclatural type to be
ancestral, the reduction of the bands has
been caused by the encroachment of the
brown or blackish-brown scales on both sides
of the bands. This is most easily discerned in
the median band. The inner side of the band
in c. catilina crosses the wing nearer to the
20
Zoological New York Zoological Society
[34: 4: 1949]
point where vein C112 forks from the cubital
stem than it does in c. angustilineata. Similar
results are obtained if one measures basally
from the point where vein Cui forks from the
cubital stem. In both c. catilina and c. angus-
tilineata, but particularly in the latter, the
encroachment of the ground color on the
yellow bands may be seen. The brown scales
on the margins of the bands are duller and
lighter than the surrounding ground color.
This is variable, being more evident in some
specimens than others and occurring indis-
criminately along the length of the bands. It
is most noticeable and frequent on the me-
dian bands. The white spots along the outer
margin of the forewing are evident to vary-
ing degrees. Their place is taken by the
brown or blackish-brown scales making up
the background. However, all specimens have
a streak of white scales on the outer side of
the yellow spot which terminates the apical
band in cell Ms. The spots in cells Mi and M2
appear to vanish first since they are faintly
discernible in only one specimen.
The hindwings of angustilineata, as in
catilina, are yellow hyaline for half the length
of the wings from the base with a large
yellow spot beyond in cells Mi and M2. Peri-
copis c. angustilineata differs in that this
large yellow spot is separated from the basal
patch by a distance approximately twice that
of catilina. The part of the basal facies that
extends into the proximal part of cell Cui is
much smaller in angustilineata than in cati-
lina. The ground color along the inner mar-
gins of the hindwings encroaches more on
the yellow basal area in angustilineata than
in catilina.
The specimens were all collected during the
day while flying. No specimens of Danaidae
or Heliconiinae were captured or seen in the
same area, though the general appearance
and flight of angustilineata is suggestive of
various members of either of the above
groups. The specimens were captured in an
area of about one hundred meters’ diameter
near the end of an unmaintained road going
to an abandoned oil well “No. 1”. The locality
is approximately ten miles west of Caripito,
State of Monagas, eastern Venezuela. The
area the specimens came from is parched
during the dry season, flooded during the wet
and characterized by numerous palms and
small to very moderate-sized seasonal trees.
Pericopis tricolora tricolora (Sulzer).
Sulzer, Gesch. Ins., t. 22, f. 5 (1776)
(Noctua) .
Three specimens taken at Kartabo; two
females on March 22 and one male on Novem-
ber 24. Recorded from eastern Peru, Ama-
zons and Guiana.
Dysschema heliconides (Swainson).
Swainson, Zool. 111. (2), 3, t. 124, f. 2
(1833) (Anthomyza).
One specimen collected at Kartabo in 1920.
Recorded from the Amazons, Guianas, Co-
lombia and Peru.
Hyalurga fenestra (Linnaeus).
Linnaeus, Syst. Ent. (ed. 10), 1, p. 505,
n. 41 (1758) (Phalaena).
One specimen collected at Kartabo which
represents a new record since the species has
only been reported from Brazil and Peru.
Hyalurga sixola Schaus.
Schaus, Ann. Mag. Nat. Hist., (8), 6, p.
206 (1910).
A total of seven specimens was taken at
Caripito; the males on March 20 (two speci-
mens), April 5, May 12, and May 16 and the
females on April 15 and July 2. Recorded
from Venezuela and Colombia.
Hyalurga m ysis (Erichson).
Erichson in Schoenburgk, Brit. Guiana,
3, p. 606 (1848) (Glaucopis).
A female from Kartabo on May 25. Has
only been found in British Guiana.
Hyalurga modesta Moschler.
Verh. Zool.-bot. Ges. Wien, 27, p. 663, t. 9,
f. 29 (1877).
One female at Kartabo on August 19.
Recorded from Guiana and Colombia.
Hyalurga partita (Walker).
Walker, List. Lep. Ins. Brit. Mus., Het. 2,
p. 335, n. 27 (1854) (Dioptis).
Four female specimens at Kartabo; two
with no date and the remaining two speci-
mens on November 21 and December 21. One
female at Caripito on March 21. Recorded
from Brazil, Guianas, Venezuela and Peru.
Goodnight : Phalangids from Rancho Grande, Venezuela
21
5.
Report on a Collection of Phalangids from Rancho Grande, Venezuela.’
Clarence and Marie Goodnight.
Department of Biology, Purdue University.
(Text-figures 1-4.)
[This is one of a series of papers resulting
from the 45th and 46th Expeditions of the De-
partment of Tropical Research of the New York
Zoological Society, made during 1945 and 1946
under the direction of Dr. William Beebe, with
headquarters at Rancho Grande in the National
Park of Aragua, Venezuela. The expeditions
were made possible through the generous coop-
eration of the National Government of Vene-
zuela and of the Creole Petroleum Corporation.
[The characteristics of the research area are
in brief as follows : Rancho Grande is located in
north-central Venezuela (10° 21' N. Lat., 67° 41'
W. Long.), 80 kilometers west of Caracas, at an
elevation of 1,100 meters in the undisturbed
montane rain forest which covers this part of
the Caribbean range of the Andes. Adjacent
ecological zones include seasonal forest, sa-
vanna, thorh woodland, cactus scrub, the fresh-
water lake of Valencia and various marine lit-
toral zones. The Rancho Grande area is gener-
ally subtropical, being uniformly cool and damp
throughout the year because of the prevalence
of the mountain cloud cap. The dry season ex-
tends from January into April. The average
humidity during the expeditions, including
parts of both wet and dry seasons, was 92.4% ;
the average temperature during the same period
was 18° C; the average annual rainfall over a
five-year period was 174 cm. The flora is marked
by an abundance of mosses, ferns and epi-
phytes of many kinds, as well as a few gigantic
trees. For further details, see Beebe and Crane,
Zoologica, Vol. 32, No. 5, 1947. Unless other-
wise stated, the specimens discussed in the pres-
ent paper were taken in the montane cloud for-
est zone, within a radius of one kilometer of
Rancho Grande.]
This paper is a report on the phalangids
collected during the 45th and 46th Expedi-
tions of the Department of Tropical Re-
search.
The species showed relationships to those
of Trinidad (Goodnight and Goodnight,
1947), many of the specimens representing
the same species. The Cosmetidae, one of the
most typical of neotropical families, is here
represented by two species, one of which is
new.
Among the Phalangodidae, the Stygnom-
matinae are represented by the wide-ranging
Zygobunus rufus (Petrunkevitch) . This was
formerly known only from Panama. The sub-
family Phalangodinae is represented by
Kalina tuberculata Goodnight and Goodnight
l Contribution No. 828, Department of Tropical Research,
New York Zoological Society.
known formerly only from Trinidad. The
Triacommatinae are represented by one new
species, Vima plana.
Among the family Gonyleptidae, three spe-
cies are represented, one of which is new.
The writers wish to express their appre-
ciation to Dr. Beebe and Mr. H. Fleming
for making this material available for their
study. Types are deposited in the collections
of the Department of Tropical Research,
New York Zoological Society, New York 60,
New York.
SUBORDER LANIATORES THORELL.
Phalangodidae Simon.
Phalangodinae Roewer.
Kalina tuberculata Goodnight and Goodnight.
Reference: Kalina tuberculata Goodnight
and Goodnight, 1947, p. 1, fig. 4.
Record: Zone 28, Rancho Grande, Vene-
zuela, 1945.
Stygnommatinae Roewer.
Zygobunus rufus (Petrunkevitch).
References: Stygnomma rufum Petrunke-
vitch, 1925, p. 62.
Zygobunus barronus Chamberlin, 1925, p.
245; Roewer, 1928, p. 546.
Stygnommatiplus rufus Roewer, 1928, p.
544.
Zygobunus barronus Goodnight and Good-
night, 1942, p. 4, figs. 10, 11, 12.
Record : Rancho Grande, Venezuela, March
4, 1945.
Tricommatinae Roewer.
Mima plana sp. nov.
(Text-figs. 1 & 2) .
Male : Dorsum with five areas, very small
paired tubercles on the 1st, 2nd, 3rd and 4th
areas. These tubercles are very small and
vary in size in different individuals. Cephalo-
thorax smooth, with a low tubercle at the
posterior lateral portion. Eye tubercle wider
than long, with low tuberculations across the
median portion. First area of the abdomen
without a median line. Boundaries of areas
indistinct, not parallel. Lateral margin of
abdominal scute smooth, without median ar-
mature. 5th area and free tergites each with
a few small granulations. Anal operculum
smooth, free sternites each with a transverse
22
Zoologica: New York Zoological Society
[34: 5
Text-fig. 1. Vima plana sp. nov. Dorsal view
of male holotype.
row of minute tubercles. Spiracle visible.
Coxae with a few small granulations; 1st
coxa with a transverse row of spines.
First leg slender, unarmed ; 2nd to 4th
legs heavier, a few scattered tuberculations
on the trochanters. Femora with longitudinal
rows of spines, remainder of legs only with
scattered hairs. 4th patella with a few apical
tubercles. Double claws smooth, without
scopula or false claw. Tarsal segments: 9-18-
8-9. Distitarsus of 1st tarsus with 3 seg-
ments, 2nd with 3 also.
Length of Legs.
I.
II.
III.
IV.
mm.
mm.
mm.
mm.
Trochanter
0.3
0.7
0.8
0.8
Femur
6.3
14.4
11.5
16.2
Patella
0.7
1.2
1.4
1.5
Tibia
4.0
10.8
6.1
8.5
Metatarsus
8.5
17.4
12.4
20.5
Tarsus
1.5
4.5
2.9
3.2
Total
21.3
49.0
35.1
50.7
Palpus with the trochanter 0.8 mm. long,
femur 1.1, patella 0.7, tibia 0.6, and tarsus
0.7. Total length, 3.9 mm. Femur armed
retrolaterally as in figure. Prolaterally fe-
mur and patella each with a median apical
spine. Tibia and tarsus armed as on retro-
lateral margin.
Proximal segment of chelicera with a dor-
sal elevation on which are scattered tuber-
cles. Distal segment greatly enlarged.
Body, chelicerae, and palpi light yellowish
with scattered black mottlings. First leg uni-
formly colored ; second leg with a white mark
at the distal end of the femur; patella black,
a white patch at the distal portion of the
tibia; third leg with a darker patella but no
white markings; fourth leg with a white
band following a black band on the distal
portion of the tibia. Legs otherwise uni-
formly dark brown to dusky.
Female: Similar in appearance to male.
Measurements in mm.: Male, total length
3.7 ; cephalothorax 1.5; width at widest por-
tion 2.6. Female, total length 7.2; cephalo-
thorax 1.7 ; width at widest portion 4.2.
Record : Male holotype from Rancho
Grande, Venezuela, July 22, 1945; paratypes
from same locality, March 22, 1945, and July
22, 1945.
Remarks: Vima plana is related to Vima
insignis Hirst. It differs from this latter spe-
cies by lacking the raised area of the dorsum
and the paired low tubercles over the eye.
Cosmetidae Simon.
Cosmetinae Cambridge.
Cynorta estebana Roewer.
Reference: Cynorta estebana Roewer,
1947, p. 18, pi. 18, fig. 66.
Record: Rancho Grande, Venezuela, Au-
gust 9, 1945.
Cynorta bromeliaca sp. nov.
(Text-fig. 3).
Male: Eye tubercle wider than long. 1st
area with a pair of enlarged tubercles, 3rd
area with a pair of robust spines which are
short and heavy at the base. Remaining areas
and free tergites without median armature.
Entire dorsum covered with small white ele-
vations. These are more numerous on the
lateral posterior portions of the scute. Each
free tergite with a transverse row of these
same tubei’culations. Anal operculum with
only a few granulations, free sternites each
with a transverse row of hair-tipped granu-
lations. Coxae and genital operculum smooth
except for scattered hairs. A few teeth on
the anterior margins of the 3rd and 4th
coxae, and a transverse row of granulations
across the 1st coxa.
Text-fig. 2. Vima plana sp. nov. Retrolateral
view of palpus of male holotype.
1949]
Goodnight: Phalangids from Rancho Grande, Venezuela
23
Text-fig. 3. Cynorta bromeliaca sp. nov. Dorsal
view of male holotype.
Legs clothed only with hairs except for a
few tuberculations at the apical portion of
the 4th patella; femora straight. Tarsal seg-
ments: 6-14-11-12. Distitarsus of both 1st
and 2nd tarsi with 3 segments. Proximal por-
tion of 1st tarsus enlarged.
Length of Legs.
I.
II.
III.
IV.
mm.
mm.
mm.
mm.
Trochanter
0.6
0.8
0.8
0.8
Femur
4.4
6.6
6.6
9.3
Patella
1.0
1.6
1.6
1.6
Tibia
2.8
8.3
3.8
5.3
Metatarsus
4.3
10.3
5.5
9.4
Tarsus
2.6
5.2
3.6
4.4
Total
15.7
32.8
21.9
29.8
Palpus with the trochanter 0.8 mm. long,
femur 1.4, patella 0.9, tibia 1.5, and tarsus
0.8. Total length, 5.4 mm. Palpus character-
istically flattened with a ventral row of teeth
on the femur.
Proximal segment of chelicera with a dor-
sal elevation on which are a few granulations.
Distal segment somewhat enlarged.
Dorsum reddish-brown, thickly covered
with white spots which are more numerous
on the lateral and posterior portions of the
scute. These form an irregular band of white
spots, with a few scattered ones in the
median area. Several white spots on the eye
tubercle. A transverse row of white spots
across each free tergite. Anal operculum
without markings. Venter, coxae, and cheli-
cerae reddish-brown with darker markings.
Legs yellowish, trochanters, and bases of fe-
mora, patellae, and tibiae reddish-brown.
Metatarsi white.
Measurements in mm.: Male, total length
6 ; cephalothorax 1.9 ; width of body at widest
portion 4.1.
Record : Male holotype from bromeliads,
Rancho Grande, Venezuela, August 8, 1946.
Remarks: This species is most nearly re-
lated to Cynorta catenulata Roewer. It differs
from this latter species by having an entirely
different pattern of white on the dorsal scute.
Gonyleptidae Sundevall.
Cranainae Roewer.
Poecilocranaus gratlosus Roewer.
Reference: Poecilocranaus gratiosus
Roewer, 1943, p. 54, pi. 7, fig. 63.
Record : Rancho Grande, March 22, 1945.
Santinezia albilineata Roewer.
Reference : Santinezia albilineata Roewer,
1932, p. 290, fig. 7.
Record: Rancho Grande, Venezuela, Au-
gust 1, 1945.
Stenostygninae Roewer.
Bunistygnellus beebei sp. nov.
(Text-fig. 4) .
Male: Dorsum smooth, cephalothorax
without a median eye tubercle. Eyes widely
separated near the posterior portion of the
cephalothorax. Between the eyes a large
rounded elevation with a short apical spine.
Elevation granular. Anterior margin of the
cephalothorax with a short anterior projec-
tion between the chelicerae and palpi. A large
vertical spine at the anterior margin near
the coxa of the palpus. A small tubercle at
the anterior lateral margin. Abdomen with
five areas. 1st area constricted in the middle,
2nd area narrow, 3rd area with a pair of
large spines, 4th and 5th areas unarmed.
Free tergites smooth, unarmed. Lateral mar-
gins of scute smooth, anal operculum smooth.
Free sternites each with a transverse row
of hair-tipped tubercles which are enlarged
into spines at the lateral margin. Spiracles
widely open. Coxae covered with hair-tipped
tubercles. A transverse row of spines across
the 1st coxa. 3rd coxa with anterior and pos-
terior teeth. 4th coxa only slightly projecting
and with a large dorsal apical spine.
Trochanters globular. 2nd and 3rd tro-
chanters tuberculate, 2nd with two small
dorsal apical spines, 3rd with a posterior
apical spine. 4th trochanter very heavy, with
a large dorsal apical spine and a lateral spine
on each side ; covered with smaller tubercles.
1st and 2nd femora clothed only with hairs.
3rd femur covered with hairs and tubercles,
ventrally with 2 long rows of spines and
with 2 dorsal apical spines. 4th femur tuber-
culate, ventrally with 2 rows of very large
spines, dorsal-apically with 2 large spines.
Remaining segments of 1st and 2nd legs
clothed only with hairs. Patella of 3rd leg
tuberculate with a large ventral apical spine.
3rd tibia with 2 ventral rows of tubercles at
the distal third, remainder of 3rd leg un-
armed. 4th patella heavily tuberculate and
with large apical spines. 4th tibia clavate,
with 2 ventral rows of spines at the apical
third. Remainder of leg clothed only with
hairs. 3rd and 4th tarsi with heavy scopulae,
double claws toothed. Tarsal segments: 7-
24
Zoologica: New York Zoological Society
[34: 5: 1949]
16-8-9. Distitarsi of both 1st and 2nd tarsi
with 3 segments.
Length of Legs.
I.
II.
III.
IV.
mm.
mm.
mm.
mm.
Trochanter
0.7
0.8
1.0
1.0
Femur
3.0
4.8
3.8
4.1
Patella
0.9
1.3
1.8
2.0
Tibia
2.0
4.3
2.6
3.3
Metatarsus
3.2
5.0
4.0
5.0
Tarsus
1.4
4.8
2.7
3.1
Total
11.2
21.0
15.9
18.5
Palpus with the trochanter 1.2 mm. long,
femur 3.6, patella 1.6, tibia 1.9, and tarsus
1.9. Total length, 10.2 mm. Coxa with scat-
tered granulations. Trochanter globular with
a dorsal elevation, with a small dorsal and
a small ventral spine. Femur curved, un-
armed except for a small basal ventral tu-
bercule. No dorsal apical or median apical
spine. Patella unarmed, tibia and tarsus each
with 5 hair-tipped spines on either side.
Tarsal claw long and curved back against
the tarsus.
Chelicera greatly enlarged, proximal seg-
ment with a dorsal elevation, with several
small tubercles dorsal and ventral. A large
retrolateral spine at the apical portion. Distal
segment huge, elevated considerably over the
proximal segment. Distal segment smooth.
Dorsum reddish-brown. Spines and eleva-
tion of the cephalothorax likewise reddish-
brown. Eyes black. Lateral portions of 1st
\i to 3rd segments with a large
white blotch. Within these
U II white areas 1 or 2 circles of
' / reddish-brown. 4th area with
/ / a transverse line of white, end-
ing in a wider area at the
^/ / lateral edge. 4th area with a
/ / posterior margin of white. A
// narrow line of white at the
y lateral margin of the scute, ex-
tending from the anterior por-
tion of the cephalothorax to
the 2nd area. A large white
spot on the lateral margins of the 3rd and
4th areas. Each free tergite with the poster-
ior margin irregularly lined in white. Last
free sternite lined with white. Venter and
coxae reddish-brown, basal portions of 3rd
and 4th legs likewise reddish-brown. Palpus,
1st and 2nd legs, and distal portion of 3rd
and 4th legs yellowish, penciled with gray.
These markings give an annulate appearance
on the metatarsi. Chelicera dark reddish-
brown, with netted dark markings.
Measurements in mm. : Male, total length
of body, 5.8 ; cephalothorax, 3 ; width of body
at widest portion, 4.
Record : Male holotype from Rancho
Grande, Venezuela, 1945.
Remarks : This species is related to Buni-
stygnellus macrochelis Roewer, differing
principally in the color markings on the dor-
sum.
References Cited.
Beebe, W., and Crane, J.
1947. Ecology of Rancho Grande, a Sub-
tropical Cloud Forest in Northern
Venezuela. Zoologica, Vol. 32, No. 5,
pp. 43-60.
Chamberlin, R. V.
1925. Diagnoses of New American Arach-
nida. Bull. Mus. Comp. Zool., Harvard
Coll., Vol. 67, pp. 211-248.
Goodnight, C. J. and Goodnight, M. L.
1942. Phalangida from Barro Colorado
Island, Canal Zone. Amer. Mus. Novi-
tates, No. 1198, pp. 1-18.
1947. Studies of the Phalangid fauna of
Trinidad. Amer. Mus. Novitates, No.
1351, pp. 1-13.
Petrunkevitch, A.
1925. Arachnida from Panama. Trans. Conn.
Acad, of Arts & Sciences, Vol. 27, pp.
51-248.
Roewer, C. Fr.
1927. Weitere Weberknechte II. Abh.
Naturw. Verein Bremen. Vol. 26, no. 3,
pp. 527-632.
1932. Weitere Weberknechte VII. Archiv.
filr Naturg. n.s., Vol. 1, no. 2, pp. 275-
350.
1943. Uber Gonyleptiden, Weitere Weber-
knechte XI., Senckenbergiana, Vol. 26,
pp. 12-68.
1947. Diagnosen neuer Gattungen und Arten
der Opiliones Laniatores (Arach.),
Senckenbergiana, Vol. 28, pp. 7-67.
Crane: Crabs of the Genus Pseudothelphusa from Venezuela
25
6.
Fresh-water Crabs of the Genus Pseudothelphusa from Rancho Grande,
Venezuela.1
Jocelyn Crane.
Research Zoologist, Department of Tropical Research
New York Zoological Society.
(Text-figures 1-3).
[This is one of a series of papers resulting
from the 45th, 46th and 47th Expeditions of the
Department of Tropical Research of the New
York Zoological Society, made during 1945, 1946
and 1948 under the direction of Dr. William
Beebe with headquarters at Rancho Grande in
the National Park of Aragua, Venezuela. The
expeditions were made possible through the gen-
erous cooperation of the National Government
of Venezuela and of the Creole Petroleum Cor-
poration.
[The characteristics of the research area are
in brief as follows: Rancho Grande is located
in north central Venezuela (10° 21' N. Lat.,
67° 41' W. Long.), 80 kilometers west of Cara-
cas, at an elevation of 1,100 meters in the undis-
turbed montane cloud forest which covers this
part of the Caribbean range of the Andes. Ad-
jacent ecological zones include seasonal forest,
savanna, thorn woodland, cactus scrub, the fresh
water Lake Valencia, and various marine littoral
zones. The Rancho Grande area is generally
subtropical, being uniformly cool and damp
throughout the year because of the prevalence
of the mountain cloud cap. The dry season ex-
tends from January into April. The average
humidity during the expeditions, including parts
of both wet and dry seasons, was 92.4%; the
average temperature during the same period
was 18° C.; the average annual rainfall over a
5-year period was 1 14 cm. The flora is marked
by an abundance of mosses, ferns and epiphytes
of many kinds, as well as a few gigantic
trees. For further details, see Beebe & Crane,
Zoologica, Vol. 32, No. 5, 1947. Unless otherwise
stated, the specimens discussed in the present
paper were taken in the montane cloud forest,
within a radius of 1 kilometer of Rancho
Grande.]
General Account.
Two species of Potamonidae, both belong-
ing to the genus Pseudothelphusa, live within
the Rancho Grande area and are common
near the laboratory at an altitude of about
3,500 feet. One, P. garmani, occurs also
farther down, near stream-beds in semi-
evergreen seasonal forest, at least to 2,000
feet. The other, the apparently new P. chacei,
is confined to the cloud forest. Each occurs
on both the Caribbean and Valencia sides of
the Cordillera.
1 Contribution No. 829, Department of Tropical Research,
New York Zoological Society. .
The general habits are similar in both.
Each is occasionally found walking along the
damp forest floor, at considerable distances
from the small mountain torrents. More
rarely the crabs are seen submerged in the
streams themselves, clinging tightly to the
rocks and moss with their spiny feet. In both
species the young are carried principally dur-
ing the dry season, in March and early April,
three females with young having been taken
during that season, as well as all of the very
small free-living young in the collection. In
contrast only one female taken during the
rains, in June, carried young. The single
female with spermatophores was captured
in September; by inference the eggs are laid
between October and February, the season
during which we have not visited the labora-
tory.
The food of both species consists largely
of insects, especially beetles. The remains of
eaten crabs are found frequently, and it
seems certain that these crustaceans form an
important food item in the diet of such local
animals as tayras and opossums.
My sincere appreciation goes to Dr. Fenner
A. Chace of the United States National Mu-
seum for his kindness in determining the
identity of P. garmani and the systematic
status of P. chacei. The specimens are in the
collections of the Department of Tropical
Research of the N. Y. Zoological Society, ex-
cept for two examples of each species which
have been deposited in the United States
National Museum. Text-figs. 3A and 3B are
the work of Miss Pamela Marmont; the re-
mainder are by Miss Louise Moore.
Field Key to the Rancho Grande Species of
Pseudothelphusa.
A. Manus of cheliped with a large tubercle
at base of fingers ; anterior part of cara-
pace slightly rough to touch; adults
large, measuring at least three inches
across, females 2% inches across still
having narrow abdomens; marsupial
young numbering between 200 and 300,
their front distinctly bilobed and con-
vex ; semi-evergreen seasonal and cloud
forests garmani
[34: 6
26
Zoological New York Zoological Society
A D
Text-fig. 1. Pseudothelphusa garmani. Adult male, length 47 mm. A, right first abdominal
appendage, extero-posterior view; B, same, extero-anterior view; C, same, postero-
internal view; D, major cheliped, external view.
AA. Manus of cheliped without tubercle at
base of fingers; anterior part of cara-
pace smooth to touch ; adults small, ma-
ture females measuring about IV2 inches
across, males less; marsupial young
numbering between 20 and 40, their
front scarcely lobed, truncate; cloud
forest only chacei, sp. nov.
Pseudothelphusa garmani Rathbun, 1898.
(Text-fig. 1, 2A) .
Reference. Pseudothelphusa garmani Rath-
bun, 1898, p. 522, text-fig. 14.
Color in life. Adults of both sexes : variable
shades of dark brown ; color of carapace uni-
formly distributed, the chelipeds, ambula-
tories and abdomen slightly lighter. Marsu-
pial young : carapace dark brown anteriorly,
paler behind; chelipeds apricot buff (Ridg-
way), deepest on upper merus, carpus and
upper half of manus ; chelae creamy or white ;
sides of carapace light brown to buff ; ster-
num and abdomen white; ischium and tro-
chanter of ambulatories buff, other segments
dark brown.
Development. Two females carried 258 and
260 young, respectively. An example, illus-
trated in Text-fig. 2A, measures 3.5 mm. in
length, 4.7 in breadth. The general form is
closely similar to that of the adult, but the
front has each of the two distinct lobes more
convex and there is no trace of a tubercle
outside the manus at the base of the fingers;
manus not swollen. No very small free-living
examples of this species were taken, but a
young female 29 mm. long has the tubercle
distinct.
Food. Three large examples all contained
comminuted black chitin, showing in one case
unmistakable beetle elytra; in addition one
stomach held soft animal matter, probably
worm tissue.
1949]
Crane: Crabs of the Genus Pseudothelphusa from Venezuela
27
Habitat and Range. Taken at Rancho
Grande from semi-evergreen seasonal and
cloud forests, between 2,000 and 3,800 feet.
Previously known also from near Caracas,
Venezuela, and from Trinidad.
Material. A total of six specimens, not
counting marsupial young, have been pre-
served. Department of Tropical Research
Cat. Nos. 4635, male, length 47 mm.; cloud
forest, Mai*. 28, 1946; 4626, female, 55 mm.,
Rancho Grande verandah, June 24, 1946, with
260 marsupial young, No. 4626a; No. 45449,
2 immature females, 29, 42.5 mm., cloud
forest, April 1, 1945. U. S. Nat. Mus. Nos.
82379, male, length 38.5, cloud forest, Mar.
18, 1946; 82380, female, with spermato-
phores, length 42.5, semi-evergreen seasonal
forest (2000 feet), Sept. 6, 1946.
Pseudothelphusa chacei sp. nov.
(Text-figs. 2B, 2C, 3).
Diagnosis : Superior margin of front dis-
tinct and tuberculate but not carinate; bi-
lobed, with median suture present; cervical
suture nearly straight; carapace scarcely
convex, smooth; exognath of maxillipeds re-
duced to a stump; manus somewhat swollen;
no tubercle at base of fingers; front low;
male matures at length of 17 mm., female
at about 23.
Description. Carapace slightly convex,
regions scarcely elevated ; gastric region
slightly less elevated than branchial; ante-
rior margins of protogastric lobes distinct
but not prominent; depressions defining an-
terior part of mesogastric region scarcely
indicated; median furrow less well defined
than in garmani ; cervical groove nearly
straight, deep, continued practically to lateral
margin. Carapace almost completely smooth
to the touch, with only a hint of microscopi-
cally fine granules in antero-lateral regions.
Antero-lateral margins with a small orbital
tooth followed by a slight gap; behind this
are about 23 to 25 fine teeth, similar and
close-set. Front low; superior margin dis-
tinct but not carinate, almost truncate, bi-
lobed, finely tuberculate, scarcely or not at
all projecting over the vertical surface; in
front view slightly depressed in middle;
lower margin sinuous, the tubercles tending
to be obsolete. Orbits nearly filled by eyes,
margins almost smooth, the superior slightly
sinuous. Maxillipeds substantially as in
garmani and simoni.
Merus of chelipeds finely rugose above, the
inner margin armed with a single line of
stout, graduated teeth, the lower and ventro-
distal margins by beaded granules. Carpus
smooth except for dorso-inner surface which
is armed by a crest of small, distinct teeth,
of which the usual large tooth is only one
exaggerated element. Major and minor
manus smooth in both sexes, save for scat-
tered microscopic granules and punctations
on outer surface and faint rugosities dor-
sally. Major manus, especially in male, defi-
nitely swollen ; upper and lower margins in
all slightly convex; sinus at base of pollex
C
Text-fig. 2. Young crabs taken from ab-
dominal pouches of female Pseudothelphusa.
A, P. garmani, carapace, length 3.5 mm.;
B, P. chacei, carapace, length 3.65 mm.
(drawn to same scale as A) ; C, same, right
cheliped, outer view, length 2.9 mm.
practically lacking; no tubercle at base of
fingers, although there may be a very slight
swelling of the margin at that point; fingers
moderately broad, slightly and irregularly
punctate, and near dorsal surface of dactyl,
tuberculate; prehensile edges practically in
contact, the teeth broad, irregular and vari-
able. Merus of all ambulatories flattened,
with upper margins convex and finely den-
ticulate ; superior margin of carpus and both
margins of manus microscopically spinulous;
dactyli slender and spined. Male abdominal
appendage illustrated (Text-fig. 3).
Color in Life. Adults variable and capable
to a certain extent of color change, the cara-
pace ranging from a dull red to dark brown.
A female with marsupial young had the cara-
28
Zoologica: New York Zoological Society
[34: 6
Text-fig. 3 [Part]. Pseudothelphusa chacei male holotype, length 17.5 mm. A, dorsal
view; B, major cheliped, external view; C, right first abdominal appendage, extero-
posterior view; D, same, extero-anterior view; E, same postero-internal view.
pace snuff brown (Ridgway) , slightly lighter
posteriorly. Chelipeds chiefly cinnamon buff
to clay color with snuff brown on dorsal
ridge; fingers pale buff; ambulatories snuff
brown with darker segment markings. Her
young were apricot buff in general coloring;
front much darker, almost black; ambula-
tories cinnamon rufous; dorsal ridges of
chelipeds apricot buff ; underparts pale buff.
Measurements. Holotype male, No. 461197 :
Length of carapace 17.5 mm.; breadth 28.5;
depth 11.5; width of front, lower margin,
7 mm.; major manus (measured along lower
margin) plus pollex 26. Para type female, No.
45105: Length of carapace 23 mm.; breadth
38.5 ; depth 15 ; width of front, lower margin,
8.4; major manus (measured along lower
margin) plus pollex 30.
Development. Two females carried 22 and
40 marsupial young, respectively. An ex-
ample, illustrated in Text-fig. 2B, measures
3.65 mm. long by 5.0 mm. broad. Compared
with the corresponding stage of P. garmarvi
the front is scarcely lobed and strongly trun-
cate, the eyes are larger and the cheliped
manus is distinctly swollen; the latter dif-
ference is carried through into the adult.
Behavior of Young. Samples of the young,
when removed from one of the mothers and
placed close to her, were almost helpless ; they
could move feebly, but did not try to climb
back into the pouch ; one, however, attempted
to grip one of her legs as she moved slowly
past.
Food. A female with young, when captured
in an open patio of Rancho Grande, was hold-
ing a large, black tenebrionid beetle (D.T.R.
No. 45,107) in her major chela, and pulling
off the legs with her left. The beetle was
uncrushed, being strong and active when
liberated. Two of three stomachs examined
contained black chitin, including small beetle
elytra, in addition to indeterminate soft ani-
mal matter; the third was empty.
Habitat and Range. Known only from
Rancho Grande, in the National Park of
Aragua, Venezuela, in the montane cloud
forest, between about 3,000 and 3,800 feet.
Affinities. Dr. Chace writes as follows re-
garding this species : “It is very close to Miss
Rathbun’s P. simoni. The male abdominal
appendages agree very well with her figure
of that species, but your specimens have a
somewhat less convex carapace and a sharply
carinate upper frontal margin which is com-
pletely lacking in P. simoni. The general ap-
pearance of your material would indicate
full specific distinction from P. simoni, but
the similarity in the male appendages sug-
gests that possibly it deserves only subspe-
cific rank.” P. simoni is known only from the
types, taken from Colonia Tovar and Caracas,
Venezuela, and from the “Antilles” (Clau-
dius).
Material. A total of 13 specimens was
taken, not including marsupial young. The
following have been designated as types:
Holotype male, Department of Tropical Re-
1949]
Crane: Crabs of the Genus Pseudothelphusa from Venezuela
29
Text-fig. B [Part]. Pseudothelphusa chacei male holotype, length 17.5 mm. A, dorsal
view; B, major cheliped, external view; C, right first abdominal appendage, extero-
posterior view; D, same, extero-anterior view; E, same postero-internal view.
30
Zoologica: New York Zoological Society
search Cat. No. 461197, length 17.5 mm.,
Rancho Grande court, March 5, 1946; para-
type female, D.T.R. No. 45105, 23 mm., with
21 young, Rancho Grande court, March 29,
1945; paratype male, United States National
Museum No. 87067, 17 mm., Water Trail,
March 15, 1945; paratype female, U.S.N.M.
No. 87067, 24 mm., Water Trail, March 7,
1945. In addition, the following were taken,
all retained in the collections of the Depart-
ment of Tropical Research: No. 45106, 1 fe-
male, length 22 mm., and 3 young, 7-16 mm.,
cloud forest, March 20-April 20, 1945; No.
4636, 1 female, length 22 mm., with 40 young,
cloud forest, March 14, 1946; No. 461198, 4
[34: 6: 1949]
young, 7-9 mm., cloud forest, March 1-15,
1946.
It gives me great pleasure to name this
species for Dr. Fenner A. Chace, Jr.
References.
Rathbun, M. J.
1898. A contribution to a knowledge of the
fresh-water crabs of America. — The
Pseudothelphusinae. Proc. U. S. Nat.
Mus., Vol. 21, pp. 507-537, (No. 1158).
1905. Les crabes d’eau douce (Potamonidae) .
Nouv. Arch. Mus. Paris, ser. 4, Vol. 7,
pp. 159-321.
Crane: Salticid Spiders; Systematics and Behavior in New Species
31
7.
Comparative Biology of Salticid Spiders at Rancho Grande, Venezuela.
Part III. Systematics and Behavior in Representative
New Species.1
Jocelyn Crane.
Research Zoologist, Department of Tropical Research,
New York Zoological Society.
(Text-figures 1-8).
[This is one of a series of papers resulting
from the 45th, 46th and 47th Expeditions of the
Department of Tropical Research of the New
York Zoological Society, made during 1945, 1946
and 1948, under the direction of Dr. William
Beebe, with headquarters at Rancho Grande in
the National Park of Aragua, Venezuela. The
expeditions were made possible through the
generous cooperation of the National Govern-
ment of Venezuela and of the Creole Petroleum
Corporation.
[The characteristics of the research area are
in brief as follows: Rancho Grande is located
in north-central Venezuela (10° 21' N. Lat.,
67° 41' W. Long.), 80 kilometers west of Cara-
cas, at an elevation of 1,100 meters in the undis-
turbed montane cloud forest which covers this
part of the Caribbean range of the Andes. Ad-
jacent ecological zones include seasonal forest,
savanna, thorn woodland, cactus scrub, the
fresh water Lake Valencia, and various ma-
rine littoral zones. The Rancho Grande area is
generally subtropical, being uniformly cool and
damp throughout the year because of the preva-
lence of the mountain cloud cap. The dry season
extends from January into April. The average
humidity during the expeditions, including
parts of both wet and dry seasons was 92.4%;
the average temperature during the same period
was 18° C.; the average annual rainfall over a
5-year period was 174 cm. The flora is marked
by an abundance of mosses, ferns and epiphytes
of many kinds, as well as a few gigantic trees.
For further details, see Beebe & Crane, Zoolog-
ica, Vol. 32, No. 5, 1947. Unless otherwise stated,
the specimens discussed in the present paper
were taken in the montane cloud forest zone,
within a radius of 1 kilometer of Rancho
Grande.]
Contents.
Page
Introduction 31
Subfamily Lyssomaninae
Lyssomanes bradyspilus sp. nov 31
Subfamily Salticinae
Semorina brachychelyne sp. nov 35
Semorina megachelyne sp. nov 38
Ashtabula furcillata sp. nov 39
Sassacu8 flavicinctus sp. nov 41
Sasmcus ocellatus sp. nov 44
Phiale flammea sp. nov 47
Mago dentichelis sp. nov 49
References r,
1 Contribution No. 840, Department of Tropical Research.
New York Zoological Society.
Introduction.
The eight species described in the present
paper have been selected from among other
Rancho Grande salticids for two reasons.
First, they represent a number of different
stages and directions in salticid evolution;
and, second, special experimental display
data and/or examples of the earliest instars
have been assembled in each. Part I of this
series (Crane, 1948.1) dealt monographically
with several species of Corythalia, while Part
II (1948.2) described the methods of study.
In the succeeding parts, which will be based
largely on Corythalia and the present group
of species, it is proposed to discuss the re-
leasing mechanisms of display, to compare
post-embryological development and, finally,
to evaluate evolutionary trends.
With the exception of Text-figure 8F,
which was drawn from life at Rancho Grande
by Mr. Kenneth Gosner, all the illustrations
are the work of Miss Louise A. Moore.
The types are deposited in the collections
of the Department of Tropical Research,
New York Zoological Society, New York 60,
N. Y.
Lysso manes bradyspilus sp. nov.
(Text-fig. 1),
Diagnosis : Retromargin of fang groove
with 6 teeth, the 2 proximal minute, none
crowded toward fang base ; basal segment of
chelicera in male with cluster of 3 to 7 dorsal
distal spines; fang toothless; no fringes on
first metatarsi, which are straight; no clus-
ter of dorsal tibial spines on palp, its distal
apophysis very small, blunt; bulb with three
strong, spinous, distal processes; epigynum
with two pairs of large rounded bodies dis-
tinct, the anterior pair the smaller and prac-
tically contiguous. Abdominal black spots
present or absent.
Color.
Color in Life : Adult male. Cephalothorax :
Integument of carapace translucent green,
without dark pigment, varying from a yel-
lowish-green, especially in recently molted
32
Zoologica: New York Zoological Society
[34: 7
examples, to apple green ( Ridgway) . Ocular
quadrangle including black eye tubercles
with varying amounts and proportions of
yellowish- or silvery white and orange-red
scale hairs, the latter usually placed anteri-
orly. AME rimmed with silvery-white; the
eyes themselves clear apple green, shifting
to black (see under Behavior) ; other eyes
black. A narrow submarginal clypeal band
of orange-red scale hairs, directed downward.
Chelicerae fangs brown. Palpal bulbs pink-
ish to orange. Integument of legs translucent
apple green, without dark pigment except
for black tarsal pads. Abdomen : Integument
translucent green, sparsely covered with
short hairs, ranging from apple green to
dull green-yellow, usually with a short, me-
dian basal stripe of darker green. Hairs
short, rather sparse, of same color as integu-
ment. Paired, subdermal black spots on pos-
terior half of abdomen present or absent,
strong or weak, rarely appearing — if at all
— until three or four days after final molt;
any number up to four pairs may develop. A
patch of white hairs often present at distal
end of dorsum.
Adult female. Differs from male as fol-
lows: Scale hairs of ocular quadrangle, in-
cluding eye tubercles, tend to be more uni-
formly yellowish- or silvery white, with the
orange-red reduced or absent, except for a
variable, sometimes conspicuous, crest be-
hind AME ; subdermal clypeal band of
orange-red absent, replaced by a band of
scant white hairs; palps completely green;
legs usually with some dark subdermal pig-
ment concentrated near joints ; this often is
confined to a single spot in antero-distal part
of first tibia. As in the case of the abdominal
spots, it develops, if at all, after the final
molt. No female seen with more than two
pairs of abdominal spots ; as in the male, they
develop slowly or not at all.
Color in Alcohol : All green fades promptly,
as usual in the genus, to yellowish-white; no
black leg or abdominal pigment remains; on
the other hand, the orange-red clypeal band
of males and the crest of females are strongly
persistent and even intensified.
Structure.
Characteristics below apply to both males
and females unless otherwise specified; per-
centages approximated; measurements of
types given on p. 34.
Carapace: Height, including tubercle of
PLE, scarcely more than half length; short
anterior part of thoracic slope almost level,
descent of posterior part moderate; width
greatest midway between PLE and pedicel,
wider in male (1.5 times height, 79% of
length), narrower in female (1.35 times
height, 71% of length) ; longitudinal thora-
cic .groove well developed, lying midway be-
tween PLE and pedicel.
Eyes: Eight eyes in four distinct rows;
all except AME elevated on low black tu-
bercles, the PME on same tubercle as ALE.
First row 87% as wide as second; length of
Text-fig. 1. (Part). Lyssomanes bradyspilus.
A-D holotype $: A, dorsal view; B, chelicera,
ventral view; C, left palp, ventral view; D,
same, ectal view. E, paratype ?: epigynum.
ocular quadrangle including AME 42% of
carapace length, length from ALE to PLE
27 % ; breadth at ALE much wider than at
PLE, 46% and 34% of length respectively;
ocular quadrangle length from ALE to PLE
only 66% of its breadth at ALE. Diameter of
AME 21% of carapace length : ratio of eyes :
AME: ALE: PME: PLE: :100: 42: 11. 5:
35. AME practically contiguous, separated
from ALE by about a tenth of their diam-
eter; PME slightly closer to ALE than to
PLE.
Clypeus: Height in males 38% of AME
diameter; 54% in females.
Chelicerae: In males, strongly produced
but of variable length, porrect, robust, diver-
gent. Length of basal segment in best devel-
oped more than half carapace length, in least
developed about half. Each with 1-2 prs. of
overlapping spines near base on medial front
1949]
Crane: Salticid Spiders; Systematica and Behavior in New Species
33
Text-fig. 1. (Part). Lyssomanes bradyspilus. A-D, holotype $: A, dorsal view; B, cheli-
cera, ventral view; C, left palp, ventral view; D, same, ectal view. E, paratype $: epigynum.
margin, and a group of 3 to 7 strong distal
spines, the number and arrangement varying
even on two sides of same individual. Fang
slender and sinuous, toothless; groove weak;
promargin with three small teeth near base,
the smallest proximal, it and the next closer
together than second and third; no tooth at
base of fang; inferior margin typically with
6 teeth in a straight row, increasing in size
distally, along entire edge of groove. The
basal one or two, however, although appar-
ently constant, are minute, delicate and
easily destroyed ; they are separated consid-
erably from each other and the distal 4,
which are quite evenly spaced. In females
the chelicerae are, of course, much shorter;
distal spine group absent; teeth closer to-
gether, tending to be evenly spaced through-
out and of more nearly equal size.
Maxillae: Parallel; width 60% of length;
distal dilation slight; external angle evenly
rounded without tubercle.
Lip: Breadth 90% of length; basal exca-
vation extending 25% of length; distal end
reaching slightly beyond middle of maxillae;
sternal suture straight.
34
Zoologica: New York Zoological Society
[34: 7
Sternum : Broadly scutiform; width 85%
of length in males, slightly less in females;
equally wide between second and third coxae ;
base of lip 60% as wide as anterior border
in males, 50% in females; posterior end a
bluntly rounded lobe extending about half-
way between fourth coxae, which are sepa-
rated by two-thirds of their diameter.
Legs : Tibial indices : Holotype male, fii’st
leg 12, fourth leg 11; paratype female, first
and fourth legs, 13. First legs of male con-
siderably elongated and enlarged. See Table
I for formula.
TABLE I.
Lyssomanes bradyspilus : Leg Formula.
12 4 3
Male holotype 3.8 3.0 3.0 2.9
1 4 2 3^
Female paratype 3.2 2.9 2.7 2.6
All legs with little hair; hairs on metatarsi
arranged clearly in dorsal and ventral rows,
but in no sense profusely enough to be called
fringes.
Spines: (Male holotype and female para-
type). First and second legs: Femur dorsal
1-1-1; prolateral and retrolateral 0-1-1. Pa-
tella 0 but with a long, slender dorsal distal
bristle. Tibia: Prolateral 1-1; retrolateral
1-1 in male, and on second female leg, 0-1 on
first female leg; ventral 2-0-2-2, not opposite,
the distal ones not terminal. Metatarsus ven-
tral only 2-2-2, not terminal. Third leg: Fe-
mur as in first and second. Patella dorsal
distal only 1. Tibia, dorsal 1-0-0-1; pro- and
retrolateral, as in first and second male; ven-
tral 0-0-2-0. Metatarsus, prolateral and re-
trolateral 1-1-0; ventral, male, 2-0-0, female
none. Fourth leg: Femur dorsal 1-1-1; pro-
and retrolateral male 0-0-1, female none. Pa-
tella as in third. Tibia dorsal as in third;
pro- and retrolateral as in first and second
male, except fourth female prolateral is 0-1;
ventral none. Metatarsus ventral only 1 (re-
tro) -0-0. In addition, there are rudiments
on third and fourth legs of distal metatarsal
spines, 2 prolaterals, 2 retrolaterals and 2
ventrals, all minute and very weak. Palpal
spines: Femur dorsal 0-1-1; pro- and retro-
lateral distal 1 ; patella, dorsal distal 1 ; tibia,
prolateral male, 0-1, female, 1-1 ; metatarsus
female, dorsal 1-0; pro and retrolateral 1-1.
Abdomen: About 3 times longer than
broad in males and young females, tapering
from level of genital groove; anal tubercle
not pronounced; vestigial colulus not indi-
cated.
Palp: Femur slightly curved; patella and
tibia nearly equal; tibia without dorsal spine
cluster; tibial apophysis scarcely more than
a truncate tubercle opposing basal ridge of
tarsus; bulb with three pointed distal pro-
cesses, variously shaped, and a distal tubercle
(see Text-fig. 1), the whole complex struc-
ture differing only in proportions and details
from Chickering’s description of the palp in
L. banksi (1946, p. 12).
Epigynum : No median notch. Two pairs of
large, rounded bodies; members of anterior
pair smaller, apparently contiguous; pos-
terior pair separated by less than half their
own diameter.
Measurements.
Male holotype. Total length in alcohol 4.7.
mm. ; carapace length 2.1 ; carapace breadth
1.6; carapace height 1.1; ocular quadrangle
length, AME to PLE .89, ALE to PLE .58;
ocular quandrangle breadth, at ALE .96, at
PLE .72; diameter AME .45, ALE .19, MLE
.05, PLE .15; clypeus height .17; basal seg-
ment chelicera 1.1; patella breadth, 1st leg
.38, 4th .24.
Leg Measurements, Male.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1st
2.3
.96
2.1
2.0
.55 7.9
2nd
1.9
.75
1.6
1.7
.41 6.4
3rd
1.8
.68
1.4
1.7
.41 6.0
4th
1.7
.65
1.5
1.9
.44 6.2
Palp
1.0
.44
.41
—
.58 2.4
Female paratype. Total length in alcohol
4.7 mm.; carapace length 2.1; carapace
breadth 1.5; carapace height 1.1; ocular
quadrangle length, AME to PLE .89, ALE
to PLE .65; ocular quadrangle breadth, at
ALE .99, at PLE .72; diameter AME .45,
ALE .19, AME .05, PLE .15; clypeus height
.24; basal segment chelicera .75; patella
breadth 1st leg .34, 4th .24.
Leg Measurements, Female.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1st
2.0
.79
1.8
1.7
.41
6.7
2nd
1.7
.75
1.4
1.5
.38
5.7
3rd
1.6
.68
1.2
1.5
.38
5.4
4th
1.9
.55
1.4
1.8
.41
6.1
Palp
.82
.44
.44 —
Behavior.
.68
2.4
Locomotion: This species is a typical run-
ner; I have never seen it jump, except in a
final short pounce upon prey. The spider
runs in brief spurts, during which the palps
hang down practically touching the ground ;
during the pause they palpate the surface.
No special use is made of the first legs, which
take an active part in running.
Courtship Display: In Stage I, the cara-
pace is held high, the first three pairs of legs
braced somewhat forward, obliquely, and the
fourth pair back; the palps hang over chel-
icerae, now and then tapping ground, while
the abdomen hangs straight down. To super-
ficial observation, the display consists only
of posing in this position, varied with occa-
sional bobbing of the carapace and twitching
of the abdomen during rising excitement.
Not until Stage II is reached, within touch-
ing distance of the female, are the first legs
raised; they are then extended to the front,
while the carapace sinks low and the abdo-
men is swung back in the horizontal position.
When the spiders are observed from their
own level, however, in a straight front view,
it is obvious that during display the rate of
1949]
Crane: Salticid Spiders; Systematics and Behavior in New Species
35
activity of the muscles controlling the an-
tero-median eyes is considerably increased;
this gives rise to a much accelerated color
“change” of the eyes, from green to black to
green again. Similar eye color shifts have
been known for many years in a few other
salticids (e.g. Bristowe 1941, p. 419 ff. and
I references ) . It is apparently caused by slight
motions of the long, cone-shaped optic
“cups,” possibly concerned with a change in
focus, or in the lateral range of vision, al-
I though the exact mechanism does not seem
to have been worked out. In Lyssomanes the
shifts take place slowly but continuously
during ordinary daily activity, and may
be observed at close range under a bi-
nocular microscope. The mechanism works
independently in the two eyes, and at a given
instant either or both eyes show any propor-
tion of green or black. To human beings, at
least, the asymmetrically rolling effect is
startling. In a dorsal view, the slight motions
of the elongate “cups” may be simultaneously
viewed through the translucent cuticle of the
carapace.
Bristowe suggests the possibility that the
color shifts may be useful in enticing prey.
However that may be, after the Rancho
Grande observations it seems to me highly
probable that acceleration of muscular ac-
tivity during display should be considered
as a definite part of the behavior pattern,
probably with an adaptive significance; its
relative value among the various sign stim-
uli has not yet been established. This entire
subject will be further considered in subse-
quent papers.
Once her attention has been attracted, the
female usually sits quietly, sagging to one
side on several folded legs; during the male’s
display, the rate of her ocular muscular ac-
tivity also is increased.
Threat Display : Males usually took no no-
tice of one another, and were induced to dis-
play only three times. During these periods,
eye color shift was not especially noted. No
differences were observed between threat
position and activity from those of court-
ship, except that the carapace and abdomen
were neither bobbed nor twitched. I never
saw the long chelicerae unsheathed, although
twice there was a brief, butting skirmish be-
fore one opponent retreated.
Habitat : Known only from the cloud forest
near Rancho Grande. Shaken from green
herbs, shrubs and low trees; one example
taken from an epiphytic bromeliad growing
twenty feet from the ground.
Affinities : This species holds its chief
characteristics in common with a number of
Lyssomanes, although their combination
seems quite distinct. L. quadrinotatus Simon,
(1900), from nearby mountains, has only
three teeth on inferior margin of fang
groove.
Material : A total of 7 adult males and 4
females have been preserved in addition to
a number of young. The following have been
designated as types:
HOLOTYPE: Male. Cat. No. 461199, De-
partment of Tropical Research, New York
Zoological Society; Portachuelo, Rancho
Grande, near Maracay, National Park of
Aragua, Venezuela; 1136 meters; cloud for-
est; March 20, 1946.
PARATYPE: Female. Cat. No. 45450,
Department of Tropical Research, New York
Zoological Society; same locality as holo-
type; July 9, 1945.
The name bradyspilus is proposed in ref-
erence to the delayed development of the
black markings after the final molt.
Semorina brachychelyne sp. nov.
, (Text-fig. 2).
Diagnosis : Small, brown, scale-less salt-
icids, carapace low, abdomen long and nar-
row with a very slight constriction near mid-
dle, first legs greatly elongated and enlarged,
extended forward and scarcely used in walk-
ing, while the abdomen is frequently ele-
vated. Chelicerae in male scarcely a fourth
length of carapace; tibial apophyses of palp
both curved.
Color.
Color in Life : Adult male. Carapace in-
tegument dark brown, without scales and al-
most without hairs, except around eyes.
AME clear ochraceous brown shifting to
black. Palps dark. First legs brown, the fe-
mur and tibia almost black, the tarsi and
sometimes the metatarsi translucent horn-
color. Other legs translucent horn. Abdomen
covered with fine dark brown hairs with a
pair of small spots of white hairs (not scales
and not shiny or iridescent) three-fifths of
distance from base to tip. In one male there
was a pair of faint pale spots near tip of
abdomen in addition to the distinct more
anterior pair.
Adult female. Carapace integument yel-
lowish-brown except sternum which is faint-
ly pinkish. Eyes surrounded by a few yel-
lowish hairs. Eyes themselves as in male.
Tibia and tarsus of palps shiny silvery
white, very conspicuous when vibrated.
Swollen tibia of first legs with a ventral
dark spot extending laterally; entire first
leg darker than the others, which are pale
translucent yellow-brown. Abdomen with a
median, slightly darker stripe giving off
three pairs of dark cross bars reaching mid-
dle of side. A median dark spot immediately
before tip of abdomen.
Color in Alcohol: The white spot(s) of the
male abdomen are pi'actically invisible, the
pattern now resembling closely that of the
female, which is little altered from life.
Structure.
The characteristics below apply to both
males and females unless otherwise speci-
fied; percentages approximated; measure-
ments of types given on p. 37.
Carapace: Height only 30% of carapace
length; postocular plateau long; thoracic
Text-fig. 2. Semorina brackychelyne. A-G, holotype $ : A, dorsal view; B. first leg,
anterior view; C, carapace, lateral view; D, palp, ventral view; E, same, ectal view;
F, same, tibial apophysis; G, chelicera, ventral view. H-I, paratype $: H, first leg,
anterior view; drawn to same scale as B; I, epigynum.
slope slightly concave; width of carapace
greatest at level of PLE, about twice height,
and 60% of carapace length. Longitudinal
groove well defined, in middle of postocular
plateau.
Eyes : Eyes occupying slightly less than
one-half length of carapace. Ocular quad-
rangle only a third as long as broad, the
sides practically parallel but with PLE very
slightly closer together than ALE. Carapace
extending moderately beyond PLE at their
level; PME median, or slightly nearer ALE
than PLE. Diameter of AME about 21% of
carapace length; ratio of eyes, holotype:
AME : ALE : PME : PLE : : 100 : 41 : 7 : 3 : 41.
AME practically contiguous, separated from
ALE, which are recurved, by about one-third
diameter of ALE.
Clypeus : Height in male only 5 to 6% of
AME diameter, in female 11 to 12%.
Chelicerae : Short, divergent, 25% of cara-
pace length in male, slightly shorter in fe-
male. Two small teeth on superior, one larger
on inferior margin.
Maxillae: Length 54% of width in male,
64% in female ; outer distal margin a blunted,
obtuse angle, not produced.
Lip : Width 55% of length in male, 78%
in female. Sternal suture straight.
Sternum: Width 56% of length in males,
53% in females. Anterior margin straight,
a little narrower than lip base, greatest
width between posterior margins of first
legs; posterior end tapering, blunt-tipped
extending between fourth coxae; the latter
separated by less than a quarter of their
thickness.
Legs : Tibial indices : Holotype male, first
leg 17, fourth leg 14; paratype female, first
leg 23, fourth leg 12. First leg in both sexes
much elongated and enlarged with the femur
and tibia especially deep (tibia depth of first
leg in male 30% of its length, in female
45%) . See Table II for formula. Hair scanty
1949]
Crane: Salticid Spiders; Systematics and Behavior in New Species
37
except as follows. In male, first tibia and
metatarsus with a short, moderately dense
ventral fringe of dark hairs, and a scantier
dorsal one of pale hairs; second tibia with a
very scant pale fringe, dorsally and ventrally,
metatarsus with a similar, slightly longer
one ventrally only; third and fourth legs
with very scanty ventral metatarsal fringes
only. Fringes of negligible development in
female.
TABLE II.
JL 4 2 3
Male holotype 2.5 1.8 1.6 1.3
1 4 2 3^
Female paratype 1.6 1.6 1.3 1.1
Spines : (From male holotype and female
paratype). Femur, dorsal 0-1-1-1 through-
out, the proximal two weak, bristle-like, es-
pecially in female. Patella spineless through-
out. Spines otherwise as follows: First leg:
Tibia, ventral only 0-2-2-2, the latter not
terminal; metatarsus, ventral only 0-2-2.
Second leg: Tibia, retro-ventral only 1-1-0;
metatarsus, male 0, female 0-2. Third leg:
Tibia and metatarsus 0. Fourth leg: Femur,
retrolateral distal in male 1, in female 0;
tibia and metatarsus 0.
Abdomen : Very elongate and tapering in
both sexes, the breadth about a third of
length, a very slight constriction near mid-
dle.
Palp: Femur practically straight; tibia
more than one-half length of patella; two
lateral tibial apophyses, the more dorsal
longer, tapering, recurved at tip, the more
ventral shorter, strongly curved antero-in-
wardly. Embolus slender and tapering. Dis-
tal part of bulb with a conspicuous, chiti-
nized, knob-like protuberance directed out-
ward.
Epigynum: An anterior pair of kidney-
shaped bodies, diverging posteriorly ; a pos-
terior pair, smaller and closer together, fol-
lowed by a pair of conspicuous small dark
spots; a broad and shallow marginal notch.
Measurements.
Male holotype. Total length in alcohol 5.3
mm. ; carapace length 2.2, breadth 1.3, height
.68; ocular quadrangle length .79, breadth
1.2; diameter AME .46, ALE .19, PME .03,
PLE .19; clypeus height .02; basal segment
of chelicera .55 ; sternum length .86, breadth
.48; patella breadth, 1st leg, .38, 4th .21; ab-
domen length 3.2, breadth .99.
Leg Measurements, Male.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
1.7
.89
1.5
.96
.48
5.5
2
1.1
.51
.79
.65
.31
3.4
3
.82
.38
.58
.68
.31
2.8
4
1.1
.51
1.0
.89
.38
3.9
Palp
.72
.24
.14
—
.62
1.7
Female paratype. Total length in alcohol
5.3 mm.; carapace length 2.2; carapace
breadth 1.3; carapace height .65; ocular
quadrangle length .79; ocular quadrangle
breadth 1.2; diameter AME .45; ALE .19
PME .03, PLE .19; clypeus height .05; basal
segment of chelicera .50; sternum length .96,
breadth .50; patella breadth, 1st leg, .32,
4th .15; abdomen, length 3.2, breadth 1.1.
Leg Measurements, Female.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
1.1
5.8
.92
.65
.38 3.6
2
.82
.48
.62
.51
.44 2.9
3
.79
.34
.44
.55
.34 2.5
4
1.0
.48
.79
.75
.48 3.5
Palp
.65
.27
.24
—
.44 1.6
Behavior.
Locomotion: The movements of this spider
in the field are absurdly reminiscent of
those of scorpions or pseudoscorpions, and
bear little resemblance to ant behavior. Their
small size, however, makes the existence of
an adaptive mimetic function extremely
questionable. They are to be counted among
the runners in the family, their progress
being a rapid sort of scurry, with short
jumps reserved for crossing gaps in the ter-
rain, or, of course, for the final stage in
catching prey. During running the palps are
vibrated continually up and down, while the
first legs are held straight out in front, the
metatarsi and tarsi curved inward; these
legs are often vibrated, scarcely or not at all
touching the ground, almost as rapidly as the
palps. Meanwhile the abdomen is frequently
elevated and waved slightly, also in the ver-
tical plane. Immature specimens show all
these characteristics in progress, and they
are typical of locomotion whether or not
another individual is present. Both abdomen
and first legs are invariably raised whenever
any obstacle is encountered.
Courtship Display: Indistinguishable
from ordinary locomotion except that the
first legs are extended at a wide angle (more
than 90%) and slightly more elevated, the
tarsi usually bent down; often the palps are
held still; there is the usual pursuit with
sidling, and the abdomen, with increasing
excitement, tends to remain elevated. Mo-
tionless posing with abdomen up and first
legs extended at the usual angle, also occurs
with excitement. In Stage II the first legs
are brought close together in front, about as
in simple locomotion. During courtship the
female vibrates her white palps rapidly, once
her attention has been gained.
Threat Display: No threat displays were
seen, although a number of attempts were
made to induce them.
Habitat: Known only from the montane
cloud forest (about 3,600 feet) around Ran-
cho Grande. Always shaken from shrubs and
low trees.
Affinities: This species differs from
Simon’s Venezuelan species, known only from
females (S. seminuda and S. iris, 1901), in
the complete lack of shining abdominal scales
in any specimens. It likewise appears dis-
tinct from Mello-Leitao’s S. lineata (1945)
38
Zoologica: New York Zoological Society
[34: 7
from the Argentine. No other species seem
to have been referred to this genus. It differs
clearly from the other Rancho Grande spe-
cies (see below) in details of the chelicerae,
palp and epigynum.
Material : A total of 5 adult males and 4
adult females have been preserved, in addi-
tion to a number of young. The following
have been designated as types :
HOLOTYPE: Male. Cat. No. 481558, De-
partment of Tropical Research, New York
Zoological Society ; Portachuelo, Rancho
Grande, near Maracay, National Park of Ar-
agua, Venezuela; 1136 meters; cloud forest;
July 15, 1948.
PARATYPE: Female. Cat. No. 461200,
Department of Tropical Research, New York
Zoological Society; Limon Gorge, Rancho
Grande, near Maracay, National Park of Ar-
agua, Venezuela; 1100 meters; lower edge
of cloud forest; April 20, 1946.
The name brachychelyne is proposed in
reference to the relatively short chelicerae.
Semorina megachelyne sp. nov.
(Text-fig. 3).
Diagnosis: Very similar to S. brachychel-
yne in general appearance. Chelicerae elon-
gated, about half carapace length in male;
tibial apophyses of palp slender and straight.
Color.
Color in Alcohol: Both sexes scaleless,
brown except for pale second, third and
fourth legs; no distinct and unvarying spots
or other markings.
Structure.
Does not differ significantly from S. brach-
ychelyne except as follows: Height of cara-
pace slightly more in male (33% of length,
instead of 30%) ; thoracic groove less dis-
tinct, transverse rather than longitudinal;
ALE and PLE slightly larger, almost one-
half diameter of AME. Ratio of eyes, holo-
type: AME : ALE :PME :PLE : : 100 : 48 :8 :
48. Clypeus even narrower, in both sexes,
about 4% of AME in male, 5.4% in female.
Maxillae and sternum both narrower with
little sexual difference in breadth.
Chelicerae: These form a major specific
difference, being long in males, the length of
the basal segment 50% of carapace length;
in females it is only 30%. They are held al-
most horizontally in both sexes, but are
more divergent in males than in females.
Tooth on inferior margin relatively larger
in males of present species than in brachy-
chelyne.
Legs : Tibial indices : Holotype male, first
leg 12, fourth leg 12; paratype female, first
leg 20, fourth leg 15. General form, propor-
tions and fringes similar to those in brachy-
chelyne. The leg formula is given in Table
III.
TABLE III.
Semorina megachelyne: Leg Formula.
14 2 3
Male holotype 2.3 1.7 1.5 1.2
14 2 3
Female paratype 1.5 1.4 1.1 1.1
Text-fig. 3. Semorina megachelyne. A-E, holotype $: A, carapace,
lateral view; B, chelicera, ventral view; C, palp, ventral view; D, same,
ectal view; E, same, tibial apophysis. F, paratype female: epigynum.
1949]
Crane: Salticid Spiders; Systematics and Behavior in New Species
39
Spines : (From male holotype and female
paratype). As in brachychelyne, except for
second leg, as follows: In male, metatarsus
retroventral 1-0, not 0; female, as in brachy-
chelyne male, except metatarsus is lr-2.
Palp : Differs from brachychelyne as fol-
lows : Both tibial apophyses are straight, the
tarsus along with its bulb is more slender,
and the coiling of the tubule within the bulb
is different.
Epigynum : The structure differs dis-
tinctly in the two species, as shown in the
figure; the more nearly spherical shape of
the four bodies is especially noticeable.
Measurements.
Male holotype. Total length in alcohol 4.7
mm. ; carapace length 2.2, breadth 1.3, height
.75; ocular quadrangle length .79, breadth
1.2; diameter AME .43, ALE .21, PME .03,
PLE .21; clypeus height .02; basal segment
of chelicerae 1.1 ; sternum length .96, breadth
.46; patella breadth, 1st leg .27, 4th .17;
abdomen, length 2.5, breadth .82.
Leg Measurements, Male.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
1.5
.82
1.4
.96
.41
5.1
2
.96
.51
.75
.62
.34
3.2
3
.82
.38
.55
.68
.27
2.7
4
1.1
.48
.92
.85
.38
3.7
Palp
.72
.31
.17
—
.58
1.8
Female paratype. Total length in alcohol
4.0 mm.; carapace length 1.7, breadth 1.0,
height .55; ocular quadrangle length .75,
breadth .96; diameter AME .36, ALE .16,
PME .03, PLE .17 ; clypeus height .09; cheli-
cera, basal segment .52 ; sternum length .79,
breadth .36; patella breadth 1st leg .21, 4th
leg .14; abdomen, length 2.3, breadth .79.
Leg Measurements, Female.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
.79
.48
.58
.44
.27 2.6
2
.62
.34
.41
.31
.24 1.9
3
.55
.31
.34
.41
.27 1.9
4
.79
.31
.62
.55
.27 2.5
Palp
.44
.17
.14
—
.34 1.1
Behavior : Locomotion as in brachychel-
yne. No displays observed.
Habitat : Known only from lower edge of
montane cloud forest, about 3,500 feet, near
Rancho Grande. Collected from tree trunks
and shrubs.
Affinities : See remarks under brachy-
chelyne.
Material: A total of 2 adult males and 5
adult females were taken, along with a num-
ber of young. The following have been des-
ignated as types:
HOLOTYPE: Male Cat. No. 461201, De-
partment of Tropical Research, New York
Zoological Society; Water Trail, Rancho
Grande, near Maracay, National Park of Ar-
agua, Venezuela; 1100 meters; lower edge
of cloud forest; May 5, 1946.
PARATYPE: Female. Cat. No. 461202.
Same data as holotype.
The name megachelyne is proposed in ref-
erence to the long chelicerae of the male.
Ashtabula furcillata sp. nov.
(Text-fig. 4) .
Diagnosis: Color in life above entirely ir-
idescent green with white dorso-lateral band
encircling carapace and abdomen ; dorsal ab-
dominal spots lacking, although sometimes
faintly indicated in alcohol; carapace low;
abdomen elongate; tibial apophysis of male
forked.
Color.
In Life: Adult male. Carapace above en-
tirely covered with iridescent scales, rich
green with bronze reflections. A white stripe
starting behind ALE, bordered narrowly on
ventral margin with black, passing imme-
diately below PME and PLE, and extending
along thorax almost to pedicel. Sides of cara-
pace naked, black with a narrow white sub-
marginal border of scales. AME narrowly
rimmed with yellowish. Clypeus black, naked.
Palps and first legs black (except pale 1st
tarsi), other legs translucent buff. Sternum
black. Abdomen covered with green scales
like those of carapace, outlined dorso-later-
ally with white, which either continues to
tip of abdomen or stops short of the tip;
a white distal median spot present or absent.
Moderate green iridescence on lower abdom-
inal sides, below white stripe; venter black.
Adult female. Like male, except sides of
carapace brown, not black ; palps light green-
yellow, not black ; first legs dark brown, not
black, the distal metatarsus and entire tar-
sus paler; other legs pale as in male, but
with greenish tinge.
In Alcohol: The green iridescence is al-
most or completely lacking, and the scales
may be largely missing, especially on the ab-
domen, where there may be faint traces of
median spots or other markings. The white
dorso-lateral bands, however, are very per-
sistent.
Structure.
Essentially as in Chickering’s description
of A. dentata Cambridge, 1901 (Chickering,
1946, p. 248). The only significant differ-
ences are as follows : Chelicerae : Large pro-
lateral tooth of basal segment of chelicera
straight, not curved; enlargement at base
of fang less distinct, a tubercle rather than
a tooth. Fringe on first leg continues onto
metatarsus. Spines: Very similar in the two
species ; the femoral prolateral distal spines
tend to be more numerous than in dentata
(first leg 2, not 1; 4th leg, n>ale, 1 not 0, but
0 in female) ; metatarsal prolaterals tend
to be fewer than in dentata (second leg 0, not
1; third leg 1, not 2) ; a weak fourth meta-
tarsal ventral distal is present in furcillata,
absent in dentata. Female furcillata as in
male, except that femoral distal spines are
reduced, about as in male dentata, and tibials
are completely absent. Palp: Tibial apophy-
sis differs radically from that of all pre-
Zoologica : New York Zoological Society
[34: 7
Text-fig. 4. Ashtabula furcillata. A-E, holotype 3 : A, carapace and abdomen, dorsal
view; B, carapace, lateral view; C, chelicera, ventral view; D, palp, ventral view;
E, same, ectal view; F, courtship display. G, paratype $: epigynum.
1949]
Crane: Salticid Spiders; Systematics and Behavior in New Species
41
viously known males — zonura Peckham,
1894, dentata Cambridge, 1901, and of den-
tichelis, sexgutta and glauca, all of Simon,
1902; in furcillata alone it is not simple, but
distally forked.
Measurements.
Male holotype. Total length in alcohol 4.2
mm. ; carapace length 1.9, breadth 1.4, height
.75; ocular quadrangle length .82, breadth
1.2; diameter AME .34; ALE .17; PME .04;
PLE .17; clypeus height .05; basal segment
of chelicera 2.4; patella breadth, 1st leg, .19,
4th .21; length of abdomen 2.3, breadth 1.1.
Leg Measurements, Male.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
1.2
.79
.99
.79
.41 4.2
2
.82
.44
.55
.48
.31 2.6
3
.79
.38
.51
.51
.31 2.5
4
.99
.51
.72
.58
.34 3.1
Palp
.68
.14
.10
—
.62 1.5
Female paratype. Total length in alcohol
3.7 mm.; carapace length 1.7; carapace
breadth 1.1; carapace height .68; ocular
quadrangle .79 ; ocular quadrangle breadth
1.1; diameter AME .33; ALE .17 ; PME .04;
PLE 17; clypeus height .03; basal segment
of chelicera .36 ; patella breadth, 1st leg .26,
4th .21 ; length of abdomen 2.0, breadth 1.0.
Leg Measurements, Female.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
.82
.55
.62
.48
.31 2.8
2
.68
.38
.44
.38
.27 2.2
3
.68
.38
.38
.41
.31 2.2
4
.85
.44
.65
.51
.31 2.8
Palp
.44
.21
.41
—
.38 1.4
Tibial indices: Holotype male, first leg 11,
fourth leg 17 ; paratype female, first leg 22,
fourth leg 19. See Table TV for formula.
TABLE IV.
A. furcillata : Leg formula.
1 4 2 3
Male holotype 2.2 1.6 1.4 1.3
1 4 2 3
Female paratype 1.6 1.6 1.3 1.3
Behavior.
Locomotion : A scurrying run, the first legs
held flat and low, straight in front of body;
both they and the palps palpate the surface
almost constantly during progress. During
pauses the first legs are usually elevated, and
they and the palps jerked rapidly up and
down. Both Ashtabula and Sassacus are mas-
ters of backward running, and both can
jump well, although they never resort to it
except in crossing gaps and in the final stage
of prey capture.
Courtship Display : Stage I. Male carapace
well elevated, abdomen swung to one side
(usually the left) , where it is held low, prac-
tically resting on ground; the spider sidles
back and forth, raising the front legs at a
wide angle and waving them up and down
in unison. The palps occasionally jerk up and
down, but hang quietly during height of dis-
play. The white abdominal stripe and its
bounding iridescence show clearly, little im-
peded by the short, pale, posterior legs. When
the attention of a female has been gained,
her pale, greenish-yellow palps jerk up and
down rapidly and almost continuously, being
conspicuous against her dark brown clypeus
and mouthparts. Stage II. Not seen.
Threat Display : Inter-male display seems
feebly developed in this species; three dif-
ferent pairs of males at various times, all in
display condition, judging by their behavior
toward females, paid little or no attention
to each other, except for some brief elevation
of the forelegs, which frequently takes place
in any situation and appears to be of an ex-
ploratory nature.
Habitat : Known only from the montane
cloud forest (about 3,600 feet) around
Rancho Grande. Always taken on herbs,
shrubs and low trees.
Affinities: Close to A. dentata; see re-
marks under Structure. It seems likely that
dentata, dentichelis and furcillata will prove
to be no more than subspecies of zonata.
Material : A total of 5 adult males and 1
adult female have been preserved. The fol-
lowing have been designated as types:
HOLOTYPE : Male. Cat. No. 461203, De-
partment of Tropical Research, New York
Zoological Society; Portachuelo, Rancho
Grande, near Maracay, National Park of
Aragua, Venezuela; 1,136 meters; cloud for-
est; June 15, 1946.
PARATYPE: Female. Cat. No. 481559,
Department of Tropical Research, New York
Zoological Society ; same locality as holotype ;
July 21, 1948.
The name furcillata is proposed in refer-
ence to the characteristic forked tip of the
palp’s tibial spine.
Sassacus ttavicinctus sp. nov.
(Text-fig. 5).
Diagnosis : Male black with yellow on cly-
peus, in paired stripes and a submarginal
band on carapace, and in transverse mark-
ings on abdomen. Female brown with obscure
ochraceous markings. Chelicera of male
strongly produced, the promargin with two
teeth, far separated, the retromargin with a
single strong tooth near distal end. Tibial
apophysis of palp strong, simple, tapering,
tip slightly recurved ; embolus curved.
Color.
Color in Life : Adult male. Cephalothorax :
Integument of carapace black, with a mod-
erate number of long bristles in ocular re-
gion, and with lemon yellow (Ridgway)
scales arranged in dense bands as follows:
A pair on carapace just below dorsal eyes,
converging slightly behind them and ending,
without meeting, halfway down thoracic
slope; a narrow submarginal band; a well-
developed band of scales and scale-hairs
42
Zoological New York Zoological Society
[34 : 7
Text-fig. 5. Sassacus flavicinctus. A-E, holotype $: A, carapace and abdomen, dorsal
view; B, carapace, lateral view; C, chelicera, ventral view; D, palp, ventral view;
E, same, ectal view. F, paratype $: epigynum.
completely covering and slightly pendent
from the narrow clypeus. Mouthparts and
legs black except as noted below; all tarsi
brown ; tibia and metatarsi of all except first
legs banded brown and black in varying pro-
portions ; all legs with small anterior patches
of yellow and white hairs and scales on some
or all of the following segments: Femur,
patella and tibia ; these markings are highly
variable. Sternum black with white hairs,
which occur also on underside of coxae. Ab-
domen : A basal semi-circular band of lemon
yellow scales continuing backward a third
of abdominal length; behind this two pairs
of short, curved bars, concave posteriorly,
of which the posterior pair may join in the
midline ; at tip of abdomen a tiny round spot,
or a short bar concave posteriorly, may be
present or absent. Center black with a tri-
angular patch of white hair, the apex pos-
terior.
Adult Female. Cephalothorax : Carapace
black with rather weak markings of ochra-
ceous brown scale-hairs as follows: Across
clypeus and completely encircling sides of
carapace and thoracic slope, absent only in j
middle of ocular quadrangle. Palps dark with
yellowish hairs. Legs banded light and dark
brown. Sternum black.
Abdomen : Dorsum with an indistinct, in-
terrupted reticulated pattern which consists
basically of an anterior basal band, followed
by several pairs of hollow bands ; the latter
do not meet in midline, but join with the pre-
ceding band by a narrow stripe just before
the center; tip of abdomen covered with
1949]
Crane: Salticid Spiders; Systematics and Behavior in New Species
43
ochraceous hairs. Venter black with a few
scattered light hairs.
All scale-hairs easily removed, and fre-
quently absent in preserved specimens.
Structure.
The characteristics below apply to both
males and females unless otherwise specified ;
percentages approximated ; measurements of
types given below.
Carapace : Height about half (female) or
slightly less than half (male) of carapace
length; anterior part of thorax flat, with a
very gentle slope, rounding into rounded
sides of cephalic part; descent of posterior
part (less than half postocular length)
abrupt, slightly concave; width of carapace
greatest a little behind PLE, 1.5 times height,
67% (male) to 75% (female) of carapace
length ; thoracic groove scarcely indicated.
Eyes: Length of ocular quadrangle about
58% as long as broad, its sides almost par-
allel, though very slightly wider at ALE than
at PLE ; carapace extending slightly beyond
PLE at their level, PME slightly nearer ALE
than PLE. Diameter of AME about 20% of
carapace length; ratio of eyes, holotype;
AME: ALE: PME :PLE : : 100 :48 :8 :44.
AME practically contiguous, separated from
ALE, which are slightly recurved, by about
an eighth of their diameter.
Clypeus : Height 12% of AME diameter.
Chelicerae : In males strongly produced,
held almost parallel to ground, divergent;
length of basal segment about three-fifths of
carapace length. Promargin with one slender
tooth at proximal end of groove and one,
long, robust, triangular, far removed, near
base of fang; slightly proximal to this on
retromargin a single large tooth. Fang slen-
der, slightly sinuous. Chelicerae of females
much shorter with a very short groove
flanked on promargin by two teeth close to-
gether, the proximal the smaller, and one
large tooth on retromargin.
Maxillae: Width about 75% of length;
outer distal edge in male more dilated and
obtusely angled than in female.
Lip: Breadth more than 90% of length;
distal end reaching slightly beyond middle
of maxillae; sternal suture curved, especi-
ally in male.
Sternum: Width 62% of length in males;
wider, about 73%, in females. Anterior mar-
gin concave, narrower than base of lip;
greatest width between first and second legs ;
posterior end bluntly pointed, extending
slightly between fourth coxae; the latter
separated by less than half their diameter.
TABLE V.
S. flavicinctus : Leg Formula.
1 4 2 3
Male holotype 1.8 1.5 1.4 1.35
2 1 3 jl
1.4
Legs : Tibial indices : Holotype male, first
leg 16, fourth leg 17.5; paratype female, first
leg 26, fourth leg 23. First femur in both
sexes enlarged, and entire first leg somewhat
thickened and elongated in male. See Table V
for formula. All legs with little hair.
Spines: (From male holotype and female
paratype). Patella without spines through-
out. First leg : Femur, dorsal 3 in distal half ;
prolateral distal 1 in male, 2 in female; tibia
ventral only lr-2-2, the two distal pairs close
together, the proximal at beginning of sec-
ond quarter of segment; metatarsus, ventral
only, 0-2-2. Second leg differs from first in
having tibia ventral lr-lr-2, (male) or
lr-0-2 (female) ; tibia prolateral, male only,
1-1 (both small) ; metatarsus, female only,
with 1 prolateral distal. Third leg, femur,
dorsal 0-1-1-1, prolateral distal 2 (male),
or 1 (female) ; tibia prolateral 0-1 (male)
or none (female) ; retrolateral 0-1; ventral
0-0-2 (male) or lp-lp-2 (female) ; meta-
tarsus prolateral distal 2, retrolateral distal
2, ventral distal 2. Fourth leg, femur as in
third; tibia prolateral none (male), or 0-1
(female) ; retrolateral 0-1 or none (variable
on two sides) ; ventral lp-0-2 or lr-lr-2 or
0-0-2 (variable on two sides) ; metatarsus
prolateral distal 0-1, sometimes in female
only 0-2, the second weak; ventral distal
only 2, on one side of female 0-2-2.
Abdomen: Ovate in both sexes, the breadth
about 70-75% of length, widest near middle.
Palp: Femur strongly curved; tibia more
than one-half length of patella; tibia with a
retrolateral apophysis which tapers to a
blunt, slightly recurved point. Embolus tap-
ering from a broad base to a curved and slen-
der tip.
Epigynum : An anterior pair of bodies well
separated, a posterior pair contiguous; mar-
ginal notch deep and narrow.
Measurements.
Male holotype. Total length in alcohol 4.51
mm.; carapace length 2.4, breadth 1.6, height
1.0; ocular quadrangle length .79, breadth
1.4; diameter AME .43, ALE .21, PME .03,
PLE .19; clypeus height .05; basal segment
of chelicera 1.37 ; patella breadth, 1st leg, .31,
4th .24.
Leg Measurements, Male.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
1.3
.85
1.0
.72
.44
4.3
2
1.1
.58
.62
.62
.41
3.3
3
1.1
.48
.55
.68
.34
3.2
4
1.2
.55
.82
.75
.34
3.7
Palp
.79
.27
.17
—
.58
1.8
Female paratype. Total length
in
alcohol
4.68 mm.; carapace length 2.05, breadth
1.54, height 1.03; ocular quadrangle length
.83, breadth 1.4; diameter AME .43, ALE
.21, PME .03, PLE .19; clypeus height .05;
basal segment of chelicera .72; patella
breadth, 1st leg, .34, 4th .26.
Female paratype
1.8
1.65 1.7
44
Zoologica: New York Zoological Society
[34: 7
1
Leg Measurements, Female.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
1.1
.65
.68
.55
.38
3.4
2
1.2
.51
.85
.72
.38
3.7
3
1.2
.51
.79
.68
.34
3.5
4
.92
.55
.55
.48
.34
2.8
Palp
.55
.24
.24
—
.38
1.4
Behavior.
Locomotion : Compared with Ashtabula,
this Sassacus is somewhat more a jumper
and walker, less a scurrier; also it palpates
the ground far less with the first legs and
palps. Compared to the spiders of the Plexip-
pus group, however, it is a poor and reluctant
jumper.
Courtship Display. Stage I. Male follows
female about, the carapace moderately ele-
vated and the first legs raised at a wide angle
to each other; frequently lowered; the ab-
domen hangs down and is trailed inconspicu-
ously from side to side with sideling. Display
tends to be in a wide semi-circle around fe-
male, once her attention has been attracted.
The long chelicerae are folded but held out
laterally (when not displaying they are held
at right angles to each other) , and the palps
extend straight out also, in contrast to their
usual resting position when they hang over
chelicerae. With increasing stimulation, zig-
zagging becomes more pronounced and a
slow rocking is involved, the carapace and
abdomen held stiffly and rocking as a unit.
Stage II is usually attained within three to
five minutes by couples of low threshold to
display stimuli, and consists of the first legs
thrust out in front, clear of the ground.
Threat Display: Stage I. Indistinguish-
able from Stage I of courtship, except that
no rocking is involved. Stage II. It is only
in the rare occurrence of this stage that the
chelicerae blades are unsheathed ; when two
opponents are practically touching the first
legs are brought upright, from the obliquely
outward display position, and simultaneously
the chelicerae blades are extended straight
out in front, at right angles to the basal seg-
ment, which is maintained in the horizontal
position typical of display. In each of the
dozen or so observed encounters that reached
this stage, one or the other male usually
backed off promptly at this point; more rare-
ly there was a brief tangle which ended with-
out apparent injury. Usually one or both
males retreated before reaching Stage II.
Habitat: Known only from the montane
cloud forest (about 3,600 feet) around
Rancho Grande. Always shaken from herbs,
shrubs and low trees.
Affinities: This species appears exceed-
ingly close to S. arcuatus Simon, 1902, from
Teffe, in the Amazon region. From the brief
description, the only apparent differences are
slight distinctions in the abdominal mark-
ings and the absence, in the present form,
of a yellow spot on the palp femur.
Material: A total of 5 adult males and 5
adult females have been preserved in addi-
tion to a number of young. The following
have been designated as types:
HOLOTYPE: Male. Cat. No. 45451. De-
partment of Tropical Research, New York
Zoological Society; Portachuelo, Rancho
Grande, near Maracay, National Park of
Aragua, Venezuela; 1,136 meters; cloud for-
est; June 1, 1945.
PARATYPE: Female. Cat. No. 45452 De-
partment of Tropical Research, New York
Zoological Society ; same locality as holotype
(with which she mated) ; July 1, 1945.
The name flavicinctus is proposed in ref-
erence to the yellow bands characterizing the
male.
Sassacus oce/fatus sp. nov.
(Text-fig. 6).
Diagnosis: Both sexes iridescent green
above, with a pair of black spots, each
crossed by a white bar, near tip of abdomen.
Chelicera of male strongly produced, the pro-
margin with two well-separated teeth in
proximal half, opposed by a single large
tooth on retromargin. Spines of first tibia
2-2-2. Tibial apophysis of palp strong, sim-
ple, tapering, tip straight; embolus tip
straight.
Color.
Color in Life : Adult male. Cephalothorax :
Integument of carapace black; ocular region
with a number of long bristles and completely
covered with iridescent green scales which
extend a little below it on sides and thoracic
region. A broad band of white hairs, starting
below PME on side of carapace, extends for-
ward across clypeus. Palps, mouthparts and
first legs jet black; other legs brown; two
narrow, conspicuous stripes of white scales
extend along anterior and posterior sides of
first patella, tibia and base of metatarsus.
These scales, although progressively fewer
posteriorly, are present on anterior sides
of all other legs, as well as on posterior sides
of second legs. Sternum black. Abdomen en-
tirely covered above, except as hereafter
noted, with iridescent green scales, larger
than those on carapace. On dorso-lateral sur-
face on each side of posterior third is a large
spot of velvety black scales, each with a nar-
row cross-bar of white scales from one-third
to two-thirds of the way to its posterior
edge. Around the entire abdomen laterally is
a narrow band of iridescent green, confluent
except in region of spot, with the dorsal
green. Venter black.
Adult female. Cephalothorax : carapace as
in male, with the addition of a narrow sub-
marginal border of white scales continuing
almost as far as pedicel. Entire face, around
eyes, with more white scales and hairs than
in male. Chelicerae black with a few white
hairs basally; palps translucent brown
barred narrowly with darker on joints, and
with a few white hairs on patellae. All legs
translucent brown except first femora, which
are almost black. Sternum black. Abdomen
as in male, except that there is a faint an-
1949]
Crane: Salticul Spiders; Systematics and Behavior in New Species
45
Text-fig. 6. Sassacus ocellatus. A-E, holotype $: A, carapace and abdomen, dorsal view;
B, carapace, lateral view; C, chelicera, ventral view; D, palp, ventral view; E, same,
ectal view. F, paratype $: epigynum.
terior band of white scales, dying out later-
ally in variable faint spots, while the white
cross-bars on the posterior black spots tend
to be on the latter’s anterior margin.
In alcohol the iridescent green completely
vanishes, the scales appearing dull yellowish
or brownish ; the abdominal black spots with
white cross-bars are discernible, but far less
distinct than in life, the anterior part of the
spot tending to disappear altogether. As us-
ual, the black integumentary areas fade to
brown.
Structure.
Essentially as in S. flavicincta except in
the following respects: carapace lower, its
height less than half carapace length in both
46
Zoologica: New York Zoological Society
[34: 7
sexes, lower in male than in female. Cheli-
cera of male even longer in some specimens,
but varying in individuals; basal segment
in holotype is 5/6 of carapace length; pro-
margin with two small teeth well separated,
along proximal half of groove; opposite their
interspace, on retromargin, is a single, much
larger, conical tooth. Tibial indices : Holo-
type male, first leg 21, fourth leg 19; para-
tvpe female, first leg 25, fourth leg 19. See
Table VI for formula.
TABLE VI.
S. ocellatus: Leg Formula.
1 4 2 3
Male holotype 1.9 1.5 1.4 1.2
14 2 3
Female paratype 1.7 1.7 1.3 1.3
Spines : As in flavicinctus, but with first
tibial ventral 2-2-2, not lr-2-2, and with
spines on posterior legs somewhat fewer,
viz.: Second leg: Male, tibia prolateral 0,
not 1-1 ; female as in flavicinctus. Third leg :
Male, femur prolateral distal 1, not 2; fe-
male, prolateral 0, retrolateral 1; tibia 0 in
both sexes, not with a few pro- and retro-
laterals and ventrals; metatarsus (both
sexes) pro- and retrolateral distals each 1,
not 2. Fourth leg (both sexes) : Femur pro-
lateral 1 not 2 ; tibia, ventral distal only lp in
male, 0 in female; metatarsus, as in third
leg, but with traces of another lateral distal
pair (very weak), similar to those in flavi-
cinctus; especially noticeable in female.
Palp: Tibial apophysis and embolus both
straight, not curved. Epigynum : Radically
different from that of S. flavicinctus (see
figure) ; marginal notch broad and shallow.
Measurements.
Male holotype. Total length in alcohol 3.3
mm.; carapace length 2.1, breadth 1.5, height
.79; clypeus height .07; basal segment of
chelicera 1.8; patella breadth, 1st leg .34,
4th leg .22; length of abdomen 2.2, breadth
1.4.
Leg Measurements, Male.
Leg
Femur Pat.
Tib.
Metat. Tarsus Total
1
1.2
.72
.92
.65
.44
3.9
2
.89
.55
.62
.58
.38
3.0
3
.82
.44
.51
.48
.38
2.6
4
.99
.51
.68
.65
.38
3.2
Palp
.82
.55
.14
—
.58
2.1
Female
paratype. Total length
in
alcohol
5.0 mm. ; carapace length 1.7, breadth 1.3,
height .72 ; clypeus height .10 ; basal segment
of chelicera .58; patella breadth, 1st leg .31,
4th leg .21 ; length of abdomen 3.3, breadth
2.1.
Leg Measurements, Female.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
.85
.62
.62
.48
.34 2.9
2
.68
.48
.41
.38
.31 2.3
3
.68
.41
.41
.44
.31 2.3
4
.89
.48
.65
.55
.37 2.9
Palp
.44
.21
.21
—
.34 1.2
Behavior.
Locomotion: About midway between Ash-
tabula and S. flavicinctus. Its usual progress
is a rapid scurry, jumping only when neces-
sary, the first legs held forward, usually
scarcely touching the ground, the palps held
just clear of it. During the infrequent
pauses, the first legs and palps are raised in
the air and waved up and down ; after which
both sets of appendages sometimes palpate
the ground itself.
Courtship Display: Stage I. Carapace
scarcely elevated, first legs held up at about
right angles to each other, and brought to
ground again during the jerking, zig-zag
approach to female. The long chelicerae are
sheathed, the palps hanging quietly over
them in the normal resting position, except
for occasional vibration. Approach to the fe-
male is often quick and direct after the pre-
liminary zig-zags. The most interesting
phase may or may not be included; it con-
sists of posing for a few moments, motion-
less, the legs elevated, and the abdomen
twisted slightly to one side or the other; once
the female was seen to perform the same mo-
tion, although that courtship was not com-
pleted. The relatively short abdomen was
never swung far to the side as in the elon-
gate Ashtabula, and the black, white-barred
terminal spot could not have been in full
view. In the single courtship which ended
in actual mating, this phase was altogether
omitted. Stage II. This was often reached
within three minutes ; in one case mating fol-
lowed five minutes after display began. It
did not differ from that of flavicinctus.
Threat Display: True fighting frequently
takes place in this species and even when
inter-male display ends in mere threat, the
chelicerae are always more or less un-
sheathed, which never happens in courtship.
The behavior otherwise is similar except that
I observed little or no trace of the side-
swinging of the abdomen. During actual bat-
tle the first legs are raised directly overhead,
and the palps extended laterally, wide-
spread, out of the way; the wide-open cheli-
cerae are opposed to those of the opponent.
The two may then push back and forth for
seconds, until one of the pair retreats or is
bitten.
Habitat: Known only from the montane
cloud forest (about 3,600 feet) around
Rancho Grande. Always shaken from herbs,
shrubs and low trees.
Affinities: The abdominal markings are
somewhat similar to those of S. aurantiacus
Simon, 1902, from Para, Brazil, known only
from the briefly described female. The pres-
ent species has a full set of 2-2-2 spines on
the first tibia, instead of lp-2-2, in both sexes.
Material: A total of 11 adult males and 9
adult females have been preserved in addi-
tion to a number of young. The following
have been designated as types:
HOLOTYPE : Male. Cat. No. 461204, De-
partment of Tropical Research, New York
1949]
Crane: Salticid Spiders; Systematics and Behavior in New Species
47
Zoological Society; Portachuelo, Rancho
Grande, near Maracay, National Park of
Aragua, Venezuela; 1,136 meters; cloud for-
est; March 27, 1946.
PARATYPE: Female. Cat. No. 481560,
■ Department of Tropical Research, New York
Zoological Society; same locality as holo-
type; July 17, 1948.
The name ocellatus is proposed in refer-
ence to the eye-like abdominal markings.
Phiale Homme a sp. nov.
(Text-fig. 7).
Diagnosis : All carapace bands in both
sexes creamy yellow. Male : Carapace mark-
ings broad, including submarginal and cly-
peal bands and mid-dorsal stripe; no spots
near PME. Abdomen above bright rufous
with white markings; median spot absent,
although a faint cross-bar may be present or
absent beneath rufous scales; no terminal
hook on antero-lateral band; three terminal
spots. Palp with tibial apophysis stout, trun-
cate; bulb strongly bilobed; lateral process
of embolus shorter than and widely separated
from embolus proper. Female: Carapace
markings less extensive than in male. Ab-
domen with reddish scales ranging almost
to black; anterior abdominal band as in
male; strong, post-median cross-bar and
terminal spots present. Epigynum with two
strongly chitinized, external cross-bars.
Color.
Color in Life: Adult male. As in Chick-
ering’s description of P. aliceae in alcohol
(1946, p. 207), except as follows: Cephalo-
thorax : Integument of carapace, mouthparts,
palps and first legs (except metatarsus and
tarsus) black, not dark brown; integument
of other legs translucent, medium brown. All
carapace scale-hair bands distinctly buffy
yellow; anterior eyes rimmed with rust; cly-
peus with a strong band of creamy yellow
scalerhairs, instead of only “a fringe of yel-
lowish bristles;” palp femur with dorsal
scale-hair patch as in aliceae; a patch of
white-scale hairs on proximal anterior face
of first metatarsus and tarsus; variable num-
bers and arrangements of similar scales,
diminishing posteriorly, on other segments
of other legs. Abdomen: Dorsum in full sun-
light often matches the flame scarlet of Ridg-
wayj other individuals tend to orange rufous.
As in aliceae, white markings consist of a
simple anterior band extending dorso-later-
ally more than halfway to spinnerets, and
ending without a hook-shaped inward curve
Text-fig. 7. Phiale fiammea. A-E, holotype $: A, carapace and abdomen, dorsal view;
B, carapace, lateral view; C, palp, ventral view; D, same, ectal view; E, same, tibial
apophysis. F, G, paratype ?: F, carapace and abdomen, dorsal view; G, epigynum.
48
Zoologicn: New York Zoological Society
[34: 7
(as is characteristic of P. dybowskii, for ex-
ample) ; usually it ends abruptly; sometimes
there is a very slight inward curve. The
“narrow, light-colored central bar” of aliceae
is invisible in live specimens though it some-
times shows in preserved examples, beneath
the rufous scales. Three small white terminal
markings, in the form of spots or short bars,
as in aliceae; carapace stripe easily rubbed,
often small in preservative.
Adult female. Exceedingly variable, both
in the pattern of white and dark scales, and
in the vividness of the reddish abdominal
markings ; the individuals are separated with
difficulty in pattern from at least two other
species occurring typically on the lower
slopes of the same mountain range. They dif-
fer from the male as follows : Cephalothorax :
buff stripe and bands of carapace — median,
submarginal and clypeal — much less exten-
sive; sparse rusty hairs usually present on
and around ocular quadrangle; anterior eyes
rimmed with yellowish-white, not rust; some
buff hairs on face below ALE; palps trans-
lucent buffy yellow, not black, and lacking
buff scales; first legs black only on femur
and patella; white hairs and scales of all legs
reduced or absent. Abdomen : Red of dorsum
exceedingly variable, practically always less
bright than in male, sometimes almost black.
A strong post-median, black-bordered cross-
bar of white scales always present, but of
variable length and breath, sometimes con-
fluent with ends of anterior dorso-lateral
band, which is as in male; posterior spots
present as in male, but of more variable
size and shape, sometimes partly confluent.
Structure.
This species is so close to P. aliceae (known
only from holotype male) that no significant
structural differences emerge from a com-
parison of Chickering’s description with our
species, except for minor spine and palp dif-
ferences as given below. The females are
closely similar to the males in structure, ex-
cept for the usual leg differences, and for
the absence of the small hooked maxillary
process.
Spines (both sexes) : Differ from aliceae
as follows: First leg, Female: Patella pro-
lateral 0, not 1. Second leg, both sexes : Tibia
prolateral as in first (1-0-1, not 1-1-1), ven-
tral apparently consistently lr-2-2, not vari-
able; metatarsus male, prolateral distal 0,
not 1, but this spine present in female. Third
leg (female only) : Femur prolateral distal
only 2, not 1-2, retrolateral 1, not 2; tibia
dorsal 0, not 1 ; metatarsus with slight ir-
regularities on one side of paratype female
only, retrolateral 0-1-2, not 1-1-2, ventral
lp-lp-2, not 0-2-2. Fourth leg: Femur (both
sexes) prolateral and retrolateral distal re-
spectively 0 and 1, not each 2; male tibia as
on right side of aliceae holotype, female dor-
sal 0, not 1.
Palp\ Differs from that of aliceae in its
relatively greater breadth and in the char-
acter of lateral process of embolus; in
flammea the two parts are much farther
apart, though connected by a thin, horny
plate ; also, the lateral process is much short-
er than embolus proper, and scarcely curved
distally.
Epigynum : Confusing, as usual in this
genus, on account of the frequent secretion
of gummy matter which obscures and dis-
torts the structure. Always distinct, how-
ever, are two strongly chitinized transverse,
lip-like structures, one between the two pairs
of subdermal bodies and one near posterior
border.
Measurements.
Male holotype. Total length in alcohol 4.6
mm. ; carapace length 2.7, breadth 1.9, height
1.1; clypeus height .19; basal segment of
chelicera .89; patella breadth. 1st leg, .41,
4th .28; length of abdomen 1.9, breadth 1.4.
Leg Measurements, Male.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
1.8
1.1
1.7
1.1
.68
6.4
2
1.3
.75
.89
.79
.48
4.2
3
1.5
.79
.85
1.1
.55
4.8
4
1.6
.75
1.2
1.3
.55
5.4
Palp
.85
.24
.24
—
.82
2.2
Female paratype. Total length in alcohol
5.1 mm.; carapace length 2.5, breadth 1.7,
height 1.1 ; clypeus height .07 ; basal segment
of chelicera .85; patella breadth, 1st leg .40,
4th .31 ; length of abdomen 2.6, breadth 1.7.
Leg Measurements, Female.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
1.2
.85
.89
.65
.51
4.1
2
1.1
.65
.65
.62
.48
3.5
3
1.3
.72
.79
.82
.55
4.2
4
1.4
.72
.99
1.1
.62
4.8
Palp
.65
.27
.31
—
.55
1.8
Tibial indices : Holotype male first leg 15,
fourth leg 14; paratype female, first leg 23, ,
fourth leg 18. See Table VII for formula, i
TABLE VII.
P. flammea: Leg Formula.
4 3 2
Male holotype 2.4 2.0 1.8 1.5
J. 1 3_ 2
Female paratype 1.9 1.6 1.7 1.4
Locomotion : Primarily a runner, although
jumps are undertaken over gaps without
hesitation. The first legs take little part in
locomotion and are habitually waved up and
down during the pauses.
Courtship Display : Stage I. Carapace ele-
vated high; abdomen hangs down, usually
touching ground and leaving a silk thread.
First legs raised at 45° angle with each other
and the ground. Female approached in zig-
zag spurts, as the carapace is rocked from
side to side, sinking alternating almost to
the ground, from right to left. Palps irregu-
larly vibrated up and down. Pursuit of fe-
male plays an important part in early stages,
1949]
Crane: Salticid Spiders; Systematics and Behavior in Neiv Species
49
but once female’s attention is gained, she
usually watches with first legs elevated and
palps vibrating rapidly.
Stage II. Male abruptly crouches almost
on ground, when two inches or less from
female; his legs far outstretched in front,
almost parallel, he approaches her directly
with crawling motion, the palps vibrating in
unison and entire body quivering. The re-
markable point about Stage II in this species
is that it begins at such a relatively long
distance from the female.
Threat Display. As in Stage I of court-
ship, except that the palps are held quiet
most of the time, the creamy yellow patch of
the curved femur continuing that of the cly-
peus in an unbroken line. When approach is
very close the chelicerae are opened and the
first legs spread more widely, often actually
touching those of the opponent. The bouts
are always brief and I have never seen dam-
age inflicted.
Habitat : Known only from the montane
cloud forest (about 3,600 feet) around
Rancho Grande. Always taken on herbs,
shrubs or small trees.
Affinities : The closeness of this species to
P. aliceae has already been noted. When ade-
quate material is taken from intermediate
localities, it seems likely that the distinc-
tions will prove to be of only subspecific im-
portance.
Material : A total of 14 adult males and
20 adult females have been preserved. The
following have been designated as types :
HOLOTYPE: Male. Cat. No. 481561, De-
partment of Tropical Research, New York
Zoological Society; Portachuelo, Rancho
Grande, near Maracay, National Park of
Aragua, Venezuela; 1,136 meters, cloud for-
est; July 25, 1948.
PARATYPE : Female. Cat. No. 45453, De-
partment of Tropical Research, New York
Zoological Society; same locality as holo-
type; July 26, 1945.
The proposed name flammea refers to the
color of the male dorsum.
Maqo dentichelis sp. nOV.
(Text-fig. 8).
Diagnosis: Carapace of unrubbed individ-
uals with a median white stripe enclosing a
central black spot. Male chelicera with tooth
on external border; four or five teeth on in-
ferior margin; two or three teeth, plus a
series of denticles, on superior margin ; tibia
of palp with three unequal apophyses;
epigynum with a median, rounded, super-
ficial, pale anterior body.
Color.
Color in Life: Adult male. Cephalothorax :
Carapace integument black, practically
naked except for a conspicuous median stripe
of white scales enclosing, behind level of
PLE, a central black spot. The stripe begins
behind AME, or near level of PME, widens
to encompass the spot, then narrows once
more, ending at or behind middle of thorax.
White of spot region sometimes extending
laterally as a short cross-bar. Sparse chest-
nut and black hairs scattered on ocular
quadrangle near dorsal eyes, and around
AME. The wide clypeus is black and com-
pletely naked; palps, mouthparts and first
pairs of legs black, except for leg tarsi. These
and entire third and fourth legs translucent
brown, variably and faintly banded with
darker near ends of segments. Palps and all
legs, especially first two, with inconspicuous
white scale-hairs on antero-dorsal surfaces
near joints. Sternum black. Abdomen: Pat-
tern of dorsum very variable, formed chiefly
of short hairs or scale hairs, brown mixed
with gray and white areas. Usually a white
lyre-shaped anterior marking — a strongly
curved band with a short median basal stripe
— is distinct; this is followed by several pairs
of faint chevrons and some white lateral
streaks and spots. The most constant mark-
ings are a pair of white terminal spots. Ven-
ter black with a pair of pale faint longi-
tudinal stripes in middle; buff hairs rather
thickly scattered over entire surface.
Adult female. Dorsal markings very simi-
lar to those of male, but posterior abdominal
spots less distinct and more variable. Palps
pale, translucent horn; first and second legs
banded, not black; white scale-hairs on ap-
pendages almost or completely absent,
though short yellowish hairs sometimes pre-
sent near joints.
In alcohol, the distinctive markings usu-
ally disappear from both sexes.
Structure.
The characteristics below apply to both
males and females unless otherwise speci-
fied ; percentages approximated ; measure-
ments of types given on p. 51.
Carapace : Height 57 % of carapace length ;
profile rises behind AMF, gently convex, to
PLE; anterior half of thorax descends very
gently, posterior half abruptly; widest at
level of PLE, 1.3 times height, 73% of
length; total length of eye group slightly
more than half carapace length. A distinct
longitudinal thoracic groove, centering at
level of posterior margin of PLE.
Eyes: Length of ocular quadrangle about
two-thirds of breadth, its sides almost paral-
lel but width at ALE slightly greater than
at PLE ; carapace extending well beyond
PLE at their level; PME slightly closer to
ALE than to PLE. Diameter of AME 23%
of carapace length; ratio of eyes, holotype:
AME : ALE : PME : PLE :: 100:46:14:40.
AME practically contiguous, separated from
ALE, which are slightly recurved, by about
a tenth of their diameter.
Clypeus : Height 52% of AME diameter in
male, 28% in female.
Chelicerae: Not produced, vertical, paral-
lel. Length of basal segment less than 30%
of carapace length. Male with a strong tooth
about middle of external border. Promargin
50
Zoologica: New York Zoological Society
[34: 7
Text-fig. 8. Mago dentichelis: A-E, holotype $: A, carapace and abdomen, dorsal view;
B, carapace, lateral view; C, chelicera, ventral view; D, palp, ventral view; E, same,
ectal view; F, threat display. G, paratype $: epigynum.
with two (rarely three) moderate-sized teeth
at proximal angle, the distal the larger;
distal to these is a series of minute granular
teeth, numbering three or more. Inferior
margin usually with four, sometimes five,
contiguous, well developed teeth.
Maxillae : Less than twice as long as wide,
outer distal angle little dilated.
Lip: Length and breadth similar; pos-
terior margin slightly convex, about equal
in breadth to anterior margin of sternum.
Sternum: Breadth three-fourths of length
in male, two-thirds in female, widest at an-
terior margin of third leg. Anterior border
concave, posterior broad and convex, ending
before anterior half of fourth coxae; pos-
terior half of latter separated by about an
eighth of their diameter.
Legs: Tibial indices: Holotype male, first
leg 17, fourth 29; paratype female, first leg
23, fourth 16. First femur, patella and tibia
moderately enlarged, less so in second leg.
See Table VIII for formula. All legs with
little hair.
19491
Crane: Salticid Spiders; Systematics and Behavior in New Species
51
TABLE VIII.
M. dentichelis : Leg Formula.
1 4 3 2
Male holotype 2.1 2.0 1.9 1.9
4 3 1 2
Female paratype 1.9 1.8 1.7 1.6
Spines: First leg: Femur dorsal 0-1-1-1,
prolateral distal only 2; patella prolateral
only 1 or 0; tibial prolateral 1-0-1 (both
weak), or 0-0-0; retrolateral 0; ventral,
lr-lr-2, or 2-2-2; metatarsus ventral only
2-2. Second leg: Femur dorsal 0-1-1-1, pro-
lateral distal only 2, retrolateral female only
1; patella prolateral 1 or 0; tibia prolateral
1-1-1 or 1-0-1, retrolateral 0, ventral lr-2-2;
metatarsus ventral only 2-2. Third leg:
Femur dorsal 0-1-1-1, prolateral 1 or 2, retro-
lateral 1 or 0; patella prolateral 1, retro-
lateral 1 ; tibia prolateral 1-1, retrolateral
1-1-1; ventral lp-0-2; metatarsus prolateral
1-1, retrolateral 1-2, ventral 2-2. Fourth
leg: Femur dorsal 0-1-1-1, prolateral 1, re-
trolateral 1 ; patella prolateral 1, retrolateral
1 ; tibia prolateral 1-1-1 or 1-1, retrolateral
1-1-1, ventral lp-2; metatarsus prolateral
1-1, retrolateral 1-1-2, ventral lp-2.
Abdomen: Rather narrowly ovate, widest
near middle.
Palp: Femur slightly curved, tibia about
70% length of patella; tibia with three apo-
physes, one small and ventral, one long and
tapering, external to the first, and the third
still larger, sinuously tapering, dorso-lateral.
Embolus short and simple.
Epigynum: A large, rounded, median,
whitish anterior area, followed by a variable
arrangement of four or five subdermal, near-
median tubules, related to two less distinct,
well separated oval bodies.
Measurements.
Male holotype: Total length in alcohol 5.2
mm.; carapace length 2.6, breadth 1.9, height
1.5; total length of eye group 1.4; ocular
quadrangle length 1.1, breadth 1.7 ; diameter
AME .60, ALE .28, PME .09, PLE .24; cly-
peus height .31 ; basal segment of chelicera
.99; sternum length .99, breadth .75; ab-
domen length 2.6, breadth 1.5; patella
breadth, 1st leg, .41, 4th .39.
Leg Measurements, Male.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
1.6
.96
1.4
.99
.58
5.5
2
1.4
.89
1.1
.92
.58
4.9
3
1.6
.75
1.1
1.1
.48
5.0
4
1.5
.68
1.2
1.2
.62
5.2
Palp
.89
.38
.27
—
.68
2.2
Female paratype: Total length in alcohol
5.3 mm.; carapace length 2.5, breadth 1.8,
height 1.4; total length of eye group 1.4; ocu-
lar quadrangle length 1.1, breadth 1.6;
diameter AME .55, ALE .26, PME .09, PLE
.24; clypeus height .15; basal segment of
chelicera .79; sternum length .96, breadth
.65; abdomen length 2.8, breadth 2.0; patella
breadth, 1st leg .40, 4th .28.
Leg Measurements, Female.
Leg
Femur
Pat.
Tib.
Metat. Tarsus Total
1
1.3
.79
.96
.72
.38
4.2
2
1.3
.79
.85
.65
.44
4.0
3
1.4
.79
.92
.89
.51
4.5
4
1.5
.65
1.1
1.1
.58
4.9
Palp
.68
.41
.34
—
.38
1.8
Behavior.
Locomotion: Not specially observed in this
species; however, another Mago (unde-
scribed) as well as Hypaeus sp. are both ex-
cellent jumpers. In these the repeated pat-
tern of ordinary progress is a deliberate walk
for two or three centimeters followed by a
series of short jumps; the first legs take ac-
tive part in the walking and jumping, and
are never raised except during display.
Courtship Display: Stage I. Carapace ele-
vated only enough so that the motionless,
hanging palps clear the ground; first legs
raised at a wide angle to each other (about
135°), the other legs extending far side-
wards, the second pair slightly forward. Pos-
ing in this attitude is extended, but at in-
tervals the first legs wave alternately up
and down. Meanwhile the abdomen, which
is held horizontally clear of the ground, is
occasionally vibrated briefly up and down.
Stage II. First legs extend to front, usu-
ally not befoi-e female thrusts her first legs
momentarily forward. Carapace and legs of
male, in addition to the abdomen, twitch
and jerk before he touches her.
Threat Display: Much more active than
courtship, and in several respects quite dis-
tinct. Stage I : Carapace held moderately
low, the abdomen either straight out as in
courtship, or relaxed downward for silk at-
tachment. First legs held with femur bent
obliquely up, the other segments out; from
that joint the two legs are waved up and
down, usually in unison with each other,
sometimes alternately. The palps hang down
outside the closed chelicerae, as in courtship.
Stage II. The tempo and span of waving
increases, the first legs almost meeting over-
head at peak of display. Series of waves are
punctuated by the rapid rubbing together of
the first and second tarsi of each side, the
second legs are braced somewhat forward,
as in courtship, and are occasionally lifted
briefly from the ground during waving.
Stage III. The two males oppose each other
closely, the first legs straight overhead, prac-
tically or completely touching, the palps
swung obliquely out, and the chelicerae
opened wide and knocking against each other
for seconds at a time. I have seen this stage
reached only twice, no injury being inflicted
either time. Only when one was retreating
did the abdomen twitch very briefly, as in
courtship.
Habitat: Known only from the montane
cloud forest (about 3,600 feet) around
52
Zoologica: New York Zoological Society
[34: 7: 1949]
Rancho Grande, taken from vines on tree
trunks, herbs and shrubs. Several specimens
collected on upper Rancho Grande verandah,
many yards from vegetation.
Affinities: Apparently related to Simon’s
briefly described longidens and acutidens
from Brazil, although distinct in details of
white markings, distal dentition of cheli-
cerae and presence of three apophyses on
palpal tibia.
Material: A total of 6 adult males and 11
adult females have been preserved in addi-
tion to a number of young. The following
have been designated as types:
HOLOTYPE: Male. Cat. No. 45454, De-
partment of Tropical Research, New York
Zoological Society; Portachuelo, Rancho
Grande, near Maracay, National Park of
Aragua, Venezuela; 1,136 meters; cloud for-
est; June 6, 1945.
PARATYPE: Female. Cat. No. 45455.
Taken near holotype, same locality and date.
The name dentichelis is proposed in refer-
ence to the large outer tooth of the chelicera.
References.
Beebe, W., and Crane, J.
1947. Ecology of Rancho Grande, a Sub-
tropical Cloud Forest in Northern
Venezuela. Zoologica, Vol. 32, No. 5,
pp. 43-60.
Bristowe, W. S.
1941. The Comity of Spiders. London, printed
for the Ray Society. Vol. 2.
Cambridge, F. 0. P.
1901. Arachnida. Araneidea and Opiliones,
Vol. II, in Biologia Centrali- Ameri-
cana.
Chickering, A. M.
1946. The Salticidae (Spiders) of Panama.
Bull. Mus. Comp. Zool., Harvard Coll.,
Vol. 97.
Crane, J.
1948.1 Comparative biology of salticid spiders
at Rancho Grande, Venezuela. Part I.
Systematics and life histories in Cory-
thalia. Zoologica, Vol. 33, No. 1, pp.
1-38.
1948.2 Comparative biology of salticid spiders
at Rancho Grande, Venezuela. Part II.
Methods of Collection, Culture, Obser-
vation and Experiment. Zoologica, Vol.
33, No. 9, pp. 139-145.
Mello-Leitao, C. de
1945. Aranas de Misiones, Corrientes y Entre
Rios. Rev. Mus. La Plata (n.s.), 4,
Zool., 29, pp. 213-302.
Peckham, G. W. and E. G.
1894. Spiders of the Marptusa group of the
family Attidae. Occ. Pap. Nat. Hist.
Soc. Wisconsin, Vol. 2, 1892-1895, pp.
85-141.
Simon, E.
1900. Descriptions d’especes nouvelles de la
famille des Attidae. Ann. Soc. Entom.
France, Vol. 69, pp. 27-61.
1901. Descriptions d’especes nouvelles de la
famille des Salticidae. Ann. Soc.
Entom. France, Vol. 70, pp. 66-76.
1902. Descriptions d’arachnides nouveaux de
la famille des Salticidae (Attidae).
Ann. Soc. Entom. Belgique, Vol. 46,
pp. 24-54.
Beebe: Swifts of Rancho Grande, and Their Migration
53
8.
The Swifts of Rancho Grande, North-central Venezuela,
with Special Reference to Migration.1
William Beebe.
Department of Tropical Research, New York Zoological Society.
(Plate I; Text-figures 1-3).
[This is one of a series of papers resulting
from the 45th, 46th and 47th Expeditions of the
Department of Tropical Research of the New
York Zoological Society, made during 1945, 1946
and 1948, under the direction of Dr. William
Beebe, with headquarters at Rancho Grande in
the National Park of Aragua, Venezuela. The
expeditions were made possible through the
generous cooperation of the National Govern-
ment of Venezuela and of the Creole Petroleum
Corporation.
[The characteristics of the research area are
in brief as follows: Rancho Grande is located
in north-central Venezuela (10° 21' N. Lat.,
67° 41' W. Long.), 80 kilometers west of Cara-
cas, at an elevation of 1,100 meters in the undis-
turbed montane cloud forest which covers this
part of the Caribbean range of the Andes. Ad-
jacent ecological zones include seasonal forest,
savanna, thorn woodland, cactus scrub, the
fresh water lake of Valencia, and various ma-
rine littoral zones. The Rancho Grande area is
generally subtropical, being uniformly cool and
damp throughout the year because of the preva-
lence of the mountain cloud cap. The dry season
extends from January into April. The average
humidity during the expeditions, including
parts of both wet and dry seasons, was 92.4% ;
the average temperature during the same period
was 18° C.; the average annual rainfall over a
5-year period was 174 cm. The flora is marked
by an abundance of mosses, ferns and epiphytes
of many kinds, as well as a few gigantic trees.
For further details, see Beebe & Crane, Zoolog-
ica, Vol. 32, No. 5, 1947. Unless otherwise stated,
the specimens discussed in the present paper
were observed or taken in or over the montane
cloud forest zone, within a radius of 1 kilometer
of Rancho Grande.]
Contents.
Page
Introduction 53
Streptoprocne zonaris albicincta (Cabanis, 1862) .... 64
Chaetura brachyura brachyura (Jardine, 1846) 57
Chaetura cinereiventris lawrencei Ridgway, 1893 58
Chaeturella rutila brunneitorques Lafresnaye, 1844 ... 58
Cypseloides cherriei Ridgway,1893 59
Cyp8eloide8 cryptus Zimmer, 1945 60
Aeronaute g montivagus montivagus (d’Orbigny and
Lafresnaye, 1837) 61
Panyptila cayennensis (Gmelin, 1789) 61
1 Contribution No. 841, Department of Tropical Research,
New York Zoological Society.
Introduction.
In all of South America there have been
recorded (Peters, 1940) nine genera of
swifts, divided into twenty species and a total
of thirty-three kinds, including subspecies.
In Venezuela Mr. William H. Phelps informs
me there are six genera, of thirteen species
of twenty kinds, if we include subspecies. Of
these Venezuelan birds, within an area of
less than one square kilometer with its cen-
ter at Rancho Grande, I have recorded eight
species of five genera; roughly eighty per
cent, of the genera and sixty per cent, of the
total Venezuelan species of swifts.
Late in the year 1937 Dr. Alexander Wet-
more (Wetmore, 1939) spent some time col-
lecting birds near Rancho Grande, and I
quote the following notes concerning the
swifts.
“In Tropical America swifts are tantaliz-
ing birds usually seen out of range ... On
November 4 at Rancho Grande several (Chae-
tura brachyura ) circled out of range. This
species appears very black as it flies over-
head, so that at first glance it suggests the
black swift ( Nephoecetes niger), but a sec-
ond look distinguishes it by the shorter, light-
colored tail. The specimen taken, a male,
measures as follows: Wing 118.7, tail 29.0,
culmen from base 5.8, tarsus 11.8 mm.”
Concerning Streptoprocne zonaris albi-
cincta, he writes, “While I was collecting in
Portachuelo above Rancho Grande on Novem-
ber 3, 6 and 10, groups of these large swifts
dashed at intervals through the pass at light-
ning speed with a great rushing of wings.
Occasionally I observed them circling in air.”
These are, I believe, the only published
notes on swifts in this restricted area.
My thanks go to Mr. William H. Phelps for
the loan of skins of rare swifts, to Dr. Neal
Weber for names of ants in the food of birds
taken in 1948, and to Dr. J. Bequaert for the
name of the feather fly found on Aeronautes.
The three text-figures are the work of Miss
Louise A. Moore. The photographs were
taken by Miss Jocelyn Crane.
54
Zoological New York Zoological Society
[34: 8
Streptoprocne zonaris albicincta
(Cabanis, 1862).
Giant White-collared Swift.
Species Range : Southern Mexico and the
Greater Antilles, south over northern South
America to British Guiana, north Matto
Grosso and Peru; vertically to more than ten
thousand feet in the Andes.
Subspecies Range: Five subspecies are
recognized, of which albicincta occurs at
Rancho Grande. Its range is extensive, from
Honduras south to British Guiana, northern
Matto Grosso and Peru, together with the
islands of Granada and Trinidad. In Vene-
zuela, Mr. Phelps records it as inhabiting the
northern mountains.
Field Characters for Sight Identification :
The most unmistakable species, distin-
guished by great size and white nuchal col-
lar. It measures eight to nine inches in
length, as compared to the five-inch average
of the seven other species. Panyptila is the
only other Rancho Grande swift with a white
collar, but is about half the size of albicincta,
and has a deeply forked tail. In young giant
swifts the collar is reduced and indistinct in
flying birds.
Occurrence: February 22 and September
9 are the earliest and the latest dates of our
occupancy of Rancho Grande throughout
three years. On both dates I recorded giant
swifts within sight of the laboratory. Seldom
did a day pass between these extremes when
one or more did not come into view. Soon
after we opened the station I ceased keeping
detailed notes on these birds, as their visits
seemed governed by no regularity.
They commanded attention under four sep-
arate conditions: (1) Almost daily either
singly, but usually in small flocks, they
hawked in the sky after insects, or (2) they
flew headlong through Portachuelo Pass, low
over the trees. (3) They entered rarely into
the diet of a pair of resident bat falcons,
Falco albigularis, and (4) on nights of storm,
rain or neblina they occasionally struck
against the windows of our lighted labora-
tory. Throughout the seven months during
which we carried on our observations, there
was no marked period of absence or extreme
scarcity of these swifts. The breeding period
must have occurred throughout part of this
time but it was not noticeable in the rai'ity
or abundance of individuals or flocks.
When it became evident that Portachuelo
Pass was used as a migrating flyway on an
unprecedented scale by other birds and by
insects, I watched and noted these passing
swifts for a period of several weeks, to see
if there was any definite factor or sequence
in their numbers or movements.
The daily, circling, feeding birds whose
general direction was indefinite, varied their
elevation, high or low, according to the volant
stratification of edible insects. This proved
to be definitely associated with the southward
migration of insects of many orders through
the pass. The swifts often joined flocks of
swallows and even of large dragonflies where,
on clear days, the migrants offered rich feed-
ing in the area of the pass. At times of dense
fog, high winds or lowering of temperature,
the lessening or cessation of migration was
correlated with a total absence of giant
swifts. At Kilometer 15, a few kilometers
south of the pass, I frequently saw flocks of
these birds feeding high in air as I passed
in the car; and to the north at Kilometer 30,
six or eight pairs of the swif ts were occasion-
ally seen hawking about. Beyond these limits
I saw no swifts.
Giant swifts are supposed to be normal in-
habitants of strictly tropical regions. At
Kartabo, British Guiana, at practically sea-
level, I found them commonly in good-sized
flocks, feeding on flying insects, especially
in June, July and early August. During this
season, mating flights of ants and termites
were frequent.
From March 14 to July 17 I noted the fol-
lowing groups of giant swifts passing on
twenty-three days at full speed south through
the pass, all between 7 and 8:30 A.M. 1, 16,
6, 2, 11, 4 and 16, 3 and 7, 21, 12, 8, 16,
4, 1, 3, 1, 2, 14, 5, 4, 22, 19, 7, 5. All were in
a terrific hurry, flying headlong, mostly low,
their whistling wings just clearing the upper
branches of bushes and trees. Throughout
this period there were only five records of
birds going north in early morning and few
in numbers, 2, 6, 1, 9, 1. On June 24 at 3 P.M.
64 swifts rushed past over my head, headed
full speed northward through the pass, just
ahead of the onrolling fog.
On June 6, at five in the afternoon, a com-
pact flock of 200 to 210 birds, at a moderate
height, circled northward, giving the impres-
sion of a leisurely, non-feeding migration.
On August 1, closely intermingling with
about five thousand Argentine martins,
Phaeoprogne tapera fusca, about 300 giant
swifts accompanied the other birds, all at
high speed. On August 8, 24 swifts passed
low, going north through the pass.
The assumption of the northward return
every afternoon and evening of these swifts
through Portachuelo Pass seems justified be-
cause of the number of birds which long after
dark on nights of storm or fog struck against
the windows of our laboratory. These acci-
dents occurred from 7:20 to 10:45 P.M.
Fourteen birds struck in this way on eleven
nights, April 9, 12, 18, May 4, 16, 23, June
10, 27, July 3, 4 and 6. On three nights two
birds appeared. Four of the swifts which
crashed the windows were skinned, three
others were sexed, and the remaining seven
escaped. All examined were males, and. of
those examined, only the two birds which
struck on April 9 were in full breeding con-
dition.
Reviewing the records through the pass,
it seems reasonable to assume a daily migra-
tion from some more northerly sleeping or
breeding place, south to a feeding area, with
the return very late in the afternoon or in
the evening.
1949]
Beebe: Swifts of Rancho Grande, and Their Migration
55
On May 4, four of these swifts fearlessly
attacked a bat falcon, the male of the pair
whose nesting we were watching. In connec-
tion with the attacking and repulsing of this
hawk by the the swifts, we were interested
to see the same individual falcon on three
separate occasions return to his lofty perch
with a dead swift. This is a remarkable feat
when we realize that the latter is only about
one-quarter less in size than the hawk. In an
active flight dive the falcon could strike and
capture any small bird it selected, but on a
level the swifts were superior.
In the Santa Marta mountains of Colom-
bia, about 575 kilometers west of Rancho
Grande and at an altitude of 1,500 meters,
this giant swift has been found nesting
(Todd and Carriker, 1922). The account is
as follows (p. 245) : At the coffee plantation
of Cincinnati, “on March 19, 1917, a colony
of this large swift was discovered nesting
in a shallow cavern behind a waterfall. The
place was absolutely inaccessible, so that no
idea of the number of nests could be had.
Only one nest, which happened to be near the
top, was secured, together with the occu-
pants, . . . which had been stunned by the
blasting, and proved to be an adult female
and two recently hatched young. The nest re-
sembled very closely that of the Chimney
Swift, being composed of twigs fastened to-
gether with saliva. The birds entered and
left the cavern by dashing through the cur-
tain of water falling over the front of it. The
altitude of the site was about 4,300 feet.”
Data on Collected Specimens.
For comparison I have included data con-
cerning a female of this species taken many
years ago at Kartabo, British Guiana.
hundred winged females of small
ants.
31188 : Stomach crammed with ants.
(Weber).
Homoptera: Cicadellidae.
Diptera: fly fragments.
Hymenoptera: parasitic sp.
Hymenoptera: Pheidole sp.
Hymenoptera: Atta sexdens Linn.
Four gasters and a hind wing frag-
ment. A species known from Ciudad
Bolivar, Venezuela, and south. Found
in Eastern British Guiana, but ap-
parently not in Venezuelan Guiana,
the Orinoco Delta, N. W. District
(B.G.) ; in these places replaced by
A. cephalotes Linn.
Aztecal : wings.
Camponotus ( Myrmobrachys ) sp.
Same as I took at 1,020 meters in
Rio Porce, Colombia.
Camponotus ( Tanaemyrmex ) sub-
stitutaus Emery. Distribution : Cen-
tral America to Paraguay. I have the
same form from Kartabo, B. G.
Camponotus ( Myrmobrachys )
crassus Mayr. Distribution: South
America.
529 : Three beetles, three wasps, one hem-
ipteron, three membracids, one tipu-
lid, and upwards of two hundred
female ants of six species.
It is significant that although swifts 30447
and 30452 struck the laboratory within six
minutes of each other, yet their food was
quite distinct, indicating very different feed-
ing territories. Yet they were headed for the
pass, focusing upon a sixty-foot-wide bottle-
neck.
30447
male
Lgth.
205
Wing
194
30452
male
196
188
31135
male
202
195
31188
male
210
200
529
female
190
190
The relative discrepancies between length
and weight are accounted for by the food.
The stomach, with contents, of No. 31188
weighed 22.6 grams. Gross food content may
be expressed as follows: 30447, crammed
with ants; 30452, moderately filled; 31135,
empty, after a day in cage; 31188, crammed
with ants; 529, only about one-fourth filled
with insects. Without exception, all the ants
in the food were winged females.
Detailed Food.
30447 : At least 800 ants of an undetermined
species of Azteca.
30452 : Several hundred females of Dolicho-
derus ( Monads ) debilis Emery, and
Crematog aster ( Orthocrema ) sp. A
single female Solenopsis geminata
edwardi Forel.
31135: Five female Atta sp. More than four
Tail
Grams
Weight
Extent
Date
64
105.8
501
July
3, 1945
61
96
482
July
3, 1945
67
68.5
500
April
9, 1948
70
109.5
July
6, 1948
60
77.2
July
10, 1919
Individual Characters.
I find the following recorded concerning
the Kartabo female, No. 529:
Parasites: Only a few bete rouge on the
head feathers.
Colors: Bill black, face pale medici blue,
iris light brownish-olive, legs and feet vina-
ceous slate.
Eyelid: Quite bare above. Below, a line of
fifteen small feathers along rim. At posterior
end of eye a small group of a dozen feathers,
arranged in several rows.
Oilgland: Elongated, blunt, tapering, bare.
Wing Graph: Primaries
10th — 152 mm. 5th — 111mm.
9th— 155 “ 4th— 98 “
8 th— 149 “ 3rd— 86 “
7th— 139 “ 2nd— 75 “
6th— 126 “ 1st— 62 “
56
Zooloffica: New York Zoological Society
[34: 8
Secondaries
1st — 49 mm.
2nd— 50 “
3rd— 52 “
4th— 54 “
5th — 53 mm,
6th— 53 “
7 th— 49 “
8th— 42 “
Scalation : Front of tarsus with an indis-
tinct irregular line of ten, fleshy scales down
the inner aspect. Inner, rear and outer sides
of tarsus, bare, wrinkled skin, with no trace
of scales.
Text-fig. 1. Streptoprocne zonaris albicincta Text-fig. 2. Streptoproc7ie zonaris albicincta
(Cabanis). Scalation of tarsus, front and side (Cabanis). Tongue.
views.
Palate : The palatine fissure begins well
toward the front of the roof of the mouth,
and divides in front. It is rather long (12.5
mm.) and is guarded by about a dozen pairs
of teeth. Four-fifths of the way back there
is a double-curved or angular transverse row
of teeth, extending out at right angles on
each side, with about twenty teeth on each
side. The guardian, denticulated flaps end
openly posteriorly, and just beyond is the
very small tubal fissure. At the extreme pos-
terior of the roof of the mouth is a transverse
row of about twenty weak teeth.
Tongue : Narrow for a swift, and small for
the size of the mandibular area. Greatest
width of tongue 5.7 mm., length 10; greatest
width of mandible 20 mm., length 26. Tongue
fleshy, channelled toward the tip, sides some-
what sinuate, tapering slowly to two blunt
tips. The two cornua are lined along the edge,
both on inner and outer sides, with strong
teeth. Smaller ones are scattered over the
posterior surface of the tongue itself.
Glottis : A narrow ellipse on a low flat
area, with inconspicuous unarmed rim. Pos-
teriorly, there is an irregular transverse row
of teeth, all large, flattened, sub-equal in size
and numbering about fifteen on each side.
Beyond these there arises a second irregular
row. Most of the teeth, in a cleared condition,
show stout, parallel-sided bases, and slender
tips.
Syrinx : Swift No. 31135 (KOH No. 2589) .
Male. April 9, 1948. There is little change in
the posterior tracheal rings except that the
last eight are slightly narrowed with more
even edges. The syrinx is a wide tracheo-
bronchial collar of bone. Anteriorly the upper
margin is level except in the center where
an irregular, rounded projection overlies a
segment of the last tracheal ring. This, like
the rest of the syringeal collar is ossified and
coarsely fenestrated. The anterior vertical
width of the collar is 1.6 mm., its lateral,
front to back, length is 3.3 mm. The lower
border of the collar is formed by the closely-
applied, strongly arched, upper bronchial
1949]
Beebe: Swifts of Rancho Grande, and Their Migration
57
Text-fig. 3. Streptoprocne zonaris albicincta (Cabanis). Syrinx, front
and side views.
semiring. This semiring dips far down in
front, forming an acute angle with the pes-
salus. The triangular space within this angle,
as far anterior as the collar, is ossified, but
without fenestration.
Posteriorly the tracheal rings are appre-
ciably wider, with little more intervening
membrane showing than the median, open
notches. Two of the rings anastomose. The
two lowermost rings are narrow and even as
to outline.
Posteriorly, the syringeal collar is similar
to its anterior half, with the difference that
this aspect is flat and the median anterior
projection is less pronounced. There is a faint
but distinct indication that the present ossi-
fied syringeal collar was originally composed
of two rings. The entire ventral syringeal
aspect, bounded by the lower border of the
collar, the pessalus and the first semiring is
of course membranous. The free ends of the
anterior semirings, joined by the tympani-
form membrane, narrow rapidly posteriorly,
until the ninth onwards become almost com-
plete rings, thus forming the end of the
membrane. There are about twenty-three
bronchial rings, the second, third and fourth
being somewhat longer, projecting slightly
into the inner profile of the bronchi. From
the eighteenth bronchial ring onwards there
is a gradually increasing disintegration of
the rings within the lung tissue, a thinning
and irregular anastomosing of adjoining
rings.
A drawing and description of the syrinx
of the female No. 529, made thirty years ago,
are similar to that of the present male except
that posteriorly, the median protuberance is
considerably larger, and extends forward
over the last three tracheal rings.
Chaetura brachyura brachyura
(Jardine, 1846).
Short-tailed Swift.
Species Range-. Same as that of the sub-
species below, with the addition of the Lesser
Antilles.
Subspecies Range: Northern Venezuela
and the Guianas to Trinidad and Tobago,
south through eastern Ecuador and Peru to
Matto Grosso and Para.
Field Characters for Sight Identification :
This is the smallest of the Rancho Grande
swifts (length 100 mm.). On the wing it ap-
pears totally black, with conspicuous pale
brownish-gray rump, tail-coverts and tail.
The absence of gray on the underparts dis-
tinguishes it from the slightly larger cinere-
iventris.
Occurrence : By far the commonest swift
at Rancho Grande, becoming really abundant
after the rains began.
On clear days numbers were often seen
feeding with swallows and other swifts, espe-
cially when migration of various orders of
insects was in full swing. On partly cloudy
days or when fog drifted up the lower val-
leys, these short-tailed swifts would swing
through the pass in small or larger numbers.
There was none of the rather regular,
southward, morning shift of the giant swifts.
We saw this species every month from March
to August, and on almost every clear day. It
often flew in twos and threes, or again in
flocks of considerable size. For three con-
secutive days, we were able to identify an
individual trio. Two of the birds had recog-
nizable gaps in their primaries due to molt,
and these marked birds hawked on the north
side of the pass, low in the gorge, throughout
a three-day flight of termites, Coptotermes
testaceus. On the third day an onrushing
mass of dense fog drove these swifts away
in the wake of eight turkey vultures has-
tening toward the upper zone of clear sun-
shine. The swifts did not return, and the
next day the insect flight had ceased.
On July 9, 1948, we observed an unusual
flocking, a migration of sorts ; a fairly com-
pact mass of considerably more than four
58
Zoologica: New York Zoological Society
[34: 8
hundred of these swifts, circling, not feeding.
As they approached the pass from the south,
the flight changed to a more direct movement,
and when siphoning through, all circling
ceased, and the rush of wings was like a loud
wind as the birds passed low and at great
speed. The northern valley was partly filled
with fog and the birds rose slowly above it,
and before they passed from view, were
again circling high in air as before. The gen-
eral effect was of a maze of inorganic units,
without volition, sucked by a wind through
the narrow notch, and then sprayed out in a
slower movement as the draught lessened.
As a matter of fact, there was little or no
breeze in the pass itself.
A spectacular coincidental sight of the
same type of flocking of the same species is
recorded in my notes on an identical July 9,
but thirty-two years before, at Kalacoon,
British Guiana. The note reads as follows:
“An enormous flock of short-tailed swifts
appeared over the forest at 9:30 this morn-
ing. There were certainly more than a
thousand birds, all flying in a great circle,
gradually attaining higher and higher alti-
tude. They were massed so closely together
that there seemed hardly room for any move-
ment of the outspi’ead, crescentic wings. The
relative slowness of movement and the fre-
quent effortless gliding indicated an upward
surge of air. Through the glasses a scatter-
ing of equally small white-rumped swifts,
Chaetura spinicauda, was clearly seen. The
whole company vanished very high up and
drifting southward.”
Although these swifts showed little fear
of the pair of bat falcons nesting near the
laboratory, yet at least seven individuals
fell victims to these hawks, and an eighth
mangled swift found in the road near the
nesting tree indicated an additional dropped
item of diet. In this, as in other species, the
swifts were able to evade the raptores when
on the same aerial level. It was only when
the hawks could go into a vertical dive that
their speed made them almost unavoidable.
Two short-tailed swifts crashed against
the laboratory windows on successive eve-
nings, May 16 and 17, but one was able to
fly away. The other was a male, breeding,
with the stomach quite empty.
Chaetura cinereiventris lawrencei
Ridgway, 1893.
Gray-breasted Swift.
Species Range: The range of the eight
recognized subspecies extends from Nicara-
gua, Grenada, Tobago and Trinidad south
to Venezuela, Colombia, Ecuador, Peru, cen-
tral Brazil and Bolivia.
Subspecies Range: lawrencei occurs in
Grenada, Tobago, Trinidad and the moun-
tains of northern Venezuela.
Field Characters for Sight Identification:
A small swift. Black, except for rump and all
underparts, which are pale gray. When as-
sociated with brachyura and seen from above
at a distance, the birds are much alike, the
brownish shade of the rump of brachyura
being hardly distinguishable. From below,
the gray of cinereiventris instantly sets it
apart.
Occurrence : This is not common at Rancho
Grande, but was observed occasionally
throughout May, June and July. Only once,
on June 9, did an individual come to the lab-
oratory windows. It clung for five minutes
to the sill but evaded all efforts at capture.
On May 9, the male bat falcon caught a
gray-breasted swift and plucked it. The fe-
male then gave it to her nestlings who tore
it apart and ate it. As they pulled it apart I
could distinctly see the mass of small ants
which filled the stomach, together with two
large abdomens of Atta queens which were
eaten by the young birds.
Chaeturella rutlla brunneitorques
Lafresnaye, 1844.
Chestnut-collared Swift.
Species Range : Central Mexico, south
through northern South America to Peru,
the Guianas and Trinidad.
Subspecies Range: Southeastern Mexico,
south to Colombia, Venezuela, Ecuador and
Peru.
Field Characters for Sight Identification:
A medium-sized swift, about five inches in
length. It is the only species marked with
chestnut; throat, breast and collar. This
color is especially distinct when the birds are
silhouetted against the foliage of the moun-
tain jungle, but with glasses is conspicuous
even when they are high in the sky.
Occurrence: The chestnut-collared swift
was third in order of abundance at Rancho
Grande, surpassed only by Streptoprocne
and Chaetura brachyura. It was frequently
seen shuttling back and forth through the
pass, or hawking about on days of insect mi-
gration, associating with giant swifts or
with swallows, mostly single birds or in small
flocks. Occasionally they would race back and
forth through the pass, yet the diurnal ob-
servations I was able to make showed no cer-
tain regularity of north or southward shift.
On the other hand sixteen birds struck
against the laboratory windows and on three
occasions, when no swifts actually flew
against the glass, individuals were seen flut-
tering about among the bats within the area
of illumination. This would indicate a daily,
crepuscular northward migration, as in
Streptoprocne, also perhaps to some roosting
or breeding colony.
Support of the probability of such a mi-
gration is furnished by a chestnut-collared
swift (31129) taken eight kilometers east
of Rancho Grande. A reliable assistant, Pe-
dro Infante, shot this bird January 8, before
my ai'rival, on the Choroni road which par-
allels that from Maracay to Ocumare. He re-
ported this swift as shot from a group of
thirty to forty which, throughout October,
November and December assembled every
evening and spent the night clinging in a
1949]
Beebe: Swifts of Rancho Grande, and Their Migration
59
' compact mass to the vertical side of a rocky
, cliff near the road. All left at dawn. Thus we
have evidence of a roosting colony at the
same elevation as the pass, and, from the
point of view of a volant swift, only a short
distance away.
At Rancho Grande the meteorological con-
ditions which induced the appearance of the
swifts at the lighted windows were, high
but dense clouds; low neblina fog with or
without wind ; precipitation, whether drizzle
or pelting rain and with or without light-
ning. No birds ever came on clear nights,
whether moon or starlighted, and no bird
after 10 P. M.
There was considerable variation in the
amount of chestnut on the plumage of these
swifts, but the typical pattern was rich
chestnut throat and breast with a wide col-
lar extending over nape and hind neck. Two
adult males had the chestnut reduced to a
pectoral tinge, and several swifts on the
wing were intermediate between these ex-
tremes. The only female examined (31143),
an adult, showed no pigmental difference
from a full-plumaged male, except that the
under tail coverts were strongly edged with
white.
Coleoptera spp., including a ceram-
bycid.
Hymenoptera : Camponotus sp.,
fragmentary remains of several
hundred.
31132: Half the meal composed of ants
(Weber) .
Coleoptera spp.
Hymenoptera: Camponotus sp.,
fragmentary.
31143: Dominant food, hundreds of small
flying ants (Weber).
Hemiptera spp.
Coleoptera: cucurlionid.
Hymenoptera: Pheidole sp.
Hymenoptera: Very small dolicho-
derine fragments.
Cypseloides cherriei Ridgway, 1893.
White-spotted Swift.
Former Records and Species Range : Two
swifts taken on Volcan de Irazu, central
Costa Rica, were described by Ridgway in
1893. The type was thought to be a male;
the second bird was uncertainly sexed as a
female. The most noticeable character was
“a large, sharply defined spot of silky white
Data on Chestnut-collared Swifts
Lgth.
Wing
30382
male
126
123
30382a
male
30448
male
133
128
Four swifts escaped
31128
male
115
120
One swift escaped
31131
male
130
120
31132
male
128
120
Four swifts escaped
31143
female
122
125
Tail
Grams
Weight
Extent
Date
45
25
May 14, 1945
43
22.9
320
May 14, 1945
July 3, 1945
36
292
July 3, 1945
Mar. 23, 1948
43
21.5
295
Mar. 23,1948
April 3, 1948
45
21.5
302
April 3, 1948
43
19.5
305
April 3, 1948
April 24, 1948
Detailed Food.
30382: Many winged females of Campono-
tus (T anaemyrmex) coruscus F.
Smith; and Solenopsis geminata
edwardi Forel.
30448: A series of winged females of Cre-
matogaster ( Orthocrema ) sp.
31128: Stomach crammed with a mass of
winged ants and small beetles.
(Weber) .
Homoptera: Cicadellidae.
Coleoptera spp., including a carabid.
Hymenoptera: parasitic sp.
Hymenoptera : Crematogaster ?
wing.
Hymenoptera: Solenopsis geminata
Fabr., widespread in the northern
neotropics and replaced in the
south by saevissima F. Smith,
from the interior of British Gui-
ana (Courantyne) and Brazil.
Camponotus sp.
31131: Food dominantly flying ants
(Weber).
Hemiptera: wing.
on each side of the forehead, immediately
over the lores, and a short streak of the
same color immediately behind the eye.” A
third specimen was reported (Zimmer, 1945)
in a collection of birds from Colombia. This
was taken at San Gil, Santander, and was
also questionably sexed as a female.
This then, on February 26, 1948, was the
summation of our knowledge of the white-
spotted swift. The Costa Rican birds came
from the same 10th degree of north latitude
as Rancho Grande, but 1,800 kilometers
west; whereas the Colombian swift, from 6
degrees, 33 minutes north latitude, was 750
kilometers to the southwest of our labora-
tory.
Field Characters for Sight Identification :
The white-spot is a five-inch swift, appear-
ing uniformly black, with a conspicuous,
round, white spot between beak and eye.
These spots stand out strongly whenever
these birds are seen head-on in flight or
from the side, giving a rather fantastic im-
pression of a slightly misplaced pair of bril-
liant eyes.
Occurrence : At 9.30 o’clock in the evening
60
Zoologica: New York Zoological Society
[34:8
of February 26, 1948, a white-spotted swift
came to the windows of Rancho Grande and
was caught. From this time until June 13
we captured or recorded eight others, making
a total known of this unusual species of
twelve individuals. The details of the nine
Rancho Grande birds are as follows.
31125: Female not breeding. February
26, 1948. Length 132, wing 123, tail 132, ex-
tent 308 mm. Black above, sooty brown be-
low; supra-loral spot and small post-ocular
patch white; small feathers along edge of
wrist and front of wing white-edged; trace
of white on chin. Ovary small but distinct.
At 10 P.M. this bird fluttered against a
bedroom window. It was later found and cap-
tured on the Rancho Grande porch, fluttering
confusedly around the electric light. The eve-
ning was one of dense fog, with a strong
breeze blowing from the southwest.
Food: A mass of rather comminuted flying
ants. (Weber).
Coleoptera spp.
Hymenoptera: Highly fragmentary Cam-
ponotus sp., forming most of the con-
tents.
31133: Male, not breeding, April 4, 1948.
Length 120, wing 120, tail 39, extent 305 mm.
Weight 22.5 grams.
Frontal spots large and pure white, begin-
ning on lores with only a few feathers be-
tween them and nostrils, and extending back
over eye frame, and on a narrow line to mid-
way over eyes, thus approaching the post-
ocular spot. This latter forms the posterior
border of the feather circle around the bare
area on the lower lid. When the eye is closed
and this lid drawn up, the spot is directly
behind the eye. When the eye is open it is
behind and below eye. Wrist edge of wing and
rim featherlets all have distinct white edges.
The chin is grayish-white.
The bird was caught at 8:30 P.M. as it
clung to the vertical electric light wire de-
pending from the ceiling of the porch outside
the laboratory. There was sufficient fog to
hide the stars and the recent wind had died
down. The air was cool, 62 degrees Fahren-
heit, sufficient to keep all moths away.
Food: Flying ants. (Weber).
Coleoptera spp.
Hymenoptera: Camponotus sp.
Syrinx: C. cherriei, No. 31133 (KOH No.
2588) differs from Streptoprocne zonaris al-
bicincta in there being three, instead of two,
rows of post-glottid teeth. The syrinx proper,
although completely ossified, shows distinctly
its composition of three rings. The median
anterior protuberance is directly connected
with a slight, posterior, cartilaginous projec-
tion of the lowermost free tracheal ring.
31134: Female, not breeding. April 5,
1948. Length 137, wing 127, tail 51, extent
310 mm. Weight 25.5 grams.
Large white preocular spots almost join
white chin. Postocular and white wing edges
well developed. Flew against laboratory win-
dows at 7 :30 P.M. in dense cold fog.
Food : Stomach crammed with insects, one-
half of which were ants (Weber).
Hemiptera spp.
Coleoptera spp.
Hymenoptera: parasitic sp.
Hymenoptera: Camponotus sp. fragmen-
tary.
Hymenoptera: dolichoderine wings.
April 5. At 7 :45 two more white-spots
came to the porch but both escaped. At 8:10
another bird came and went. In all, the men-
tal white was almost absent.
April 11. In dense, drenching fog a white-
spot came to my bedroom window at 9:30
P.M. and clung out of reach to the rough
surface of a cement pillar. Eye-white as
usual, with more on the chin than in any bird
hitherto seen.
May 10. Male bat falcon caught a white-
spot, held it for three minutes, with the
dangling head in full view. Hardly any men-
tal white, but very large and fluffed out eye
spots. After plucking it he gave it to the
female who fed her young.
June 13. Female bat falcon brought a swift
to her perch, and had begun plucking it when
the young male flew up, took it and ate it.
Cypseloides cryptus Zimmer, 1945.
Tropical Black Swift.
Former Records and Species Range: In
1945 a new species of swift was described
(Zimmer, 1945). The type came from the
Rio Tavara, Peru. Only four other specimens
were known, taken at the following localities :
British Guiana (Kaieteur Falls), Venezuela
(Mt. Auyan-tepui, and Saroropan-tepui) ,
and Costa Rica (San Pedro).
Only a single individual of this species was
seen at Rancho Grande in 1948. The two
which were taken in 1946 have already been
reported by me (Beebe, 1947), and I here
repeat several paragraphs.
Field Characters for Sight Identification :
This five-inch black swift would show no
definite characters in flight except the gen-
eral black coloration. The grizzled and vari-
able dull whitish of the lores and chin could
hardly be detected.
Occurrence: On April 20, 1946, a female of
this swift crashed against the laboratory
windows at Rancho Grande at 8 :30 o’clock in
the evening and was stunned. It is No. 30,634,
female, not breeding, fairly fat, weight 40.2
grams. Length 120, wing 137, tail 48, extent
355 mm. The stomach was filled with winged
female Azteca ants.
On April 21, the following evening, at the
same time, a second bird killed itself against
the identical window. This is No. 30,640, fe-
male, not breeding, considerable fat, weight
35.8 grams. Length 138, wing 130, tail 50
mm. First primary in each wing half grown.
Food: A great quantity of Crematogaster
and Azteca flying ants.
April 12, 1948. At 10:15 A.M. the male bat
falcon swung up to his perch in the top of
the candelo tree, with a swift. Through the
1949]
Beebe: Swifts of Rancho Grande, and Their Migration
61
20-power glasses I could see every detail and
in every respect of the cephalic pale color it
seemed to be this species. There was no trace
of the supraloral white spots of cherriei, and
the area around the base of the beak showed
the pale grizzled appearance so apparent in
both of the specimens taken in 1946. I could
have had no more certain evidence if the
bird had been in my hand. It was slowly and
thoroughly plucked and as the female did not
appear, the male proceeded to eat the eighth
known individual of Cypseloides cryptus.
Aeronautes montivagus mantivagus
(d’Orbigny and Lafresnaye, 1837).
White-breasted Swift.
Species and Subspecies Range : Mountains
of northern Venezuela, Peru and Bolivia.
Field Characters for Sight Identification:
A small five-inch swift. Easily distinguished
by great extent of ventral white, no nuchal
collar, and almost square tail.
Occurrence : Known at Rancho Grande
laboratory from a single specimen which flew
into the porch in dense neblina, and at about
9:15 P.M. was caught as it crouched in a
corner.
31142: Adult male, breeding, testes 8.5
mm. April 23, 1948. Length 120, wing 110,
tail 40, extent 266 mm. Weight 20 grams.
Food : Many flying ants and small cucurli-
onid beetles (Weber).
Hymenoptera: ponerine and Camponotus
ant fragments.
Parasites: This swift was strongly in-
fested with parasites, one of which (48375)
was a giant feather fly with bright green ab-
domen. In addition, there were several Mallo-
phaga, and a number of bete rouge.
Dr. J. Bequaert has kindly identified the
large feather fly, and sends me the following
note:
“The fly is Brachypteromyia neotropica J.
Bequaert. This was described from a single
male, taken from the same host species, at
Galipan, close to Pico Avila, Estado Mirando,
2,000 meters elevation, Venezuela.2 Yours is
the second specimen known, also a male. The
description, with figure, is in Psyche, 49.
(1942) published in 1943, p. 113. The only
other species of the genus, Brachypteromyia
fimbriata (Waterhouse), is North American,
on the swifts Aeronautes saxatilis and
Nephoecetes niger.”
In addition to the single captured specimen
of this swift, we have three other records.
June 19, 1948. Six white-breasted swifts
flew, one after the other, through the pass
at 10 A.M. They were headed south and not
flying very fast. While still in sight two of
the birds veered aside from their direct
flight and caught insects.
June 21, 1948. A compact flock of twelve
of these swifts swung south through the pass
at 8:05 in the morning. They flew very low,
just skimming the trees.
2 Pico Avila is in the immediate neighborhood of Caracas,
about 100 kilometers due east of Rancho Grande.
Three of this species were caught by the
male bat falcon; on April 4, June 10 and 19,
1948.
Panyptila cayennensis (Gmelin, 1789).
Fork-tailed White-collared Swift.
Species Range: Southeastern Nicaragua,
south over Colombia, Ecuador, Venezuela,
Tobago, Trinidad and the Guianas to Bahia
and Sao Paulo.
Field Characters for Sight Identification:
A five-inch swift, unmistakably fork-tailed,
black except for white eye-spots and flank-
spots, chin, throat and collar.
Occurrence: 30439: A male flew against
the windows of the laboratory and was badly
injured. July 1, 1945. Length 123, wing 125,
tail 57 mm.
Food: Small species of flying ants.
(This specimen was overlooked in the
paper on Avian Migration at Rancho Grande,
Beebe, 1947).
June 8, 1948. Six fork-tailed swifts hawk-
ing about early in the morning with three
blue and white swallows, over the compound
of Rancho Grande.
June 12, 1948. A swift of this species
caught and eaten by male bat falcon.
Summary.
At Rancho Grande a total of eight species
of swifts were collected or observed, out of
the thirteen species recorded from Venezuela
as a whole. The types of observation resolve
into: flocks feeding at various altitudes,
others migrating through Portachuelo Pass
usually low down, or striking against the
lighted windows of the laboratory on nights
of fog or rain. Finally a number of the birds
were caught by a male bat falcon.
Throughout twenty months of residence
during three years no swift was seen to
alight, nor was there at Rancho Grande first-
hand proof of breeding or roosting colonies,
although the latter were indicated as a result
of various activities.
Observations in adjoining areas, both
higher up the surrounding mountains, and
down to four hundred and forty-five meters
on the Maracay plain, showed a relative
dearth or absence in comparison with their
numbers in the square kilometer whose cen-
ter was Rancho Grande and the pass. The
obvious explanation of this concentration of
swifts, by day and night, must be the same
as that of many other organisms, both ver-
tebrates and invertebrates; viz., the contin-
ual procession on migration of countless
numbers of insects representative of almost
every order, traversing the sixty-foot-wide
pass, from north to south, on every clear
day throughout the rainy season. This abun-
dant and ever renewed source of food was
obviously a focusing factor of prime impor-
tance.
A second reason for the abnormal numbers
of species and individuals was the use of the
pass by several of the species on daily mi-
gration from a presumed breeding or roost-
62
Zoologica : New York Zoological Society
[34: 8: 1949]
ing place to a trans-pass feeding-area. It is
difficult otherwise to account for the forty-
four specimens of all eight species taken or
observed on black nights of poor visibility.
Of twenty-two specimens sexed, seventeen
were males, five females.
The presence throughout their breeding
season of a pair of bat falcons, close to
Rancho Grande, revealed an interesting rela-
tionship between these birds and the swifts.
The latter showed little fear of the small fal-
cons when these were perched, and giant
swifts did not hesitate to attack and drive
off the male hawk. Yet I recorded nineteen
individuals and seven out of the eight spe-
cies of swifts as entering into the diet of
the hawks. When high in the sky, a power
dive attack of the male falcon rendered es-
cape impossible on the part of the swifts, but
when the birds met at horizontal levels, the
speed and dodging ability of the small birds
rendered them safe.
The flocking habits of these Venezuelan
swifts are, in some ways, suggestive. Single
birds were very rare, and pairs were not
often seen. But, especially in mid rainy sea-
son months, trios were common. Even in
flocks of twenty to fifty, feeding in midair,
subdivision into trios was often evident. If
the same rule of a single surviving young
holds in tropical swifts as in many other
tropical birds, these trios probably repre-
sented the season’s families. Larger flocks in
rapid movement were too infrequent to war-
rant definite classification or object.
Literature Cited.
Beebe, William & Jocelyn Crane
1947. Ecology of Rancho Grande. Zoologica,
32: 43-59.
Peters, J. L.
1940. Check-list of Birds of the World. IV:
1-291.
Ridgway, R.
1893. Description of Two Supposed New
Species of Swifts. Proc. U. S. Nat.
Mus., XVI: 43-44.
Todd, W. E. C. & M. A. Carriker, Jr.
1922. The Birds of the Santa Marta Region
of Colombia. Annals of the Carnegie
Museum, XIV : 3-582.
Wetmore, A.
1939. Observations on the Birds of Northern
Venezuela. Proc. U. S. Nat. Mus., 87:
No. 3073, 173-260.
Zimmer, J. T.
1945. A New Swift from Central and South
America. Auk, 62 : 586-592.
EXPLANATION OF THE PLATE.
Plate I.
Two migrant swifts which came to the electric
lights of Rancho Grande on nights of rain or fog.
Fig. 1. Giant White-collared Swift. Strepto-
procne zonaris albicincta (Cabanis).
Fig. 2. Chestnut-collared Swift. Chaeturella
rutila brunneitorques Lafresnaye.
BEEBE.
PLATE I.
FIG. 2.
THE SWIFTS OF RANCHO GRANDE. NORTH-CENTRAL VENEZUELA, WITH
SPECIAL REFERENCE TO MIGRATION.
Hertlein & Strong: Mollnsks of Mexico and Central America
63
Q
Eastern Pacific Expeditions of the New York Zoological Society. XL.
Mollusks from the West Coast of Mexico and Central America. Part VII.1
Leo George Hertlein & A. M. Strong.
Calif ornia Academy of Sciences.
(Plate I) .
[This is the fortieth of a series of papers deal-
ing 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)
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 63
Superfamily Tellinacea 64
Family Tellinidae 64
Genus Tellina Linnaeus 64
Subgenus Tellinella Morch 64
Tellina ( Tellinella ) cumingii Hanley 65
Tellina (Tellinella) zacae Hertlein & Strong, sp.
nov 65
Subgenus Scrobiculina Dali 66
Tellina ( Scrobiculina ) ochracea Carpenter ... 66
Tellina ( Scrobiculina ) viridotincta Carpen-
ter 66
Subgenus Moerella Fischer 67
Tellina ( Moerella ) amianta Dali 67
Tellina ( Moerella ) arenica Hertlein & Strong,
sp. nov 68
Tellina ( Moerella ) erythronotus Pilsbry &
Lowe 69
Tellina (Moerella) felix Hanley 70
Tellina ( Moerella ) macneilii Dali 70
Tellina (Moerella) paziana Dali 71
Tellina (Moerella) recurvata Hertlein & Strong,
sp. nov 71
Tellina ( Moerella ) suff usa Dali 72
Tellina (Moerella) tabogensis Salisbury 72
Subgenus EuryteUina Fischer 73
Tellina (EuryteUina) eburnea Hanley 73
Tellina (EuryteUina) inaequistriata Donovan . 74
Tellina (EuryteUina) laceridens Hanley 75
Tellina (EuryteUina) mantaensis Pilsbry &
Olsson 75
Tellina ( EuryteUina ) panamanensis Li 76
Tellina (EuryteUina) planulata Sowerby .... 76
Tellina (EuryteUina) prora Hanley 77
Tellina (EuryteUina) regia Hanley 78
Tellina (EuryteUina) rubescens Hanley 78
Tellina (EuryteUina) simulans C. B. Adams.. 79
Subgenus Tellinidella Hertlein & Strong, subgen.
nov 79
Tellina (TeUinidella) purpureus Broderip &
Sowerby 80
Subgenus Macaliopsis Cossmann 81
Tellina (Macaliopsis) lyra Hanley 81
Tellina ( Macaliopsis ) lyrica Pilsbry & Lowe . . 81
Subgenus Merisca Dali 82
TeUina (Merisca) crystalling Spengler 82
Tellina (Merisca) proclivis Hertlein & Strong,
sp. nov 83
Tellina (Merisca) reclusa Dali 84
Subgenus Scissula Dali 84
TeUina ( Scissula) cognata C. B. Adams 84
TeUina (Scissula) nicoyana Hertlein & Strong,
sp. nov 85
Tellina (Scissula) virgo Hanley 86
Subgenus Phyllodina Dali 86
Tellina (Phyllodina) pristiphora Dali 86
Subgenus Phyllodella Hertlein & Strong, subgen.
nov 87
1 Contribution No. 842, Department of Tropical Research,
New York Zoological Society. .
Tellina ( Phyllodella ) insculpta Hanley 87
Subgenus Elliptotellina Cossmann 87
TeUina (Elliptotellina) pacifica Dali 87
Genus Tellidora Morch in H. & A. Adams 88
Tellidora burneti Broderip & Sowerby 88
Genus Macoma Leach 88
Subgenus Macoma s.s 88
Macoma (Macoma) nasuta Conrad 88
Subgenus C ymatoica Dali 89
Macoma (Cymatoica) undulata Hanley 89
Subgenus Psammacoma Dali 89
Macoma (Psammacoma) elongata Hanley. ... 89
Macoma (Psammacoma) lamproleuca Pilsbry
& Lowe 90
Macoma (Psammacoma) panamensis Dali .... 91
Macoma (Psammacoma) panamensis spectri
Hertlein & Strong, subsp. nov 91
Subgenus Psammotreta Dali 92
Macoma (Psammotreta) aurora Hanley 92
Macoma (Psammotreta) pads Pilsbry & Lowe 92
Subgenus Macoploma Pilsbry & Olsson 93
Macoma (Macoploma) medioamericana Olsson 93
Genus Apolymetis Salisbury 93
Apolymetis cognata Pilsbry & Vanatta 93
Apolymetis dombei Hanley 94
Genus StrigiUa Turton 95
Strigilla cicercula Philippi 95
StrigiUa costulifera Morch 95
StrigiUa disjuncta Carpenter 96
Strigilla lenticula Philippi 96
Introduction.
This is the seventh of a series of papers
dealing with collections of mollusks taken on
the Templeton Crocker Expedition (1936)
and the Eastern Pacific Zaca Expedition
(1937-1938). The general plan of presenta-
tion followed in the present contribution is
that mentioned in Part II of this series of
papers2. Formal headings and keys are given
for 51 species and subspecies of the Tellini-
dae collected by the expeditions of 1936 and
1937-1938. Occasionally additional species
are included in the keys for convenience but
in such cases it is indicated which species do
not occur in the present collection.
Acknowledgment is due Dr. G. Dallas
Hanna, Curator, Department of Paleontology
of the California Academy of Sciences, Mr.
A. G. Smith, Research Associate of the same
institution, and Dr. A. Myra Keen, Stanford
University, California, for assistance and
suggestions. Acknowledgment is also due
Miss Viola Bristol, Curator of Mollusks, San
Diego Society of Natural History, for the
loan of specimens. The photographs used for
illustrations on the plate were prepared by
Mr. Frank L. Rogers.
2 Hertlein, L. G., and Strong, A. M. Eastern Pacific
Expeditions of the New York Zoological Society. XXIII.
Mollusks from the West Coast of Mexico and Central
America. Part II. Zoologica, New York Zool. Soc., Vol. 28,
Pt. 3, December 6, 1943, pp. 149-168, pi. 1. See especially
pp. 149-150.
64
Zoologica: New York Zoological Society
[34:9
Superfamily Tellinacea.
Family Tellinidae.
Papers by Dali3 and Salisbury4 dealing
with the Tellinidae are very useful in a study
of West American members of this family.
Key to the Genera of the
Family Tellinidae.
A. Shell with lateral teeth in one or both
valves
a. Exterior with oblique, flexuous, divari-
cating striae; orbicular Strigilla
aa. Exterior usually without, sometimes
with, oblique but not flexuous, divari-
cating striae
b. Dorsal margins serrate; trigonal;
very inequivalve and compressed
Tellidora
bb. Dorsal margins not serrate or oc-
casionally so only posteriorly; usu-
ally elongate Tellina
B. Shell without lateral teeth
a. Suborbicular to subtrigonal; subequi-
lateral; moderately inflated; broad
submedian concavity in right valve
Apolymetis
aa. Subtrigonal or elongate; posterior end
produced and narrowed ; moderately
compressed; sometimes inequivalve
Macoma
Genus Tellina Linnaeus.
The present collection from the tropical
eastern Pacific contains a good representa-
tion of the species of Tellina which occur in
that region. Many of these shells are conspic-
uous because of their beautiful red color,
especially when observed upon the beach.
Morch long ago mentioned that red Tellinas
are characteristic of tropical American
waters.
Key to the subgenera of Tellina.
A. Posterior area with simple concentric
lamellae or smooth
a. Surface obliquely grooved Scissula
aa. Surface not obliquely grooved
b. Beaks anteriorly directed
Macaliopsis
bb. Beaks not anteriorly directed
c. Right anterior lateral distant
from the beak
d. Smooth, polished
Tellina s.s.5
dd. Strong concentric sculpture
Tellinella
3 Dali, W. H. Synopsis of the Family Tellinidae and of
the North American species. Proc. U. S. Nat. Mus., Vol. 23,
[No. 1210], November, 1900, pp. 285-326, pis. 2-4.
4 Salisbury, A. E. On the Nomenclature of Tellinidae,
with Descriptions of new species and some remarks on
Distribution. Proc. Malacol. Soc. London. Vol. 21, Pt. 2,
July, 1934, pp. 74-91, pis. 9-14.
5 Not represented in the present collection.
cc. Right anterior lateral extendi
close beneath or near beak
e. Shell usually exceeding
25 mm. in length
f. Resilium external;
shell elongate
g. Thick; right pos-
terior lateral
strong
Eurytellina
gg. Thin; right pos-
terior lateral
weak; fine reticu-ji
late sculpture
Tellinidella
ff. Resilium internal;
shell thin, high
Scrobiculina
ee. Shell not exceeding 25
mm. in length (usuallyi
not exceeding 20 mm.)
h. Sculpture'
chiefly of con-
centric lines of
growth (ex-
cept on poste-
rior area)
Moerella
hh. Sculpture
chiefly of con-,
centric lamel-
lae (over en-
tire shell) ; tri-
gonal; strong,
very narrow
posterior flex-
ure Merisca
B. Posterior area with plate-like foliations \
or posterior end with strong radial sculp-
ture
a. Pallial sinus free or confluent with '
pallial line for not more than one-third 1
its length
b. Posterior area with plate-like fo-
liations Phyllodina
bb. Posterior end of shell with strong
radial sculpture ; very small
Elliptotellina
aa. Pallial sinus confluent with pallial line
for entire length Phyllodella
Subgenus Tellinella Morch.
Tellinella Gray, Morch, Cat. Conch. Yol-
di, Fasc. 2, 1853, p. 13. [Species originally
cited under Tellinella include antoni Phil-
ippi, interrupta Solander, pulchella Lamarck,
rostrata Linnaeus, virgata Linnaeus, and
several others]. — Dali, Bartsch & Rehder,
Bernice P. Bishop Mus., Bull. 153, July 25,
1938, p. 187. Type : Tellina virgata Linnaeus.
Type (designated by Stoliczka, Mem. Geol.
Surv. India, Palaeont. Indica, Ser. 6, Vol.
3, 1870, pp. XVII, 116) : Tellina virgata
Linnaeus [Syst. Nat., ed. 10, 1758, p. 674.
“Habitat in O. Indico.” Illustrated by Hanley,
)| .949]
Hertlein & Strong: Mollusks of Mexico and Central America
65
a Thes. Conch., Vol. 1, 1846, p. 228, pi. 63, fig.
212. Indian Ocean].
Key to the species of TellineUa.
A. Ornamented by radial stripes or spots of
chocolate or purple; distance separating
pallial sinus from anterior adductor im-
pression 5 mm. or more cumingii
B. Ornamented by radial stripes or bands
of golden-orange ; distance separating pal-
lial sinus from anterior adductor impres-
sion not exceeding 2 mm. zacae
Tellina I Tellinella I cumingii Hanley.
Tellina cumingii Hanley, Proc. Zool. Soc.
London, September, 1844, p. 59. “Hab. Gua-
comayo, Central America; in coral sand.”
— Hanley, Thes. Conch., Vol. 1, 1847, p. 223,
pi. 58, fig. 72. “Guacomayo and America.”
— M. Smith, Panamic Mar. Shells (Tropical
Photogr. Lab., Winter Park, Florida), 1944,
p. 64, fig. 847. Lower California to Panama.
[Not the record “Red Sea?”].
Type Locality : Guacomayo, Central Amer-
ica.
Range : Magdalena Bay, Lower California,
to the Gulf of California and south to Gor-
gona Island, Colombia.
Collecting Stations : Mexico: Tangola-
Tangola Bay (196-D-14, 15), 5 fathoms,
crushed shell; Costa Rica; Culebra Bay;
Cedro Island (213-D-4), 5 fathoms, mud; off
Ballena Bay, Gulf of Nicoya (213-D-15), 40
fathoms, mud.
Description: Shell elongate, narrow,
rather compressed, subrostrate, somewhat
biangulated and bent to the right posterior-
ly; yellowish-white with radiating brown
or chocolate or purplish streaks or spots;
sculptured with moderately fine, close, con-
centric lamellae which become coarser and
elevated on the posterior portion of the shell;
hinge with two cardinal teeth in each valve,
the right posterior and left anterior card-
inals grooved, and two equidistant laterals
in each valve; pallial sinus rather wide (in
young shells rounded but in the adult oblique-
ly pointed at the end), projecting forward
about two-thirds the distance between the
two adductor impressions and for about two-
thirds its length confluent with the pallial
line ; interior white or pale yellow or a com-
bination of the two.
Large specimens of this species attain a
length of 55 mm. or more.
Tellina interrupta Wood6, which ranges
from North Carolina to Brazil, is a similar
species. Tellina strophia Dali, in the Mio-
cene of Florida, also is somewhat similar
to T. cumingii.
a Tellina interrupta Wood, General Conch., 1815, p. 146,
2 , J}g" 3- Inhabits the Indian and American Seas.”
(Also edit. 1835. I Regarding the dates of issue of this
book see Pritchard & Gatliff, Proc. Roy. Soc. Victoria,
£°1- 16 (N.S.) , Pt. 1, September, 1903, p. 114; Iredale,
roc. Malacol. Soc. London, Vol. 15, Pts. 2 and 3, December,
1922. p. 91],
Not Tellina interrupta Solander, Portland Cat., 1786.
DP. 31, 72, 105. Nomen nudum.
Distribution: A few specimens of this
species were taken by the expedition off
western Mexico and Costa Rica. The species
also is known to occur in the Pleistocene of
Magdalena Bay, Lower California.
Tellina ITellinellal zacae Hertlein & Strong,
sp. nov.
Plate I, Figs. 12, 13, 17.
Shell of moderate size, elongately ovate,
umbos a little posterior to the center, white,
with golden-orange radiating bands of vary-
ing width; anterior end elliptically rounded,
posterior end rather pointed (but acutely
rounded at the extremity) and slightly bent
toward the right, the ventral margin is
broadly curved; right valve with an elevated,
rounded, curved ridge radiating from the
posterior side of the umbo to the posterior
ventral margin and there is a correspond-
ing depression in the left valve; posterior
to the ridge the shell is depressed and an-
terior to the ridge there is a slight sinus;
the early part of the shell is ornamented
with fine, rounded, concentric threads, these
on the adult shell become stronger and slight-
ly irregular and somewhat lamellated pos-
teriorly; ribs separated by interspaces of
about the same width or in some instances
slightly narrower, very fine concentric
threads are present in the interspaces; a
short but rather stout ligament on a narrow
nymph is present posterior to the umbos;
hinge with two cardinals in each valve, the
right posterior and the left anterior ones
grooved, the other two are more slender,
there are two laterals in each valve, those
on the right valve are strong, those on the
left low and fused with the margin; pallial
sinus long, rounded at the end and extending
about three-fourths the length of the shell;
interior whitish, the external rays showing
through the shell. Dimensions of the holo-
type: length, 33.4 mm.; height, 15.2 mm.;
convexity (both valves together) , 7.8 mm. ;
pallial sinus extends anteriorly 25 mm. from
the posterior end of the shell.
Holotype (California Acad. Sci. Paleo.
Type Coll.), from Station 136-D-l, Arena
Bank in the Gulf of California, Lat. 23° 29'
N., Long. 109° 25' W., dredged in 45 fathoms
(82 meters), mud. Paratypes were dredged
in the same general region at Station 136-D-
31-32, Lat. 23° 24' 30" to 23° 28' N., Long.
109° 24' to 109° 23' 30" W., in 35-42 fathoms,
in sand, calcareous algae and weeds. Other
specimens were dredged at Station 150-D-
12, Gorda Banks in the Gulf of California,
Lat. 23° 02' N., Long. 109° 28' W., in 80-90
fathoms, sand.
The shell of this new species is in general
features similar to that of Tellina cumingii
but it differs in several details. In the pres-
ent species the posterior area on the left
valve possesses a deep well-developed groove
corresponding to a ridge in the opposite
valve, while in T. cumingii the correspond-
ing area on the left valve is somewhat flat-
66
Zoologica: New York Zoological Society
[34: !
tened and dorsally bounded by a fine incised
radial line. The pallial sinus of the new spe-
cies is more evenly rounded at the anterior
end and extends much nearer (about three-
fourths the length of the shell) the anterior
adductor impression than that of T. cum-
ingii. Furthermore in the color pattern the
beautiful radial orange stripes on a white
ground attain greater width than the purple
stripes which occur on a yellowish-white
ground on T. cumingii. The specimens of the
new species in the present collection do not
attain the size of Tellina cumingii.
Tellina zacae bears a resemblance to Tel-
lina crassiplicata Sowerby as illustrated by
Dali, Bartsch & Rehder7 from Hawaii, but
is narrower anteriorly and less broadly trun-
cated posteriorly. The original illustration
of Tellina crassiplicata8 does not show any
radial stripes.
Subgenus Scrobiculina Dali.
Scrobiculina Dali, Proc. U. S. Nat. Mus.,
Vol. 23, No. 1210, November, 1900, p. 290.
“Type, Scrobicularia viridotincta Carpen-
ter.”
Schumacheria Cossman, Rev. Crit. de Pa-
leozool., Vol. 6, No. 1, January, 1902, p. 52.
New name for Scrobiculina Dali, not Scro-
biculinus Monterosato, 1884.
Type (by original designation) : Scrobi-
cularia viridotinca Carpenter.
Cossman proposed the name Schumacheria
to replace Scrobiculina Dali, 1900, because
of the prior name Scrobiculinus Montero-
sato, 1884.
A strict interpretation of the present In-
ternational Rules of Zoological Nomencla-
ture (Article 36), allows the retention of
Scrobiculina Dali.
Key to the species of Scrobiculina.
A. Color of umbos ochraceous ochracea
B. Color of umbos yellowish-green
viridotincta
Tellina IScroblculinal ochracea Carpenter.
Tellina ( Peronaeoderma ) ochracea Car-
penter, Ann. & Mag. Nat. Hist., Ser. 3, Vol.
13, April, 1864, p. 312. Cape St. Lucas. Re-
print in Smithson. Miscell. Coll., No. 252,
1872, p. 210.
Tellina ( Scrobiculina ) ochracea Carpen-
ter, Dali, Proc. U. S. Nat. Mus., Vol. 23, 1900,
p. 302. “Cape St. Lucas to the Gulf of Cali-
fornia.”
Type Locality : Cape San Lucas, Lower
California.
Range-. Cape San Lucas to the Gulf of
California.
Collecting Station: Mexico: Arena Bank,
7 Tellina crassiplicata Sowerby, Dali, Bartsch & Rehder,
Bernice P. Bishop Mus., Bull. 153, July 25, 1938, p. 187,
pi. 48, figs. 5-8. Various localities in Hawaii cited, also
Midway Island and Ocean Island.
8 Tellina crassiplicata Sowerby, Conch. Icon., Vol. 17,
Tellina, April, 1869, species 332, pi. 56, figs. 332a, 332b.
“Hab. Sandwich Islands.”
Gulf of California (136-D-6), 45 fathoms ;
mud, Area conglomerate.
Description: Two somewhat worn ant
broken specimens in the present collectioi
answer to the description of Tellina ochracea
They are colored a light sulphurous yellow
strongest near the beaks and fading gradu
ally toward the margins. Dali (1900), statec
that the species was very similar to Telliru
viridotincta Carpenter, differing only ii
color. From the ranges given by Dali it woulc
seem that Tellina viridotincta is a mort
southern shell while T. ochracea is a specie^
of the Gulf of California. The green tip of
the beak of T. viridotincta may not be a con-
stant character because Stearns and Pilsbrj
& Lowe have recorded it well within tht
range of T. ochracea. If the color differences
do not prove to be constant, viridotincta
being the older name, should take precedence
Distribution: A few specimens referred
to this species were dredged by the expedi-
tion on Arena Bank in the Gulf of California
in 45 fathoms.
Tellina IScroblculinal viridotincta Carpenter.
? Scrobicularia virido-tincta Carpenter,
Proc. Zool. Soc. London, November 11, 1856,
p. 160. “Hab. in Sinu Panamensi, una cum.
?S. producta; legit T. Bridges. Sp. un. in
Mus. Cuming.”
Macoma viriditincta Carpenter, Stearns)
Proc. U. S. Nat. Mus., Vol. 17, 1894, p. 156.
La Paz; various localities in the Gulf of
California.
Tellina ( Scrobiculina ) viridotincta Car-
penter, Dali, Proc. U. S. Nat. Mus., Vol. 23, i
1900, p. 302. Lower California to Panama.,
Tellina viridotincta Carpenter, Pilsbry &
Lowe, Proc. Acad. Nat. Sci. Philadelphia,
Vol. 84, 1932, p. 133. Espiritu Santo Island;
La Paz.
Type Locality: Bay of Panama.
Range: Gulf of California to Panama.
Collecting Station: Mexico: Port Guatulco
(195-D-6), 3 fathoms, sand, algae, crushed
shell.
Description : Shell large, high, oval, beaks
posterior to the center, anterior end ellip-
tically rounded, ventral margin broadly
rounded, posterior end tapering, subtrunc-
ately rounded and bluntly pointed at the ex-
tremity; white, umbos greenish-yellow; pos-
terior area set off by a low umbonal ridge
and anterior to this there is a low, broad,
radial depression; valves ornamented with
fine, rather regular, concentric threads and
striae, which are more crowded and irregular
on the posterior area on which, especially on
the right valve, there is usually one and some-
times more fine radial ridges ; the concentric
sculpture is crossed by very fine, faint, ra-
diating striae over much of the disk; the
pallial sinus extends about three-fourths the
length of the shell, highest somewhat an-
terior to the posterior adductor impression
then sloping downward anteriorly where, at
about 5-8 mm. from the pallial line, it is sub-
949]
Hertlein & Strong: Mollusks of Mexico and Central America
67
ngularly or bluntly rounded and then joins
he pallial line; a large sunken ligament is
ituated upon a nymph; hinge with two car-
inals in each valve, the right posterior and
jft anterior ones grooved ; right valve with
wo well-developed laterals, left valve with
. weak anterior lateral and the posterior
ateral very slight or obsolete ; interior white
Hth blotches of greenish-yellow, especially
oward the dorsal portion of the shell.
A specimen from the Gulf of California
n the Henry Hemphill collection of the Cali-
'ornia Academy of Sciences, measures:
ength, 62 mm.; height, 44 mm.; convexity
both valves together), 14.5 mm.; pallial
;inus extends anteriorly 46 mm. from the
)osterior end of the shell. The species attains
i greater size than this.
As mentioned by Dali, about the only dif-
ference between Tellina viridotincta and T.
)chracea is in color.
Distribution: One rather worn pair of
calves of this species was dredged in 3
?athoms at Port Guatulco, Mexico.
Subgenus Moerella Fischer.
Moerella Fischer, Man. de Conchyl., Fasc.
LI, June 15, 1887, p. 1147. Sole species,
rellina donacina Linnaeus.
Type (by monotypy) : Tellina donacina
Ldnnaeus. Recent, seas of Europe. Illustrated
>y Bucquoy, Dautzenberg & Dollfus, Moll.
Mar. Roussillon, Vol. 2, Fasc. 25, March,
L898, p. 648, pi. 91, figs. 13, 14, and vars.
L5-19. Mediterranean. Also other localities
:ited.
In the present paper nine species and sub-
species have been referred to the subgenus
Xloerella. Some of these, in earlier publica-
;ions, have been referred to Angulus Megerle
/on Miihlfeld. The type of Angulus desig-
lated by Gray, 1847, is Tellina lanceolata
Linnaeus, a species in which the hinge is
said to possess a right anterior lateral but
acking all other laterals. Salisbury, 1934,
stated that only two Recent species, T. lan-
:eolata, the type, and T. armata Sowerby,
were referable to Angulus. However, in the
explanation to his plates five species are re-
ferred to Angulus. It is unfortunate that this
well known supraspecific group name should
be applicable to so few species.
All the species in the present paper which
have been referred to Moerella, although
varying somewhat in shape, possess a right
posterior lateral tooth of varying strength.
This lateral occurs just below a socket which
is present just below the ventral end of the
nymph upon which the ligament is situated.
The results of our studies which have led
us to place the following group of species
under the subgenus Moerella, are in agree-
ment with the conclusions reached by Gard-
ner9 with regard to the east American
Miocene and Pliocene species formerly re-
ferred to Angulus.
9 Gardner, J., U. S. Geol. Surv., Prof. Paper Ht-E,
1928, p. 195; U. S. Geol. Surv., Prof. Paper 199-A, 1943,
p. 94.
Key to the species of Moerella.
A. Posterior end triangular or bluntly
pointed
a. Posterior end the longer, tapering,
pointed; white, yellowish or pinkish
suffusa
aa. Anterior end the longer
b. Posterior dorsal margin sinuous;
ivory white tabogensis
bb. Posterior dorsal margin straight
or slightly curved
c. Ventralmargin strongly curved ;
posterior end blunt; white
paziana
cc. Ventral margin very gently
curved
d. Length more than twice the
height; white, occasionally
yellowish amianta
dd. Length less than twice the
height
e. Area anterior to poste-
rior umbonal angulation
strongly depressed; usu-
ally pink on dorsal mar-
gins erythronotus
ee. Area anterior to poste-
rior umbonal angulation
not depressed or only
faintly so; red or pink
zoned with white
f. Umbos moderately
inflated; posterior
end very short
macneilii
ff. Umbos rather com-
pressed; posterior
end more attenuated ;
rose red felix
B. Posterior end broad, obliquely truncated
a. Pale rose color arenica
aa. White ; posterior end more abruptly
truncated recurvata
Tellina l Moerella I amianta Dali.
Tellina ( Moerella ) amianta Dali, Proc.
U. S. Nat. Mus., Vol. 23, No. 1210, November,
1900, pp. 303, 317, pi. 3, fig. 12. “Dredged
in 14 fathoms, sand, off Cape Tepoca, Lower
California, near the head of the Gulf, by the
U. S. Fish Commission at station 3019.”
Type Locality: Off Cape Tepoca, Lower
California, near the head of the Gulf of
California, in 14 fathoms, sand.
Range: Gulf of California to Colombia.
Collecting Station: Mexico: Santa Inez
Bay, Gulf of California (145-D-l, 3), 4-13
fathoms, sand; Nicaragua: Corinto (200-D-
19), 12-13 fathoms, mangrove leaves, sand,
also on shore in beach drift; Costa Rica:
Port Parker (203-D-l, 3), 12-15 fathoms,
sandy mud, shelly mud, crushed shell.
Description: Shell small, elongated, ante-
riorly produced and rounded, the posterior
end shorter, obliquely truncated and rather
68
Zoologica: New York Zoological Society
pointed at the extremity; sculptured with
fine, close, concentric thi-eads which toward
the posterior angulation become somewhat
irregular, sharper, and in the adult shell two
or three ribs coalesce to form low raised
lamellae on the posterior area; hinge with
two cardinals in each valve, the right pos-
terior and left anterior ones grooved, right
valve with a strong fairly close anterior
lateral and a small posterior lateral; pallial
sinus extends near to, but does not touch,
the anterior adductor impression, confluent
with the pallial line below; color white or
partly salmon yellow.
A large right valve dredged in Santa Inez
Bay in the Gulf of California in 4-13 fath-
oms, measures: length, 14 mm.; height, 6.8
mm.; convexity (one valve), 2.2 mm.
The small size, elongate form which is
much produced anteriorly, and fine, close,
concentric sculpture are characteristic fea-
tures of this species.
Specimens dredged by the expedition in
Santa Inez Bay in the Gulf of California are
typical of the species. Specimens collected at
Corinto, Nicaragua, in beach drift, and
dredged in 12-13 fathoms agree in general
characters with T. amianta except that they
are thinner. Many small specimens dredged
in 12-13 fathoms off Port Parker, Costa Rica,
in which the hinge, concentric sculpture and
other general characters agree well with
T. amianta, appear to represent the young
of that species. Some are less elongate in
proportion to the height as compared to
typical T. amianta but this appears to be
somewhat variable among these young shells.
Many small specimens in the collections of
the California Academy of Sciences, dredged
in the Gulf of California, are similar to
T. amianta. Some are white, some white with
pink radial streaks or dots, others yellowish-
white. These bear a similarity to one of the
illustrations given by Sowerby10 (pi. 47, fig.
278d) under the name of Tellina silicula
Deshayes. That species was originally de-
scribed by Deshayes11 with the type locality
“W. Columbia.” Salisbury12 stated that the
type specimen of T. silicula Deshayes is
referable to Tellina rhomboides Quoy &
Gaimard, a species which occurs in the wes-
tern Pacific region. According to Iredale13
Tellina clathrata Deshayes is the correct
name for that species. Sowerby’s figures
278a, b, c, appear to be referable to it but
it seems possible that his figure 278d might
be referable to a young T. amianta.
Distribution : This species was dredged in
Santa Inez Bay, Gulf of California, in 4-13
fathoms, at Corinto, Nicaragua, in 12-13
10 Sowerby, G. B., Conch. Icon., Vol. 17, Tellina, October,
1868, species 278, pi. 47, fig. 278d. "Hah. W. Columbia.”
[Not figs. 278a, b, c.].
11 Tellina silicula Deshayes, Proc. Zool. Soc. London for
1854, (issued May 16, 1855), p. 363. "Hab. Columbia. Coll.
Cuming.”
12 Salisbury, A. E., Proc. Malacol. Soc. London, Vol. 21,
Pt. 2, 1934, p. 89. See also Lamy, E., Bull. Mus. Nat. Hist.
Nat. (Paris), Vol. 24, No. 2, 1918, p. 116.
13 Iredale, T., Mem. Queensland Mus., Vol. 9, Pt. 3,
June 29, 1929, p. 266.
[34:
fathoms, also in beach drift on shore, and a | ,>rred
Port Parker, Costa Rica, in 12-15 fathoms I. eT
It also has been recorded as occurring in th
Pleistocene at Magdalena Bay, Lower Cali |
fornia.
Tellina IMoerellal arenica Hertlein & Strong! tie
The
sp. nov.
Plate I, Figs. 5, 11.
Tellina carpenteri Dali, Packard, Univ\
Calif. Publ. Zool., Vol. 14, No. 2, Septembe:
12, 1918, p. 276, pi. 25, figs. 10a, 10b. ,
“in 68 fathoms just south of the Faralloi
Islands.” —I. S. Oldroyd, Publ. Puget Sount
Biol. Sta., Vol. 4, 1924, pi. 41, figs. 10a, 10b
(Copies of Packard’s figures). —I. S. Old
royd, Stanford Univ. Publ. Univ. Ser. Geol
Sci., Vol. 1, 1924, pi. 44, figs. 10a, 10b
(Copies of Packard’s figures).
Not Tellina carpenteri Dali, 1903.
Description : Shell of moderate size, trans-
versely ovate, thin, compressed, the anterioi
end slightly the longer, color creamy white
salmon pink and rose in concentric zones of
irregular width ; anterior dorsal margin
sloping gently convexly from the beaks, an-
terior end rounded, ventral margin broadly| !
rounded, posterior dorsal margin gently slop-
ing, slightly concave, posterior end obliquely
truncated; posterior area defined only by a
rounded umbonal angulation ; sculptured
with concentric lines of growth and by some-
what irregularly spaced fine, shallow, con-
centric grooves which on the posterior area
become deeper and give rise to sublamellate
sculpture; hinge of right valve with two
cardinal teeth, the posterior one bifid, an
anterior lateral is fairly close to the cardinals
and there is a distant posterior lateral below
a socket; left valve (paratype) with a bifid
anterior cardinal and traces of a posterior
cardinal (some valves with a thin posterior
cardinal lamella) ; pallial sinus highly tri-
gonal back of the beak then sloping some-
what irregularly to a position below and well
separated from the anterior adductor im-
pression, the end rounded then bending pos-
teriorly for a very short distance where it
becomes confluent with the pallial line; in
terior white and pink. Dimensions of the
holotype : length, 24.5 mm. ; height, 15 mm. ;
convexity (one valve) , 2.5 mm. ; pallial sinus
extends anteriorly 19 mm. from the posterior
end of the valve.
Holotype, a right valve (Calif. Acad. Sci
Paleo. Type Coll.), dredged at Station 136-
D-20 in Lat. 23° 30’ N., Long. 109° 26’ W.,
in 43 fathoms, mud, on Arena Bank, at the
south end of the Gulf of California. One ad
ditional specimen, a paratype, was dredged
at the same locality. Three small single
valves were dredged in the channel east of
Cedros Island at Station 126-D-17, in 40
fathoms. Paratypes were dredged by the
Templeton Crocker Expedition in 1932, near
Puntarenas, Costa Rica.
Range: Farallon Islands, California, to
Panama.
1949]
Hertlein & Strong: MollusJcs o) Mexico and Central America
69
This beautiful species has often been re-
ferred to Tellina carpenteri Dali14, a distinct
species, which has a smaller shell and a gen-
erally more northern distribution. Dali15 cit-
ed T. carpenteri as occurring in the Gulf of
Panama at a depth of 182 fathoms. It seems
probable that that record may be referable
to the present species.
The form illustrated under the name of
T. carpenteri by Packard, 1918, which was
dredged in 68 fathoms just south of the
Farallon Islands, the illustrations of which
were reproduced by I. S. Oldroyd, appears
to be identical with the present specimens.
This new species differs from T. carpenteri
in the much greater size, irregular concen-
tric zones of color and in the stronger con-
centric grooves. It differs from Tellina re-
curvata Hertlein & Strong [_—T. recurva
Dali, 1900, not of Deshayes, 1844], in the
much more obliquely truncated posterior end
and in the pink coloration. The shell of Tel-
lina arenica differs from that of T. tabo-
gensis in the larger size, pink color and in
that the posterior dorsal margin is slightly
concave or nearly straight rather than flexu-
ous.
Tellina IMoerellal erythronotus Pilsbry &
Lowe.
Tellina ( Angulus ) erythronotus Pilsbry &
Lowe, Proc. Acad. Nat. Sci. Philadelphia,
Vol. 84, May 21, 1932, p. 94, pi. 12, fig. 7.
“Panama, east of the city” (type) . Also from
Montijo Bay, Panama.
Type Locality : Panama, east of the city.
Range : Magdalena Bay, Lower California,
to the Bay of Panama.
Collecting Stations: El Salvador: Mean-
guera Island, Gulf of Fonseca (199-D-l), 16
fathoms, sand, mud, crushed shell ; La Union,
Gulf of Fonseca (199-D-8-16, 19-25), 3-16
fathoms, mud, mangrove leaves ; Nicaragua:
Monypenny Point, Gulf of Fonseca (199-D-
2-6), 4-7 fathoms, mud.
Description: Shell donaciform, moderately
thin, somewhat compressed, creamy white
often tinged with pink around the margins,
opalescent; anteriorly elongated and round-
ed, ventrally gently rounded and posteriorly
slightly embayed due to the presence of a
shallow depressed area which occurs anterior
14 Angulus variegatus Carpenter, Rept. Brit. Assoc. Adv.
Sci. for 1863 (issued August, 1864) , p. 611. “Mont., Cat. Is.,
20-60 fm. ; rare (Neeah Bay, Swan).” Also pp. 627, 639.
Reprint in Smithson. Miscell. Coll., No. 252, 1872, pp.
97, 113. 125. —Carpenter, Ann. & Mag. Nat. Hist., Ser. 3,
Vol. 14, December,1864, p. 423. “Hab. Neeah Bay (Swan) ;
Monterey and Catalina Island, 20-60 fathoms, rare
(Cooper).” Reprint in Smithson. MisceU. Coll., No. 252,
1872, p. 235.
Not Tellina variegata Gmelin, Syst. Nat., ed. 13, 1790,
p. 3237.
Tellina (Angulus) carpenteri Dali, Proc. U. S. Nat. Mus.,
Vol. 23, No. 1210, November, 1903, pp. 303, 320. New
name for Angulus variegatus Carpenter, 1864, not Tellina
variegata Gmelin, 1790. “Strait of Juan de Fuca to Lower
California.” Illustrated by I. S. Oldroyd, Publ. Puget
Sound Biol. Sta., Vol. 4, 1924, p. 51, pi. 10, fig. 4. —I. S.
Oldroyd, Stanford Vniv. Publ. Univ. Ser. Geol. Sci., Vol.
1, 1924, p. 166, pi. 29, fig. 2. San Pedro, California.
15 Tellina ( Angulus ) carpenteri Dali, Dali, Bull. Mus.
Comp. Zool., Vol. 43. No. 6, October, 1908, p. 421. “U. S. S.
‘Albatross’, station 3355, Gulf of Panama, in 182 fathoms,
mud, bottom temperature, 54°. 1 F. U. S. N. Mus. 122,
934.”
to the posterior umbonal ridge; posteriorly
triangular and somewhat bluntly pointed,
with nearly straight dorsal margin; pallial
sinus long not quite touching the anterior
adductor impression, along the base wholly
confluent with the pallial line; hinge of right
valve with a grooved anterior and a bifid
posterior cardinal, the anterior lateral strong
and close to the cardinals, the posterior lat-
eral farther removed but not distant; left
valve with a grooved anterior and a thin
posterior cardinal lamina, anterior lateral
represented by a triangular projection of
the margin, a similar posterior projection
faintly developed or absent.
One of the largest specimens in the present
collection measures : length, 25 mm. ; height,
14.5 mm. ; convexity (both valves together) ,
7 mm.
The specimens in the present collection
show all the characters of Tellina erythrono-
tus mentioned by Pilsbry & Lowe.
The shell of this species appears to be
very similar to that of Tellina hiberna Han-
ley10 which also has been cited as occurring
at Panama. However, the shell of T. ery-
thronotus appears to be somewhat more
elongate in comparison to the figures of T.
hiberna given by Hanley and Salisbury. The
posterior dorsal margin of Hanley’s species
is said to be first convex then concave, but
there is no marked convexity on the margin
of T. erythronotus. Furthermore there is
nothing in Hanley’s description regarding
pink or prismatic colors such as often can
be observed on T. erythronotus.
Tellina puellula Salisbury17 (=T. puella C.
B. Adams, not Hanley) , is another closely re-
lated species. According to Pilsbry & Lowe it
is larger, relatively higher and thicker than
T. erythronotus.
Tellina hiberna was described in 1844 and
it seems possible that either T. erythronotus
or T. puellula, both described later, may be
referable to it, but we have not the type spec-
imens available to enable us to make any def-
inite decision on this question.
Tellina pananiensis Philippi18, from Pana-
ma, was described as white with the umbos
red both exteriorly and interiorly but it was
said to lack lateral teeth.
Distribution: Tellina erythronotus was
dredged quite abundantly in the Gulf of Fon-
seca at a depth of 3 to 16 fathoms on a bottom
of mud and mangrove leaves. The present
16 Tellina hiberna Hanley, Proc. Zool. Soc. London, De-
cember, 1844, p. 148. “Hab. Panama and Bay of Quayaquil ;
six to eleven fathoms, in sandy mud : Cuming.” —Hanley,
Thes. Conch., Vol. 1, 1846, p. 282, pi. 57, fig. 53. Original
localities cited.
Tellina ( Angulus ) hiberna Hanley, Salisbury, Proc.
Malacol. Soc. London, Vol. 21, Pt. 2, 1934, p. 91, pi. 13,
figs. 7, 8, 9. [Illustrations of holotype and paratypes].
17 Tellina puellula Salisbury, Proc. Malacol. Soc. London,
Vol. 21, Pt. 2, July, 1934, p. 86. A new name for Tellina
puella C. B. Adams, Ann. & Lyceum Nat. Hist. New York,
Vol. 5, July 1852, pp. 507, 646 (separate, pp. 283, 322).
“Panama.” Not Tellina puella Hanley, Proc. Zool. Soc.
London, February, 1845, p. 165. “Hab. Senegal. Cuming,
Metcalfe.”
18 Te'linu yanamensis Philippi, Zeit. f. Malakozool..
Jahrg. 5, No. 11, 1848, p. 176. "Ad Panama legit frater
E. B. Philippi.”
70
Zoologica: New York Zoological Society
[34:9
records of occurence furnish a long extension
north of the known range of the species. The
noi’thernmost occurrence of this species
known to us is that based upon a specimen in
the collections of the California Academy of
Sciences which was collected by C. R. Orcutt
at Magdalena Bay, Lower California.
Tellina IMoerellal felix Hanley.
Plate I, Fig. 1.
Tellina felix Hanley, Proc. Zool. Soc. Lon-
don, September, 1844, p. 71. “Hab. Panama;
sandy mud, six to ten fathoms.” — Hanley,
Thes. Conch., Vol. 1, 1846, p. 281, pi. 57,
fig. 52. “Panama.”
Type Locality : Panama, in 6 to 10 fathoms,
sandy mud.
Range: Mazatlan, Mexico, to Zorritos,
Peru.
Collecting Station: Nicaragua: Potosi and
Monypenny Point, Gulf of Fonseca; Corinto
(200-D-8, 9, 17, 19), 6-13 fathoms, sand,
mangrove leaves, also beach drift.
Description: Shell small, elongate, anteri-
or end much the longer, rounded, the posteri-
or end very short and obtusely obliquely sub-
truncated, anterior dorsal and ventral mar-
gins nearly parallel, exterior and interior of
a glossy rose red color; a posterior area is
set off by an umbonal angulation; valves
ornamented with fine, regular, concentric
striae; hinge of right valve with two cardi-
nals, the posterior one grooved, a strong,
high, elongate anterior lateral extends al-
most to the beak and a small posterior lateral
is present just beyond and below the posteri-
or end of the ligamentary area; left valve
with a well-developed anterior cardinal, a
thin posterior cardinal lamella is almost
fused to the posterior margin and a pointed
projection of the nymph represents an an-
terior lateral; pallial sinus not quite reach-
ing the anterior adductor impression and
along the base, except for a short distance,
it is confluent with the pallial line.
A specimen from the Gulf of Fonseca in
the Henry Hemphill collection in the Cali-
fornia Academy of Sciences measures ap-
proximately : length 17.2 mm. ; height, 9.4
mm.; convexity (both valves together), 4.3
mm.
The shell of this little species is character-
ized by the beautiful glossy rose red color,
weakly inflated valves, nearly parallel anteri-
or dorsal and ventral margins, strong right
anterior lateral, and by the short, obliquely
subtruncated posterior end.
The shorter posterior end, more gently
sloping anterior dorsal margin and deep red
color are features which serve to separate
Tellina felix from T. tabogensis Salisbury.
The longer, stronger, right anterior lateral
and more steeply sloping posterior dorsal
area just below the beaks are features sepa-
rating T. felix from T. carpenteri Dali.
Carpenter19 mentioned that Tellina puella
19 Carpenter, P. P., Proc. Zool. Soc. London, 1863, p. 366.
Reprint in Smithson. Miscell. Coll., No. 262, 1872, p. 202.
C. B. Adams20 [ —T . puellula Salisbury] is
“not unlike T. felix.” The remarks of Pilsbry
& Lowe seem to indicate that the species de- ;
scribed by Adams possesses a higher, heavier
shell than T. felix.
Distribution: Specimens of this species
were dredged off Nicaragua in 6-13 fathoms
and also were taken in the beach drift. It has
been recorded with doubt, as occurring in the
Miocene21 of Peru, and definitely in the
Pliocene of Ecuador. This species has been
cited as occuring in the Red Sea but, as men-
tioned by Lamy22, it does not occur in that
region.
Tellina IMoerellal macneilii Dali.
Tellina ( Angulus ) macneilii Dali, Proc.
U . S. Nat. Mus., Vol. 23, No. 1210, November,
1900, pp. 303, 318, pi. 3, fig. 7. “Obtained at
Guaymas, Mexico, by W. H. Dali.”
Type Locality: Guaymas, Mexico.
Range : Guaymas, Mexico, to the Gulf of
Nicoya, Costa Rica.
Collecting Station: Costa Rica: Cedro Is-,
land to off Ballena Bay, Gulf of Nicoya (213- '
D-11-17), 35-40 fathoms, mud.
Description : Shell small, solid, inequilat-
eral, the anterior end longer, rounded, the
posterior end quite short, depressed, bluntly
pointed ; color deep rosy, slightly zoned, and
paler toward the basal margin ; surface
closely, sharply concentrically striated, the
posterior dorsal area feebly imbricate, with
a little obscure radial striulation ; valves
moderately full, flatfish toward the middle
of the disk; hinge strong, normal; internal
ray obscure; pallial sinus long, nearly reach- I
ing the anterior adductor scar, wholly con- j
fluent below. Lon. 12.5, alt. 7.6, diam. 3.5 mm.
(Original description).
One pair and two single valves dredged in
the Gulf of Nicoya agree well with Dali’s de-
scription and illustration. The largest speci-
men measures approximately: length 11.5
mm. ; height, 7.3 mm. ; convexity (both valves
together) , 4.2 mm. These are identical with
specimens identified as T. macneilii in the
Lowe collection in the San Diego Society of
Natural History.
The hinge of this species is similar to that
of T. felix Hanley.
Compared to T. felix, the valves of T. mac-
neilii are more inflated in proportion to their
size, less elongate, the posterior end is more
abruptly truncated and the anterior dorsal
margin slopes more steeply. These same char-
acters (except the slope of the anterior dor-
sal margin) as well as the pink color likewise
serve to separate this species from T. tabo-
gensis.
Tellina guaymasensis Pilsbry & Lowe23 is
20 See footnote No. 17, p. 69.
21 Tellina (Eury tellina) cf. felix Hanley, Olsson, Bull.
Amer. Paleo., Vol. 19, No. 68, June 30, 1932, p. 123, pi. 14,
fig. 8. “Tumbez formation, Que Tucillal at Zorritos." Peru.
Miocene.
22 Lamy, E., Bull. Mus. Nat. Hist. Nat. (Paris), Vol.
24, No. 2, 1918, p. 119 (footnote).
23 Tellina ( Angulus ) guaymasensis Pilsbry & Lowe, Proc.
Acad. Nat. Sci. Philadelphia, Vol. 84, May 21, 1932, p. 94,
pi. 16, fig. 7. "Guaymas.” Mexico.
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
71
quite similar to T. macneilii but the shell
appears to be a little more depressed medially
and is in greater part white with streaks of
pink.
The present specimens agree well with the
brief description of Tellina deshayesii Car-
penter24 from the Bay of Panama. It was
said to resemble Tellina similis Sowerby but
much more inequilateral. However the spe-
cific name proposed by Carpenter is unten-
able in any case because of the prior use of
that combination of names, Tellina deshaye-
sii, by Hanley25.
Distribution : Specimens of Tellina mac-
neilii were taken by the expedition in the
Gulf of Nicoya in 35-40 fathoms. This record
represents an extension south of the known
range of this species.
Tellina IMoerellal paziana Dali.
Tellina ( Moerella ) paziana Dali, Proc. U.
S. Nat. Mus., Vol. 23, No. 1210, November,
1900, pp. 303, 318, pi. 3, fig. 8. “Dredged in
26 V2 fathoms, near La Paz, Lower Califor-
nia, by the U. S. Fish Commission, at station
2823.”
Type Locality : Near La Paz, Lower Cali-
fornia, Mexico, in 26*4 fathoms.
Range : Gulf of California to Cedro Island,
Costa Rica.
Collecting Stations : Mexico : Port Guatulco
(195-D-2, 7, 10, 11), 3-5 fathoms, sand, gr.
sand, crushed shell, dead coral ; Tangola-Tan-
gola Bay (196-D-6, 7, 13, 14, 15), 5-12.8
fathoms, sand, crushed shell; Nicaragua:
Corinto (200-D-8, 9, 19,) , 6-13 fathoms, man-
grove leaves, also on the beach ; Costa Rica :
Cedro Island, Gulf of Nicoya (213-D-1-10) ,
4-10 fathoms, mud, sand, crushed shell.
Description : Shell small, thin, white, con-
vex, the anterior end slightly longer, round-
ed, the posterior end bluntly pointed ; surface
finely concentrically sculptured by the incre-
mental lines, covered with a very delicate de-
hiscent pale straw-colored epidermis; hinge
well-developed, a minute but distinct anteri-
or left lateral present; interior polished, only
about half the lower portion of the pallial sin-
us confluent, the anterior part not reaching
the adductor. Lon. 10.2, alt. 7, diam. 3.5 mm.
(Original description).
The shell of this species is usually small,
about 10-15 mm. in length. A large right
valve in the present collection from Corinto,
Nicaragua, measures approximately: length,
17.5 mm.; height 14 mm., convexity (one
valve) , 3.3 mm. ; pallial sinus extends for-
ward 12.8 mm. from the posterior margin.
The pallial sinus in this species ascends
to a high rounded point slightly posterior to
a line vertical with the beaks, then descends
obliquely. The end is rounded and well sepa-
rated from the anterior adductor impression.
24 Tellina deshayesii Carpenter, Proc. Zool. Soc. London ,
November 11, 1856, p. 160. “Hab. in Sinu Panamensi ;
legit T. Bridges. Sp. un. in Mus. Cuming.**
25 Tellina deshayesii Hanley, Proc . Zool. Soc . London,
December, 1844, p. 148. “Hab. Red Sea? Mus. Cuming,
Deshayes.”
Tellina paziana somewhat resembles T.
meropsis Dali. It differs from that species in
the more elongate outline, the anterior end
is longer in proportion to the length, the pos-
terior end is much more bluntly pointed and
the pallial sinus is separated from the an-
terior adductor impression by a much wider
space. According to Dali, “This differs from
the young of Scrobiculina viridotincta Car-
penter, which in outline it resembles, by be-
ing less polished, more inflated, and without
the deep-set resilium.”
Distribution: This species was taken by
the expedition from off western Mexico to the
Gulf of Nicoya, Costa Rica, in 3-13 fathoms,
also on the beach. This is an extension south
of the known range of this species.
Tellina IMoerellal recurvata Hertlein &
Strong, sp. nov.
Plate I, Figs. 2, 3, 4, 8.
Tellina ( Angulus ) recurva Dali, Proc. U.
S. Nat. Mus., Vol. 23, No. 1210, November,
1900, pp. 304, 320, pi. 3, fig. 4. “Dredged near
the head of the Gulf of California in 24 fath-
oms, mud, off Point Fermin.”
Not Tellina recurva Deshayes, Proc. Zool.
Soc. London for 1854 (issued May 16, 1855),
p. 361. “Hab. Australia.”— Hedley, Proc.
Linn. Soc. New South Wales, Vol. 38, Pt. 2,
1913, p. 272. Hedley stated: “I failed to find
this unfigured Australian species in the Brit-
ish Museum. It is recommeded that the name
be treated as lost and unrecognizable.”
Description : Shell small, translucent
white, polished, rather compressed, beaks
very low, the anterior end the longer; anteri-
or dorsal margin gently curved, ventral mar-
gin gently rounded, posterior dorsal margin
depressed below the beak, sloping gently
along the ligamentary area (about 2 mm. on
the type), the end obliquely sloping and
roundly truncated ; a weak posterior umbonal
angulation present; surface of valves with
concentric, chiefly incremental sculpture, the
posterior area with low lamellae ; hinge plate
arched anterior to the beaks; right valve
with two cardinals, the posterior one
grooved, anterior lateral strong and fairly
close to the cardinals, a posterior lateral oc-
curs below a socket at about the end of the
ligamental area; left valve with two car-
dinals, the posterior one sloping posteriorly,
a faint projection of the margin represents
an anterior lateral; pallial sinus subtri-
angular, extending forward about three-
fourths the length of the shell but well sepa-
rated from the anterior adductor impres-
sion, along the base confluent with the pal-
lial line. Dimensions of the type: length, 12
mm.; height, 7.5 mm.; convexity (both
valves together), 2.9 mm.
Holotype (Calif. Acad. Sci. Paleo. Type
Coll.), from Loc. 23802 (C.A.S.), San Luis
Gonzaga Bay, Gulf of California. Paratypes
from the same locality. Dredged by the ex-
pedition at the following stations: Mexico:
4 miles SSW. of Maldanado Point (192-D-
72
Zoologica: New York Zoological Society [34: 9
1), 26 fathoms, mud; Port Guatulco ( 195-
D-20), 23 fathoms, mud; Costa Rica: Port
Parker (203-D-l), 15 fathoms, sandy mud,
crushed shell.
Range: Point Firmin, Lower California,
near the head of the Gulf of California, to
Port Parker, Costa Rica.
Some of the shells of this species are iri-
descent. Dali pointed out that the shell of this
species somewhat resembles that of young
Macoma yoldif ormis but is more blunt pos-
teriorly and the hinge of course is different.
The more abruptly truncated posterior end
and white color are features separating the
present species from T. arenica. Compared
to T. tabogensis the outline of T. recurvata
is less elongate, the anterior dorsal margin
is more arcuate and the posterior dorsal
margin is not flexuous. Compared to T.
buttoni Dali, the shell of the present species
is less attenuated both anteriorly and pos-
teriorly, the posterior dorsal margin just
below the beaks slopes more gently and it
lacks the strong anterior ray internally
which is so conspicuous in Dali’s species.
Distribution: A few specimens of this
species were dredged by the expedition off
western Mexico in 23-26 fathoms and off
Port Parker, Costa Rica, in 15 fathoms. The
present record of the occurrence of this shell
off Costa Rica is an extension south of the
known range of the species.
Tellina IMoerellal suffusa Dali.
Tellina ( Angulus ) suffusa Dali, Proc.
U. S. Nat. Mus., Vol. 23, No. 1210, November,
1900, pp. 303, 319, pi. 3, fig. 10. “Collected at
San Ignacio Lagoon, Lower California, by
Henry Hemphill.”
Type Locality: San Ignacio Lagoon, west
coast of Lower California.
Range: San Ignacio Lagoon, Lower Cali-
fornia, to Corinto, Nicaragua.
Collecting Station: Nicaragua: Corinto,
beach.
Description: Shell cuneate, very thin, con-
vex, blunt in front, pointed behind, the pos-
terior end slightly longer, pinkish, yellowish,
or translucent white in color ; surface rather
strongly, closely, and irregularly concen-
trically striate, with an unusually large and
wide lunular impression, but no escutcheon
to speak of ; hinge normal, delicate ; interior
polished; the pallial sinus high, well sepa-
rated from the anterior adductor, though
there seems to be no trace of a ray in the
specimens examined. Lon. 13.5, alt. 9.2, diam.
4.7 mm. (Original description).
The unusually large lunular area, short,
blunt anterior end and pointed posterior end
are features characteristic of this little
species.
Several single valves from Corinto, Nica-
ragua, the largest measuring 11 mm. in
length, agree well with Dali’s description
and illustration of Tellina suffusa. They like-
wise appear to be identical with specimens
of that species from Magdalena Bay in the
Hemphill collection of the California Acad-
emy of Sciences.
Tellina pumila Hanley,26 described from
Chile, is somewhat similar in outline but the
posterior end is narrower.
Distribution: Specimens of this species
were taken by the expedition only in beach
drift at Corinto, Nicaragua. This is a con-
siderable extension south of the known range
of this species.
Tellina IMoerellal tabogensis Salisbury. '
Tellina ( Angulus ) panamensis Dali, Proc.
U. S. Nat. Mus., Vol. 23, No. 1210, November,
1900, p. 319, pi. 3, fig. 3. “Types.— No. 108557,
U.S.N.M., dredged in 30 fathoms in Panama
Bay by the U. S. Fish Commission, at station
2799.”
Not Tellina panamensis Philippi, Zeit. f.
Malakozool., Jahrg. 5, No. 11, 1848, p. 175.
“Ad Panama legit frater E. B. Philippi.”
Tellina tabogensis Salisbury, Proc. Mala-
col. Soc. London, Vol. 21, Pt. 2, July, 1934,
p. 86. A new name for Tellina ( Angulus )
panamensis Dali, 1900, not Tellina panamen-
sis Philippi, 1848.
Type Locality: Panama Bay, in 30 fath-
oms.
Range : Gulf of California to Santa Elena
Bay, Ecuador.
Collecting Stations: El Salvador: Mean-
guera Island, Gulf of Fonseca (199-D-l), 16
fathoms, sand, mud, crushed shell ; La Union
(199-D-13), 6 fathoms, mud; Nicaragua:
Corinto (200-D-19), 12-13 fathoms, man-
grove leaves ; Costa Rica : Cedro Island, Gulf
of Nicoya (213-D-1-10) , 4-10 fathoms, mud,
sand, crushed shell.
Shell small, thin, ivory white, polished,
rather compressed, flexuous behind, the an-
terior end much the longer, produced and
rounded, posterior end with the ligament
rather deeply inset, margin obliquely de-
scending to a rather blunt point; surface
smooth or marked only by incremental lines,
except near the basal margin, where there
are a few incised lines with wider inter-
spaces, not quite in harmony with the lines
of growth; posterior dorsal area minutely
concentrically rippled; hinge normal, deli-
cate ; pallial sinus large, not reaching the ad-
ductor, mostly confluent below; the elevated
ray absent or obsolete. Lon. 9, alt. 5.25, diam.
2.5 mm. (Original description of Tellina
panamensis Dali).
Fresh specimens exhibit on the surface a
lovely iridescent glow (Dali).
A large series of specimens of this species
dredged off El Salvador by the expedition
have been compared with a series dredged
at Acapulco, Mexico, in the H. N. Lowe Col-
lection in the Museum of the San Diego So-
ciety of Natural History and with specimens
26 Tellina pumila Hanley, Proc. Zool. Soc. London, Sep-
tember, 1844, p. 69. “Hab. Valparaiso: sandy mud, from
seven to thirty fathoms.” —Hanley, Thes. Conch., Vol.
1, 1846, p. 279, pi. 57, fig. 41. “Valparaiso.”
Tellina (Angulus) pumila Hanley, Salisbury, Proc.
Malacol. Soc. London, Vol. 21, Pt. 2, July, 1934, p. 91,
pi. 13, figs. 3 and 4. [Illustrations of holotype and para-
type].
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
73
in the California Academy of Sciences. It
appears from this study that Dali’s type, 9
mm. in length, is a juvenile shell and that the
species reaches a much larger size. A large
right valve in the present collection dredged
off El Salvador, measures : length, 18.3 mm. ;
height, 10 mm.; convexity (one valve), 2.6
mm.
The most striking features of this species
are the ivory white color, iridescence in fresh
shells and the flexuous posterior dorsal mar-
gin. This flexuosity is caused by the convex-
ity of an area on the posterior dorsal margin
just postei'ior to the ligamentary area. Some
specimens are faintly tinted with pink on the
umbonal area. Variation in the thickness
and in the flexuosity of the posterior dorsal
margin can be observed in a series of speci-
mens. These features are most pronounced
in the larger shells. The present specimens
as well as a series in the Collections of the
California Academy of Sciences from Pana-
ma Bay, a series in the same collection from
Santa Elena Bay, Ecuador, collected by
Woodbridge Williams, and a series in the H.
N. Lowe collection from Acapulco, Mexico,
show gradation from small, thin shells with
a slightly flexuous posterior dorsal margin to
fairly thick shells with strong flexuous pos-
terior margins.
The hinge of this species is similar to that
of Tellina felix except that the right anteri-
or lateral is shorter and in the left valve the
inner margin back of the beak is slightly
thickened and some large specimens show
traces of a left posterior lateral. The ivory
color, longer posterior end and more convex
posterior dorsal margin easily serve to sepa-
rate this species from T. felix.
Tellina hiberna Hanley27 appears to be a
very similar shell. The illustrations of that
species published by Hanley and Salisbury
indicate that the shell is more abruptly slop-
ing posteriorly, that there is a constricted
area just anterior to the posterior umbonal
ridge and anterior to this the shell is more
expanded than in T. tabogensis.
Distribution : This species was dredged by
the expedition off Meanguera Island, El Sal-
vador, in the Gulf of Fonseca, where it oc-
curred abundantly in 6 to 16 fathoms, off
Corinto, Nicaragua, in 12-13 fathoms, and in
the Gulf of Nicoya, Costa Rica, in 4-10 fath-
oms. It occurs north to the Gulf of California
and south to Ecuador.
Subgenus Eurytellina Fischer.
Key to the species of Eurytellina.
A. Shell white or brownish
a. Pallial sinus touching the anterior ad-
ductor impression
b. Very elongate; adult valves with a
median depressed area ventrally
planulata
27 See TeUina hiberna Hanley, Thes. Conch., Vol. 1, 1846,
p. 282, pi. 67, fig. 63. “Panama ; Bay of Guayaquil.” —Sal-
isbury, Proc. Malacol. Soc. London, Vol. 21, Pt. 2, 1934,
p. 91, pi. 13, figs. 7, 8, 9. [Illustrations of holotype and
para types].
bb. Shell high; without a median de-
pressed area ventrally laceridens
aa. Pallial sinus not touching anterior ad-
ductor impression
c. Sculpture coarse, about 1 rib
per millimeter panamanensis
cc. Sculpture fine, about 3 ribs per
millimeter eburnea
B. Shell entirely or partly some shade of pink
or red
a. Concentric sculpture of even strength
over shell
b. Pallial sinus confluent with a por-
tion of the posterior margin of the
anterior adductor impression
rubescens
bb. Pallial sinus not touching anterior
adductor impression
c. Concentric sculpture decussated
by radial striae princeps2S
cc. Concentric sculpture not decus-
sated, radial striae very fine or
absent
d. About 10 concentric grooves
per millimeter prora
dd. About 2 or 3 concentric
grooves per millimeter
e. Right valve with de-
pressed area anterior to
posterior umbonal angu-
lation simidans
ee. Right valve without de-
pressed area anterior to
posterior umbonal angu-
lation
f. Posterior dorsal area
with strong concen-
tric sculpture
mantaensis
ff. Posterior dorsal area
with weak concentric
sculpture or of
growth lines only
ecuadoriana 28
Tellina I Eurytellina I eburnea Hanley.
Tellina eburnea Hanley, Proc. Zool. Soc.
London, September, 1844 p. 61, “Hab. Tum-
bez, Peru ; in soft sandy mud, five fathoms.”
—Hanley, Thes. Conch., Vol. 1, 1846, p. 241,
pi. 58, fig. 91. Tumbez, Peru. — Sowerby,
Conch. Icon., Vol. 17, Tellina, 1867, species
60, pi. 13, fig. 60. Original locality cited.
28 Not represented in the present collection.
aa. Concentric sculpture of unequal
strength over shell
g. Posterior third of shell with coarse
concentric lamellae giving way to
fine striae anteriorly
inaequistriata
gg. Posterior third of shell smooth or
nearly so in one or both valves, an-
teriorly sculptured with distant
concentric grooves regia
74
Zoologica: New York Zoological Society
[34: 9
Tellina ( Peronaeoderma ) eburnea Hanley,
Morch, Malakozool. Blatter, Bd. 7, 1860, p.
186. Sonsonate, El Salvador.
Type Locality : Tumbez, Peru, in 5 fath-
oms, sandy mud.
Range : Gulf of California to Tumbez,
Peru.
Collecting Stations: Guatemala: 7 miles
west of Champerico (197-D-l, 2), 14 fath-
oms, mud; El Salvador: La Libertad (198-D
-1,2), 13-14 fathoms, mud ; Nicaragua : Cor-
into, beach; Panama: Gulf of Chiriqui (221-
D-l-5) , 35-40 fathoms, sandy mud.
Description : Oblong, solid, opaque, rather
inequivalve, convex, whitish, glossy, clearly
inequilateral, with strong deep concentric
sulci, which usually become obsolete in one of
the valves, and which diverge and become
elevated on passing the flattened space at the
upper edge of the more convex valve ; ventral
edge very slightly convex, curving obliquely
upward anteriorly; posterior side much the
shorter, subcuneiform; the ligamental edge
straight, and abruptly sloping; ligament
shoi't and prominent ; fold and umbonal ridge
almost obsolete; inside pure white, teeth as
in punicea. (Hanley, Thes. Conch., 1846).
The largest specimen in the present collec-
tion measures : length 28.3 mm. ; height, 17.8
mm. The present specimens agree well with
the younger stages of a specimen of T. ebur-
nea in the collection at Stanford University
which was collected in Ecuador by Stanley
Herold, which measures : length, 47.5 mm. ;
height, 29.8 mm.; convexity (both valves to-
gether), 12.6 mm.
The concentric sculpture on the posterior
dorsal areas becomes stronger and raised
after crossing the umbonal ridge. This fea-
ture is emphasized by Hanley and Sowerby
but it is not very pronounced on the present
specimens.
Tellina eburnea is very similar to T. alter-
nata Say, an east American species.
Tellina laplata Pilsbry & Olsson29, describ-
ed from the Pliocene of Peru, is also very
similar to T. eburnea. Compared to T. laplata
the present specimens do not show such
strong raised sculpture on the posterior dor-
sal areas nor is the sculpture as strong on the
left anterior dorsal area as that shown in the
illustrations by Pilsbry & Olsson.
The shell of Tellina eburnea is higher in
proportion to the length as compared to that
of T. simulans, furthermoi'e the color is pure
white both exteriorly and interiorly. The
general character of the pallial sinus is simi-
lar to that of T. simulans. It is somewhat
higher behind and extends forward almost to
but does not quite touch the anterior adduc-
tor impression and is confluent with the pal-
lial line below. The hinge is similar to that of
T. simulans except that the right posterior
lateral is less distant from the cardinals. The
sculpture is much finer and the pallial sinus
29 Tellina (Eurytellina) laplata Pilsbry & Olsson, Proc.
Acad. Nat. Sci. Philadelphia, Vol. 93, September 9, 1941,
p. 67, pi. 15, figs. 1-5. "Canoa formation, Punta Blanca.”
Ecuador. Pliocene.
extends nearer the anterior adductor impres-
sion than that of T. panamanensis Li.
Distribution: Specimens here referred to
Tellina eburnea were dredged by the expedi-
tion in 13-40 fathoms, from off Guatemala
and El Salvador, in the Gulf of Chiriqui,
Panama, and were taken on the beach at Cor-
into, Nicaragua.
Tellina I Eurytellina I inaequistriata Donovan.
Plate I, Fig. 18.
Tellina inaequistriata Donovan, Nat. Hist.
Brit. Shells, Vol. 4, 1802, pi. 123 [two figs.].
“A very rare species of Tellina communi-
cated to Da Costa after his Conchology was
published, and therefore not noticed in that
work. It has been found by the late Dr. Pul-
teney we believe on the coast of Dorsetshire.”
— Chenu, Bibl. Conchyl., Ser. 1, Vol. 1, 1845,
p. 82, pi. 32, figs. 7, 8. [French translation of
Donovan’s work on Shells]. — Hanley, Thes.
Conch., Vol. 1, 1846, p. 238, pi. 57, fig. 58; pi.
58, fig. 80. “Bay of Guayaquil; (Cuming).” —
Forbes & Hanley, Hist. Brit. Moll., Vol. 1, |
1853 (issued August 1, 1848), p. 314. “In- '
habits the Bay of Guayaquil; was introduced
by Donovan, who only surmised that it had
been taken by Dr. Pulteney on the Dorset
coast.”
Tellina sanguinea Wood, Gen. Conch., 1815,
p. 159, pi. 44, fig. 2. “This shell is in the cabi-
net of Dr. Coombe.” [No locality cited]. —
Wood, Index Test., 1825, p. 18, pi. 4, fig. 27.
Also ed. 1828. Locality unknown. Also ed. by
Hanley, 1842-1856, p. 23, pi. 4, fig. 27.
“Guayaquil.” [States that fig. 80 in Thes.
Conch, represents T. sanguinea ]. — Hanley,
Cat. Rec. Biv. Shells, 1843, p. 67.
Tellina ( Eurytellina ) leucogonia Dali,
Proc. U. S. Nat. Mus., Vol. 23, No. 1210, No-
vember, 1900, p. 317, pi. 4, fig. 5. “Type. —
No. 102182, U.S.N.M., from the Gulf of Cali-
fornia, Stearns collection.”
Type Locality : Bay of Guayaquil, Ecuador
( according to Hanley and Forbes & Hanley) .
[Erroneously cited from the coast of Dorset-
shire, England, by Donovan].
Range: Gulf of California to the Bay of
Guayaquil, Ecuador. Caribbean (Dautzen-
berg) .
Collecting Stations : Mexico : Santa Cruz
Bay (195-D-19-21) , 17-18 fathoms, mud, gr.
mud, crushed shell; Tangola-Tangola Bay
(196-D-13), 10 fathoms, gr. sand, crushed
shell; Costa Rica: Port Parker (203-D-3),
12 fathoms, shelly mud.
Description: Shell ovate, compressed and
rather flattish, rosy, very finely striated
transversely : the striae fewer and larger at
the anterior [posterior] end (Donovan).
Shell elongate, moderately thick, some-
what compressed, glossy, subequilateral, red
or orange-red; a well-defined fairly broad
posterior area is set off by a rounded post-
umbonal ridge; posterior dorsal margin
slightly rounded, sloping downward and
slightly expanded along the ventral half ; the
ornamentation consists of concentric striae
which, especially on the right valve, are very
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
75
strongly developed and widely spaced on
about the posterior third of the shell then
change abruptly to fine striae anteriorly;
hinge normal for the subgenus; the pallial
sinus does not quite touch the anterior ad-
ductor impression but is separated by a nar-
row area, wholly confluent with the pallial
line below and, in general features, similar
to that of T. simulans and T. prora.
A right valve in the present collection from
Santa Cruz Bay, Mexico, measures : length,
23 mm. ; height, 12.5 mm. It agrees well with
Hanley’s plate 50, figure 80. Other specimens
in the collection are similar but show the
strong sculpture only slightly developed an-
terior to the angulation.
Some of the small shells, especially left
valves, almost lack strong concentric sculp-
ture anterior to the posterior area; in such
cases they may be ornamented only by faint
grooves slightly out of harmony with the
incremental lines.
A series of specimens in the H. N. Lowe
collection of the San Diego Society of Natu-
ral History, varying in length from approxi-
mately 8.5 mm. to 26.2 mm. and identified by
Lowe as Tellina leucogonia Dali, are identical
with the present specimens. The present ser-
ies together with Lowe’s specimens show all
variations from young smooth shells up to a
large right valve with the typical sculpture
of T. inaequistriata. We are therefore inclin-
ed to consider Dali’s species as identical with
T. inaequistriata.
The very distinct concentric sculpture,
changing from coarse to fine at about the
posterior third of the shell, serves to sepa-
rate this form from similar west American
species of the genus.
Tellina waylandvaughani Maury30, de-
scribed from the Miocene of Santo Domingo,
is a similar species.
Distribution : Specimens of this species
were dredged by the expedition in 10 to 18
fathoms from off western Mexico and Costa
Rica. The present records of occurrence show
the range of this species to extend from the
Bay of Guayaquil north to the Gulf of Cali-
fornia. Dautzenberg31 cited Tellina inaequi-
striata as occurring in the Caribbean region
at the Island of Martinique and in Venezuela.
He stated that he could detect no differences
which would serve as a basis for separating
the Caribbean shells from those illustrated
under that name from the Bay of Guayaquil.
Tellina I Eurytellina I laceridens Hanley.
Tellina laceridens Hanley, Proc. Zool. Soc.
London, September, 1844, p. 61. “Hab. Tum-
bez, Peru; soft sandy mud, five fathoms.”
“Var. testa, magis trigona . . .” “Hab. chir-
iqui, West Columbia ; sandy mud, three fath-
oms.” —Hanley, Thes. Conch., Vol. 1, 1846,
P- 243, pi. 61, figs. 168, 176. [Not. pi. 66,
SO Tellina waylandvaughani Maury, Bull. Amer. Paleo.,
Vol. 5. No. 29, 1917, p. 386 (222) , pi. 64 (38) , figs. 7 and 8.
‘‘Zone G. Rio Gurabo at Los Quemados.” Santo Domingo.
Miocene.
81 Dautzenberg, P., Mem. Zool. Soc. France, Vol. 13
1900, p. 260.
fig. 258.]. Original localities cited. — Sower-
by, Conch. Incon., Vol. 17, Tellina, 1867, spe-
cies 104, pi. 20, fig. 104. Tumbez, Peru.
Type Locality : Tumbez, Peru, in 5 fath-
oms, soft, sandy mud.
Range: Realejo [near Corinto], Nicara-
gua, to Tumbez, Peru.
Collecting Stations: Nicaragua: Corinto,
beach ; Costa Rica : Port Culebra ; Colombia :
Gorgona Island.
Description: Shell elongately trigonal,
beaks nearly central but slightly anteriorly
placed, posterior end obliquely truncated;
sculpture of fine, concentric grooves which
in places are irregular, the umbonal region,
and often the posterior area, relatively
smooth ; white, sometimes with a pinkish
spot on the umbonal region ; ligament large,
exterior; hinge with cardinals grooved, es-
pecially the right posterior cardinal which
appears ragged due to 7 to 10 grooves; the
pallial sinus touches the anterior adductor
impression just above the base; interior
white and yellow and in large specimens
with somewhat granular areas and salmon
pink spots.
A specimen collected at Gorgona Island,
Colombia, measures: length, 53 mm.; height,
34.8 mm.; convexity (both valves together),
11.6 mm.
Some of the characters in which this spe-
cies differs from Tellina panamanensis Li
are: the finer concentric sculpture, larger
smooth umbonal area, regular sculpture
along the posterior dorsal margin and in
that the pallial sinus touches the anterior
adductor impression.
Distribution: A few specimens of this
species were collected by the expedition in
the beach drift at Corinto, Nicaragua, one
at Port Culebra, Costa Rica, and one at Gor-
gona Island, Colombia. It also has been re-
corded as occurring in beds of Pliocene age
in Panama.
Tellina lEurytellinal mantaensis Pilsbry &
Olsson.
Tellina ( Eurytellina ) mantaensis Pilsbry
& Olsson, Nautilus, Vol. 56, No. 3, January,
1943, p. 80, pi. 8, figs. 1-4. “Manta, Ecuador.”
Type Locality: Manta, Ecuador.
Range : Gulf of Chiriqui, Panama, to Man-
ta, Ecuador.
Collecting Station : Panama : Gulf of Chir-
iqui (221-D-1-5), 35-40 fathoms, sandy mud.
Description : Shell elongated, rather com-
pressed, the anterior end slightly the long-
er, posterior end obliquely subtruncated ;
sculpture of flat concentric ridges which are
separated by narrower grooves; the poster-
ior area is usually ornamented by waved
raised threads and with faint radial lines
on one or the other valve; hinge of right
valve with a strong anterior lateral adja-
cent to the cardinals and a more distant pos-
terior lateral, the laterals are smaller in the
left valve; the pallial sinus reaches almost
to the anterior adductor impression and in
this character is similar to T. simulans,
76
Zoologica: New York Zoological Society
[34: 9
and is confluent with the pallial line below;
fresh specimens are rose colored, more
deeply on the umbos, and somewhat brown-
ish colored ventrally.
Two valves in the present collection agree
well with the illustrations of T. mantaensis
Pilsbry & Olsson. The larger specimen meas-
ures approximately 21 mm. in length and
15 mm. in height.
The shell of this species differs from that
of Tellina simulans in that it is more elon-
gated and the dorsal margins slope more
gently, especially posteriorly where the area
is somewhat flattened. There also are dif-
ferences in the details of the hinges of the
two species. The right posterior cardinal of
T. mantaensis reaches almost to the ventral
margin of the hinge plate which beneath
that tooth is strongly indented. In T. simu-
lans the corresponding tooth reaches little
more than halfway to the ventral margin
of the hinge plate which at that point is
gently rounded.
The strong concentric sculpture on the
posterior dorsal area and more gently slop-
ing anterior dorsal margin are features
which serve to sepai'ate this species from
T. ecuadoriana Pilsbry & Olsson.
Distribution: Two single valves of this
species were taken by the expedition in 30-
40 fathoms on a bottom of sandy mud in the
Gulf of Chiriqui, Panama. This is an exten-
sion north of the known range of the species.
Tellina I Eurytellinal panamanensis Li.
Tellina panamanensis Li, Bull. Geol. Soc.
China, Vol. 9, No. 3, October, 1930, p. 262,
pi. 5, fig. 32. “ ‘Brought up by marine dredge
from depths varying from 10. ft. to 40. ft.
in the mud at the mouth of the Rio Grande
near La Boca about one mile from the main-
land in Panama Bay.’ ” “Horizon : Gatun
formation.” —Pilsbry, Nautilus, Vol. 58, No.
4, April, 1945, p. 145.
Tellina ( Eurytellina ) panamanensis Li,
Pilsbry, Proc. Acad. Nat. Sci. Philadelphia,
Vol. 83, November 13, 1931, p. 436, pi. 41,
figs. 4, 5, 6. A Recent shell from Panama
Bay.
Tellina liana Hertlein & Strong, Nautilus,
Vol. 58, No. 3, January, 1945, p. 105.
“Dredged off Meanguera Island, El Salva-
dor, in the Gulf of Fonseca, in 16 fathoms.”
Type Locality : Mouth of Rio Grande near
La Boca about 1 mile from the mainland
in Panama Bay, 10-40 feet, mud.
Range: Tenacatita Bay, Mexico, to the
Bay of Panama.
Collecting Stations : Mexico : Tenacatita
Bay (183-D-2), 30 fathoms, muddy sand;
Port Guatulco (195-D-20, 21), 18-23 fath-
oms, mud; Tangola-Tangola Bay (196-D-17,
18), 23-30 fathoms, mud; El Salvador: Me-
anguera Island, Gulf of Fonseca (199-D-l),
16 fathoms, sand, mud, crushed shell.
Description: Shell similar in general out-
line to that of Tellina laceridens but more
steeply truncated posteriorly ; the left valve
is more convex and overlaps the right along
the right anterior dorsal margin; a faint /
broad median concavity is present toward
the ventral margin of the valves ; color gray-
ish-white covered by a thin ochraceous per- i
iostracum; the concentric sculpture con-
sists of ridges (about 1 per millimeter)
which on their upper portions are flat, on
the lower sloping, crossed by fine weak ra-
dial striae; on the right valve a ridge or
angulation occurs from beak to base pos-
teriorly; on the posterior dorsal area the
concentric sculpture is usually sinuated,
sometimes bent back, due to the presence |
(although sometimes absent) of a median
radial convexity; hinge with two cardinals
and two laterals in each valve, the right an-
terior cardinal grooved, the posterior with
about four sulcations, the anterior lateral
close to the cardinals, the posterior lateral
distant about one-third the length of the i
posterior dorsal margin ; left valve with the
anterior cardinal grooved, the posterior car- i
dinal a thin lamella, laterals weak; pallial
sinus highest beneath the beaks, descending
to a broadly rounded or blunt point which
is well separated from and lower than but j
posterior to the anterior adductor impres- ,
sion, along the base confluent with the pal- t
lial line. Dimensions of a typical specimen : I
length, 44.9 ; height, 29 mm. ; convexity .}
(both valves together), 13 mm.; pallial
sinus extends anteriorly 34.5 mm. from pos-
terior end of shell.
The shell of this species differs from that
of Tellina laceridens in the more steeply 't
sloping posterior dorsal margin, more con- 1
vex valves, much coarser sculpture which, ■
especially on the left valve, is sinuated on
the posterior dorsal area, in the smaller
smooth area at the beaks, less crenated car- u
dinal teeth and in that the pallial sinus does '
not touch the anterior adductor impression r
but is separated from it by a considerable
distance. The much coarser sculpture and the
much greater distance between the pallial
sinus and the anterior adductor impression
are features separating it from T. eburnea.
Distribution : Specimens of Tellina pan- I
amanensis were dredged by the expedition
from Tenacatita Bay, Mexico, to Meangu-
era Island, El Salvador, in 16-30 fathoms.
This record of Tenacatita Bay, Mexico, is
an extension north of the known range of
the species. It also has been recorded as oc-
curring in the Pliocene of Ecuador.
Tellina (Eurytellinal planulata Sowerby.
Plate I, Fig. 22.
Tellina planulata Sowerby, Conch. Icon., i
Vol. 17, Tellina, June, 1867, species 186, pi.
33, fig. 186. “Hab. — ?”
Type Locality: Gulf of Dulce, Costa Rica
(here designated as type locality). No lo-
cality originally cited.
Range: La Libertad, El Salvador, to the
Gulf of Dulce, Costa Rica.
Collecting Stations: El Salvador: La Lib-
ertad (198-D-2), 14 fathoms, mud; Costa
1949]
Hertlein & Strong: Mollusks of Mexico and. Central America
77
Rica: Gulf of Dulce; Golfito Bay, Gulf of
Dulce.
Description: Shell elongate, the posterior
end the longer, fairly thick, white, similar
to Tellina laceridens in general features but
longer in proportion to the height; some-
times with a low broad medial depression
toward the ventral margin; posterior dor-
sal area set off by a slight angulation ; sculp-
ture consisting of very fine, shallow, some-
what irregularly spaced concentric grooves ;
right valve with two grooved cardinals, an-
terior lateral close to the cardinals but the
posterior lateral distant about half the
length of the posterior dorsal margin; left
valve with a grooved anterior cardinal and
posterior to this two small, thin, laminae,
latex'als small; the pallial sinus projects an-
teriorly and barely touches the base of the
anterior adductor impression, along the base
it is confluent with the pallial line.
A left valve measures: length, 59.2 mm.;
height, 33 mm. ; convexity (one valve) , 6 mm.
Several single valves from the Gulf of
Dulce agree so closely with Sowerby’s de-
scription and figure of Tellina planulata that
we have assigned our specimens to that spe-
cies. This species was originally described
without information as to the locality from
which it came. Paetel32 cited the species as
occurring at “Sitka,” Alaska, but we have not
seen any specimens from that region which
appear to be referable to it. Other than Pae-
tel’s record the species apparently has not
been recognized as occurring elsewhere. We
therefore have designated the Gulf of Dulce
as type locality. Sowerby stated that it is
“A much larger and flatter shell than Tel-
lina eburnea, with closer grooves and no
transverse ridges on the dorsal margin of the
overlapping valve.” Those differences are
true with regard to the present specimens.
The pallial sinus in the present specimens
touches the anterior adductor impression
whereas in T. eburnea it is separated from
the corresponding impression by a narrow
space. Sowerby stated with regard to the
posterior end: “terminal margin notched.”
This latter feature is not pronounced on our
specimens but they are not perfectly pre-
served.
The present specimens closely resemble
Tellina ecuadoriana Pilsbry & 01sson32a de-
scribed from Ecuador. A single valve of that
species collected by the senior author at Cor-
into, Nicaragua, is in the collections of the
California Academy of Sciences.
Compared to Tellina ecuadoriana the pos-
terior dorsal margin of T. planulata slopes
a little more steeply and the posterior area,
especially on the right valve, appears to be
a little narrower than the corresponding
32 Paetel, Fr„ Cat. Conchyl.-Samm]., Vierte Neube-
arbeitung (Berlin: Verlag von Gebruder Paetel), Abt
3, 1890, p. 49.
32a Tellina ( Eurytellina ) ecuadoriana Pilsbry & Olsson,
Proc. Acad. Nat. Sci. Philadelphia, Vol. 93, September 9,
1941, p. 67, pi. 15, figs. 6, 7, 8. Canoa formation, Punta
Blanca. Ecuador, Pliocene. Also Recent, Santa Elena.
Ecuador (type), also at Canoa and Callo, the port of
Jipijapa.
area on the species described by Pilsbry &
Olsson. The shell of the present species is
white rather than rose red with whitish
zones. Furthermore the pallial sinus in the
present specimens touches the anterior ad-
ductor impression while in T. ecuadoriana
the two are separated by a narrow space.
Distribution : Several valves of this spe-
cies were collected by the expedition on the
beach in the Gulf of Dulce, Costa Rica. One
small specimen with both valves was dredged
in 14 fathoms off La Libertad, El Salvador.
Tellina lEurytellina) prora Hanley.
Tellina prora Hanley, Proc. Zool. Soc. Lon-
don, September, 1844, p. 61. “Hab Porto St.
Elena, West Columbia; sandy mud, six fath-
oms; and Salango, West Columbia, sandy
mud, nine fathoms.” — Hanley, Thes. Conch.,
Vol. 1, 1846, p. 243, pi. 60, fig. 152. Original
localities cited. — Sowerby, Conch. Icon.,
Vol. 17, Tellina, 1866, species 90, pi. 18,
fig. 90. Original localities cited. — Salisbury,
Proc. Malacol. Soc. London, Vol. 21, Pt. 2,
July, 1934, p. 86. “The shell ranges through
the south of the Panamic and north of the
Peruvian areas.”
Tellina cibaoica Maury, Li, Bull. Geol. Soc.
China, Vol. 9, No. 3, 1930, p. 261, pi. 4, fig. 30.
Dredged in Panama Bay. Referred to the
Gatun formation, Miocene. According to
Pilsbry {Proc. Acad. Nat. Sci. Philadelphia,
Vol. 83, 1931, p. 430), Li’s record was based
upon “A left valve of Tellina prora Hanley”. . .
Not Tellina cibaoica Maury, 1917. Santo
Domingo, Miocene.
Type Locality: Santa Elena, Ecuador, in
6 fathoms, sandy mud (here designated as
type locality). Salango, Ecuador, in 9 fath-
oms, sandy mud, also originally cited by
Hanley.
Range: Mazatlan, Mexico, to the Bay of
Guayaquil, Ecuador.
Collecting Stations: Mexico: Tangola-
Tangola Bay (196-D-17), 23 fathoms, mud;
Guatemala: 7 miles west of Champerico
( 197-D-l, 2) , 14 fathoms, mud ; El Salvador :
La Libertad (198-D-2), 14 fathoms, mud;
Meanguera Island, Gulf of Fonseca (199-
D-l), 16 fathoms, sand, mud, crushed shell;
La Union, Gulf of Fonseca (199-D-15), 6
fathoms, mud ; Nicaragua : Potosi and Mony-
penny Point; Costa Rica: Port Parker (203-
D-3), 12 fathoms, shelly mud; Panama: Ba-
hia Honda (222-D-5), 11 fathoms, mud,
shells, leaves.
Description: Shell ovately trigonal,
smooth, polished, a posterior area set off
by an angulation, colored rose pink with
whitish concentric bands; sculpture of very
fine closely spaced (about 10 per millimeter)
incised concentric striae; hinge with two
grooved cardinals in each valve, the left pos-
terior one very narrow, two laterals in each
valve, those in the left valve much the
smaller; pallial sinus highest in middle part
of shell, usually separated from the anterior
adductor impression by about a millimeter
78
Zoological New York Zoological Society
[34:9
but sometimes almost, but not quite, touch-
ing, the end blunt and almost in line verti-
cally below the posterior side of the adductor
impression, along the base confluent with
the pallial line.
A large specimen dredged off Guatemala
measures approximately: length, 46.4 mm.;
height, 17.5 mm.; convexity (both valves
together), 12 mm.
The shell of this species differs from that
of Tellina rubescens Hanley in that the pos-
terior slope is more gently inclined, the con-
centric incised striae are much finer and
much more closely spaced and the pallial
sinus does not touch the anterior adductor
impression. The very much finer and more
closely spaced concentric sculpture easily
serves to separate the species from T. sim-
ulans C. B. Adams.
The record of Tellina prora from the Cape
Verde Islands in the Atlantic cited by Roche-
brune33 can be referred to some other spe-
cies. Tellina (Eury tellina) trinitatis Tom-
lin34, described from Colon Harbor on the
east side of the isthmus of Panama, is said
to be similar to T. prora.
Distribution: This species was collected
by the expedition off western Guatemala,
El Salvador and Nicaragua, in 6-14 fath-
oms on a muddy bottom. It also has been re-
corded as occurring in the Pliocene of Ecua-
dor.
Tellina I Eurytellina! regia Hanley.
Tellina regia Hanley, Proc. Zool. Soc. Lon-
don, September, 1844, p. 61. “Hab. Real Lle-
jos, Central America; in coarse sandy mud,
seven fathoms.” — Hanley, Thes. Conch.,
Vol. 1, 1846, p. 240, pi. 60, fig. 140. Original
locality cited.
Type locality: Real Llejos [near Corinto],
Nicaragua, in 7 fathoms, coarse, sandy mud.
Range : Known only from the type locality
and vicinity.
Collecting Station: Nicaragua: Corinto,
beach drift.
Description: Oblong, thin, rather com-
pressed, almost inequivalve, subequilateral;
extremely glossy, both externally and in-
ternally of a deep subpellucid purplish crim-
son; the surface marked with distant con-
centric grooves, which posteriorly become
obsolete in one or both of the valves; the
ventral edge nearly straight, subretuse in
the middle; anterior side slightly shorter,
its extremity obtusely rounded ; posterior
extremity almost biangulated; dorsal mod-
erately and almost equally sloping on
either side of the beaks, nearly straight pos-
teriorly; umbonal ridge and flexure nearly
obsolete; the ligament rather prominent;
teeth as in punicea. (Hanley, Thes. Conch.,
1846).
S3 Peronaeoderma prora Hanley, Rochebrune. Nouv.
Arch. Mus. d’Hist. Nat. (Paris), Ser. 2, Vol. 4, 1881, p. 258.
“Hab.— Rade de Saint-Vincent.” Cape Verde Islands.
34 Eurytellina trinitatis Tomlin, Jour. Conch., Vol. 18,
No. 11, July, 1929, p. 310. “Hab. Colon Harbour, not un-
common, dead but very fresh in 6 f.”
A left valve from Corinto, Nicaragua
measures approximately: length, 17 mm.
height, 10 mm.; convexity (one valve), 1
mm.
The outline as well as the other features
of the present specimen are similar to thost
described for Tellina regia which came from
the same general vicinity. The concentric
ornamentation of the shell is like that de-
cribed for T. regia, namely, distant concen-
tric grooves which become obsolete poste
riorly.
Tellina regia differs from Tellina rubes
cens in that the concentric grooves become
obsolete posteriorly, the dorsal margins do
not slope so steeply, and the pallial sinus
does not touch the anterior adductor impres-
sion. It differs from Tellina princeps in lack
ing radial striae.
Distribution: Only one valve referred to
this species was found in the beach drift
at Corinto, Nicaragua.
kPr
liter
Ike i
Lov
Tellina I Eurytellina} rubescens Hanley.
Tellina rubescens Hanley, Proc. Zool. Soc.\ m
London, September, 1844, p. 60. “Hab. Pan- PP
ama and Tumbez; in sandy mud.” — Hanley, !IB
Thes. Conch., Vol. 1, 1846, p. 242, pi. 60, fig. jj1
153. Tumbez, and Panama (Cuming). — $!’
Sowerby, Conch. Icon., Vol. 17, Tellina, 1866, v
species 93, pi. 18, fig. 93. “Hab. Tumbez and A
Panama, Peru.”
Type Locality: Panama in sandy mud
(here designated as type locality). Tumbez,
Peru, in sandy mud, also originally cited. S(
Range: Tenacatita Bay, Mexico, to Turn- ;
bez, Peru.
Collecting Stations: Mexico: Tenacatita ‘
Bay; El Salvador: La Union, Gulf of Fon-
seca (199-D-12), 5 fathoms, mud; Nicara-
gua: Potosi and Monypenny Point, Gulf of D
Fonseca; Costa Rica: Port Parker.
Description: Shell trigonally ovate,
smooth, polished, colored exteriorly and in-
teriorly by light and darker concentric bands ;
of rose pink; posterior dorsal margin slop-
ing steeply, the area set off by an angulation ; |
sculpture consists of fine concentric grooves
(about 2 per millimeter) and between these
finer concentric striae, the whole crossed
by very fine submicroscopic radial striae;
hinge with two cardinals and two laterals
in each valve, the right anterior cardinal
usually grooved, the posterior bifid, left an-
terior grooved, the posterior one thin, lat-
erals in left valve weak ; anterior end of pal-
lial sinus confluent with the lower posterior
margin of the anterior adductor impression
from a point just above the base to approx-
imately the middle of the base of the im-
pression, confluent with the pallial line be-
low.
A large specimen from the Gulf of Fon-
seca measures : length, 43 mm. ; height, 37
mm.; convexity (both valves together), 9.3
mm.
The shell of Tellina rubescens differs from
that of T. prora Hanley in that it is higher
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
79
in proportion to the length, the posterior
dorsal margin slopes more steeply, the in-
cised concentric sculpture is more widely
spaced and the pallial sinus is confluent with
a portion of the posterior side of the ante-
rior adductor impression. These same char-
acters of proportionate height to length and
that of the pallial sinus serve to separate
T. rubescens from T. simulans C. B. Adams,
| a species in which the concentric sculpture
is much more strongly developed.
Distribution : This species was taken by
the expedition along the west coast of Mex-
ico and Central America but at no place abun-
dantly. It also has been recorded by Arnold,
1903, as occurring in the upper Pleistocene
of San Pedro, California. Some of the rec-
ords of the occurrence of this species at San
Ignacio Lagoon and at Magdalena Bay,
Lower California, are referable to T. simu-
lans.
Tellina I Eurytellina I simulans C. B. Adams.
Tellina simulans C. B. Adams, Ann. Ly-
ceum Nat. Hist. New York, Vol. 5, July, 1852,
pp. 508, 546 (separate pp. 284, 322). “Pan-
ama.” Also cited from Xipixapi, Ecuador,
in sandy mud at 10 fathoms. — Romer, Neues
Syst. Conchyl.-Cab. Martini-Chemnitz, Bd.
10, Abt. 4, Tellina, 1872, p. 99, pi. 25, figs.
4, 5.
Tellina punicea Born, Carpenter, Cat.
Mazatlan Shells, August, 1855, p. 35. Maz-
atlan, Mexico. Also earlier records cited. —
Sowerby, Conch. Icon., Vol. 17, Tellina, 1866,
species 53, pi. 12, fig. 53. “Hab. Xipixapi,
W. Columbia; in sandy mud, ten fathoms;
H. Cuming.”
Not Tellina punicea Born, Test. Mus. Caes.
Vind., 1780, p. 33, pi. 2, fig. 8. “Patria ig-
nota.” [Now believed to be a Caribbean spe-
cies. See Gardner, U. S. Geol. Surv., Prof.
Paper 142 -E, 1928, p. 193].
Tellina costaricana Olsson, Li, Bull. Geol.
Soc. China, Vol. 9, No. 3, October, 1930 p.
262, pi. 4, fig. 31. Dredged in Panama Bay
at mouth of Rio Grande River in 10-40 ft.
“Occurrence : Gatun Stage, Banana River,
Costa Rica.” “Horizon: Gatun formation.”
This record is based upon “Two valves of
Tellina simulans C. B. Ad.,” Panama, Recent,
according to Pilsbry ( Proc . Acad. Nat. Sci.
Philadelphia, Vol. 83, 1931, p. 430).
Type Locality : Panama.
Range : Scammon Lagoon, Lower Califor-
nia, to the Gulf of California and south to
Tumbez, Peru.
Collecting Stations: Mexico: Santa Inez
Bay, Gulf of California; Tenacatita Bay;
17 miles SE. X E. of Acapulco (189-D-3),
13 fathoms, mud; Nicaragua: Corinto (200-
D-ll, 19), 8-13 fathoms, sand, mangrove
leaves; Costa Rica: Culebra Bay; 1 mile
south of Golfito.
Description: Shell elongately oval, pos-
terior dorsal margin obliquely sloping, the
end obliquely truncated and slightly bent to
the right, rose pink with whitish concentric
bands; posterior area set olf by an angula-
tion anterior to which is a shallow broad
depressed area on the right valve, the pos-
terior area bears a narrow, shallow, radial
furrow which is especially noticeable on the
left valve; sculpture of regular, deep, con-
centric grooves (about 2 or 3 per millimeter)
and very fine submicroscopic radial striae;
hinge with anterior laterals close to the car-
dinals, posterior laterals distant, weaker in
left valve; the pallial sinus does not quite
touch the anterior adductor impression, the
end is blunt and almost in line vertically
below the posterior margin of the adducto**
impression, along the base it is confluent
with the pallial line.
A large specimen of this species in the
Henry Hemphill Collection of the California
Academy of Sciences from Magdalena Bay,
Lower California, measures: length, 48.8
mm.; height, 29 mm.; convexity (both valves
together), 11 mm.
This species, in some instances, has been
cited in the earlier literature under the
names of Tellina punicea Born and T. ru-
bescens Hanley.
The shell of this species differs from that
of T. rubescens in the greater length in pro-
portion to the height, in the much deeper,
sti'onger and more widely spaced concentric
sculpture and in that the pallial sinus does
not touch the anterior adductor impression.
Tellina simulans is similar to T. angulosa
Gmelin ( T . punicea of some authors), an
east American species ; in fact Carpenter and
others considered the two to be identical.
According to Adams T. simulans differs from
the east American species in that “its fur-
rows are deeper and are continued over the
flexure, without change of depth ; the inter-
spaces are less flattened, and the lateral teeth
are nearly obsolete.” A comparison of spec-
imens of T. simidans with a series of T. an-
gulosa collected by F. M. Anderson at Car-
tagena Bay, Colombia, shows differences be-
tween the two. The west coast shells are more
pointed posteriorly, there is a low depressed
area anterior to the posterior angulation on
the right valve, and the concentric grooves
along the posterior dorsal margin bend more
acutely upward than on the east coast shells.
Tellina princeps Hanley, described from
Tumbez, Peru, is a distinct species posses-
sing a large, red, subequilateral shell with a
gently sloping posterior dorsal margin and
the concentric sculpture is crossed by strong
radial striae.
Distribution: This species was taken by
the expeditions although not abundantly,
from Santa Inez Bay, in the Gulf of Califor-
nia to Culebra Bay, Costa Rica. It ranges
south to Peru. It also is known to occur in
the Pleistocene at San Ignacio Lagoon and
at Magdalena Bay, Lower California.
Subgenus T ellinidella Hertlein & Strong,
subgen. nov.
Type: Tellinides purpureus Broderip &
Sowerby.
Shell elongate, compressed, very thin,
80
Zoologica : New York Zoological Society
[34 : 9
with a strong posterior angulation; orna-
mented with fine concentric granulated
ridges which are crossed by impressed ra-
dial striae forming reticulate sculpture;
hinge as in Eurytellina but with a very
small right anterior lateral and a weak pos-
terior lateral, in the left valve the laterals
are faint or obsolete.
The general outline and hinge of this
new subgenus are similar to those of Eury-
tellina. The weak distant right posterior lat-
eral and the very thin shell, which is orna-
mented by both concentric and radial sculp-
ture, are characteristic features of the type
species of T ellinidella.
Dali, 1900, placed Tellina purpureus Brod-
erip & Sowerby in the subgenus Tellinides
Lamarck33. That name was proposed by La-
marck for a genus of Tellina with the sole
species T. timorensis as type. No illustra-
tions accompanied that work. Dubois36 dis-
cussed Lamarck’s genera but he did not il-
lustrate the type species of Tellinides. Nei-
ther did Delessert, 1841, include T. timor-
ensis among his illustrations of the types
of Lamarck’s shells. Hanley37 later gave il-
lustrations showing two views of the ex-
terior of right valves of T. timorensis. Phil-
ippi38 also illustrated a species under that
name and gave views of both the exterior
and interior of the right valve and an um-
bonal view of both valves. His illustrations
agree well with Lamarck’s description of
the species. Bertin39, 1878, stated that 4
type specimens of Lamarck’s species were
present in the collections of the Museum of
Natural History in Paris. In his synonymy
of the species he included references to Han-
ley’s figures 158 and 172, Philippi’s figure
3, as well as illustrations of Romer’s, 1812,
plate 34, figures 4-6.
It appears then that these figures may
be considered to represent authentic speci-
mens of T. timorensis. The right valve has
two diverging cardinal teeth, a close ante-
rior lateral and a weak close posterior lat-
eral. The left valve is said to lack laterals.
The valves are not flexed posteriorly and
the posterior umbonal fold or ridge is broad-
ly rounded and nearly obsolete.
The hinge of the right valve of Tellina
purpureus has two diverging cardinal teeth,
the posterior one strong and bifid or grooved.
The posterior lateral is distant from the
cardinals. A small lamina sometimes occurs
near the margin above the small anterior
lateral and sometimes the margin is some-
35 Tellinides Lamarck, Hist. Nat. Anim. s. Vert., Vol. 5,
July, 1818, p. 535. Type, Tellina timorensis, p. 536. “Habite
l'ocean des grandes Indes ou austral, pres de Timor.”
36 Dubois, C., Epit. Lamarck’s Arrang. Test., 1824, p. 68.
37 Tellina ( Tellinides ) timorensis Lamarck, Hanley, Thes.
Conch., Vol. 1, 1846, p. 292, pi. 61, figs. 158 and 172. Isle
of Negros, Philippines, etc.
38 Tellina timorensis (Tellinides) Lamarck, Philippi,
Abbild. u. Beschreib. Conchyl., Bd. 2, Heft 4, Tellina,
August, 1846, p. 90 (22), pi. 4, fig. 3. Timor, Philippine
Islands, Sumatra, etc.
39 Bertin, V., Nouv. Arch. Mus. Hist. Nat. (Paris), Ser.
2, Vol. 1, 1878. p. 283. [He mentioned that T. timorensis is
represented in the collection by specimens including “par
4 individus ordinaires de Timor (types de Lamarck)."]
what projecting just over the posterior lat-
eral. Furthermore the shell has a strong,
angular posterior umbonal fold exteriorly
and the concentric sculpture on the shell is
decussated by radial grooves giving it a fine-
ly granulose character. These characters on
the present shell are so different from those
of T. timorensis that we propose a new sub-
genus Tellinidella with Tellina purpureus
Broderip & Sowerby as type.
Tellina ITellinidellal purpureus Broderip &
Sowerby.
Tellinides purpureus Broderip & Sowerby,
Zool. Jour., Vol. 4, No. 15, January, 1829,
p. 363. “Hab. ad littora Oceani Pacifici.” —
Sowerby, Zool. Beechey’s Voy., 1839, p. 153,
pi. 42, fig. 2. “Inhabits the sandy shores of
the Pacific Ocean.”
Tellina ( Tellinides ) purpurascens Brod-
erip & Sowerby, Hanley, Thes. Conch., Vol.
1, 1846, p. 295, pi. 62, fig. 194. “Real Leijos
[Llejos] Central America (Cuming).”
Not Tellina purpurascens Gmelin, Linn, i
Syst. Nat., ed 13, Vol. 1, Pt. 6, 1790, p. 3237.
Habitat not cited. Ref. to Lister, Conch., pi.
391, fig. 230. Also “B” and “Y,” ref. to Gual-
tieri, Test., pi. 77, figs. L? and M? Tellina
purpurata is described on p. 3243. Hab. not
cited. Ref. to Gualtieri, Test., pi. 77, fig. L.
Tellina broderipii “Desh. ms. (teste
Cum.)” Carpenter, Cat. Mazatlan Shells, Au-
gust, 1855, p. 32. “Mazatlan.”
Tellina purpurascens Broderip & Sowerby,
Sowerby, Conch. Icon., Vol. 17, Tellina, 1867,
species 103, pi. 20, fig. 103. Same locality as
given by Hanley.
Type Locality : Real Llejos [near Corinto],
Nicaragua (here designated as type local-
ity). Shores of Pacific Ocean originally cit-
ed.
Range : Altata, Mexico, in the Gulf of Cal-
ifornia, to Colombia.
Collecting Stations: Mexico: Tenacatita
Bay; Sihuatanejo Bay; Nicaragua: Corinto,
beach drift.
Description: Shell ovately oblong, sub-
equilateral, thin, compressed, colored a beau-
tiful purplish-rose with the dorsal margins
white; sculpture of close, decussating con-
centric and radial striae; posteriorly a low
radial furrow is present near the dorsal mar-
gin which anteriorly is bounded by a low
carina; right valve with two cardinals, the
posterior one the larger and bifid, and a very
close, small, anterior lateral above which
there is sometimes a small lamina near the
margin, and a posterior lateral distant near-
ly half the length of the posterior dorsal
margin, above this tooth the margin is
sometimes somewhat projecting; left valve
with laterals obsolete; the end of the pal-
lial sinus is considerably lower than and pos-
terior to the anterior adductor impression,
the base is confluent with the pallial line.
A right valve from Tenacatita Bay, Mex-
ico, measures : length, 49.4 mm. ; height, 26.4
mm.; convexity (one valve), 4 mm.; pallial
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
81
sinus extends forward 36 mm. from the pos-
terior end of the valve. The species attains
a greater size than this specimen.
The original name for this species, given
by Broderip & Sowerby in 1829, was Tellin-
ides purpureas. Dali40 apparently considered
that combination of names preoccupied due
to the fact that Tellinides is now considered
to be a subgenus of Tellina, also the fact
that Dillwyn41 had referred to a Tellina pur-
purea.
However, Dillwyn in a footnote, in re-
marking on certain of Gmelin’s species, re-
ferred to Gmelin, page 3243, and on that
page of Gmelin’s work the species was cited
as Tellina purpurata .. It thus appears that
Dillwyn’s spelling of “purpurea” was a mis-
print and not a renaming of Gmelin’s spe-
cies. If this view is accepted, there then ap-
pears to be no valid reason for rejecting the
original name given the west American spe-
cies by Broderip & Sowerby. The name Tel-
lina purpurascens which Hanley used for
this species is preoccupied by Tellina pur-
purascens Gmelin, 1790. The name Tellina
broderipii attributed to Deshayes was ap-
plied to the west American form by Carpen-
ter.
The shell of this species bears a resem-
blance to that of Tellina princeps Hanley42
but is thinner, narrower, the posterior dor-
sal margin slopes more gently, the radial
striae are stronger and the right posterior
lateral is weaker.
Distribution: A few specimens of this
species, nearly all right valves, were taken
by the expedition along the coast of west
Mexico and in the beach drift at Corinto,
Nicaragua.
Subgenus Macaliopsis Cossmann.
Key to the species of Macaliopsis.
A. Beaks directed strongly anteriorly; con-
centric lamellae about 1 per mm. .... lyra
B. Beaks directed only slightly anteriorly;
concentric lamellae, lower, fewer, usually
about 3 per mm. lyrica
Tellina I Macaliopsis! lyra Hanley.
Tellina lyra Hanley, Proc. Zool. Soc. Lon-
don, September, 1844, p. 68. “Hab. Tumbez,
Peru.”— Hanley, Thes. Conch., Vol. 1, 1846,
p. 271, pi. 62, fig. 187. Tumbez, Peru. — Sow-
erby, Conch. Icon., Vol. 17, Tellina, Septem-
ber, 1867, species 203, pi. 36, fig. 203. Tumbez,
Peru.
Type Locality: Tumbez, Peru.
Range : Lower California to Tumbez, Peru
(Dali).
40 Dali, W. H., Proc. U. S. Nat. Mus., Vol. 23, 1900, p. 302.
■U Dillwyn, L. W., Descript. Cat. Rec. Shells, Vol. 1,
1817, p. 72, footnote.
42 Tellina princeps Hanley, Proc. Zool. Soc. London, Sep-
tember, 1844, p. 62. “Hab. Tumbez, Peru ; soft sandy mud.
five fathoms.” —Hanley, Thes. Conch., Vol. 1, 1846, p. 238,
pi. 63, fig. 206. “Tumbez, Peru; (Cuming).” —Sowerby,
Conch. Icon., Vol. 17, Tellina, 1867, species 135, pi. 25, fig.
135. Tumbez, Peru. —Salisbury, Proc. Malacol. Soc. London,
Vol. 21, Pt. 2, 1934, p. 91, pi. 9, fig. 4. (Illustration of
type] .
Collecting Stations: Guatemala: 7 miles
west of Champerico (197-D-l, 2), 14 fath-
oms, mud; El Salvador: La Libertad (198-
D-2), 14 fathoms, mud,
Description: Shell elliptic, thin, compress-
ed, slightly longer anteriorly, dull white
exteriorly and interiorly, beaks curved for-
ward ; anterior dorsal margin excavated be-
low the beaks, anterior end rounded, ventral
margin broadly rounded, the posterior dorsal
margin nearly straight or very broadly curv-
ed and sloping posteriorly directly from the
beaks ; a deep smooth lunule and escutcheon
present; the sculpture consists of regular,
thin, sharp, raised, concentric ribs which are
separated by much wider interspaces (about
1 mm. wide) but which become narrower
toward the ventral margin, a narrow pos-
terior area is set off by an umbonal carina;
hinge of right valve with a strong grooved
triangular posterior and a thin anterior car-
dinal near the margin, a strong anterior and
posterior lateral present; left valve with a
grooved anterior and a thin posterior cardi-
nal tooth and lateral triangular projections
of the nymph ; the pallial sinus extends for
about three-fifths the length of the shell, sub-
trigonal above and highest just posterior to
a line vertical with the beaks, then descend-
ing and narrowly elliptically rounded anteri-
orly then bending posteriorly and for about
two-thirds of its length confluent with the
pallial line.
The largest specimen in the present collec-
tion, a left valve, measures : length, 50 mm.
height, 35 mm.; convexity (one valve), 6
mm.; pallial sinus extends anteriorly 29 mm.
from the posterior end of the valve.
The strongly anteriorly directed beaks,
more convex anterior dorsal margin which
is excavated beneath the beaks, higher and
more widely spaced concentric sculpture and
shorter and more trigonal pallial sinus are
features separating this species from Tellina
lyrica Pilsbry & Lowe.
Tellina protolyra Anderson43 is a similar
species but it is smaller, less elongate, more
inflated and the posterior dorsal margin is
straighter than in the present species.
Tellina ( Macaliopsis ) aequizonata Pilsbry
& Olsson44, described from the Pliocene of
Ecuador, is said to be much larger, more
strongly sculptured and more circular in out-
line than T. lyra.
Distribution: A few specimens, mostly
single valves, of this interesting species were
dredged in 14 fathoms off Guatemala and
El Salvador on a mud bottom.
Tellina I Macaliopsis! lyrica Pilsbry & Lowe.
Tellina ( Macaliopsis ) lyrica Pilsbry &
Lowe, Proc. Acad. Nat. Sci. Philadelphia,
43 Tellina protolyra F. M. Anderson, Proc. Calif. Acad.
Sci., Ser. 4, Vol. 18, No. 4, March 29, 1929, p. 174, pi. 21,
figs. 2, 3. "From Loc. 267-B, C. A. S., horizon M-N, of the
Tubera group, Colombia; Miocene.”
44 Tellina ( Macaliopsis ) aequizonata Pilsbry & Olsson,
Proc. Acad. Nat. Sci. Philadelphia, Vol. 93, September 9,
1941, p. 68, pi. 14. fig. 7. “Jama formation, Puerto Jama.”
Ecuador. Pliocene.
82
Zoologica: New York Zoological Society
[34 : 9
Vol. 84, May 21, 1932, p. 94, pi. 10, figs. 4, 4a.
“Guaymas in about 20 fathoms.”
Type Locality : Guaymas, Mexico, in about
20 fathoms.
Range: Gulf of California to the Gulf of
Chiriqui, Panama.
Collecting Stations: Mexico: Santa Inez
Bay in the Gulf of California (143-D-3, 4),
25-35 fathoms, mud, crushed shell, sand ; El
Salvador: La Libertad (198-D-2), 14 fath-
oms, mud ; Panama : Gulf of Chiriqui (221-D-
1-5) , 35-40 fathoms, sandy mud.
Description : Shell transversely oval, beaks
turned slightly forward, anterior end broadly
rounded, posterior dorsal margin broadly
curved and rather steeply sloping; a slight
angulation sets off a narrow posterior area ;
lunule smooth, slightly sunken ; escutcheon
deeply sunken and bounded by a high keel;
sculpture consists of fine, close, concentric
ribs (about 3 per millimeter) which are nar-
rower than the interspaces ; right valve with
a simple anterior and grooved posterior
cardinal and well-developed laterals, the nos-
terior one more distant; left valve with a
grooved anterior and thin posterior lamella-
like cardinal, anterior lateral weak, posterior
lateral represented by a triangular projec-
tion of the margin ; the pallial sinus is high-
est posteriorly, rounded at the end and
extends forward about four-fifths the length
of the shell and is separated from the ante-
rior adductor impression bv a considerable
distance, for over half its length along the
base it is confluent with the pallial line.
A large right valve from Santa Inez Bay
in the Gulf of California measures; length,
40.2 mm.; height, 29.6 mm.; convexity (one
valve). 6.5 mm.; pallial sinus extends for-
ward 31 mm. from the posterior end of the
valve.
The exterior of fresh valves shows a bril-
liant iridescence of spectral colors due to the
fine growth lines between the concentric
ridges acting as a diffraction grating.
The specimens in the present collection
show the differences pointed out by Pilsbry
& Lowe between this species and Tellina lyra.
They stated : “Related to T. lyra Hanley, but
relatively longer, with the beaks less, only
very slightly, turned forward, the dorsal
margin in front of them less convex, and the
concavity runnning to the posterior basal
extremity wider.” The concentric ribbing on
T. lyrica is much finer and more closely
spaced than that on T. lyra.
Distribution: This species was dredged in
Santa Inez Bay in the Gulf of California, in
25-35 fathoms, off La Libertad, El Salvador,
in 14 fathoms and in the Gulf of Chiriqui,
Panama, in 35-40 fathoms, in mud and sandy
mud. It also has been recorded as occurring
in the Pliocene of Ecuador.
Subgenus Mer/sco Dali.
Key to the species of Merisca.
A. Interspaces with minute radial striae
a. Rostrum attenuated posteriorly; con-
centric lamellae about 1 per mm.
crystallina
aa. Rostrum not attenuated posteriorly ;
concentric lamellae about 5 per mm.
reclusa
B. Interspaces without radial striae
proclivis
Tellina l Merisca I crystallina Spengler.
Tellina crystallina Chemnitz, Neues Syst.
Conchyl.-Cab. von Martini-Chemnitz, Bd. 11,
1795, p. 210, pi. 199, figs. 1947, 1948. “Es ist
diese Muschel an der Nordamericanischen
Kiiste bey Newport auf Rhode-Island ge-
funden worden.” (Spengler). — Spengler,
Skr. Nat. Selsk. (Copenhagen), Vol. 4, No.
2, 1798, p. 113. “Fra Newport Long-Island.”
Ref. to Chemnitz, pi. 199, figs. 1947, 1948.—
Wood, General Conch., 1815, p. 149. [No lo-
cality cited]. Ref. to Chemnitz, pi. 199, figs.
1947, 1948. — Hanley, Thes. Conch., Vol. 1,
1846, p. 270, pi. 57, fig. 43. “St. Elena, West
Columbia.”— Olsson, Bull. Amer. Paleo., Vol.
9, No. 39, 1922, p. 421 (249), pi. 29 (26), fig.
10. Banana River, Costa Rica. Miocene. Also
Recent.
Tellina ( Merisca ) crystallina Wood, Dali,
Proc. U. S. Nat. Mus., Vol. 23, 1900, pp. 293,
302, 311, pi. 2, fig. 10. Sullivans Island, South
Carolina, to Cartagena, Colombia. Lower
California to Panama. — M. Smith, Panamic
Mar. Shells (Tropical Photogr. Lab., Winter
Park, Florida), p. 64, fig. 834. Lower Cali-
fornia to Guayaquil, Ecuador. Also West
Indies.
Tellina schrammi Recluz, Joum. de Con-
chyl., Vol. 4, December, 1853, p. 152, pi. 6,
figs. 7, 8. . . “habite la rade de la Pointe-a-
Pitre (Guadeloupe) : elle a ete draguee sur
un fond vaseux au fond de cette rade.”
Type Locality: Newport, Rhode Island.
Range: Scammon Lagoon, Lower Cali-
fornia, to the Gulf of California and south to
Guayaquil, Ecuador. Also Atlantic, from
Charleston, South Carolina, to Cartagena
Bay, Colombia.
Collecting Stations: Mexico: Tangola-
Tangola Bay, on beach; Nicaragua: Corinto
(200-D-19), 12-13 fathoms, mangrove leaves,
also on beach.
Description: Shell rather small, roundly
trigonal, posteriorly rostrate, somewhat at-
tenuated and with a sharp flexure, thin,
white; sculpture of strong, raised distant,
concentric ribs separated by much wider in-
terspaces (about 1 mm. wide) although the
spacing varies with various specimens, and
where the lamellae cross the flexure they are
V-shaped; the interspaces contain fine con-
centric striae (sometimes about 8) and faint
radial striae; right posterior and left ante-
rior cardinals grooved, lateral teeth in right
valve well developed, in the left obsolete or
nearly so; pallial sinus high behind then
descending and sometimes touching the base
of the anterior adductor impression, wholly
confluent with the pallial line below.
The specimens in the present collection are
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
83
small but a large valve in the collections of
the California Academy of Sciences, from
Mazatlan, Mexico, measures : length, 23 mm. ;
height, 17.4 mm. ; convexity (one valve) , 3.4
mm.
Chemnitz originally described and illus-
trated Tellina crystallina and cited Spengler
as the authority for the locality, Newport,
Rhode Island. The International Committee
on Zoological Nomenclature has recently
ruled against acceptance of the specific
names proposed by Chemnitz. Spengler next
described the species. He referred to Chem-
nitz’s description and illustrations and gave
the same locality as Chemnitz. However, the
species is not known with certainty to occur
north of Charleston, South Carolina. One
might venture the opinion that since
Spengler was a citizen of Denmark, the type
specimens secured by him originally might
have come from the Danish West Indies.
Salisbury45 recently stated that the east
and west American shells referred to this
species should be considered as separate spe-
cies. So far as we have been able to deter-
mine, there is no reason to differ from
Dali’s46 conclusion that “The specimens from
the two oceans are absolutely similar, and
differ no more than individuals from either
sea among themselves.” Tellina schrammi
Recluz, described from the island of Guade-
loupe in the West Indies, appears to be iden-
tical with T. crystallina.
Tellina errati Pilsbry & Johnson47, de-
scribed from the Miocene of Santo Domingo,
is identical or a very similar form. Maury,48
1917, stated that fossils from Santo Domingo
were identical with Recent shells found on
the beach on that island.
Tellina ( Merisca ) sancti-dominici
Maury,46 also described from the Miocene of
Santo Domingo, is a similar species. Tellina
martensi Lynge,50 described from the East
Indies, was compared to T. crystallina, but
the west American species is longer with a
more rostrate form.
Distribution : A few specimens of this spe-
cies were collected by the expedition on the
beach at Tangola-Tangola, Mexico, and
others were dredged in 12-13 fathoms at
Corinto, Nicaragua. It also occurs from Mio-
cene to Recent in the Caribbean region and
has been recorded as occurring in the Plio-
45 Salisbury, A. E., Proc. Malacol. Soc. London, Vol. 21,
Pt. 2, July, 1934, p. 83.
40 Dali, W. H., Proc. U. S. Nat. Mus., Vol. 23, 1900, p.
302.
41 Tellina ( Merisca ) errati Pilsbry & Johnson, Proc.
Acad. Nat. Sci. Philadelphia, Vol. 69, May 5, 1917, p. 201.
Santo Domingo, Oligocene [Miocene]. —Pilsbry, Proc. Acad.
Nat. Sci. Philadelphia, Vol. 73, Pt. 2, 1922, p. 425, pi. 41,
fig. 7. [Figure of type].
48 Maury, C. J., Bull. Amer. Paleo., Vol. 5, No. 29, 1917,
pp. 387 (223) -388 (224), pi. 64 (38), fig. 4.
49 Tellina ( Merisca ) sancti-dominici Maury, Bull. Amer.
Paleo., Vol. 5, No. 29, Pt. 1, April 7, 1917, p. 388 (244),
pi. 64 (38), fig. 11. “Bluff 3. Cercado de Mao.” Santo
Domingo, Miocene.
50 TeUina ( Merisca ) martensi Lynge, Kyi. Dansk.
Vidensk. Skr., Ser. 7, Nat. og Math., Bd. 5, 1909, p. 195,
pl. 3, figs. 40-42. "South of Koh Kut, 17-20 fathoms, mud
(%). Gulf of Siam. Singapore, 2-3 fathoms, coral reef;
5/2."
cene of Ecuador and Pleistocene of Magda-
lena Bay, Lower California.
Tellina IMeriscal proclivis Hertlein & Strong,
sp. nov.
Plate I, Figs. 6, 7, 14.
Tellina declivis Sowerby, Conch. Icon.,
Vol. 17, Tellina, March, 1868, species 261,
pl. 44, fig. 261. “Hab.— ?”— I. S. Oldroyd,
Stanford Univ. Publ. Univ. Ser. Geol. Sci.,
Vol. 1, 1924, p. 165 (under section Merisca).
Catalina Island, California, to Panama.—
Strong & Hertlein, Allan Hancock Pac. Ex-
ped., Vol. 2, No. 12, 1939, p. 184. Bahia
Honda, and off Taboga Island, Panama.
Not Tellina declivis Conrad, Jour. Acad.
Nat. Sci. Philadelphia, Ser. 1, Vol. 7, 1834,
p. 131. “Locality, Yorktown, Va.” Tertiary.
[Miocene].
Tellina ( Merisca ) declivis Sowerby, Dali,
Proc. U. S. Nat. Mus., Vol. 23, 1900, p. 301.
“Cerros Island, Lower California, to the Gulf
of California.”
Type Locality : Magdalena Bay, Lower
California, Mexico.
Range : Cedros Island, Lower California,
to Panama.
Collecting Stations: Mexico: 4 miles SSW.
of Maldanado Point (192-D-l), 26 fathoms,
mud; PortGuatulco (195-D-20), 23 fathoms,
mud; Tangola-Tangola Bay (196-D-17), 23
fathoms, mud; Costa Rica: Port Parker
(203-D-l, 3), 12-15 fathoms, sandy mud,
crushed shell, shelly mud.
Description: Shell small, subtrigonal,
white, beaks subcentral and elevated; ante-
rior dorsal margin sloping, anteriorly
rounded and inflated, posterior dorsal mar-
gin steeply sloping, the end pointed, wedge-
shaped, subcompressed; a well-developed and
sunken lunular area and long escutcheon
present; right valve with a broad flexure,
the left with a shallow radial depression fol-
lowed by an angulation; sculpture of fine,
regular, concentric lamellae, about 3 or 4 per
millimeter on adult shells; hinge of right
valve with two strong cardinals, the poste-
rior one grooved, and two laterals, left valve
with a strong grooved anterior and weaker
posterior cardinal, the latter close to the
margin, the posterior cardinal in each valve
grooved, pallial sinus high in the middle
then descending near to but not touching the
anterior adductor impression, then bending
posteriorly and for about half its length con-
fluent with the pallial line. Dimensions of
the type: length, 9 mm.; height, 7.8 mm.;
convexity (both valves together), 4.8 mm.
Holotype (Calif. Acad. Sci. Paleo. Type
Coll.), from Loc. 20299 (C.A.S.), Magdalena
Bay, Lower California, Mexico; Charles R.
Orcutt collector. Paratype, from station
196-D-17, Lat. 15°45' N., Long. 96°05'34" W.,
Tangola-Tangola Bay, Mexico, dredged in
23 fathoms, mud.
One of the largest valves in the present
collection measures 9.3 mm. in length. A
large right valve dredged about 5 miles west
of Mazatlan, Mexico, by the Templeton
84
Zoological New York Zoological Society
[34: 9
Ci'ocker Expedition of the California Acad-
emy of Sciences in 1932, measures: length,
11.4 mm.; height, 9.8 mm.; convexity (one
valve, 3 mm.
The use of the combination of names, Tel-
lina declivis by Conrad, 1834, makes it nec-
essary to pi'opose a new name which is based
on a new type specimen for the west Amer-
ican shell described under that name by Sow-
erby in 1868 which is here named Tellina
proclivis.
The shell of Tellina proclivis differs from
that of T. reclusa in the more steeply sloping
dorsal margins, in lacking radial sculpture,
in that the pallial sinus does not touch the
anterior adductor impression and along the
base it is confluent with the pallial line for
only about one-half its length. It differs
from T. meropsis in the more steeply sloping
dorsal margins, more pointed posterior end
and in the narrower, radially depressed area
posteriorly.
Distribution : Specimens of this species,
mostly single valves, were dredged by the
expedition off western Mexico and Costa
Rica in 12-26 fathoms, mostly on a mud bot-
tom. Dali51 cited this species as occurring
north to Catalina Island, California, but
Burch52 indicated that it is questionable
whether it is a member of the fauna of Cali-
fornia.
Tellina IMeriscal reclusa Dali.
Tellina ( Merisca ) reclusa Dali, Proc. U.S.
Nat. Mus., Vol. 23, No. 1210, November,
1900, pp. 301, 315, pi. 3, fig. 2. “Types.— No.
105513, U. S. N. M., from San Ignacio La-
goon, Lower California, Hemphill. Also off
Lower California, in lat. 30° 28', by the U. S.
Fish Commission, at Station 3019, in 14
fathoms, Gulf of California.’’
Tellina reclusa Dali, E. K. Jordan, Contrib.
Dept. Geol. Stanford Univ., Vol. 1, No. 4,
1936, p. 145. Magdalena Bay and San Ignacio
Lagoon, Lower California, Pleistocene. Re-
cent from San Ignacio Lagoon to the Gulf of
California.
Type Locality : San Ignacio Lagoon,
Lower California.
Range : San Ignacio Lagoon, Lower Cali-
fornia, to the Gulf of California and south
to Bahia Honda, Panama.
Collecting Stations: Mexico: Arena Bank,
Gulf of California (136-D-18, 21, 22), 40-45
fathoms, mud ; Santa Inez Bay, Gulf of Cal-
ifornia (142-D-3; 145-D-l, 3), 4-40 fathoms,
sand, weed; Port Guatulco (195-D-2, 3),
3-3.5 fathoms, sand, crushed shell, Tangola-
Tangola Bay (196-D-14-15) , 5 fathoms,
crushed shell; Nicaragua: Corinto (200-D-
8, 9), 6-6.6 fathoms, mangrove leaves, also
on beach.
Description : Shell small, moderately con-
vex, subtrigonal, posteriorly wedge-shaped
and flexed, the posterior end keeled dorsally;
51 Arcopagia declivis Sowerby, Dali, U. S. Nat. Mus.,
Bull. 112, 1921, p. 45 (under section Merisca). Catalina
Island, California, to Panama.
52 Burch, J. Q., Min. Conch. Club South Calif., No. 45,
sculpture of fine, close (about 5 per mm.),
little elevated, sharp, concentric lamellae
which are separated by wider interspaces
which are faintly, radiately striated; lunu-
lar area smooth, escutcheon long, narrow,
deep; hinge normal for the subgenus; pallial
sinus high behind then descending and usu-
ally, although not always, touching the base
of the anterior adductor impression and
wholly confluent with the pallial line below.
A specimen from Tangola-Tangola Bay,
Mexico, measures: length, 20 mm.; height,
15.5 mm.; convexity (one valve), 3.6 mm.
Dali stated that “this species is notable
for the rasp-like quality of its surface to
the touch.”
Compared to Tellina proclivis the shell of
T. reclusa is longer in proportion to the
height, the dorsal margins slope less steeply
and the interspaces are ornamented by fine
radial sculpture. The shell of T. reclusa is
more elongate in outline but the posterior
end is less attenuated and flexed and the con-
centric sculpture is more closely spaced than
that of T. crystallina. The coarser sculpture
and the fact that the pallial sinus usually
touches the anterior adductor impression are
features separating T. reclusa from T. mer-
opsis Dali53.
Tellina ( Merisca ) lintea Turton, which
occurs in the Caribbean region from Florida
to Brazil, is very similar to T. reclusa. Pris-
tipagia gemonia Iredale,54 an Australian
species, also is a somewhat similar shell.
Distribution: This species was collected
by the expeditions from Santa Inez Bay in
the Gulf of California, to Corinto, Nicara-
gua, on the beach and dredged at depths of
3-45 fathoms. It is also known to occur in
the Pleistocene of Lower California.
Subgenus Scissula Dali.
Key to the species of Scissula.
A. Shell large, length exceeding 25 mm.
a. Posterior end obliquely truncated;
white or tinged with brown cognata
aa. Posterior end tapering and roundly
pointed ; pale rose color nicoyana
B. Shell small, length not exceeding 25 mm.;
very thin, glassy
a. Oblique striae widely spaced, coarse,
very oblique virgo
aa. Oblique striae closer, finer, less oblique
and approaching lines of growth
varilineata 55
5S Not represented in the present collection.
Tellina IScissulal cognata C. B. Adams.
Tellina cognata C. B. Adams, Ann. Lyceum
Nat. Hist. New York, Vol. 5, July, 1852, pp.
503, 545 (separate pp. 279, 321). “Habitat.
— Panama.”
53 Tellina ( MoereUa ) meropsis Dali, Proc. U. S. Nat.
Mus., Vol. 23, No. 1210, November, i900, p. 317, pi. 3,
fig. i. “San Diego, California.”
3- Pristipagia gemonia Iredale, Rec. Australian Mus..
Vol. 19, No. 5, April 7, 1936, p. 281, pi. 21, fig. 6. “Habitat.
Hertlein & Strong: Mollusks of Mexico and Central America
85
1949]
Psammobia casta Reeve, Conch. Icon., Vol.
10, Psammobia, June, 1857, species 55, pi. 8,
fig. 55. “Hab. Guatemala.”
Not Tellina casta Hanley, Proc. Zool. Soc.
London, September, 1844, p. 63. “Hab. Singa-
pore ; sandy mud.”
Tellina tenuilineatus Li, Bull. Geol. Soc.
, China, Vol. 9, No. 3, October, 1930, p. 262,
pi. 5, fig. 33. “Brought up by marine dredge
from depths varying from 10. ft. to 40. ft. in
1 the mud at the mouth of the Rio Grande near
La Boca about one mile from the mainland
in Panama Bay.” “Horizon : Probably Gatun
formation.” According to Pilsbry this record
is based upon “Two pieces of a left valve of
‘Tellina’ cognata C. B. Ad. Compared with
Adams’ type” {Proc. Acad. Nat. Sci. Phila-
delphia, Vol. 83, 1931, p. 431).
Type Locality : Panama.
Range : Mazatlan, Mexico, to Panama.
Collecting Stations: Guatemala: 7 miles
west of Champerico (197-D-l, 2), 14 fath-
oms, mud; El Salvador: La Libertad (198-
D-l, 2), 13-14 fathoms, mud; Meanguera
Island, Gulf of Fonseca (199-D-l), 16 fath-
oms, sand, mud, crushed shell; Nicaragua:
Monypenny Point, Gulf of Fonseca (199-D-
4, 5, 6), 4-7 fathoms, mud; Costa Rica: 13
miles S. X E. of Judas Point (214-D-l), 42
fathoms, mud, shell; Panama: Gulf of Chiri-
qui (221-D-1-5), 35-40 fathoms, sandy mud.
Description: Shell oblong, anterior end
obliquely rounded, posterior end obliquely
truncated, gaping, a depressed posterior area
is delimited by a rounded umbonal angula-
tion, white or with a reddish or pale brown-
ish tinge; sculpture consists of minute un-
equal radiating striae and concentric grooved
striae which cross the lines of growth
obliquely and become weak or absent on the
posterior third of the shell; lateral teeth
obsolete; pallial sinus highest posteriorly
then descending gently and extending ante-
riorly for a little more than three-fourths the
length of the shell but well separated from
the anterior adductor impression; basally,
for about one-half its length, it is confluent
with the pallial line.
A specimen from the Gulf of Fonseca, in
the present collection, measures : length, 45.3
mm. ; height, 30 mm. ; convexity (both valves
together), 13.5 mm.; pallial sinus extends
forward 36.4 mm. from the posterior end of
the shell. A large left valve dredged in the
Gulf of Chiriqui, Panama, measures: length,
49.5 mm. ; height, 32 mm. ; convexity (one
valve) , 7.5 mm.
This species is unlike any other along the
Pacific Coast of the Americas. It bears only
a general resemblance to the east American
species T. similis Sowerby.
The present species appears to be referable
to the subgenus Scissula Dali, 1900, type
Tellina decora Say, rather than to other
supraspecific groups in which the shell bears
oblique striations such as Scissulina Dali,
1924, type, T. dispar Conrad; Jactellina
Iredale, 1929, type, T. obliquaria Deshayes;
Obtellina Iredale, 1929, type, T. bougei
Sowerby; and Loxoglypta Dali, Bartsch &
Rehder, 1938, type, T. obliquilineata Conrad.
Distribution: This species was dredged by
the expedition from Guatemala to Panama
in 4-42 fathoms. It occurred rather abund-
antly off Champerico, Guatemala, in 14 fath-
oms, and in the Gulf of Chiriqui, Panama, in
35-40 fathoms, mostly on a muddy bottom.
It also has been recorded as occurring in the
Pliocene of Ecuador.
Tellina IScissula) nicoyana Hertlein & Strong,
sp. nov.
Plate I, Figs. 23, 24, 25, 26.
Shell elongately ovate, thin, gently inflated,
nearly equilateral, color, pale rose; anterior
dorsal margin slightly rounded, gently slop-
ing anteriorly then rounding abruptly to
the elliptically rounded anterior end which
merges into the broadly rounded ventral
margin; posterior end tapering, roundly
pointed; the posterior dorsal margin slopes
gently from the beaks with a convexity in
the middle portion ; the ornamentation con-
sists of fine concentric lines of growth ; these
are crossed by oblique striae which begin on
the anterior end and continue for about two-
thirds the length of the shell but are absent
on the posterior end where there are a few
vague submicroscopic radiating striae; hinge
of right valve with two grooved cardinals
and, close by, an anterior lateral, and distally
a posterior socket below which is a weak
lateral; left valve with a grooved anterior
cardinal and a thin lamella-like posterior
cardinal, also a short anterior lateral or pro-
jection of the nymph and a faint posterior
lateral; pallial sinus extending about four-
fifths the length of the shell and separated
from the anterior adductor impression by
a considerable distance, high and rounded
beneath the beaks then sloping and broadly
undulating downward then turning abruptly
downward to the pallial line with which it is
confluent below; interior beautifully pale
yellowish-rose and with faint submicroscopic
radiating striae. Length, 34.4 mm. ; height,
19 mm.; convexity (both valves together),
7.8 mm. ; pallial sinus extends forward about
27 mm. from the posterior end of the shell.
Holotype and paratypes (Calif. Acad. Sci.
Paleo. Type Coll.), dredged in Ballena Bay,
Gulf of Nicoya, Costa Rica; also dredged in
the same vicinity at Station 213-D-ll, 17, in
Lat. 9° 44' 52" to 9° 42' 00" N., Long. 84° 51'
25" to 84° 56' 00" W., in 35 fathoms, mud.
This beautiful species appears to be dis-
tinct from any described shell. It apparently
has some characters in common with Tellina
delicatula Deshayes36, the type of which has
never been illustrated, but differs in that
the shell is nearly equilateral, rather than
strongly inequilateral, and the hinge is quite
different from that described by Deshayes.
It also lacks the dark irregular lines crossing
the oblique striae on that species mentioned
5tJ Tellina delicatula Deshayes, Proc. Zool. Soc. London
for 1854 (issued May 16, 1855), p. 363. “Hab. Mazatlan.
Coll. Cuming.”— Bertin, Nouv. Arch. Mus. Hist. Nat.
(Paris), Ser. 2, Vol. 1, 1878, p. 290. Coast of Mazatlan.
86
Zoologica: New York Zoological Society
[34:9
by Carpenter57. It is interesting to note that
Maury has described Tellina ( Scissula )
cercadica 58 from the Miocene of Santo Do-
mingo, which species, she stated, is very
similar to a Recent shell from Panama Bay
in the Newcomb collection which was labeled
Tellina delicatula Deshayes. The Miocene
shell illustrated by Maury is not at all similar
to the present species but is more similar to
Tellina ( Scissula ) cognata C. B. Adams from
Panama.
Tellina IScissulal virgo Hanley.
Tellina virgo Hanley, Proc. Zool. Soc. Lon-
don, December, 1844, p. 143. “Hab. — ? Mus.
Cuming.” “Allied to the Iris of Say, but
much larger.” — Hanley, Thes. Conch., Vol 1,
1846, p. 284, pi. 57, fig. 42. “Chiriqui, West
Columbia.” — Sowerby, Conch. Icon., Vol. 17,
Tellina, 1867, species 207, pi. 37, fig. 207.
“Hab. Chiriqui, West Indies.” — Pilsbry &
Olsson, Nautilus, Vol. 56, No. 3, January,
1943, p. 79 (in text), pi. 8, fig. 5. Panama and
west coast of northern South America to
Puerto Pizarro, Peru.
Tellina ( Fabulina ) virgo Hanley, Salis-
bury, Proc. Malacol. Soc. London, Vol. 21, Pt.
2, July, 1934, p. 91, pi. 13, figs. 5 and 6. [Illus-
trations of holotype and paratype].
Type Locality: Chiriqui, west Panama
(here designated as type locality). No lo-
cality cited originally.
Range: Magdalena Bay to the Gulf of
California and south to Puerto Pizarro, Peru.
Collecting Stations: Nicaragua: Corinto
(200-D-11-13, 15), 1-8 fathoms, mangrove
leaves, also on beach.
Description: Shell small, ovately elongate,
very thin, glassy, transparent, obtusely an-
gulated posteriorly, colored pink or white;
sculpture consists of fine striae which cross
the shell obliquely but are absent on the pos-
terior area; the pallial sinus projects ante-
riorly about four-fifths the length of the
shell but does not quite touch the anterior
adductor impression, along the base it is con-
fluent with the pallial line.
One of the largest valves collected on the
beach at Corinto, Nicaragua, measures:
length, 20.2 mm.; height, 12 mm.; convexity
(one valve), 2 mm.
The spacing of the oblique lines on this
shell seems to vary somewhat but on the
specimens which we have observed these lines
are always more widely spaced and cross the
shell at a greater inclination than those of
the similar Tellina ( Scissula ) varilineata
Pilsbry & Olsson59.
Distribution: This species was collected
by the expedition only at Corinto, Nicaragua,
on the beach and dredged in 1 to 8 fathoms.
57 Carpenter, P. P., Cat. Mazatlan Shells, September,
1855, p. 37.
58 Tellina ( Scissula ) cercadica Maury, Bull. Amer. Paleo.,
Vol. 5, No. 29, Pt. 1, April 7, 1917, p. 388 (224), pL 64
(38), fig. 9. "Bluff 3, Cercado de Mao.” Santo Domingo,
Miocene.
59 Tellina ( Scissula ) varilineata Pilsbry & Olsson, Nau-
tilus, Vol. 56, No. 3, January, 1943, p. 79, pi. 8, fig. 6.
Type, "from Puerto Bucaro, Province of Los Santos,
Panama.” Also ranges south to Puerto Pizarro, Province
of Tumbez, Peru.
Subgenus Phyllodina Dali.
Tellina IPhyllodinal pristiphora Dali.
Tellina ( Phyllodina ) pristiphora Dali,
Proc. U. S. Nat. Mus., Vol. 23, No. 1210, No-
vember, 1900, pp. 302, 316, pi. 4, fig. 14.
“Dredged near La Paz, Lower California,
in 26V? fathoms.”
Tellina pristiphora Dali, Pilsbry & Lowe,
Proc. Acad. Nat. Sci. Philadelphia, Vol. 84,
1932, p. 132. Dredged in 20 fathoms, Man-
zanillo; Acapulco, Mexico.
Type Locality: Near La Paz, Lower Cali-
fornia, in 26V2 fathoms.
Range : Santa Inez Bay, Gulf of California,
to Puntarenas, Costa Rica.
Collecting Stations: Mexico: Arena Bank,
Gulf of California (136-D-4, 7, 9, 15, 18, 20-
22, 28, 32), 40-85 fathoms, mud, crushed
shell, muddy sand, sand; Santa Inez Bay
(146-D-l), 35 fathoms, mud, crushed shell;
Manzanillo (184-D-2), 30 fathoms, gravelly
sand ; Port Guatulco ( 195-D-21 ) , 18 fathoms,
mud; Costa Rica: Port Parker (203-D-3),
12 fathoms, shelly sand.
Description: Shell of medium size, ovate,
compressed, nearly equilateral, beaks low,
anterior end rounded, posterior dorsal mar-
gin sloping, the end blunt, surface yellowish-
white, somewhat chalky; sculpture consists
of evenly spaced concentric lamellae which
are separated by wider and finely concen-
trically striated interspaces, the lamellae are
somewhat obsolete medially; on the pos-
terior area, which on the right valve is set
off by an angulation and on the left valve by
a groove, the lamellae develop small squarish
or rounded elevated foliations, along the
anterior dorsal margin similar foliations
present a serrated character; lunule and
escutcheon elongate; two cardinal teeth in
each valve, the right posterior and left an-
terior ones grooved, two large laterals in the
right valve and two small ones in the left,
the anterior ones closest to the cardinals;
pallial sinus highest behind, then tapering,
the end pointed, extending toward the an-
terior adductor impression for more than
one-half the length of the shell, free and
ascending from the pallial line except for a
short distance posteriorly; interior white
and in large specimens yellowish which be-
comes pale salmon in the anterior dorsal area.
A large specimen dredged on Arena Bank
in the southern portion of the Gulf of Cali-
fornia measures: length, 35.8 mm.; height,
23 mm.; convexity (both valves together),
10.2 mm. ; pallial sinus extends forward 21.5
mm. from the posterior end of the shell.
Tellina fluctigera Dali60, described from
the Gulf of Panama, the type of which has
not been illustrated, appears to be a very
similar form.
Tellina dodona Dali and T. leptalea Gard-
ner are representative of the subgenus Phyl-
lodina in the Miocene of Florida.
60 Tellina l Phyllodina) fluctigera Dali, Bull. Mus. Comp.
Zool., Vol. 43, No. 6, October. 1908, p. 419. Dredged "in
the Gulf of Panama, in 182 fathoms, mud, bottom tem-
perature 54°.l F.”
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
87
Distribution : Specimens of Tellina pristi-
phora were dredged from Santa Inez Bay, in
the Gulf of California, to Port Parker, Costa
Rica, in 12-85 fathoms. The present record of
occurrence at Costa Rica is an extension
south of the known range of this species.
Subgenus Phyllodella Hertlein & Strong,
subgen. nov.
Shell elongate, compressed, moderately
thin, equilateral, both sides sloping nearly
equally, anterior end rounded, posterior end
roundly obliquely truncated, a posterior area
is set off by a weak posterior umbonal angu-
lation; sculpture consists of fine close con-
centric threads which are crossed by fine
radial striae; on the posterior area the con-
centric sculpture becomes squamose with
fine plate-like scales; right valve with two
grooved cardinals, a close anterior lateral
and a small distant posterior lateral; left
valve with a grooved anterior cardinal, a thin
posterior cardinal and a weak anterior lat-
eral, no posterior lateral ; pallial sinus long,
almost touching the anterior adductor im-
pression, along the base wholly confluent
with the pallial line.
The character of the hinge, the pallial
sinus, and concentric sculpture anterior to
the posterior angulation in this subgenus are
similar to those of Eurytellina. The character
of the posterior area, ornamented with squa-
mose foliations, resembles that of Phyllodina
Dali.
Tellina I Phyllodella I insculpta Hanley.
Tellina insculpta Hanley, Proc. Zool. Soc.
London, September, 1844, p. 70. “Hab. Chiri-
qui, West Columbia; sandy mud, three fath-
oms.” — Hanley, Thes. Conch. Vol. 1, 1846,
p. 289, pi. 60, fig. 136. “Chiriqui, W. Colum-
bia.” — Sowerby, Conch. Icon., Vol. 17, Tell-
ina, 1867, species 208, pi. 37, fig. 208. “Hab.
Chiriqui, West Columbia.”
Type Locality. Chiriqui, west Panama, in
three fathoms, sandy mud.
Range : Champerico, Guatemala, to Santa
Elena Bay, Ecuador.
Collecting Stations: Guatemala: 7 miles
west of Champerico (197-D-2), 14 fathoms,
mud ; El Salvador: La Libertad (198-D-l, 2),
13-14 fathoms, mud.
Description: Shell elongate, compressed,
thin, white, equilateral, dorsal margins slop-
ing, anterior end rounded, posterior end an-
gulated; sculpture of fine, close, equidistant,
concentric ribs (about 4 per mm.), both ribs
and interspaces are crossed by fine submicro-
scopic radiating striae; posterior area set
off by a weak angulation and on this area the
surface is roughened by small scales or inter-
rupted delicate lamellae; right valve with
two grooved cardinals and an anterior lateral
so closely situated near the beaks as to re-
semble a cardinal, and a small posterior lat-
eral ; left valve with a grooved anterior car-
dinal, a thin posterior cardinal and a faint
anterior lateral; pallial sinus rather high
behind then descending, end blunt and al-
most, but not quite, touching the posterior
basal margin of the anterior adductor im-
pression, along the base it is confluent with
the pallial line.
The largest specimen in the collection
measures: length, 33.6 mm.; height, 18.3
mm.; convexity (both valves together), 5.8
mm.
The only other shell described from west
American waters that bears much resem-
blance to this species appears to be Tellina
( Phyllodina ) fluctigera Dali, the type of
which has not been illustrated. According to
Dali’s description the concentric sculpture of
his species is more widely spaced (about 2
lamellae per mm. whereas there are about 4
per mm. in the present species) and the pallial
sinus is entirely free from the pallial line
rather than confluent with it.
Distribution: Specimens of Tellina in-
sculpta were dredged by the expedition west
of Champerico, Guatemala, in 14 fathoms,
and at La Libertad, El Salvador, in 13 fath-
oms, on a muddy bottom. A small specimen of
this species in the collections of the Califor-
nia Academy of Sciences was collected by
Woodbridge Williams in Santa Elena Bay,
Ecuador. These records extend the known
range of the species both to the north and
to the south.
Subgenus Elliptotellina Cossmann.
Tellina lElliptotellinal pacihca Dali.
Tellina ( Elliptotellina ) pacifica Dali, Proc.
U. S. Nat. Mus., Vol. 23, No. 1210, November,
1900, pp. 302, 316, pi. 3, fig. 9. . . . “dredged
in Panama Bay, in 18 fathoms, sand.”
Type Locality: Panama Bay, in 18 fath-
oms, sand.
Range: Santa Inez Bay, in the Gulf of
California, to the Bay of Panama.
Collecting Stations: Mexico: Santa Inez
Bay, Lower California, in the Gulf of Cali-
fornia (145-D-1-3), 4-13 fathoms, sand;
Manzanillo (184-D-2), 30 fathoms, gravelly
sand; Port Guatulco (195-D-9, 19), 7-17
fathoms, gr. mud, gr. sand, crushed shell;
Tangola-Tangola Bay (196-D-6, 7), 6-7 fath-
oms, sand, crushed shell.
Description: Shell small, oval, both ends
rounded, the anterior the longer, moderately
convex, yellowish-white with a rose colored
spot near each end of the hinge margin ;
sculpture of fine concentric grooves sepa-
rated by wider interspaces, these on the pos-
terior half (and sometimes faintly to the an-
terior third) of the shell are crossed by deep,
angular, radial grooves which serrate the
posterior ventral margin and between which
are narrow interspaces ; hinge of right valve
with two cardinals, the anterior one a large
subtriangular mass and the posterior one
smaller and faintly grooved, two laterals
present; left valve with two cardinals, the
anterior one grooved, the laterals obsolete;
pallial sinus ascending, extending anteriorly
about one-half or more the length of the shell,
rounded at the end and, except for a short
88
Zoologica : New York Zoological Society
[34: 9
distance posteriorly, free from the pallial
line.
A specimen from Tangola-Tangola Bay,
Mexico, measures approximately : length, 7.8
mm. ; height, 4.4 mm. ; convexity (both valves
together) , 2.5 mm. A left valve from Port
Guatulco, Mexico, measures 8.2 mm. in
length.
This species differs from Tellina ameri-
cana Dali, 1900, a similar east American
species, in that the radial sculpture is strong-
er and is present farther anteriorly, the
pallial sinus is longer and the color is said
to be somewhat brighter.
Tellina cymobia WoodTing''1', described
from the Miocene of Jamaica, is a very simi-
lar species.
Distribution-. The discovery of the occur-
rence of Tellina pacifica in Santa Inez Bay
in the Gulf of California is a long extension
north of the known range of the species.
Genus Tellidora Morch in H. & A. Adams.
Tellidora burneti Broderip & Sowerby.
Tellina burneti Broderip & Sowerby, Zool.
Jour., Vol. 4, No. 15, January, 1829, p. 362,
pi. 9, fig. 2. “Hab. ad Mazatlan, in Aestuario.”
“Found in the Estuary of Mazatlan, among
the shoals of large Pinnae which are left dry
at low water.” — Hanley, Thes. Conch., Vol.
1, 1846, p. 271, pi. 58, fig. 99. “Salango, W.
Columbia.” — Sowerby, Conch. Icon., Vol. 17,
Tellina, 1867, species 199, pi. 35, figs. 199a,
199b. West Colombia. — M. Smith, Panamic
Shells (Trop. Photogr. Lab., Winter Park,
Florida), 1944, p. 65, fig. 843. Lower Cali-
fornia to Ecuador.
Type Locality: Mazatlan, Mexico, in the
estuary among shoals of large Pinnae at
low water.
Range: Soledad, Lower California, to the
Gulf of California and south to Salango,
Ecuador.
Collecting Stations: Mexico: Port Gua-
tulco (195-D-17), 6 fathoms, sand; El Sal-
vador: Meanguera Island, Gulf of Fonseca
(199-D-l), 16 fathoms, sand, mud, crushed
shell; Nicaragua: Corinto, beach.
Description: Triangular, rather thin, in-
equivalve, subequilateral, compressed, sub-
nacreous, white ; the convex valve with obso-
lete distant dilated concentric grooves, which
are most visible in front and towards the
elevated acute and curving beaks (one of
which projects over the other) ; the flat valve
with the elevated fine rather irregular and
close concentric striae, which become obso-
lete posteriorly ; ventral edge arcuated ; dor-
sal edges strongly sloping on either side, and
armed with large tooth-like projections, the
front one greatly incurved, the hinder nearly
straight; ligament minute; dorsal slopes ex-
cavated; fold and flexure distinct; lateral
teeth distinct, rather remote, and subequi-
distant. (Hanley, Thes. Conch., Vol. 1, 1846,
p. 271).
61 Tellina. ( Elli ptotellina ) cymobia Woodring, Carnegie
Inst. Washington, Publ. 366, May 20, 1925, p. 174, pi. 24,
figs. 14-16. Bowden, Jamaica, Miocene.
The pallial sinus is rather high and pro-
jects anteriorly more than half the length of
the shell.
A large valve in the present collection from
the Gulf of Fonseca, measures: length, 49
mm. ; height, 40.5 mm. ; convexity ( one
valve), approximately 3.4 mm.
There is some variation in the concentric
sculpture of this species but the shape and
ornamentation are so characteristic that it
cannot be confused with any other west
American shell.
Tellidora cristata Recluz62 is a similar spe-
cies which occurs from Miocene to Recent in
the Caribbean region. Tellidora ( Tellipiura )
peruana Olsson, 1944, has been described
from the Cretaceous of Peru.
Distribution: This species was collected
by the expedition off west Mexico, in the
Gulf of Fonseca off El Salvador and at Co-
rinto, Nicaragua. It also has been recorded
as occurring in the Pleistocene at Magdalena
Bay, Lower California, and Panama.
Genus Mocomo Leach.
Key to the subgenera of Macoma.
A. Shell subtrigonal, beaks subcentral
a. Shell ornamented with slightly oblique
corrugations; small; thin Cymatoica
aa. Shell ornamented with concentric
striae only Macoma s.s.
B. Shell elongate, beaks posteriorly situated ;
posterior end much the shorter
a. Resilium internal and shorter than the
ligament Psammotreta
aa. Resilium external and about as long as
the ligament
b. Posterior area granulated
Macoploma
bb. Posterior area not granulated
Psammacoma
Subgenus Macoma s.S.
Macoma I Macoma l nasuta Conrad.
Tellina nasuta Conrad, Jour. Acad. Nat.
Sci. Philadelphia, Ser. 1, Vol. 7, 1837, p. 258.
“Inhabits coast of California near Sta.
Diego.”
Macoma nasuta Conrad, I. S. Oldroyd,
Stanford Univ. Publ. Univ. Ser. Geol. Sci.,
Vol. 1, 1924, p. 174, pi. 45, figs, la, lb, lc, Id.
Kodiak Island and Cook Inlet, Alaska, to
Scammon Lagoon, Lower California. Also
Miocene, Pliocene and Pleistocene of Cali-
fornia. — Grant & Gale, Mem. San Diego Soc.
Nat. Hist., Vol. 1, 1931, p. 365, pi. 20, figs. 11a,
lib. Earlier records cited. ? Oligocene and
Miocene to Recent.
62 Lucina cristata Recluz, Rev. Zool., Soc. Cuvierienne ,
1842, p. 270. “Hab.: — Trouvee sur le banc de Campeche i
par M. Cosmao, commandant la station navale du Mexique.”
—Recluz, Mag. de Zool. (par Guerin-Meneville) , 1843, Moll., |
pi. 60, p. 1, figs. 1-5. Original locality cited.
TeUina cristata Recluz, Sowerby, Conch. Icon., Vol. 17,
Tellina, 1868, species 291, pi. 49, figs. 291a, 291b. “Hab. — V*
— M. Smith, East Coast Mar. Shells (Edwards Bros., Ann
Arbor, Michigan), 1937, p. 60, pi. 54, figs. 6a, 6b. West
Florida to Trinidad, West Indies.
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
89
Type Locality : Coast near San Diego, Cali-
fornia.
Range: Kodiak Island and Cook’s Inlet,
Alaska, to Cape San Lucas, Lower California.
Collecting Station : Mexico: Cape San
Lucas, Lower California.
Description: A single right valve of this
well known strongly flexed shell, measuring
81 mm. in length and 42.8 mm. in height, was
taken by the expedition at Cape San Lucas,
Lower California.
The present specimen might be referable
to Macoma kelseyi but it appears to be inter-
mediate in characters between that form and
M. nasuta. Dali described Macoma kelseyi™
from the “Pleistocene” of San Diego, Cali-
fornia.
It was said to differ from M. nasuta by its
greater size, thicker, flatter shell and in that
the pallial sinus in the right valve bends pos-
teriorly before coalescing with the pallial
line rather than joining it at a right angle.
Study of a series of specimens of Macoma
nasuta and of M. kelseyi reveals that there is
some variation in the pallial sinus and it
may be open to question whether Macoma
kelseyi is a distinct species, subspecies, or
merely a very large M. nasuta.
Distribution: The discovery of the occur-
rence of Macoma nasuta at Cape San Lucas,
Lower California, is an extension south of
the known range of the species. It also is
known to occur as a fossil in California from
Oligocene or Miocene to Recent.
Subgenus Cymatoica Dali.
Macoma t Cymatoica) undulata Hanley.
Tellina undulata Hanley, Proc. Zool. Soc.
London, September, 1844, p. 72. “Hab. St.
Elena, West Columbia; sandy mud, six fath-
oms.” — Hanley, Thes. Conch., Vol. 1, 1846,
p. 310, pi. 59, figs. 107, 107*. “St. Elena, West
Columbia.” — Sowerby, Conch. Icon., Vol. 17,
Tellina , 1867, species 119, pi. 23, figs. 119a,
119b. “St. Elena, West Columbia.”
Cymatoica occidentalis Dali, Proc. U. S.
Nat. Mus., Vol. 12, No. 773, 1889 (issued
March 7, 1890) , p. 272, pi. 10, fig. 11. Dredged
“ ... in latitude 24° 18' N., longitude 110° 22'
W., off the coast of Lower California, in 26V2
fathoms, fine sandy mud.”
Tellina ( Cymatoica ) undulata Hanley,
Dali, Proc. U. S. Nat. Mus., Vol. 23, No. 1210,
1900, p. 309. “Gulf of California, south to
St. Elena, West Colombia.”
Type Locality: Santa Elena, Ecuador, in
6 fathoms, sandy mud.
Range : Off the west coast of Lower Cali-
fornia in Lat. 24° 18' N., Long. 110° 22' W.,
to the Gulf of California and south to Santa
Elena, Ecuador.
Collecting Stations: Mexico: Santa Inez
Bay, Gulf of California (145-D-l, 3), 4-13
fathoms, sand; Port Guatulco (195-D-19,
63 Macoma kelseyi Dali, Trans. Wagner Free Inst. Sci.,
Vol. 3. Pt. 5, December, 1900, p. 1052, pi. 49, fig. 7. “Pleis-
tocene of San Diego, California, obtained in the City Park
by Dr. R. E. C. Stearns.” According to Dr. W. P. Wood-
ring (oral communication ) , the type of this species came
from Pliocene beds.
20), 17-21 fathoms, gr. mud, crushed shell,
mud; Santa Cruz Bay (195-D-21), 18 fath-
oms, mud; Tangola-Tangola Bay (196-D-6,
7), 6-7 fathoms, sand, crushed shell; El Sal-
vador: Meanguera Island, Gulf of Fonseca
(199-D-l), 16 fathoms, sand, mud, crushed
shell; Costa Rica: Port Parker (203-D-3),
12 fathoms, shelly mud.
Description: Shell small, thin, oblong,
beaks subcentral, bluntly pointed and flexed
to the right posteriorly, ornamented with
small, rounded, undulating riblets which do
not coincide with the incremental lines of
growth except partially so on the posterior
fold.
A right valve from the Gulf of Fonseca,
measures: length, 16.4 mm.; height, 9 mm.;
convexity (one valve), 2.6 mm.
Macoma orientalis Dali04, described from
the Antilles, is a very similar species.
Distribution: Specimens of this species
were dredged from the Gulf of California
to Costa Rica, in 4 to 21 fathoms.
Subgenus Psammacoma Dali.
Key to the species of Psammacoma.
A. Shell moderately thick; telliniform
lamproleuca
B. Shell thin; sometimes iridescent
a. Posterior end narrow; length usually
not exceeding 35 mm.
b. Very narrow and thin panamensis
bb. Higher and thicker spectri
aa. Posterior end wide; length usually
exceeding 35 mm elongata
Macoma (Psammacoma) elongata Hanley.
Tellina elongata Hanley, Proc. Zool. Soc.
London, December, 1844, p. 144. “Hab. Chi-
quiqui [Chiriqui], West Columbia; in sand
at three fathoms.’-Hanley, Thes. Conch.,
Vol. 1, 1846, p. 302, pi. 62, fig. 199. “Chiriqui,
West Columbia.”— Sowerby, Conch. Icon.,
Vol. 17, Tellina, 1867, species 137, pi. 25, fig.
137. “Hab. Chiriqui, West Columbia.”
Type Locality: Chiriqui, Panama, in 3
fathoms, sand.
Range : Lower California (Lat. 30° 36' N.)
to Panama (Dali). Caribbean region (Daut-
zenberg) .
Collecting Stations: Mexico: Santa Cruz
Bay (195-D-21), 18 fathoms, mud; Acapulco
(189-D-4), 28 fathoms, mud; El Salvador:
Meanguera Island, Gulf of Fonseca
(199-D-l), 16 fathoms, sand, mud, crushed
shell; La Union (199-D-8-10, 12-14, 22), 3-6
fathoms, mud, mangrove leaves; Nicaragua:
Monypenny Point (199-D-2-6) , 4-7 fathoms,
mud; Costa Rica: Cedro Island, Gulf of
Nicoya (213-D-l, 10), 8-10 fathoms, mud; 14
miles S. X E. of Judas Point (214-D-l, 4),
42-61 fathoms, mud, shell, rocks.
Description : Shell elongate, thin, smooth,
64 Cymatoica orientalis Dali, Proc. U. S. Nat. Mas., Vol.
12, 1889, No. 773, (issued March 7, 1890), p. 273, pi. 10,
fig. 12. "Hab.-Samana Bay, Santo Domingo, in 16 fathoms!
mud .... Also found at the same depth at Cardenas, Cuba.
90
Zoologica: New York Zoological Society
[34:9
white, sometimes iridescent outside, white
within ; anterior end rounded, posterior end
produced and bluntly truncated; a subme-
dian radial, depressed area present on the
ventral half of the shell; ornamented with
fine concentric lines of growth which are
coarser along the posterior dorsal area, and
along the posterior umbonal ridge there are
faint irregular oblique striations that do not
coincide with the lines of growth ; hinge with
two cardinals in each valve (on a large speci-
men the right anterior one sometimes some-
what roughened anteriorly), the posterior
tooth slightly cleft, the left anterior cardinal
slightly cleft, sometimes slightly indented
at the base, on some specimens there are
grooves along the margin back of the pos-
terior tooth; pallial sinus higher behind and
rounded in front, projecting forward about
two-thirds the length of the shell and along
the base for about half its length confluent
with the pallial line.
A specimen dredged southwest of Maldan-
ado Point, Mexico, measures approximately:
length, 47.4 mm.; height, 25.5 mm.; convex-
ity (both valves together), 13 mm.; pallial
sinus extends forward 30 mm. from the pos-
terior end of shell.
Tellina lamproleuca Pilsbry & Lowe is a
somewhat similar species but the median de-
pressed area is less developed, the shell is
thicker and more telliniform and the poste-
rior end is generally narrower.
Macoma gatunensis Toula, 1908, described
from the Miocene Gatun beds at Panama, is
very similar but is more excavated beneath
the beaks. The shell illustrated by Olsson65
under the name of Macoma guatunensis
Toula appears to be very similar to the pres-
ent species; in fact Olsson mentioned that
he could detect no differences between the
fossil form and Recent specimens of M. elon-
gata. Macoma falconensis H. K. Hodson, de-
scribed from the Miocene of Venezuela, also
is a somewhat similar form as is Macoma
hosfordensis Mansfield66 which was de-
scribed from the upper Miocene of Florida.
The posterior area of Macoma elongata is
smooth in comparison to somewhat similar
species assigned to Macoploma Pilsbry &
Olsson in which the posterior area is orna-
mented with granules.
Distribution : Specimens of Macoma elon-
gata were dredged by the expedition from
off western Mexico to Costa Rica, in 3 to
61 fathoms, mostly on a muddy bottom.
Dautzenberg67 cited this species as occurring
at the island of St. Lucie in the West Indies
and in the Gulf of Paria and Gulf of Mara-
caibo, Venezuela. We have not seen specimens
from east American waters.
65 Olsson, A. A., Bull. Amer. Paleo., Vol. 5, No. 39, Pt. 2,
June 21, 1922, pi. 29 (26), fig. 13. Banana River, Costa
Rica. Miocene.
Macoma I Psammacoma! lamproleuca Pilsbry &
Lowe.
leen
ill
Tellina lamproleuca Pilsbry & Lowe, Proc. n,
Acad. Nat. Sci. Philadelphia, Vol. 84, May 21,
1932, p. 93, pi. 11, figs. 6 and 7. “Corinto,
Nicaragua.” Also Panama.
Macoma parthenopa Pilsbry & Lowe, Proc. ■'
Acad. Nat. Sci. Philadelphia, Vol. 84, May 21,
1932, p. 144, pi. 11, figs. 6 and 7. “Corinto,” ,
type. Also Panama.
Macoma lamproleuca Pilsbry & Lowe, Pils-
bry & Olsson, Proc. Acad. Nat. Sci. Philadel-
phia, Vol. 93, 1941, p. 69. Jama formation,
Puerto Jama, and Canoa formation, Punta
Blanca, Ecuador, Pliocene. Also Recent from
Panama to Zorritos, Peru.
Type Locality: Corinto, Nicaragua.
Range : Santa Inez Bay, Gulf of Calif or- ,
nia, to Zorritos, Peru. | jj
Collecting Stations: Mexico: Santa Inez [j
Bay, Gulf of California (142-D-4), 40-50 p0!
fathoms, sand; Guatemala: 7 miles west of m,
Champerico (197-D-l, 2), 14 fathoms, mud; p.]
El Salvador : La Libertad (198-D-l, 2) , 13-14 pa,
fathoms, mud; Meanguera Island (199-D-l), in
16 fathoms, sand, mud, crushed shell.
Description: Shell elongate, telliniform, I |j
fairly thick, white under a buff perio- ja
stracum, somewhat inequilateral, the ante- ,:
rior end the longer, dorsal margins sloping j .j
and nearly straight, anterior end broadly
rounded, posterior end bluntly truncated; ;
surface ornamented with irregular lines of
growth and wrinkles, stronger on the poste- j
rior area; sometimes in the right sometimes
in the left valve the posterior area bears a
median radial depression; a medial flatten-
ing from the umbos to the ventral margin
often present; lunular area lanceolate and
concave; hinge with two cardinals in each
valve, the right posterior and left anterior
ones bifid, no laterals present; the pallial
sinus extends about four-sevenths the length
of the shell, it is usually higher in the middle,
rounded at the end and joins the pallial line
at an acute angle and for a little over half
its length is confluent with the pallial line.
The largest specimen in the present col-
lection, a left valve, measures : length, 72.8 !>
mm.; height, 40.5 mm.; convexity (one
valve), 10.5 mm.; pallial sinus extends for-
ward 41 mm. from posterior end of shell.
The shell of this species is similar to that
of Macoma elongata but the dorsal margins i
slope more steeply, especially anteriorly, the
posterior extremity is usually narrower, the
shell is thicker and the hinge is heavier.
Macoma falconensis H. K. Hodson68, de-
scribed from the Miocene of Venezuela, is a
similar species.
Distribution: This species was taken off
Guatemala and El Salvador in 13 to 16 fath-
oms, usually on a muddy bottom. It also has
66 Macoma ( Psammacoma ) hosfordensis Mansfield, Flor-
ida Geol. Surv.. Bull. 8, 1932, p. 142, pi. 30, figs. 6, 10, 12,
13. “Type locality: Station 3671, 2 miles north of Hosford,
Liberty County, Fla.” Choctawhatchee, upper Miocene.
67 Dautzenberg P., Mem. Zool. Soc. France, Vol 13,
1900, p. 263.
68 Macoma ( Psammacoma ) falconensis H. K. Hodson,
Bull. Amer. Paleo., Vol. 16, No. 59, October t, 1931, p. 16,
pi. 6, figs. 1, 6, 7. “Holotype Locality—. 5 kilometers north
and 350 meters west of Urumaco, in Rio Codore, District of
Democracia, Falcon.” Venezuela. Miocene. Also other locali-
ties.
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
91
been recorded as occurring in the Pliocene
of Ecuador.
Macoma f Psammacoma! panamensis Dali.
Macoma ( Psammacoma ) extenuata var.?
panamensis Dali, Proc. U. S. Nat. Mus., Vol.
23, No. 1210, November, 1900, p. 310. “Pan-
ama.”
Macoma ( Psammacoma ) panamensis
Dali, Proc. U. S. Nat. Mus., Vol. 23, No. 1210,
November, 1900, p. 324, pi. 4, fig. 3. . . .
“dredged in 33 fathoms, sand, in Panama
Bay.”
Type Locality : Panama Bay, in 33 fath-
oms, sand.
Range : Gulf of California to Panama.
Collecting Stations: Mexico: Tenacatita
Bay (183-D-3), 40 fathoms, sandy mud; 4
miles SSW. of Maldanado Point (192-D-3),
38 fathoms, mud; Tangola-Tangola Bay
(196-D-18), 30 fathoms, mud; Costa Rica:
Port Parker (203-D-3), 12 fathoms, shelly
mud; 14 miles S. X E. of Judas Point (214-
D-l, 4,), 42-61 fathoms, mud, shell, rocks;
Panama: Gulf of Chiriqui (221-D-l, 5), 35-
40 fathoms, sandy mud.
Description : Shell very elongated, slender,
thin, moderately convex, right valve slightly
flattened posterior to the center, inequilat-
eral, the anterior end the longer, posterior
end produced and subrostrate, yellowish-
white ; ornamented with fine concentric
striae which are heavier on the posterior
area; hinge with two teeth in each valve,
the right posterior and left anterior ones
cleft; pallial sinus rounded at the anterior
end which projects forward about five-
eights the length of the shell, and for about
one-half its length confluent with the pallial
line; interior of valves sometimes show ob-
scure striations near the margins.
A right valve from the Gulf of Chiriqui,
Panama, measures: length, 31 mm.; height,
14.4 mm.; convexity (one valve), 3.5 mm.;
pallial sinus extends forward 20 mm. from
the posterior end of the valve.
Macoma extenuata Dali,69 described from
the Gulf of Mexico, is a very similar species.
Macoma panamensis canalis Olsson, de-
scribed from the Miocene of the Canal Zone,
is higher in proportion to the length.
Compared to Macoma elongata Hanley,
the shell of M. panamensis is generally nar-
rower and more elongate and the posterior
end is narrower.
Distribution: This species was dredged by
the expedition from off Mexico to the Gulf
of Chiriqui, Panama, in 12-61 fathoms,
mostly on a muddy or sandy mud bottom.
Macoma I Psammacoma I panamensis spectri
Hertlein & Strong subsp. nov.
Plate I, Figs. 9, 10, 16.
Shell elongate, fairly thick, white, some-
69 Macoma ( Psammacoma ) extenuata Dali, Proc. XJ. S.
Nat. Mus., Vol. 23, No. 1210, November, 1900, p. 314, pi. 2,
fig. 7. Dredged “between the delta of the Mississippi and
Cedar Keys, Florida, in 32 fathoms, sand.” Cited on p. 300
as “Macoma (Cydippina) extenuata.”
what flexed posteriorly ; left valve moderately
inflated, the right flattened a little posterior
to the middle ; posterior dorsal margin slop-
ing, anterior end the longer, the dorsal mar-
gin slightly convex and gently sloping, that
of the right overlapping the left for about
a third of its length ; anterior end well
rounded, ventral margin only slightly curved,
posterior end produced, roundly truncated,
on each valve a distinct angulation extending
from near the beaks to the lower end of the
truncation ; exterior surface showing distinct
resting stages between which there are many
very fine concentric striations and micro-
scopic radial striae most distinct just ante-
rior to the posterior angulation ; fresh speci-
mens are often iridescent; interior shining
white; two cardinal teeth in each valve, the
right posterior and left anterior ones bifid,
lateral teeth lacking; pallial sinus subangu-
late above, highest at a point nearly vertically
below the beaks, end elliptically rounded and
extending forward about two-thirds the
length of the shell and along the base for
more than half its length confluent with the
pallial line. Holotype, a left valve, measures :
length, 34.4 mm.; height, 18.5 mm.; con-
vexity, 5.3 mm. ; pallial sinus extends ante-
riorly 21.5 mm. from the posterior end of
the shell.
Holotype, left valve, and paratype, a right
valve (Calif. Acad. Sci. Paleo. Type Coll.),
dredged in Santa Inez Bay, Gulf of Cali-
fornia, Station 143-D-3, Lat. 26° 57' N.,
Long. 111° 56' W., in 35 fathoms (64 meters) ,
mud, crushed shell.
Three additional specimens were dredged
at the same locality. Other specimens were
dredged in the same general area in Santa
Inez Bay at Station 143-D-l, 4, in 25-29 fath-
oms, mud, crushed shell, weed, sand; speci-
mens were dredged on Arena Bank, Gulf of
California, at Station 136-D-2, 45 fathoms,
mud, Area conglomerates. About a dozen
single valves, probably from the Gulf of
California, are without information as to
locality. One valve from the Gulf of Chiriqui,
Panama, Station 221-D-1-5, 35-40 fathoms,
sandy mud, appears to belong to this sub-
species.
The growth lines of these specimens are
so fine that when fresh specimens are ex-
amined at an angle they act as a grating and
produce beautiful iridescent spectral colors.
The shell here described as a new subspe-
cies is similar to that of Macoma panamensis,
but it is higher in proportion to the length
and so far as known it is generally restricted
to a more northern range. Some young speci-
mens are very similar to Dali’s species and
perhaps additional specimens may show the
form here described as new to be without
significance, but the greater height, irides-
cent color and generally more northern dis-
tribution appear, at the present time, to be
sufficient reasons for separating it as a dis-
tinct subspecies.
This new subspecies is very similar to
92
Zoological New York Zoological Society
[34: 9
Macoma panamensis canalis Olsson,70 de-
scribed from the Miocene Gatun beds of
Panama, which also is higher in proportion
to the length as compared to M. panamensis.
The shell of the present subspecies is higher
and a little less elongate than Olsson’s sub-
species. Furthermore adult shells are mod-
erately thick whereas those of Olsson’s sub-
species were described as very thin and
fragile.
Subgenus Psammotreta Dali.
Key to the species of Psammotreta.
A. Umbos orange-red; shell subrectangular,
narrow aurora
B. Umbos white shading to ochraceous ; shell
proportionately larger and higher pads
Macoma (Psammotreta) aurora Hanley.
Tellina aurora Hanley, Proc. Zool. Soc.
London, December, 1844, p. 147. “Hab. Pan-
ama ; soft sandy mud, ten fathoms : Cuming.”
— Hanley, Thes. Conch., Vol. 1, 1846, p. 301,
pi. 58, fig. 76. Panama.
Macoma ( Psammacoma ) aurora Hanley,
Salisbury, Proc. Malacol. Soc. London, Vol.
21, Pt. 2, July, 1934, p. 91, pi. 11, fig. 4. Fig-
ure of lectotype.
Type Locality: Panama, in 10 fathoms,
soft sandy mud.
Range: Gulf of California to Boca de Pan,
Peru.
Collecting Stations: Mexico: Santa Cruz
Bay (195-D-21), 18 fathoms, mud; Tangola-
Tangola Bay (196-D-14, 15), 5 fathoms,
crushed shell; Nicaragua: Corinto (200-D-8,
9), 6-6.6 fathoms, mangrove leaves.
Description: Shell elongate, longer ante-
riorly, the end rounded, posterior end bluntly
truncated, posterior area set off by an um-
bonal angulation; resilium internal and
partly separated from the ligament; shell
white or yellowish-white with the umbonal
area both exteriorly and interiorly orna-
mented with orange red which grades into
yellow anteriorly; two cardinals in each
valve, the right posterior and left anterior
ones faintly cleft; on some large right valves
there is a small denticle-like projection of
the nymph at the ventral end of the resilium;
the pallial sinus extends forward about two-
thirds the length of the shell to the anterior
edge of the orange red area but is separated
from the anterior adductor impression by
considerable space ; along the base for about
half its length it is confluent with the pallial
line.
A left valve of this species from Tangola-
Tangola Bay, Mexico, in the present collec-
tion, measures 28 mm. in length. A specimen
from Panama in the collections of the Cali-
fornia Academy of Sciences, measures:
length, 27.5 mm.; height, 17.3 mm.; convex-
ity (both valves together), 8.2 mm.; pallial
TO Macoma panamensis Dali var. canalis Olsson, Bull.
Amer. Paleo., Vol. 9, No. 39, Pt. 2, June 21, 1922, p. 429
( 2B7 > , pi. 29 (26), fig. 11. “Gatun Stage; Mt. Hope, C. Z."
sinus extends forward 19 mm. from the pos- 1
terior end of the shell.
The elongate, subrectangular form and :<
orange red radial area on the umbonal region
are characteristic features of this Macoma.
Macoma hesperus Dali,71 an unfigured spe- fii
cies described from the Gulf of Panama, was
said to bear a resemblance to M. aurora but
with a narrower shell and a different hinge.
Distribution: A few specimens of this spe- ^
cies were dredged in 5-18 fathoms off west
Mexico and Corinto, Nicaragua.
0
Macoma ( Psammotreta ) pads Pilsbry & Lowe. 0
Macoma pads Pilsbry & Lowe, Proc. Acad.
Nat. Sci. Philadelphia, Vol. 84, May 21, 1932, '
p. 95, pi. 10. figs. 1, la, 2, 3. “La Paz, Lower
California.”
Type Locality : La Paz, Lower California.
Range : Gulf of California to Golfito, Gulf t
of Dulce, Costa Rica.
Collecting Stations : Mexico : Port Gua-
tulco; Guatemala: 7 miles west of Champer- S
ico (197-D-2) , 14 fathoms, mud ; Costa Rica :
Cedro Island, Gulf of Nicoya; Golfito.
Description: Shell resembling that of Ma- n
coma aurora but larger and higher in pro-
portion to the length, usually whiter and t
tinted with yellow and on some specimens .1
the umbonal area is of an ochraceous salmon
color. Two cardinal teeth in each valve, the
right posterior and left anterior ones bifid,
the left posterior cardinal narrow and near
the posterior margin ; pallial sinus high be-
neath the beaks then descending to a bluntly
rounded end and for about half its length
confluent with the pallial line.
A specimen in the present collection from
Golfito, Costa Rica, measures: length, 34.6
mm.; height, 21.8 mm.; convexity (both
valves together) , 9.9 mm. A large right valve
from Cedro Island in the Gulf of Nicoya,
measures : length, 52.6 mm. ; height, 34 mm. ;
convexity (one valve), 6.6 mm.; pallial sinus
extends forward 35 mm. from the posterior
end of the shell.
Small specimens of this species are very
similar to those of Macoma aurora but seem
to differ constantly in the particular men-
tioned above. As pointed out by Pilsbry &
Lowe the relation of height to length varies
considerably in different specimens.
Illustrations of Macoma plebeia Hanley72
indicate a shell which is more roundly oval
with a more rounded venti'al margin poste-
riorly. According to the illustration of that
species given by Pilsbry & Lowe the pallial
sinus is more broadly rounded at the end and
is confluent with the pallial line for a shorter
distance than that of Macoma pads.
Tl Macoma ( Psammacoma ) hesperus Dali, Bull. Mus.
Comp. Zool., Vol. 43, No. 6, October, 1908, p. 421. Dredged
in the “Guif of Panama, in 182 fathoms, mud, bottom
temperature 54°. 1 F.’’
Tellina plebeia Hanley, Proc. Zool. Soc. London, De-
cember, 1844, p. 147. “Hab. Real Llejos, Central America;
sandy mud, seven fathoms.” —Hanley, Thes. Conch., Vol. 1,
1846, p. 299, pi. 60, fig. 151. Original locality cited. [Not
pi. 59, fig. 129. “Senegal”].
Macoma plebeia Hanley, Pilsbry & Lowe, Proc. Acad.
Nat. Sci. Philadelphia, Vol. 84, 1932, p. 95 (in text), pi.
10, fig. 5. “Real Llejos ( = Corinto).“
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
93
Macoma mazatlanica Deshayes73 is very
similar to M. pads but is more tapering pos-
teriorly.
Distribution : Specimens of Macoma pads
were taken by the expedition off west Mexico,
Guatemala and Costa Rica. This is an exten-
sion south of the known range of the species.
Subgenus Macoploma Pilsbry & Olsson.
Macoma I Macoploma ) medioamerlcana Olsson.
Macoma ( Macoploma ) medioamericana
Olsson, Bull. Amer. Paleo., Vol. 27, No. 106,
December 25, 1942, p. 196 (44), pi. 17 (4),
fig. 8. “Pliocene. Quebrada Pehitas, Costa
Rica.”
Type Locality : Quebrada Penitas, Costa
Rica. Pliocene.
Range: Arena Bank, Gulf of California,
to Panama.
Collecting Stations: Arena Bank, Gulf of
California (136-D-21), 45 fathoms, mud; El
Salvador: La Libertad (198-D-2), 16 fath-
oms, mud; Costa Rica: off Ballena Bay, Gulf
of Nicoya (213-D-11-17), 35-40 fathoms,
mud.
Description: Shell elongate, moderately
thin, general characters much like those of
Macoma elongata but with the anterior dor-
sal margin more steeply sloping and with
the posterior area ornamented with strong
concentric laminae and earthy granules.
The largest specimen in the collection from
the Gulf of Nicoya, measures approximately :
length, 101 mm. ; height, 54 mm. ; convexity
(both valves together), 24 mm.
Some of the specimens in this collection
agree so closely with Olsson’s description and
illustration of Macoma medioamericana that
we have referred them to his species. There
is variation in the specimens and some might
be equally well referred to Macoma ( Maco-
ploma) ecuadoriana Pilsbry & Olsson74 de-
scribed from the Pliocene of Ecuador.
According to Olsson, M. medioamericana is
proportionately longer and has coarser and
more earthy granulations on the posterior
submargins in comparison to M. ecuadoriana.
It is obvious from a study of a series of
specimens that there are several very closely
related variable species, including Macoma
elongata, M. panamensis, M. lamproleuca, M.
ecuadoriana and M. medioamericana. The
presence of granulation on the posterior
areas appears to be the only certain criterion
separating the species of Macoploma from
some of the others. The granules are present
on specimens in the present collection after
they have attained a length of about 20 to
25 mm. It is very difficult to separate speci-
mens of a smaller size from those of M.
elongata or M. lamproleuca.
,3 Tellina mazatlanica Deshayes, Proc. Zool. Soc. London
for 1854 (issued May 16, 1855), p. 359. ”Hab. Mazatlan.
Coll. Cuming:.” — Sowerby, Conch. Icon., Vol. 17, Tellina,
October, 1868, species 320, pi. 54, fig. 320. “Hab. Mazatlan.
Coll. Cuming.”
74 Macoma ( Macoploma ) ecuadoriana Pilsbry & Olsson,
Proc. Acad. Nat. Sci. Philadelphia, Vol. 93, September 9,
1941, p. 69, pi. 19, fig. 5. ‘‘Canoa formation. Punta Blanca.”
Ecuador, Pliocene.
Distribution: Specimens of this species
were dredged by the expedition at depths of
16 to 45 fathoms from Arena Bank, Gulf of
California, La Libertad, El Salvador, and
off Costa Rica in the Gulf of Nicoya, mostly
on a muddy bottom. The species also is known
to occur in the Pliocene of Costa Rica. The
present records of occurrence reveal for the
first time that this species is living at the
present time in west American waters.
Genus Apolymeth Salisbury.
Key to the species of Apolymetis.
A. Posterior end broadly rounded
a. Shell thick, hinge broad biangulata 73
aa. Shell thin, hinge narrow cognata
B. Posterior end tapering, truncated
a. Hinge fairly broad; anterior adductor
impression elongately oval; pallial si-
nus for a little less than half its length
confluent with pallial line dornbei
aa. Hinge very narrow; anterior adductor
impression very long and narrow; pal-
lial sinus confluent with pallial line for
only a very short distance
asthenodon':>
Apoly metis cognata Pilsbry & Vanatta.
Lutricola cognata Pilsbry & Vanatta, Proc.
Washington Acad. Sci., Vol. 4, September 30,
1902, p. 556, pi. 35, fig. 5. “From Tagus Cove,
Albemarle.” Galapagos Islands.
Apolymetis cognata Pilsbry & Vanatta,
Pilsbry & Lowe, Proc. Acad. Nat. Sci. Phila-
delphia, Vol. 84, 1932, pp. 96 (in text), 133.
Galapagos Islands, also Taboga Island, Pan-
ama, and Corinto, Nicaragua.
Type Locality : Tagus Cove, Albemarle
Island, Galapagos Islands.
Range : Magdalena Bay, Lower California,
to Paita, Peru.
Collecting Station: Nicaragua: Corinto
(200-D-19) , 12-13 fathoms, mangrove leaves.
Description: Shell rounded-quadrate, mod-
erately compressed, bent to the right poste-
riorly, rather thin, gray white. Sculptured
with irregular growth wrinkles and low ra-
dial striae, covered with an extremely minute
secondary radial striation. Beaks median,
worn at the tip. Anterior margin rounded;
posterior margin subtruncate, the basal
margin straightened, sinuous; pallial sinus
ample. Length of left valve, 41 mm., alt. 34
mm., diameter 8 mm. (Original description) .
Closely related to L. alta Conr., but higher,
shorter, of a more quadrate shape. The
anterior end of the pallial sinus is more
rounded, and its upper margin is not sinu-
ous. L. excavata Sowb. is a more wedge-
shaped shell (Pilsbry & Vanatta).
This species differs from the generally
more northern A. biangulata Carpenter in
the much narrower, weaker hinge, in the
more flaring and less steeply sloping poste-
rior area, and in the generally thinner shell.
75 Not represented in the present collection.
94
Zoologica : New York Zoological Society
[34:9
Specimens referable to the present species
from the Gulf of California and southward
have in some instances, formerly been re-
ferred to “Tellina” excavata Sowerby76 which
was originally described without informa-
tion as to locality. As pointed out by Pilsbry
& Vanatta the illustration given by Sowerby
indicates a shell with the posterior end
wedge-shaped. Pilsbry & Lowe later pointed
out that there seemed to be no characters by
which “Tellina” excavata could be separated
from Apolymetis dombei Hanley. They rec-
ognized only four west American species of
the genus, Apolymetis alta Conrad [=bian-
gulata Carpenter], A. asthenodon Pilsbry &
Lowe, A. cognata Pilsbry & Vanatta and A.
dombei Hanley. However for some unex-
plained reason, probably an oversight, they
cited (p. 195) A. excavata Sowerby from
Mazatlan, Mexico. In later publications,
Lowe77 cited “Metis excavata Sowerby” as
occurring at Punta Penasco, Mexico, in the
Gulf of California, and Pilsbry & Olsson78
recorded “Apolymetis excavatus Sowerby”
from the Pliocene of Ecuador. No illustra-
tions were given of the shells representing
those records so we are uncertain which
species was represented. Specimens from the
coast of the mainland appear to be identical
with those from the Galapagos Islands, the
type locality of A. cognata.
Distribution : A single small right valve
of this species, measuring about 20 mm. in
length, was taken by the expedition at
Corinto, Nicaragua, in 12-13 fathoms.
Apolymetis dombei Hanley.
Tellina dombei Hanley, Proc. Zool. Soc.
London, December, 1844, p. 144. “Hab. Pan-
ama ; twelve fathoms, sandy mud.”— Hanley,
Thes. Conch., Vol. 1, 1846, p. 323, pi. 62, fig.
182. Panama. Variety, pi. 64, fig. 222. Tum-
bez, Peru.
Tellina dombeyi Hanley, Proc. Zool. Soc.
London for 1844 (issued February, 1845) , p.
195 (index).— Sowerby, Conch. Icon., Vol. 17,
Tellina, 1867, species 169, pi. 30, fig. 169.
“Hab. Panama, Tumbez, Peru.”— Romer,
Syst. Conchyl.-Cab. von Martini und Chem-
nitz, Bd. 10, Abt. 4, Tellina, 1871, p. 205, pi.
39, figs. 7-9. Various localities cited from the
Gulf of California to Tumbez, Peru.
Psammobia sp., Li, Bull. Geol. Soc. China,
Vol. 9, No. 3, 1930, p. 262, pi. 5, fig. 32.
Dredged in Panama Bay in 10-40 ft. “Prob-
ably Gatun formation.” Pilsbry {Proc. Acad.
Nat. Sci. Philadelphia, Vol. 83, 1931, p. 431)
stated that Li’s record was based on “A good
but bleached valve of Apolymetis dombei
( Hanley).”
Type Locality. Panama, in 12 fathoms,
sandy mud.
Range : Gulf of Fonseca to Tumbez, Peru.
Collecting Stations: Nicaragua: Potosi
76 TeUina excavata Sowerby, Conch. Icon., Vol. 17,
Tellina, March, 1867, species 138, pi. 26, fig. 138. "Hab.— ?”
77 Lowe, H. N., Trans. San Diego Soc. Nat. Hist., Vol. 8,
No. 6, 1936, p. 28.
78 Pilsbry, H. A., and Olsson, A. A., Proc. Acad. Nat. Sci.
Philadelphia, Vol. 93, 1941, p. 70.
and 5 miles SSW. of Monypenny Point, Gulf
of Fonseca, beach; Costa Rica: 1 mile S. of
entrance to Golfito Bay, Gulf of Dulce, beach.
Description: Shell ovately subtrigonal,
moderately thick, rather smooth, white but
often with umbonal area reddish-orange ex-
teriorly and sometimes interiorly; the ante-
rior end the longer, rounded and somewhat
obliquely produced; a flexure or depressed
radial area is present anterior to the poste-
rior umbonal ridge; posterior end sloping,
subtriangular and subtruncated, area set off
by an umbonal angulation; hinge with two
cardinals, the right posterior bifid, the left
anterior grooved, no laterals present; the
pallial sinus is high and subangulate in the
middle then sloping down even with but well
separated from the anterior adductor im-
pression and for a little less than one-half its
length confluent with the pallial line.
The shell of this species is somewhat vari-
able in outline. Some specimens agree almost
exactly with Hanley’s original figure, others
are more trigonal. A large right valve from
the Bay of Panama in the collections of the
California Academy of Sciences, measures :
length, 66 mm. ; height, 51 mm. ; convexity
(one valve), 14 mm.
Romer, 1871, and Stearns, 1891, pointed
out the resemblance between “Tellina”
dombei and “ Tellina ” excavata Sowerby79
which was described without information as
to the locality from which it came. Later
Pilsbry & Lowe, 1932, stated that they knew
of no difference separating these two species.
Compared to Apolymetis dombei, A. asthe-
nodon Pilsbry & &Lowe80 was described as
possessing a more elongate shell with a very
narrow hinge, a narrower, longer anterior
adductor impression. Judging from the
illustrations, the pallial sinus is confluent
with the pallial line for a very short distance
if at all.
The shell of Apolymetis dombei differs
from that of Macoma grandis Hanley81 which
was originally described from Tumbez, Peru,
and was taken at Corinto, Nicaragua, by the
Templeton Crocker Expedition, 1932, in the
stronger, broader, posterior umbonal fold
and much longer pallial sinus. Judging from
the illustration of Macoma gubemaculum
Hanley,82 originally described from Real
79 TeUina excavata Sowerby, Conch. Icon., VoL 17.
TeUina, March, 1867, species 138, pi. 26, fig. 138. “Hab. — T”
Soot-Ryen ( Nyt. Mag. for Naturvid., Bd. 70 (Meddel.
Zool. Mus. Oslo. No. 27), 1932, p. 321, pi. 2, fig. 10) illus-
trated a shell under the name of Apolymetis excavata Sow-
erby from Floreana (Charles) Island, Galapagos group.
He remarked on the variability of the outer form revealed
by different specimens, some of which were elongated,
other shorter and higher.
80 Apolymetis asthenodon Pilsbry & Lowe, Proc. Acad.
Nat. Sci. Philadelphia, Vol. 84, May 21, 1932, p. 96, pi. 11,
figs. 1-3. "Panama, on the beach (D. E. Harrower, J.
Zetek).” Type. Also at La Union, Gulf of Fonseca, El
Salvador.
81 TeUina grandis Hanley, Proc. Zool. Soc. London, De-
cember, 1844, p. 141. "Hab. Tumbez, Peru.” —Hanley.
Thes. Conch., Vol. 1, 1846, p. 327, pi. 65, fig. 247. Tumbez,
Peru.
82 TeUina gubemaculum Hanley, Proc. Zool. Soc. London,
December, 1844, p. 142. “Hab. Real Llejos, Central Amer-
ica ; in sandy mud, seven fathoms.” —Hanley, Thes. Conch.,
Vol. 1, 1846, p. 326, pi. 62, fig. 186. "Real Lejos, Central
America (Cuming).”
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
95
Llejos, Nicaragua, the anterior dorsal mar-
gins slope more abruptly than those of A.
j dombei.
Distribution-. Specimens of Apolymetis
dombei were taken by the expedition on the
beaches in the Gulf of Fonseca, Nicaragua,
and Gulf of Duke, Costa Rica. We have not
seen specimens from north of Nicaragua. It
has been reported as ranging south to Peru.
It also has been recorded as occurring in
beds of upper Pliocene age in the Galapagos
Islands. “Apolymetis cf. A. dombei (Hanley)”
has been cited by Stewart83 as occurring in
upper Pliocene beds in the Kettleman Hills,
San Joaquin Valley, California.
Genus Strigilla Turton.
Key to the species of Strigilla.
A. Shell equilateral; small, white lenticula
B. Shell inequilateral, elongated posteriorly
a. Striae much more widely spaced an-
teriorly cicercula
aa. Striae equally or more closely spaced
anteriorly
b. Shell thick; convex; roundly trun-
cated anteriorly costulifera
bb. Shell thin ; flatter ; expanded an-
teriorly disjuncta
Strigilla cicercula Philippi.
Plate I, Fig. 19.
Tellina cicercula Philippi, Zeit. f. Malako-
zool., Jahrg. 3, February, 1846, p. 19.
“Patria: Mazatlan.”
Tellina dichotoma Philippi, Zeit. f. Mala-
kozool., Jahrg. 3, February, 1846, p. 20.
“Patria: Mazatlan.”
Tellina ervilia Philippi, Zeit. f. Malako-
zool., Jahrg. 3, February, 1846, p. 20.
“Patria: Mazatlan.”
Strigilla maga Morch, Malakozool. Blatter,
Bd. 7, December, 1860, p. 189. “Sonsonate.”
El Salvador.
Strigilla interrupta Morch, Malakozool.
Blatter, Bd. 7, December, 1860, p. 190. “Son-
sonate.” El Salvador.
Strigilla circercula Philippi, Dali, Proc.
U. S. Nat. Mus., Vol. 23, No. 1210, 1900, p.
305. Gulf of California to Panama.
Type Locality: Mazatlan, Mexico.
Range : Gulf of California to Ecuador.
Collecting Station: Nicaragua: Corinto
(200-D-ll, 19, also beach), 8-13 fathoms,
sand, mangrove leaves.
Description : Shell small, usually less than
1 cm. in length, rounded, elongated poste-
riorly, polished, white, with pink color usu-
ally confined to the umbonal region; sculp-
tured with fine radial striae which on the an-
terior area are curved and very much more
widely spaced.
A large right valve of this beautiful little
Apolymetis cf. A. dombei (Hanley), Stewart, U. S.
Geol. Surv., Prof . Paper 195, 1940 (issued June 7, 1941),
p. 93, pi. 32, fig. 2. Siphonalia zone in North Dome, Kettle-
man Hills, San Joaquin Valley, California. Etchegoin
formation, upper Pliocene.
species, dredged off Corinto, Nicaragua, in
13 fathoms, measures: length, 9.5 mm.;
height, 8.5 mm. ; convexity (one valve) , 2.5
mm.
The smaller size, pink colored umbonal
area, and much wider spaced sculpture on the
anterior area, are characters which serve to
separate the shell of Strigilla cicercula from
that of the young of S. costulifera Morch.
Strigilla pisiformis Linnaeus, which oc-
curs in the Caribbean region, is a very simi-
lar species.
Distribution: This species was collected
by the expedition at Corinto, Nicaragua, on
the beach and dredged at depths of 8 to 13
fathoms.
Strigilla costulifera Morch.
Plate I, Fig. 15.
Tellina carnaria Linnaeus, Hanley, Thes.
Conch., Vol. 1, 1846, p. 260 (in part), pi. 56,
fig. 38 [West Colombia record only].
Not Tellina carnaria Linnaeus. Caribbean
region.
Tellina ( Strigilla ) fucata Gould, Proc.
Boston Soc. Nat. Hist., Vol. 4, November,
1851, p. 91. Pacific coast [No exact locality
cited].— Gould, Boston Jour. Nat. Hist., Vol.
6, October, 1853, p. 399, pi. 16, fig. 4. “In-
habits Mazatlan.”
Not Tellina fucata Hinds, Zool. Voy. Sul-
phur, Moll., Pt. 3, 1844, p. 67, pi. 21, fig. 4.
Strigilla costulifera Morch, Malakozool.
Blatter, Bd. 7, December, 1860, p. 189. “Son-
sonate.” El Salvador.
Tellina chroma Salisbury, Proc. Malacol.
Soc. London, Vol. 21, Pt. 2, July, 1934, p. 84.
New name for Tellina ( Strigilla ) fucata
Gould, 1851, not Tellina fucata Hinds, 1844.
Type Locality : Sonsonate, El Salvador.
Range : Magdalena Bay, Lower California,
to the Gulf of California and south to
Ecuador.
Collecting Stations: Mexico: Tenacatita
Bay, beach; Sihuatanejo Bay, beach; Nica-
ragua: Corinto (200-D-16), 4-7 fathoms,
mangrove leaves.
Description: Shell suborbicular, the ante-
rior side much the shorter, moderately thick,
glossy, pink, rose or white ringed with pink
or carmine ; umbos smooth ; the ornamenta-
tion consists of striae, anteriorly flexuous,
forming an angle with those on the central
area, which usually radiate obliquely poste-
riorly; on the posterior slope the striae are
fine and usually meet at an acute angle; a
lunule present; hinge with a bifid cardinal
and two laterals in each valve; one or two
thickened rays often present interiorly; the
pallial sinus touches the anterior adductor
impression and is confluent with the pallial
line below ; the interior is often of a beautiful
deep red or carmine color.
Dali pointed out that the shell of this spe-
cies is very inconstant in details of sculpture,
color, and in the presence or absence of a
smooth radial streak on each valve.
A very large specimen of this species from
Magdalena Bay, Lower California, in the
96
Zoologica : New York Zoological Society
Henry Hemphill collection of the California
Academy of Sciences, measures approxi-
mately: length, 25 mm.; height, 23 mm.;
convexity (both valves together), 10.3 mm.
The specimens in the present collection are
smaller.
Strigilla costulifera is very similar to the
east American S. carnaria Linnaeus. The
west American shell is often somewhat more
rounded and the striae appear to be less
numerous.
Distribution: Specimens of Strigilla cos-
tulifera were taken by the expedition on the
beaches along western Mexico and dredged
in 4 to 7 fathoms at Corinto, Nicaragua.
Strigilla disjuncta Carpenter.
Plate I, Fig. 20.
Strigilla disjuncta Carpenter, Proc. Zool.
Soc. London, November 11, 1856, p. 160.
“Hab. In Sinu Panamensi ; legit. T. Bridges.”
Type Locality : Bay of Panama.
Range : Corinto, Nicaragua, to Panama.
Collecting Station: Nicaragua: Corinto
(200-D-ll), 8 fathoms, sand.
Description: S. testa satis magna, alba,
tenui, planata; inaequilaterali, postice pro-
ducta; marginibus dorsalibus subrectis, ad
angulam 120°, aliis bene arcuatis; lineis in-
crementi vix monstrantibus ; lineis undulatis
exillimis, antice concentricis, umbones versus
ascendentibus, sinu angustiore; dein ad mar-
ginem ventralem rapide descendentibus;
dein subito, angulo acuto, circiter 20° postice
rursus ascendentibus; lineis angularum in
valva utraque haud convenientibus ; margine
postico sinuato, sculptura postea fortiore;
margine antico quoque sinuato; lunula dis-
tincta, sinuata ; ligamento subelongato ; dent
card, valva altera uno parvo et uno magno
bifido; altera uno parvo bifido; dent. lat.
acutioribus, haud distantibus. Long. 1.35,
lat. 1.54, alt. .54 poll. (Original description).
“Allied to S. sincera Hanl. ; remarkable for
its large size and very fine markings, and
named from the lines of markings in the two
valves not agreeing at the edges.” (Car-
penter) .
Compared to Strigilla costulifera Morch,
the shell of S. disjuncta is usually larger and
the shell is thinner, flatter, more produced
anteriorly, the beaks are more projecting and
beneath them the dorsal margin is more con-
cave. It is generally white or white tinged
with pink.
A large specimen collected at Panama by
James Zetek measures: length, 36 mm.;
height, 31 mm.; convexity (both valves to-
gether) , 13.8 mm.
[34:9:1949]
This species has been cited from wesl
American waters under the name oi
Strigilla sincera Hanley84 which, according
to Hedley85, is an Australian species. Salis-
bury80 in 1934 stated that Strigilla sincera
occurs in the Panamic area but he did not
mention Hedley’s remarks on that species.
Distribution: Two single valves of this
species were dredged by the expedition at
Corinto, Nicaragua, in 8 fathoms, and one
valve was taken on shore. This is an exten- •
sion noi'th of the known range of the species.
Strigilla lenticula Philippi.
Plate I, Fig. 21.
Tellina lenticula Philippi, Zeit. f. Malako 3.
zool., Jahrg. 3, February, 1846, p. 19. “Pa-
tria: Mazatlan.”
Strigilla serrata Morch, Malakozool. Blat-
ter, Bd. 7, December, 1860, p. 189. Central t’
America.
Strigilla lenticula Philippi, Dali, Proc.
V. S. Nat. Mus., Vol. 23, No. 1210, 1900, p. .
305. Cape San Lucas to Central America.
Type Locality: Mazatlan, Mexico.
Range : Cape San Lucas, Lower California,
to Corinto, Nicaragua.
Collecting Stations: Nicaragua: Corinto
(200-D-ll, 17, 19, also on beach), 7-13 fath-
oms, sand, mangrove leaves.
Description: Shell small, about 8 mm. in
length, rounded, inflated, white, sculptured
with well-developed, oblique, incised striae
which posteriorly develop small chevron-
shaped sinuations.
A large right valve of this species, dredged
off Corinto, Nicaragua, in 13 fathoms, meas-
ures approximately: length 8 mm.; height,
7 mm. ; convexity (one valve) , 2.7 mm.
The rounded form, white color, and slight
development of V-shaped sinuations in the
sculpture posteriorly are characters which
easily serve to separate this species from
others of the genus in west American waters.
Strigilla flexuosa Say, an east American
species, is a very similar form.
Distribution: This little species was taken
by the expedition only at Corinto, Nicaragua,
on the beach and at depths of 7-13 fathoms.
84 Tellina sincera Hanley, Proc. Zool. Soc. London, April,
1844, p. 68. “Hab.— ? Mus. Cuming, Metcalfe.” —Hanley,
Thes. Conch., Vol. 1, 1846, p. 261, pi. 60, fig. 144. [Not
the record “N. W. Coast of America, (Dr. Sinclair) ,”
according to Hedley].
85 Strigilla sincera Hanley, Hedley, Proc. Linn. Soc. New
South Wales, Vol. 38, Pt. 2, 1913, p. 272. Strigilla grossiana
Hedley, 1908, was considered by Hedley to be a synonym
of S. sincera Hanley.
86 Salisbury, A. E., Proc. Malacol. Soc. London, Vol. 21,
Pt. 2, July, 1934, p. 89.
Hertlein & Strong: Mollusks of Mexico and Central America
97
949]
EXPLANATION OF THE PLATE.
Plate I.
i
#
"IG. 1. Tellina ( Moerella ) felix Hanley. Hy-
potype, right valve, from Monypenny
Point, Gulf of Fonseca, Nicaragua.
Length, 17 mm.; height, 10 mm. View
of exterior. P. 70.
'hG. 2. Tellina ( Moerella ) recurvata Hertlein
& Strong, sp. nov. Holotype, right
valve, from Loc. 23802 (C.A.S.), San
Luis Gonzaga Bay, Lower California,
in the Gulf of California. Length, 12
I mm., height, 7.5 mm. P. 71
Fig. 3. Tellina ( Moerella ) recurvata Hertlein
& Strong, sp. nov. Holotype. View of
the exterior of the specimen shown in
Fig. 2.
Fig. 4. Tellina ( Moerella ) recurvata Hertlein
& Strong, sp. nov. Holotype, left valve.
View of the exterior of the left valve
of the specimen shown in Figs. 2 and 3.
Fig. 5. Tellina ( Moerella ) arenica Hertlein
& Strong, sp. nov. Holotype, right
valve, from Station 136-D-20, Lat. 23°
30' N., Long. 109° 26' W., Arena Bank,
Gulf of California, dredged in 43
fathoms (78 meters), mud. View of
interior. P. 68.
Fig. 6. Tellina ( Merisca ) proclivis Hertlein
& Strong, sp. nov. Holotype, left valve,
from Loc. 20299 (C.A.S.), Magdalena
Bay, Lower California, Mexico. Length,
9 mm.; height, 7.8 mm. View of in-
terior. P. 83.
Fig. 7. Tellina ( Merisca ) proclivis Hertlein
& Strong, sp. nov. Holotype. View of
the exterior of the specimen shown in
Fig. 6.
Fig. 8. Tellina ( Moerella ) recurvata Hertlein
& Strong, sp. nov. Paratype, right
valve, from the same locality as the
holotype shown in Figs. 2, 3 and 4.
Length, 11.5 mm.; height, 7 mm.
Fig. 9. Macoma ( Psammacoma ) panamensis
spectri Hertlein & Strong, subsp. nov.
Paratype, right valve, from Station
143-D-3, Lat. 26° 57' N., Long. 111° 56'
W., Santa Inez Bay, Lower California,
in the Gulf of California, dredged in
35 fathoms (64 meters), mud, crushed
shell. Length, 33.8 mm. ; height, 18 mm.
P. 91.
Fig. 10. Macoma ( Psammacoma ) panamensis
spectri Hertlein & Strong, subsp. nov.
Paratype. View of the interior of the
specimen shown in Fig. 9.
riG. 11. Tellina ( Moerella ) arenica Hertlein
& Strong, sp. nov. Holotype. View of
the exterior of the specimen shown in
Fig. 5.
riG. 12. Tellina ( Tellinella ) zacae Hertlein &
Strong, sp. nov. Holotype, right valve,
from Station 136-D-l, Lat. 23° 29' N.,
Long. 109° 25' W., Arena Bank, Gulf
of California, dredged in 45 fathoms
(82 meters), mud. Length, 33.4 mm.;
height, 15.2 mm. P. 65.
*'ig. 13. Tellina ( Tellinella ) zacae Hertlein &
Strong, sp. nov. Holotype, left valve.
View of the interior of the left valve of
the specimen shown in Fig. 12.
Fig. 14. Tellina ( Merisca ) proclivis Hertlein
& Strong, sp. nov. Holotype. Right
valve of the specimen shown in Figs.
6 and 7.
Fig. 15. Strigilla costulifera Morch. Hypotype,
right valve, from Loc. 4798 (C.A.S.),
Lower California; Henry Hemphill
Coll. Length, 25 mm.; height, 22 mm.
P. 95.
Fig. 16. Macoma ( Psammacoma ) panamensis
spectri Hertlein & Strong, subsp. nov.
Holotype, left valve, from the same
locality as the paratype shown in Figs.
9 and 10. Length, 34.4 mm.; height,
18.4 mm. P. 91.
Fig. 17. Tellina ( Tellinella ) zacae Hertlein &
Strong, sp. nov. Holotype. View of the
exterior of the specimen shown in Fig.
13. P. 65.
Fig. 18. Tellina ( Eurytellina ) inaequistriata
Donovan. Hypotype, right valve, from
Station 195-D-21, Lat. 15° 44' 45" N.,
Long. 96° 06' 55" W., Santa Cruz Bay,
Mexico, dredged in 18 fathoms (33
meters), mud, crushed shell. Length,
23 mm.; height, 12.5 mm. P. 74.
Fig. 19. Strigilla cicercula Philippi. Hypotype,
right valve, from Station 200-D-19,
Lat. 12° 28' 03" N., Long. 87° 12' 39"
W., Corinto, Nicaragua, dredged in 12-
13 fathoms (22-24 meters), mangrove
leaves. Length, 9.6 mm.; height, 8.5
mm. P. 95.
Fig. 20. Strigilla disjuncta Carpenter. Hypo-
type, right valve, from Panama; James
Zetek Coll. Length, 36 mm.; height,
31.3 mm. P. 96.
Fig. 21. Strigilla lenticula Philippi. Hypotype,
right valve, from the same locality as
the specimen shown in Fig. 19. Length,
7.9 mm.; height, 7.0 mm. P. 96.
Fig. 22. Tellina ( Eurytellina ) planulata Sow-
erby. Hypotype, left valve, from 1 mile
south of entrance to Golfito Bay, Costa
Rica. Length, 51.8 mm.; height, 29.5
mm. P. 76.
Fig. 23. Tellina ( Scissula ) nicoyama Hertlein
& Strong, sp. nov. Holotype, right
valve, from Ballena Bay, Costa Rica.
Length, 34.4 mm. ; height, 19 mm. View
of hinge. P. 85.
Fig. 24. Tellina ( Scissula ) nicoyana Hertlein
& Strong, sp. nov. Holotype, left valve.
View of hinge.
Fig. 25. Tellina ( Scissula ) nicoyana Hertlein
& Strong, sp. nov. Holotype, right
valve. View of the exterior of the speci-
men shown in Fig. 23. P. 85.
Fig. 26. Tellina ( Scissula ) nicoyana Hertlein
& Strong, sp. nov. Holotype, left valve.
View of the exterior of the specimen
shown in Fig. 24. P. 85.
All the specimens illustrated on this plate are
in the type collection of the Department of
Paleontology of the California Academy of
Sciences.
-
HERTLE1N & STRONG.
PLATE I.
MOLLUSKS FROM THE WEST COAST OF MEXICO AND CENTRAL AMERICA.
Gudger : Fishes in Ranks
99
10.
Fishes That Rank Themselves Like Soldiers on Parade.
E. W. Gudger.
American Museum of Natural History.
(Plate I; Text-
INTRODUCTION.
I have read that some mammals, such as
the American bison and the antelopes of
South Africa, advance, wheel and deploy in
something like military order. But it is
doubtful if they form in ranks with heads
in a real military formation. We all know
that migrating birds fly in fair formation
and some writers have alleged that some
aquatic birds will fish in synchronous order.
But so far as known, no one seems to have
produced photographic evidence of this lat-
ter alleged behavior.
For reptiles we apparently have no photo-
graphic evidence, but there are at least two
written accounts that surely establish the
matter of ranked formation. The first of
these is from the pen of the old naturalist,
William Bartram. In 1774 (Travels through
North and South Carolina, Georgia, and . . .
Florida, etc. London, 1792, p. 118), while
ascending the St. Johns River in eastern
Florida, he found great numbers of huge and
very aggressive alligators. Some of these
threatened attacks on his little boat, when
he sought to go into a lagoon off the river
to catch some fish for his supper. He says —
“I . . . made good my entrance into the
lagoon, though not without opposition from
the alligators who formed a line across the
entrance but did not pursue me into it.” Here
the alligators were ranked in a line, ap-
parently waiting for the fishes to try to get
out into the main stream, when the alligators
too would get their supper.
This account is counterbalanced by a par-
allel description of what C. R. S. Pitman (A
Game Warden Among his Charges, London,
1931, p. 248) saw just below Murchison Falls
on the Nile River in East Central Africa,
where crocodiles are found in incredible
numbers. “Looking from above on a still
evening, one will be struck by the regular
formation taken up by row after row of
crocodiles, like ships of war, with intervals
of about 50 feet between each crocodile [and
those on either side] and 300 feet between
the rows, which extend from bank to bank
and for about two miles down stream.” But
let a fish come down and all is wild confusion
figures 1 & 2).
and struggle of the neighboring crocodiles
to get it or at least a portion of it.
The more we know about animals, the more
we find them doing unusual and unexpected
things. It is not safe offhandedly to contra-
dict accounts by non-scientific observers of
unusual behavior not otherwise physically
impossible — in fishes, as well as in other ani-
mals. No article has been found in this search
describing regimented fishes and bearing
such a title as that at the head of this report.
However, there is widely scattered evidence
that fishes do “fall in and form ranks.” Un-
fortunately, I have never had the opportunity
to see fishes take on a military formation
but various reputable observers have, and
their cumulative evidence will now be set
forth chronologically.
Fishes Ranked Like Soldiers on Parade.
The earliest account of ranked fishes found
in this search is by a writer in The New
Monthly Magazine, 1820, part II, p. 137,
who signs himself “Amateur.” This account
is also found in Thomas Boosey’s “Anecdotes
of Fish and Fishing,” London, 1887, p. 123.
“Amateur”, in writing of the exploits of
one Darcey of Oxford, an expert swimmer
and diver, who caught fishes with his hands
in a deep hole well-known to Oxonians, makes
the following statement : —
The report that Darcey made, was that many
of these fish [barbel] lay with their heads
against the bank, in parallel line, like horses
in their stalls. They were not disturbed at his
approach, but allowed him to come close and
select the finest.
In talking over this unusual matter with
my long-time friend, the late Dr. John L.
Peters, an upstate New York man, he told
me that in his boyhood he had seen this very
thing in a stream in Ulster County. At my
request he prepared a statement of his ob-
servations, which is the earliest American
evidence that has come to hand.
In 1907 or 1908, while trout-fishing in the
headwaters of Woodland Stream in Ulster Co.,
New York, my attention was called to the
100
Zoologica : New York Zoological Society
[34: 10
peculiar formation of some brook trout after
a disturbed pool had quieted down. They seemed
to line up in a formation as if some military
officer among them had got them ready for a
parade. This I saw more than once since I used
to go out of my way to watch the trout in this
pool. When disturbed, they would scatter, but
when things quieted down, they would again
take on their military formation.
This pool was just below a little rapid in
a narrow stream. It was about 8 feet wide,
3 feet deep at its head and about 6 or 8
inches at its shallow end. Dr. Peters drew a
little diagram (Text-fig. 1) to show how the
fishes were ranked.
/I I I I
II I II
Text-fig. 1. Diagram of trout on parade, in
Woodland Stream, Ulster County, New York,
1907 or 1908. Sketch by Dr. John L. Peters.
Chronologically our next evidence is in a
personal communication from Mr. Joshua W.
Atlee of Riverton, New Jersey. He wrote that
in October, 1911, he saw ranked fishes in a
pool in a rivulet flowing into the Bay of
Chaleur, Gulf of St. Lawrence. Carefully
pushing aside the shrubbery on the bank of
the pool, he had a clear view of it and its
piscine inhabitants, which he thought were
getting ready for spawning. Of these fishes
he noted that:
An interesting feature of the sight was the
fact that in the slowly moving water, due to the
[small] volume of the pool, the fish lay in “sar-
dine fashion” closely packed with heads up-
stream, stemming the current so as to retain
a similar relative position by the slight move-
ment of their tails and fins.
Finally on being disturbed, some left the pool
in various directions, mostly upstream; but my
guide, detouring and getting into the stream
above them, actually drove many of the trout
back into the pool, where they finally settled
down again as we had first found them.
My next evidence is from Mr. Howard B.
MacDonald of Yonkers, N. Y., a traveler and
lecturer of wide experience. A photograph
taken by him at Rotorua, New Zealand, in
1925, is reproduced as Plate I, Fig. 1. Of it
he wrote (personal communication) as fol-
lows:
Unfortunately, the photograph does not show
the fish in quite such straight lines as the other
picture you have. However, these fish I saw did
act in the same manner as the ones you are
studying. Each fish had a certain definite posi-
tion in relation to the others of this company;
and if the fish were disturbed by throwing a
stone into the water near them (as we all did)
then they would scatter, but each would return
almost immediately to his same position in the
group. This was checked and verified by obser-
vation many times and there is no doubt but
that each fish knew his correct position and
always went to it.
Here is the word of another reliable ob-
server, backed by photographic evidence. As
Mr. MacDonald says, this is not such as may
be seen in Plate I, Fig. 2, but discounting
shadows, the fish are in pairs and they are
lined up fairly well in ranked rows.
And now follows an excerpt from a per-
sonal communication from Dr. Louise M.
Perry, long a winter resident of Sanibel
Island off Fort Myers, southwest coast of
Florida. Dr. Perry, an acute observer for
many years of the habits of marine fishes in
that region, writes as follows under date of
July 26, 1926:
Naples, Florida [below Fort Myers], has a
fine pier for still fishing, and while waiting for
bites, I have repeatedly watched small schools
of snook (rovalle) 8 or 12 in a group, lying on
the sandy bottom, close together and parallel
with each other, all heading the same way and
all their tails gently moving to right and left
in perfect unison. Suddenly with a rush they
would dart into a school of minnows and play
havoc for a moment, then each would gently
settle down in its former place and position.
This performance would be repeated at fairly
regular intervals for a long time, and always
made me wonder how separate individual ac-
tions could be so perfectly synchronized. How
do they do it? What is the stimulus that keeps
all the tails waving to marching time and starts
the snook off in a simultaneous dash after the
little fish?
Specific attention is called to the fact that
all the tails of these marine fishes moved to
right and left in perfect unison. And so did
the tails of the freshwater fishes observed by
Mr. Atlee in 1911 in the rivulet flowing into
the Bay of Chaleur, Gulf of St. Lawrence.
Presumably the same purpose activated both
lots of fishes— to maintain position.
And now, also in 1926, come three accounts
of this behavior of other marine fishes at the
Galapagos Islands, from the pen of that vet-
eran observer, William Beebe, in his “Arc-
turus Adventure” (New York, 1926). On
p. 54 he states that they paid out strings with
pieces of bait and enticed three sharks along-
side their boat. Here follows his description
of the behavior of a large shark and its at-
tendants.
... by pulling in the tempting morsel two feet
in front of the eager blunt snouts, we brought
them to the surface directly under our feet, so
that we could watch the movements of the
brilliant blue pilotfish, that . . . anticipated
every movement of their huge patrons. One of
the big fellows had three of these little satellites
that unfailingly held their formation, one just
above his head, the other two in perfect align-
ment a few inches in front of his jaws. So ex-
actly synchronized are the movements . . . that
1949]
Gudger : Fishes in Ranks
101
it is impossible to tell whether the shark fol-
lows the pilotfish or the pilotfish the shark.
Again Beebe (1926, p. 183) notes that:
Two mighty schools of Xesurus laticlavius
[the yellow-tailed surgeonfish] passed me graz-
i ing slowly. When within six feet, they left off
i their eternal feeding and formed up into more
or less orderly ranks which flowed like some
enormously long sea-serpent around the iden-
tical corners of rocks where had passed the
i leaders, yards and yards in advance. Invariably
the formation of an irregular line led very close
1 to me, the closing up of ranks evidently being
connected with the presence of danger or at
least something suspicious or strange.
Further, Beebe (1926, pp. 290-291) makes
note of another synchronous action of the
! yellow-tailed surgeonfish: “Several hundred
approached swimming slowly along, when,
as if at a signal, all would stop, and over a
rather flat bottom would up-end like ducks
and begin to graze [“on the plant and animal
fodder which covers the rocks,” p. 290].
On February 27, 1929, the late Prof. M. M.
Metcalf wrote — “I am sending you some quo-
tations [copied] from an old letter. These . . .
aroused my interest at the time the observa-
tions were made some years ago.” These
observations were by another man, and lack-
ing a name and date, will be entered under
date (1929) of the covering letter. The per-
tinent quotation reads as follows :
... I saw in the clear pool below Trick Falls
in the Two Medicine River in Glacier National
Park seven trout behaving in a way that seemed
interesting. They were headed into the current
and were lying motionless in two perfectly
straight rows, four in the front row and three
in the back row, aligned as accurately as a
squad of well drilled soldiers. A moth came flip-
ping over the pool, touching the water now and
then. All the trout remained quiet, except that
the right trout in the rear row turned to the
right and backed around the left end of the
squad, caught the moth, returned around the
left end of the squad to his place at the right
end of the rear row again, and they all remained
in perfect formation for the several minutes
I watched them.
On the margin of the typed sheet is a pencil
sketch of the movements of the trout at the
right hand end of the second rank. This is
reproduced herein as Text-fig. 2, and is a
graphic presentation of the interesting ac-
tion of this particular fish.
The well-known sports magazine, Field
and Stream, for November, 1929, p. 104, has
reproduced the splendid photograph shown
in Plate I, Fig. 2. It is also reproduced (in
larger size) in the same journal for June,
1935, p. 44. But in neither issue is there any
account of the phenomenon, marvellous as it
is. The 1929 figure has this caption : “A most
extraordinary photograph of resting trout in
the Brule River, 40 miles from Duluth, Min-
nesota. Note the very unusual formation —
like soldiers on parade.” The 1935 issue has
a caption which remarks that “When great
schools of fish lie in still water, it takes a
Text-fig. 2. Trout in two ranks in Two Medicine
River, Glacier National Park. The right trout
in row two, backed out and followed the course
indicated to catch a moth dipping in the water,
and then returned to his position. Sketch fur-
nished by Prof. M. M. Metcalf, 1929.
skillful angler to interest them” — and
nothing about military formation in the ar-
ticle in which the figure is set.
Here is a priceless photograph showing
eight rows of “trout on parade.” It is appar-
ently the only one ever published of this
unique, indeed phenomenal, behavior of
fishes, and there is no word of comment be-
yond the caption. It seems incredible, but
such is the fact. However, the figure splen-
didly illustrates the accounts quoted above.
But before going further, the present writer
submits the following remarks.
Trout at rest in running water always face
upstream. In pools, especially small ones,
they are likely to do the same. In “trout
water,” such a pool always has a riffle or
rapid at its head, and just below this the
water is cooler and has more oxygen than
ordinary. Gill-breathing is much easier in
fishes facing upstream. Also, in such posi-
tion, the fish can readily snap up any edibles
coming down with the current. These would
seem to explain, in part at least, the heads-
upstream of this regiment of trout.
At first glance, in Plate I, Fig. 2, we see
scores of trout in right-left ranks — fishes on
parade — and the ranks separated by right-
left stretches of gravel swept clean of fine
detritus. Now let us recall that Mr. Atlee
found ranked trout in the stream leading into
the Bay of Chaleur, Gulf of St. Lawrence,
maintaining their positions by moving their
tails right and left in unison. Also Dr. Perry
saw marine fishes at Naples, southwest coast
102
Zoologica: New York Zoological Society
[34: 10: 1949]
of Florida, acting in similar fashion. So we
must conclude that the trout in Plate I, Fig.
2, were doing this very thing. Any given rank
of trout fans out the fine detritus under the
tails of its members. This is checked and
some of it precipitated by the bodies of the
rank of fishes just behind it — and so all the
way from the foremost rank to the hindmost.
Probably these rows of trout lie on gently
backward sloping ridges of the detritus. This
cleaning action holds best for the center of
the stream but fades out somewhat on the
edges where the current is weaker.
The collecting of data for an article on
military fishes was begun more than 20 years
ago. But the work went slowly and presently
press of other work — particularly the editing
of the Bashford Dean Memorial Volume — led
to the filing away of all material till a more
convenient time — which has just come. Dur-
ing this period of inactivity in this study,
letters came in from a few persons who had
heard of my interest in this problem. But
their statements were in very general terms,
and quite unclear. Had I taken the time to
ask for more specific accounts of what they
saw, I might have gotten additional valuable
data. Now it is too late.
However, abundant evidence is to be found
in the written accounts and in the sketches
and photographs herein to establish the fact
that various fishes, but especially trout, do
rank themselves in parade order. These data
certainly justify the title of this article. For
trout in running water, some tenable expla-
nation has been advanced. But for trout and
all other fishes, where behavior has been de-
scribed, there must be a more fundamental
universal reason. This no one has attempted
— the explanation must be left to the animal
behaviorists.
Finally, it may be said that, from the com-
ments of several friends, who know of the
work on this article, I am satified that this
curious behavior of ranked fishes, “fishes on
parade,” is not at all uncommon. In fact, it
is probably far better known than the present
writer realizes. However, it is a curious
thing that in this study there has not been
found a single article with such an indicative
title as this paper bears and it is hoped that
others, who have witnessed this curious be-
havior, will publish their observations and
thus establish this parade behavior as a
normal procedure.
EXPLANATION OF THE PLATE.
Plate I.
Fig. 1. Ranked trout in a pool at Rotorua, New
Zealand. Discounting the shadows, the
trout are seen to be roughly ranked in
pairs. Photograph by Howard B. Mac-
Donald.
Fig. 2. Resting trout in eight ranks, like sol-
diers on parade, photographed in the
Brule River, 40 miles from Duluth,
Minnesota. From Field & Stream, 1929.
GUDGER.
PLATE I
FIG. 2.
FISHES THAT RANK THEMSELVES LIKE SOLDIERS ON PARADE.
vSZa
Crandall: Seasonal Changes in Creatophora cinerea
103
11.
Notes on Seasonal Changes in Creatophora cinerea, the Wattled Starling.
Lee S. Crandall.
(Plate I).
The Wattled Starling ( Creatophora ci-
nerea), is an African member of the family
Sturnidae. It has a rather wide distribution,
extending from southwestern Arabia
through East Africa to the Cape. An inves-
tigation of seasonal changes in males of
this species is outlined in the following notes.
Sequence 1.
A young specimen of the Wattled Starling
( Creatophora cinerea) of undetermined sex,
and unknown locality, was received at the
New York Zoological Park on October 17,
1927.
During the entire period of observation
this bird, as well as the others noted, was
kept in an indoor, heated aviary from Octo-
ber to May, with access to an outdoor cage
during the intervening months.
At the time of arrival, the head and throat
were completely feathered and only the usual
narrow, bare malar streaks were visible.
In the spring of 1928, the bird showed
itself to be a male by an enlargement of the
throat wattle, although there was no evi-
dence of crown wattles and no loss of plum-
age of the head.
In the spring of 1929, the throat wattle
again became enlarged, a small area of the
forehead became bare and the crown wattles
appeared, reaching an upright maximum of
about %"•
On April 19, 1930, the feathers of the
forehead were dropping out. By May 9, the
crown and face were black and entirely bare,
except for a small tuft behind each nostril.
The occipital region was bare and yellow.
The crown wattles, completely sessile, had
no power of erection. The posterior wattle,
attached longitudinally, measured 9 mm. at
the base, expanding to a width of 11 mm.
and reaching a length of 20 mm. The anterior
wattle, set at an approximate right angle to
the longitudinal center line, overhung the
base of the bill and was overhung, in turn,
by the posterior wattle. Its width at the base
was 7 mm., its greatest width was 15 mm.
and its greatest length 11.5 mm.
The throat wattle, along the anterior mar-
gin, measured 40 mm. when drawn out with
the fingers. (As this wattle is slightly re-
tractile, an accurate measurement could not
be made.) It is bifurcated at the tip, the
right division measuring 9 mm., the left
12 mm.
At this time, the bird indulged in mild
courtship maneuvers, tossing his head so
that the crown wattles might fall on one side
or other, and singing a broken and guttural
song.
On August 25, the wattles were seen to
be shrinking and feathers of head and face
growing. This process continued until, on
September 14, the wattles were entirely with-
drawn and feathering was complete. How-
ever, the nodular crown wattles, normally
invisible, remained discernible when the
plumage was tightly compressed. Also, the
malar streaks, from which the throat wattle
had sprung, remained slightly more pro-
nounced than in an immature male or a
female.
During the following years, changes took
place as follows :
1931. April 11. Feathers dropping.
May 2. Change complete.
September 26. Feathers growing.
October 17. Change complete.
1932. March 1. Feathers dropping.
March 22. Change complete.
September 19. Head feathering, wat-
tles shrinking.
October 10. Change complete.
1933. January 4. Feathers dropping.
January 25. Change complete.
August 18. Head feathering.
September 12. Change complete.
December 1. Feathers dropping.
December 26. Change complete.
1934. September 12. Head feathering.
October 1. Change complete.
1935. February 27. Feathers dropping.
March 22. Change complete.
September 30. Head feathering.
October 22. Change complete.
December 30. P’eathers dropping.
1936. January 13. Change complete.
January 20. Feathers re-growing,
wattles shrinking.
104
Zoologica: New York Zoological Society
[34: 11
February 3. Head entirely re-fea-
thered, wattles partly retracted.
February 10. Feathers dropping
again, wattles enlarging.
February 24. Change complete,
wattles fully extended, bird singing.
September 21. Feathers growing.
October 14. Change complete.
December 21. Feathers dropping.
1937. January 8. Change complete.
September 20. Feathers growing.
October 7. Change complete.
December 20. Feathers dropping.
1938. January 3. Change complete, wattles
fully enlarged.
February 7. Feathers re-growing,
wattles shrinking.
February 21. Feathering complete.
March 7. Feathers dropping again,
wattles enlarging.
March 28. Change complete.
September 12. Feathers growing.
October 1. Change complete.
December 19. Feathers dropping.
1939. January 6. Change complete.
March 5. Observations ended by death
of subject.
Sequence 2.
Two Wattled Starlings which proved to
be males, were received on December 14,
1934, from a dealer. The locality from which
they had come was unknown. These birds
were kept together, under identical condi-
tions, during the course of the following
observations. They are designated as #1 and
#2. Both were fully feathered on arrival.
1935. #1.
#2.
1936. #1.
#2.
May 1. Feathers dropping.
May 21. Change complete, crown
and throat wattles well devel-
oped.
October 21. Feathers growing,
wattles shrinking.
November 13. Change complete.
June 5. Throat wattle enlarged,
feathers dropping.
June 24. Face and crown bare,
throat wattle pendant, no evi-
dence of crown wattles.
October 28. Feathers growing.
November 11. Change complete.
April 6. Feathers dropping.
April 26. Change complete.
October 26. Feathers growing.
November 13. Change complete.
April 27. Feathers dropping.
May 16. Change complete, crown
wattles minute.
November 2. Feathers gi'owing.
November 20. Change complete.
1937. #1. March 15. Feathers dropping.
March 31. Change complete.
October 18. Feathers growing.
November 6. Change complete.
#2. April 12. Feathers dropping.
April 29. Change complete, crown
wattles minute, throat wattle
deeply pendant.
October 25. Feathers growing.
November 14. Change complete.
1938. #1. February 28. Feathers dropping.
March 16. Change complete.
October 31. Feathers growing.
November 18. Change complete.
#2. March 28. Feathers dropping.
April 18. Change complete, crown
wattles minute.
October 2. Feathers growing.
October 21. Change complete.
1939. #1. March 6. Feathers dropping.
March 25. Change complete.
October 23. Feathers growing.
November 10. Change complete.
#2. March 20. Feathers dropping.
April 7. Change complete, crown
wattles minute.
October 23. Feathers growing.
November 15. Change complete.
1940. #1 February 26. Feathers dropping.
March 15. Change complete.
September 23. Feathers growing.
October 9. Change complete.
#2 April 8. Face feathers dropping.
April 27. Change complete, crown
wattles minute.
October 28. Feathers dropping.
November 13. Change complete.
1941. #1. March 3. Feathers dropping.
March 23. Change complete.
October 13. Feathers growing.
October 29. Change complete.
#2. April 14. Feathers dropping.
May 4. Change complete, crown
wattles minute.
October 27. Feathers growing.
November 14. Change complete.
1942. #1.
#2.
1943. #1.
#2.
March 9. Feathers dropping.
March 31. Change complete.
September 28. Feathers growing.
October 19. Change complete.
March 30. Feathers dropping.
April 20. Change complete,
crown wattles minute.
November 9. Feathers growing.
November 29. Change complete.
February 15. Feathers dropping.
March 8. Change complete.
March 15. Feathers dropping.
April 7. Change complete.
Sequence 3.
Two Wattled Starlings which appeared to
be male and female, were purchased from
Christoph Schulz on August 9, 1935. They
were reported by Schulz to have been col-
lected in Kenya. The male was in breeding
condition at the time of arrival, with head
bare and wattles well developed.
1949]
Crandall: Seasonal Changes in Creatophora cinerea
105
Throughout the course of observations on
this pair, the female showed no plumage
change and no enlargement of the bare malar
streaks, at the time the male was coming into
breeding condition. During October and
November of each year she went through a
complete body molt.
Changes in the male were noted as follows :
1935. October 22. Feathers growing.
November 13. Change complete.
1936. April 6. Feathers dropping.
April 24. Change complete, wattles
well developed.
October 5. Feathers growing.
October 27. Change complete.
1937. January 18. Feathers dropping.
February 4. Change complete.
March 1. Feathers re-growing.
March 15. Face almost completely
feathered, wattles shrunken.
March 22. Feathers dropping again,
wattles re-enlarging.
April 2. Head entirely bare, wattles
large.
October 25. Feathers growing.
November 12. Change complete.
1938. February 14. Feathers dropping.
March 3. Change complete but wattles
small.
March 28. Face re-feathering.
April 11. Face almost entirely re-
feathered.
April 18. Feathers dropping again.
May 5. Change complete, wattles
much enlarged.
October 31. Face feathers growing.
November 19. Change complete.
1939. March 27. Feathers dropping.
April 17. Change complete.
October 9. Feathers growing.
October 30. Change complete.
1940. February 26. Feathers dropping.
March 20. Change complete.
September 23. Feathers growing.
October 10. Change complete.
1941. March 24. Feathers dropping.
April 12. Change complete.
September 29. Feathers growing.
October 18. Change complete.
1942. March 2. Feathers dropping.
March 20. Change complete.
October 5. Feathers growing.
October 23. Change complete.
1943. February 15. Feathers dropping.
March 6. Change complete.
Summary.
Seasonal changes in four male and one
female specimens of the Wattled Starling
( Creatophora cinerea) have been tabulated
and described. All of these birds were kept
under identical conditions, as far as caging,
food and temperatures were concerned. It is
shown that in the males there is a seasonal
loss of the plumage of the head, accompanied
by enlargement of the crown and throat
wattles. The single female showed no en-
largement of the bare gular tracts and had
only a single annual change of plumage,
which took place at the period of regression
in the accompanying male.
The males of Sequence 1 and Sequence 3
showed occasional “false starts,” in which
newly bared heads almost immediately re-
feathered, quickly followed by a resumption
of the bare condition. Neither of the two
males described in the second series of ob-
servations showed this phenomenon.
Recorded dates of changes were estab-
lished on a visual basis. However, standards
of judgement were the same in all cases, so
that periods indicated are properly com-
parable and variations would be small.
104
Zoologica : New York Zoological Society
[34: 11
February 3. Head entirely re-fea-
thered, wattles partly retracted.
February 10. Feathers dropping
again, wattles enlarging.
February 24. Change complete,
wattles fully extended, bird singing.
September 21. Feathers growing.
October 14. Change complete.
December 21. Feathers dropping.
1937. January 8. Change complete.
September 20. Feathers growing.
October 7. Change complete.
December 20. Feathers dropping.
1938. January 3. Change complete, wattles
fully enlarged.
February 7. Feathers re-growing,
wattles shrinking.
February 21. Feathering complete.
March 7. Feathers dropping again,
wattles enlarging.
March 28. Change complete.
September 12. Feathers growing.
October 1. Change complete.
December 19. Feathers dropping.
1939. January 6. Change complete.
March 5. Observations ended by death
of subject.
Sequence 2.
Two Wattled Stai’lings which proved to
be males, were received on December 14,
1934, from a dealer. The locality from which
they had come was unknown. These birds
were kept together, under identical condi-
tions, during the course of the following
observations. They are designated as #1 and
#2. Both were fully feathered on arrival.
1935. #1.
#2.
1936. #1.
#2.
1937. #1.
May 1. Feathers dropping.
May 21. Change complete, crown
and throat wattles well devel-
oped.
October 21. Feathers growing,
wattles shrinking.
November 13. Change complete.
June 5. Throat wattle enlai'ged,
feathei's dropping.
June 24. Face and crown bai'e,
throat wattle pendant, no evi-
dence of crown wattles.
October 28. Feathers growing.
November 11. Change complete.
April 6. Feathers dropping.
Api’il 26. Change complete.
October 26. Feathers growing.
November 13. Change complete.
April 27. Feathers dropping.
May 16. Change complete, crown
wattles minute.
November 2. Feathers growing.
November 20. Change complete.
March 15. Feathers dropping.
March 31. Change complete.
October 18. Feathers growing.
November 6. Change complete.
#2. Api'il 12. Feathers dropping.
Api’il 29. Change complete, crown
wattles minute, throat wattle
deeply pendant.
October 25. Feathers growing.
November 14. Change complete.
1938. #1. Februai’y 28. Feathers dropping.
March 16. Change complete.
October 31. Feathers growing.
November 18. Change complete.
#2. March 28. Feathers dropping.
April 18. Change complete, crown
wattles minute.
October 2. Feathers growing.
October 21. Change complete.
1939. #1. March 6. Feathei's dropping.
March 25. Change complete.
October 23. Feathers growing.
November 10. Change complete.
#2. March 20. Feathers dropping.
April 7. Change complete, crown
wattles minute.
October 23. Feathers growing.
November 15. Change complete.
1940. #1 February 26. Feathers dropping.
March 15. Change complete.
September 23. Feathers growing.
October 9. Change complete.
#2 April 8. Face feathers dropping.
April 27. Change complete, crown
wattles minute.
October 28. Feathers dropping.
November 13. Change complete.
1941. #1. March 3. Feathers dropping.
March 23. Change complete.
October 13. Feathers growing.
October 29. Change complete.
#2. April 14. Feathers dropping.
May 4. Change complete, crown
wattles minute.
October 27. Feathers growing.
November 14. Change complete.
1942. #1. March 9. Feathers dropping.
March 31. Change complete.
September 28. Feathers growing.
October 19. Change complete.
#2. March 30. Feathers dropping.
April 20. Change complete,
crown wattles minute.
November 9. Feathers growing.
November 29. Change complete.
1943. #1. February 15. Feathers dropping.
March 8. Change complete.
#2. March 15. Feathers dropping.
April 7. Change complete.
Sequence 3.
Two Wattled Starlings which appeared to
be male and female, were purchased from
Christoph Schulz on August 9, 1935. They
were reported by Schulz to have been col-
lected in Kenya. The male was in breeding
condition at the time of arrival, with head
bare and wattles well developed.
19491
Crandall: Seasonal Changes in Creatophora cinerea
105
Throughout the course of observations on
this pair, the female showed no plumage
change and no enlargement of the bare malar
streaks, at the time the male was coming into
breeding condition. During October and
November of each year she went through a
complete body molt.
Changes in the male were noted as follows :
1935. October 22. Feathers growing.
November 13. Change complete.
1936. April 6. Feathers dropping.
April 24. Change complete, wattles
well developed.
October 5. Feathers growing.
October 27. Change complete.
1937. January 18. Feathers dropping.
February 4. Change complete.
March 1. Feathers re-growing.
March 15. Face almost completely
feathered, wattles shrunken.
March 22. Feathers dropping again,
wattles re-enlarging.
April 2. Head entirely bare, wattles
large.
October 25. Feathers growing.
November 12. Change complete.
1938. February 14. Feathers dropping.
March 3. Change complete but wattles
small.
March 28. Face re-feathering.
April 11. Face almost entirely re-
feathered.
April 18. Feathers dropping again.
May 5. Change complete, wattles
much enlarged.
October 31. Face feathers growing.
November 19. Change complete.
1939. March 27. Feathers dropping.
April 17. Change complete.
October 9. Feathers growing.
October 30. Change complete.
1940. February 26. Feathers dropping.
March 20. Change complete.
September 23. Feathers growing.
October 10. Change complete.
1941. March 24. Feathers dropping.
April 12. Change complete.
September 29. Feathers growing.
October 18. Change complete.
1942. March 2. Feathers dropping.
March 20. Change complete.
October 5. Feathers growing.
October 23. Change complete.
1943. February 15. Feathers dropping.
March 6. Change complete.
Summary.
Seasonal changes in four male and one
female specimens of the Wattled Starling
( Creatophora cinerea) have been tabulated
and described. All of these birds were kept
under identical conditions, as far as caging,
food and temperatures were concerned. It is
shown that in the males there is a seasonal
loss of the plumage of the head, accompanied
by enlargement of the crown and throat
wattles. The single female showed no en-
largement of the bare gular tracts and had
only a single annual change of plumage,
which took place at the period of regression
in the accompanying male.
The males of Sequence 1 and Sequence 3
showed occasional “false starts,” in which
newly bared heads almost immediately re-
feathered, quickly followed by a resumption
of the bare condition. Neither of the two
males described in the second series of ob-
servations showed this phenomenon.
Recorded dates of changes were estab-
lished on a visual basis. However, standards
of judgement were the same in all cases, so
that periods indicated are properly com-
parable and variations would be small.
106
Zoologica: New York Zoological Society
[34: 11: 1949
EXPLANATION OF THE PLATE.
Plate I.
Fig. 1. Adult $ Wattled Starling (Sequence
1) , photographed on June 13, 1933. The
head is completely bare and wattles
fully developed.
Fig. 2. The same bird, photographed on No-
vember 20, 1934. He has completed
regressive changes and is in resting
condition. The nodular crown wattles,
not normally visible at this time, are
seen because the feathers are tightly
compressed. The throat wattle has re-
ceded to the lateral malar patches.
CRANDALL.
PLATE I.
FIG. I.
FIG. 2.
NOTES ON SEASONAL CHANGES IN CREATOPHORA C1NEREA, THEWATTLED STARLING.
Beebe : Insect Migration at Rancho Grande
107
Insect Migration at Rancho Grande in North-central Venezuela.
General Account.1
William Beebe.
Director, Department of Tropical Research,
New York Zoological Society.
(Plates I & II; Text-figure 1).
[This is one of a series of papers resulting
from the 45th, 46th and 47th Expeditions of
the Department of Tropical Research of the
New York Zoological Society, made during 1945,
1946 and 1948, under the direction of Dr.
William Beebe, with headquarters at Rancho
Grande in the National Park of Aragua, Vene-
zuela. The expeditions were made possible
through the generous cooperation of the Na-
tional Government of Venezuela and of the
Creole Petroleum Corporation.
[The characteristics of the research area are
in brief as follows ; Rancho Grande is located in
north-central Venezuela (10° 21' N. Lat., 67° 41'
W. Long.), 80 kilometers west of Caracas, at an
elevation of 1,100 meters in the undisturbed
montane rain forest which covers this part of
the Caribbean range of the Andes. The migra-
tion flyway of Portachuelo Pass, which is also
the water-shed between the Caribbean and Lake
Valencia, is 200 meters from Rancho Grande.
Adjacent ecological zones include seasonal
forest, savanna, thorn woodland, cactus scrub,
the fresh-water lake of Valencia and various
marine littoral zones. The Rancho Grande area
is generally subtropical, being uniformly cool
and damp throughout the year because of the
prevalence of the mountain cloud cap. The dry
season extends from January into April. The
average humidity during the expeditions, in-
cluding parts of both wet and dry seasons, was
92.4%; the average temperature during the
same period was 18° C; the average annual rain-
fall over a five-year period was 174 cm. The flora
is marked by an abundance of mosses, ferns and
epiphytes of many kinds, as well as a few
gigantic trees. For further details see Beebe
and Crane, Zoologica, Vol. 32, No. 5, 1947. Un-
less otherwise stated, the specimens discussed
in the present paper were taken in the montane
cloud forest zone, within a radius of one kilo-
meter of Rancho Grande.]
Contents.
General Account 107
Migrating Insects 108
Migration Factors 109
Rainy Season 109
Inhibiting Conditions 109
Favoring Conditions 109
Recurrent Waves 109
Between Waves 109
Wing Condition 109
Models and Mimics 109
Specific Characteristics 109
Sexes and Breeding 109
Diurnal Sequence 109
Interpretation HO
Sight Identification 110
t Contribution No. 843, Department of Tropical Research,
New York Zoological Society.
General Account.
Throughout the first year of our occupa-
tion of the station at Rancho Grande in
north-central Venezuela, we had no idea of
the importance of Portachuelo Pass as a
migration flyway for birds and insects.
Even later on, when we came to compile a
list of thirteen life zones within our visual
radius, a fourteenth, the Aerial Zone, was
added with hesitation, having in mind the
inclusion of organisms such as humming-
birds, swifts and mayflies which spend the
major part of their lives in midair. Almost
immediately, however, the value of and need
for such a niche in our phenological pro-
gram became apparent.
If for no other reason, an Aerial Zone
was needed to accommodate the volant or-
ganisms which passed and repassed, or oc-
casionally were detected soaring in air, and
which were never to be found resting or
flying in the jungle of our immediate area
of research. Many of these creatures were
essentially tropical, occupying our sub-
tropical elevation only as a temporary route
of passage.
About 200 meters beyond Rancho Grande,
the road leads through a narrow notch in
the east-west, coastal Andean range. This
is Portachuelo Pass with an elevation of
1,136 meters, about 36 meters higher than
Rancho Grande. The flattened floor of the
pass is only about 20 meters wide, and the
shoulders on either side rise in sharp
ridges, 389 meters to the summit of Pico
Periquito on the west, and 764 meters on
the east to the top of Pico Guacamayo.
The pass is at the 22.5 kilometer mark
on the road from Maracay, and, as has been
stated, is at an elevation of 1,136 meters.
Kilometer 31, well to the north of the pass,
is 770 meters above the sea. Kilometer 15,
equally distant to the south of the pass and
close to the beginning of the lowland savan-
nas, is 760 meters above the sea. At both of
these lower stations many migrants have
been taken, en route to or on their way from
the pass.
The pass is on the real divide, shunting
the waters on its northern slope into the
108
Zoologica: New York Zoological Society
Caribbean Sea, and those on the south side
ultimately into Lake Valencia.
Fifteen orders of insects have already
been collected or observed as they passed
southward on migration, singly, in few or
in enormous numbers. Of other possible
. migrants this leaves only three orders,
Ephemerida, Embiidina and Trichoptera.
The four parasitic groups, Anopleura, Si-
phonoptera, Mallophaga and Strepsiptera,
are of course absent, although the two
latter doubtless hitch-hike through the pass
on birds and bees respectively.
The migrating orders, arranged in three
columns of relative numbers, are as follows :
Rare
Isoptera
Neuroptera
Plecoptera
Corrodontia
Thysanoptera
Dermaptera
Mecoptera
Moderate
Orthoptera
Odonata
Homoptera
Hemiptera
Abundant
Coleoptera
Lepidoptera
Diptera
Hymenoptera
Up to the date of this publication/ .Mr.
Henry Fleming has identified two hundred
and forty-five species of butterflies, and
fifty-two species of day-flying moths. Of the
single family of nocturnal moths, Sphing-
idae, we have recorded seventy-six species,
either directly migrating through the pass,
or flying about our lights at Rancho Grande.
Mr. Fleming has found only two of these
which may be classed as breeding in the
cloud forest surrounding the pass and our
laboratory.
Except for a temporary, limited, north-
ward drift of individual Phoebis ( Catopsilia )
in the early part of the rains, the movement
of all orders of insects was invariably from
north to south through the pass.
Migrating Insects.
Examples of extremes in migration will
serve to point up succeeding papers dealing
with families and species.
On April 29, 1946, I caught a solitary
butterfly at the rim of the pass. Its wing
spread was small, and it was an inconspic-
uous purplish-brown with five spots of dull
white near the tip of each fore wing. It
belonged to the family Nymphalidae. I gave
it the reference field name of Ten-spot
Brown and later found its technical name
was Eunica monima (Cramer).
On May 4, five days later, I caught sight
of several butterflies passing overhead and
at the pass itself I entered a dense swarm
of the Ten-spots. Mingled with them were
tailed nymphalids, Marpesia chiron chiron
(Fab.) in large numbers, and now and
then a swarm of large sulphurs, Phoebis
eubule marcellina (Cramer). In the dis-
tance I could see myriads of Ten-spots con-
verging on the pass. One swoop of the
net captured seven, five of which were tat-
tered and torn, the remaining two freshly
emerged.
[34: 12
Jfll
T J'r-
Text-fig. 1. Map showing location of Rancho
Grande, Portachuelo Pass and surrounding
territory.
bg to
j indwf
Two of us climbed a mound giving a view
of about half the width of the pass, and
here, facing in opposite directions, at eye
level, we averaged thirteen hundred butter-
flies in several counts of four minutes each.
At intervals throughout an hour and a half
this insect content of a limited time and
space remained fairly constant, and when
we left we knew that at the very least, two
hundred and eighty-six thousand Ten-spots
had passed close to us. An hour later the
insects were going full strength and now I
brought to bear my giant binoculars, first
twelve and then twenty powers. I began
about twenty-five feet overhead and then
refocussed slowly upward until the limit of
vision of the small insects was reached.
This, judged by horizontal tests of objects
of similar size would be about a half mile
zenithwards, and at every fractional turn
of the screw, more and more smaller-ap-
pearing butterflies fluttered into clarity.
Throughout the entire extent of vertical-
ity there was no lessening of denseness of
flying insects, and it was almost a pure
culture of Eunica and Marpesia. For many
days this particular phase of migration con-
tinued, millions upon millions coming from
some unknown source, travelling due south
to an equally mysterious destination.
Three weeks later, on May 24, there was
a resurgent migration of the same species,
all fresh insects. Their numbers far ex-
ceeded the first wave. Four of us lined up
across the entire width of the pass, with
stop-watches and counters, completely failed
ami .
rent o
m. ■
lion ::
Bid bs
my ne
two V,'
ind :
warm
K CJ
butte;
ing v
dean
April
tort
rery
Se]
wr 1
day r
leai
icon
m\
Fen
Dole
teaj
Dial
T1
liar
cfa
1949]
Beebe: Insect Migration at Rancho Grande
109
to keep up with fast enough estimate of
1 numbers, but at the minimum clocked a
thousand a second going past in the face
of a gentle breeze. In the narrow trail above
the gorge it was necessary to put on glass-
es, so dense were the crowds impinging
upon our faces.
As the other extreme, I may mention a
half hour of collecting when many species
in fewer numbers were passing. Twenty
successive specimens of butterflies resolved
into eleven species of pierids. On another
occasion thirteen individual butterflies
proved to be thirteen separate species of
ithomiids.
Non-recognition of the pass as a fly way
accounts for the small number of observa-
tions in the year 1946, and the still more
meagre and casual notes in 1945. Some time
passed before we realized that all of the
host of moths which came in windrows to
the roof lights and laboratory windows of
Rancho Grande were Portachuelo Pass mi-
grants, deflected by confusion of fog or rain.
On clear nights of star and moonlight ohr
torches and portable ultra-violet machines
revealed unbroken streams of moths of all
sizes headed up and through the pass. Other
indirect evidences were the wings, belong-
ing to great numbers of species and indi-
viduals of moths, found glued in early morn-
ing to the dew-moistened leaves of shrubs
and weeds in the pass; the remains of noc-
turnal feasts of marauding bats.
Migration' Factors.
One definite factor, which seems the
dominant stimulus of migration, is the ad-
vent of the rainy season. For example, in
1948, there was no hint whatever of migra-
tion in February. On March 1 a single torn
and bedraggled nymphalid, Marpesia chiron
chiron, struggled up to the pass and into
my net. Hardly another insect appeared for
two weeks, throughout a period mostly cold
and overcast. Then, on March 15, a day of
warm sun after several days of heavy rain,
we caught or recorded twenty species of
butterflies in considerable numbers. Succeed-
ing weeks of cold resulted in a complete
dearth or mere scattering of insects, until
April 15. From this date until August 1
there was no cessation of numbers pouring
through, varied only by irregular fluctua-
tions due to occasional days of cold rain or
very high wind.
September 9 is the latest date of any of
our three years of residence, and on that
lay migration was in full swing. From what
I can learn, the passing insects gradually
decrease throughout the succeeding two
weeks. On the authority of Dr. Francisco
Fernandez, Venezuelan Government Ento-
mologist, diurnal migration at the pass
ceased for the year by October 1. The an-
nual picture thus seems clear cut.
The following applies more particularly to
diurnal Lepidoptera, but in general is true
of all orders :
Inhibiting Conditions: Very high winds,
from twenty-five miles per hour upward;
chilly temperatures, 62° Fahr. down; dense
fog (neblina) or heavy rain; darkness.
Favoring Conditions : Calm, up to a twen-
ty mile per hour following wind; 64° Fahr.
plus; sun or thin neblina.
Recurrent Waves: These last from twenty
minutes to three weeks, and usually com-
prise few species (two to twenty), but often
large numbers of individuals. These waves
are occasionally independent of favorable
conditions, the hosts of insects banking up
in the lee of brush, waiting for good flying
weather.
Between Waves: At these times insects
tend to fly singly, and in great variety of
species.
Wing Condition: Worn and fresh speci-
mens may be present in the same wave, but
as a rule all are worn or all are freshly
emerged.
Models and1 Mimics: Some of the more
generally accepted categories of models and
mimics may appear mingled together in the
waves, or, very interestingly, there are not
infrequently pure cultures of each, confined
to waves of considerable magnitude.
Specific Characteristics: Normal specific
characteristics of flight and of choice of
habitat are maintained throughout migra-
tion. Rapid or slow flyers do not alter their
relative speeds, nor change their dodging,
zigzag or direct 'flight. The same applies as
well to high or low habitual levels of flight,
fast or slow flapping of wings. Species which
prefer to wind their way through low, thick
brush adhere to this habit en route up to and
through the pass.
Sexes and Breeding: The general rule is
the presence of both sexes, and many of the
females captured alive deposit eggs within
twenty-four hours. Rarely, attempts at mat-
ing on migration are observed, or pairs ap-
pear to be fighting as they circle rapidly in
midair. Very few loiter to feed at blossoms.
Few worn individuals stop to rest.
Diurnal Sequence: A few insects, belong-
ing to various orders other than Lepidoptera,
appear very early in the morning, for months
on end, flying past singly, but in the aggre-
gate in great numbers. Especially noticeable
among these are two species of cockchafers
( Cyclocephala spp.), a chrysomelid ( Dia -
brotica quindecimpunctata) , a small vespid
( Stalopolybia areata ) , a giant hairy scoliid
( Campsomeris ianthina ) , and a bee ( Euglos -
sa fasciata ) . The numbers of these solitary
migrants passing on the morning of June 19,
1948, typifies the numbers on every day for
the preceding two months: cockchafers, 200;
chrysomelids 150; small vespids 150; giant
scoliids 140; rufous bees 90.
Following these there comes, for an hour
or longer, a steady procession of day-flying
moths, also singly. Butterflies dominate the
remainder of the day as far as relatively
large insects are concerned. Throughout the
110
Zoologica: New York Zoological Society
[34: 12: 1949]
0-
daylight hours there is a continuous passing
of migration nekton, hosts upon hosts of
minute winged insect life. When dusk gives
way to darkness, moths and other nocturnal
insects appear, and surge through the pass.
If the night is fine, with clear moon or star-
light, all continue down Limon Gorge. If the
sky is overcast, thick with neblina or rain,
the moths leave their direct southern route
and detour in tens of thousands to our lighted
laboratory windows or white roof walls.
Interpretation: At present, before a de-
tailed study has been made of the mass of
specimens and data, and further explorations
undertaken of places of origin and destina-
tion in surrounding country, no reasonable
explanation of this wholesale annual emigra-
tion is possible.
Unlike the migration of hosts of Phoebis
males which I have observed in British Gui-
ana and elsewhere, the Portachuelo hosts are
represented by both sexes. Many of the fe-
males are ready to deposit eggs, although
they are headed away from areas rich in a
variety of plants, toward less luxuriant
savannas.
The known distribution of many species,
or especially subspecies, of butterflies such
as the Papilios, shows that their place of
origin cannot be very far away to the north
and west, but as yet we have no hint of
whence the fifteen orders are derived, or
whither they are headed. From our own ob-
servations, reinforced by the reports of reli-
able govenment official entomologists, we are
certain that little or no migration occurs
during the dry season, and that not an insect
ever returns to its natal haunts. Hence the
phenomenon is really an emigration of the
cross section of a considerable volant inverte-
brate fauna of this part of Venezuela.
Observations during three years confirm
the fact that this migration is a regular an-
nual event. It presents the inexplicable prob-
lem of a regular abstraction of an appreci-
able percentage of non-returning population
from the area of origin. This implies the
leaving behind of an adequate residual num-
ber of non-migrants to carry on the race and
to sustain future migration.
Sight Identification: When there came
recognition of migration on a relatively great
scale, our first and indeed continued impulse
was to capture as many specimens of as many
different kinds as was humanly possible. On
an early day of observation a butterfly was
taken which, in our mind, was instantly label-
led a Monarch, Danias archippus, or, if you
prefer, Danaus plexippas, a familiar of our
northern fields. Within an hour eighteen of
the same species flapped slowly past, two of
them alighting for a few seconds. Within ten
minutes there passed eight smaller, darker
red butterflies, two of which we took, which
vividly l'eminded us of our northern Danaus
berenice, commonly called the Queen. Ul-
timately these two resolved into Danaus
plexippus megalippe and Danaus eresimus
eresimus, but throughout my notes, before .
identification, they were nostalgically record- '
ed as Monarch and Queen. The important
thing is that, being easily and accurately
recognizable at a distance in flight for what
they are, I am able to record, without fear
of error, all the numbers that came within
my purview, in all my hours of observation
at the pass.
This is all by way of introducing the im-
portant question of sight identification,
which, in any research such as the present,
must play a dominant part. I based all my
field naming first, on captured and ultimately
precisely named specimens, and secondly, on
characters in flying or resting insects which
left no shadow of doubt. Although hundreds
and tens of thousands of insects passed with
only the vaguest hints of family or genus,
yet day after day familiarity introduced to
the perception characteristics of flight, pat-
tern, color and shape of wings, and general
facies, which materially increased range and
certainty of recognition.
Viewing from a distance of ten meters,
groups of species mounted in open cases,
proved an excellent check on visual aware-
ness of distinctions. A brief treatment of
sight identification will be added to each
paper dealing with families of insects.
This and following papers are intended
only as factual presentations of notes made
during three seasons of observations of mi-
gration from north to south through Por
tachuelo Pass. Hence no attempt has been
made at correlation or even mention of mi-
grations of the same or related species ob-
served by entomologists elsewhere. The in-
sects themselves will be considered group by
group in successive papers, with a final sum
mary in detail of the migration as a whole.
We hope, in future expeditions in this
same field, that data will be obtained which
will clarify the place of origin and ultimate
destination of the insect hosts, as well as the
initiating stimuli and directive factors of
their migration.
An account of the bird migration through
the pass has already appeared, treating of
sixty species divided among ten types of
migration.2
Fig.
Fig.
EXPLANATION OF THE PLATES.
Plate I.
1. Looking south toward Portachuelo
Pass from Kilometer 31.
2. Portachuelo Pass, the notch in the
distant sky-line, from farther north,
near the sea.
Plate II.
Fig. 3. Migrant insects alive but quieted by
refrigeration.
Fig. 4. Migrant moths deflected in great num-
bers, on nights of storm, from their
migration through the pass, to the
electric lights on Rancho Grande roof.
2 Zoologica. 32 (18), 1947, pp. 153-168.
BEEBE.
PLATE I
FIG. I.
FIG. 2.
INSECT MIGRATION AT RANCHO GRANDE IN NORTH-CENTRAL VENEZUELA.
GENERAL ACCOUNT.
:ebe.
PLATE II.
FIG. 4.
INSECT MIGRATION AT RANCHO GRANDE IN NORTH-CENTRAL VENEZUELA.
GENERAL ACCOUNT.
Riess, Ross, Lyerly & Birch: Behavior of Captive Lowland Gorillas
111
13.
The Behavior of Two Captive Specimens of the Lowland Gorilla,
Gorilla gorilla gorilla (Savage & Wyman).1
B. F. Riess,2 * Sherman Ross,2 S. B. Lyerly,4 and H. G. Birch."
Behavior Research Fellows (1948), New York Zoological Society.
(Plates I & II; Text-figures 1 & 2).
I. Introduction.
The field of comparative behavior research
has long been subject to two methodological
factors which have to a certain extent pre-
vented the attainment of its goal, namely the
securing of information on behavioral pro-
cesses of representative species of a wide
range of living organisms. The first factor
has been the concentration of research work-
ers on those animals which are adaptable to
the limited conditions of laboratories. This
emphasis on a few selected species has led
to the second factor, the acceptance of the
conventional laboratory as the prototype of
habitat for the species under investigation.
Both these methodological limitations have
arisen in part from the same set of circum-
stances, the relative lack of availability of
less adaptable organisms and the expense of
field studies. The increasing number of in-
vestigations under naturalistic field condi-
tions and of studies on rare specimens under
favorable conditions has provided additional
evidence of the fruitfulness of the extension
of both laboratory and field methodology to
specimens other than those generally used in
comparative behavior laboratories.
It is one of the purposes of this paper to
point to a source of data which can facilitate
not only the collection of more information
on a wider variety of animals but which can
also serve as a training facility for field-
workers and others. Within the reach of re-
searchers in most large cities there exist
collections of living animals in great variety
of species and under varied living conditions.
The reference is to the zoological parks and
exhibition areas. In many of these, natural
habitat conditions are approximated and
even the differences can be fertile sources of
comparative psycho-ecological studies.
1 The success of this project was due in large part t
the co-operation of the staff and keepers of the New Yor
Zoological Park. Particular thanks are due Mr. Fairfiel
Osborn Mr. John Tee-Van, Mr. Lee S. Crandall, Dr. L .
Goss, Keepers Reilley and Quinn, and Mr. Sara Duntoi
photographer.
2 Hunter College of the City of New York, N. Y. C.
3 Bucknell University, Lewisburg, Penna.
4 University of North Carolina, Chapel Hill N C
5 City College of New York, N, Y. C.
In 1937, Carpenter (2) published a study
of two young male mountain gorillas, Gorilla
gorilla berengei, resident at the San Diego
Zoo. In this paper he pointed to the oppor-
tunities presented by the fourteen specimens
of the largest of the great apes which he
listed as available in the zoological parks of
the world. Gorillas offer a dramatic field for
this type of research since they are, with the
exception of the orang-utan, the least studied
of the anthropoid apes. The present paper
seeks to compare Carpenter’s data with those
obtained in a study of male and female pre-
adolescent lowland gorillas, Gorilla gorilla
gorilla, in the New York Zoological Park
(Bronx Zoo). Such comparative data as can
be assembled will be helpful to workers who
seek a base line for similar investigation
elsewhere. That these investigations are pos-
sible is demonstrated in Table I which lists
the location and over-all biological indices of
the specimens now resident in various collec-
tions. The number of gorillas in the United
States is increasing. In 1937 there were only
eight specimens in this country; today there
are twenty-four.
Other than Carpenter’s study, material on
the behavior of the gorilla is found only in
a limited number of papers. Yerkes’ pioneer
work (6) with the pre-adolescent female
mountain gorilla, Congo, is known to all
students of comparative psychology. Bing-
ham’s 1932 observations (1) on gorillas in
their native habitat and Valker’s similar
study in the Gaboons (5) in 1931 are the only
other relevant research. The two lowland
gorillas in the New York Zoological Park are
included in a report of the external genitalia
published by Goss (3).
II. Subjects.
The gorillas studied at the Bronx Zoo are
Oka, female, and Makoko, male. Since this
study was made the Bronx Zoo has acquired
a young female mountain gorilla, but no
reference to it will be made in this report,
except to list it in Table I. Little is known
of the early history of Oka and Makoko. Both
animals arrived in the Zoo on September 7,
1941, at which time they weighed 20 and 28
112
Zoologica: New York Zoological Society
[34: 13
Text-fig. 1. Rate of growth of Oka and Makoko
from time of arrival at the New York Zoological
Park on September 7, 1941, until summer of
1948.
pounds respectively. Ages were estimated as
one and three years on the basis of weight
and bone structure. Text-figure 1 gives the
data on the growth of the specimens from
the time of arrival until the summer of 1948.
The female was weighed, up to June, 1948,
while on the keeper’s back, the male’s weight
being merely estimated. In 1948 a Toledo
balance was installed, on which the animals
were weighed when they voluntarily mounted
a platform.
At the present time, Oka and Makoko are
magnificent specimens. Makoko has a par-
ticularly brilliant coat and a notably promi-
nent supra-orbital ridge. Oka is less im-
pressively marked but is a splendid female
example of the species. Both animals are
active, healthy and strong. Indeed, Makoko’s
grip is so powerful that it has been necessary
to replace the %-inch steel bars of the cage
front, which he bent repeatedly. A steel hori-
zontal ladder had to be removed from his cage
when he tore it loose from its mooring. Oka
is less destructive and retains her ladder.
III. Environmental Conditions.
The gorillas are housed in individual cages
separated by a partition consisting of a solid
metal access door and a double grill of steel
bars. At the ends of each cage are doors
leading to shift cages into which the animals
are chased when the exhibition compart-
ments are cleaned or repaired. The back wall
of the living space is solid masonry with a
recessed access door. Between the glass par-
tition through which the public views the
animals and the metal barred cage fronts is
a passageway for the keepers. The internal
features of the living compartments include
a platform and a three-step staircase, with
the platforms raised two inches from the
floor, facing each other in front of the grill
between the two cages. The staircases are
at the ends near the shift cages. Oka’s cage
contains a metal horizontal ladder slung
between the rear wall and the cage front at a
height of three feet. Text-figure 2 shows the
floor plan of the gorilla exhibition space.
IV. Daily Schedule of Animal Care.
Although the animals are on exhibition
during the hours of 10:00 A.M. to 5:00 P.M.
(except on Sundays when the visiting time
is extended to 6:30 P.M.), the daily routine
is more extensive. The overhead fluorescent
lights are turned on at 8:00 A.M. when the
keepers enter the building. Between 8 :30 and
9:00 in the morning, the gorillas, each of
whom has been in its own cage all night, are
given their morning meal of skimmed milk
and raw eggs. The ingredients are mixed and
fed to each gorilla by tilting a can containing
the mixture into the subject’s mouth as the
animal protrudes its lips through the bars
of the cage front. Some half-hour to three-
quarters of an hour later, the keepers move
the gorillas to the shift cages. Separation
and enclosure in the individual cages are
effected by means of a stream of water. With
the gorillas out of the way, the cages undergo
thorough cleaning. Shortly before visiting
time, the animals are released and are fre-
quently permitted to remain together in one
of the two cages for periods up to one hour.
This opportunity for association occurs five
A
.
c •
, — r — '
B 21' 9" X 8'9" t
i
B'
_ , h-,
»n_ jstci
U“
*
A'
Text-fig. 2. Floor plan of gorilla compartments in New York Zoological Park. A, A',
passageways for keepers. B, B’, exhibition cages for Oka (female) and Makoko (male),
respectively. C, C', shift cages. D, three-step stairs. E, platforms; right-hand platform
used for weighing. Thin line indicates solid wall; dotted line indicates grill-front walls;
double line indicates glass partition.
1949J Riess, Ross, Lyerly & Birch: Behavior of Captive Lowland Gorillas 113
TABLE I.
Physical Characteristics of Gorillas in the United States.*
1
Name.
1
1
Location.
1
Form.
Sex.
Estimated
age on
September
1, 1949.
Known |
Weight. |
1
Estimated
Weight.
Oka
N. Y. Zool. Park |
gorilla
F
9 yrs.
282 lbs. |
Makoko |
N. Y. Zool. Park
gorilla
M
11 yrs.
408 lbs. |
Sumaili |
N. Y. Zool. Park
berengei
F
20 mos.
20 lbs. 1
Joanne |
Central Park Zoo, N. Y. C. |
gorilla
F
10 yrs.
| 190 lbs.
Carolyn |
Central Park Zoo, N. Y. C. |
gorilla
F
10 yrs.
j 190 lbs.
Bamboo |
Philadelphia Z. G.
gorilla
M
23 yrs.
| 435 lbs.
Massa
Philadelphia Z. G.
gorilla
M
18 yrs.
1 400 lbs.
Bushman |
Lincoln Park Zoo, Chicago |
gorilla
M
21% yrs.
542 lbs. |
Sinbad |
Lincoln Park Zoo, Chicago |
gorilla
M
20 mos.
38 lbs. |
Rajah
Lincoln Park Zoo, Chicago |
gorilla
M
29 mos. 1
1
47 lbs.
Irvin Young |
Lincoln Park Zoo, Chicago |
gorilla
M
33 mos.
52 lbs. |
Lotus
Lincoln Park Zoo, Chicago |
gorilla
F
43 mos.
1
75 lbs.
Miss Congof |
Chicago Z. P.
gorilla
F
16 yrs.
1
325 lbs.
Phil
St. Louis Z. P.
gorilla
M
10 yrs.
1
320 lbs.
Bobo
St. Louis Z. P.
gorilla
M
2 yrs.
1
1
44 lbs.
Big Boy |
Cincinnati Zoo
gorilla
M
3 yrs.
35% lbs. |
Albert |
San Diego Zoo
gorilla
M
6 mos.
9% lbs. |
Bouba |
San Diego Zoo
gorilla
F
10 mos.
12% lbs. |
Bata |
San Diego Zoo
gorilla
F
8 mos.
10% lbs. |
Phil |
Colorado Springs, Col. |
gorilla
M
4 yrs. |
|
40 lbs.
Gargantua |
Ringling Bros. Circus |
gorilla
M
17 yrs. |
550 lbst |
Mtoto
Ringling Bros. Circus
gorilla
F
1 6 yrs. |
438 lbs.f 1
* On October 31. 1949, three young specimens of Gorilla g. gorilla arrived in New York. They were a male and a
female weighing 12-14 pounds, and a female weighing about 40 pounds. At the time this paper went to press they
were still in the hands of Henry Trefflich, an animal dealer,
t Died Sept. 22, 1949.
t 1947 weight.
or six times a week and is the only occasion
on which the animals are in unrestricted
contact with each other. From the time of
separation until the main feeding of the day
at approximately 2:30 P.M., the gorillas are
unattended except insofar as the keepers
play with them while passing in front of the
cages. The typical afternoon meal consists
of carrots, celery, oranges, apples, grapes,
bananas, beets, beans, sweet-potatoes, cab-
bage, onions, cherries and other seasonable
fruits and vegetables. All food is fed raw
and supplied through the front bars of the
cages. Water, ad lib., is available to each
animal from a continuously running spigot
which empties into a trough running along
the back wall of each cage. At 5:30 P.M. the
lights are turned out and the gorillas remain
unattended until the next morning.
V. Observational Procedures.
Preliminary study of the gorillas was
started at the beginning of May, 1948. One
of the writers (B.F.R.) visited the Primate
House at the Zoo on two days of each week
and observed the gorillas for one hour. The
times selected were in the early morning
when the keepers placed the two gorillas in
one cage, and in the late afternoon, at about
3:30 P.M. following the afternoon meal. At
the later time, the animals were in separate
cages.
As a result of the work during May and
early June, a list was evolved on which both
quantitative and qualitative indices of be-
havior could be noted. The inventory con-
sisted of 42 items divided into five catego-
ries : posture and locomotion ; eating, drink-
ing and elimination ; self-oriented activity
(play?); inter-individual behavior; and
observer-oriented behavior. The sheets of the
check list were divided into 10 columns, each
of which was used for a three-minute period
of observation during the total 30 minute
duration of the observational session. Thus
it was possible to arrive at the total amount
of each of the 42 types of behavior during the
30-minute period and also to determine the
sequence of behavior during the period. Ad-
ditional space was provided for running com-
ments on the activity of the gorillas and for
additional notes.
The observers worked in pairs and rotated
the pairing so that a measure of control over
reliability of observation was possible. There
was joint discussion of the meaning of each
term on the check-list so that the observers
would agree on how to label the activity ob-
served. The problem of the animal’s reaction
to the observer was considered and it was
decided to standardize the position of the
observers. Since the gorillas are a very popu-
lar exhibit and attract large crowds of spec-
tators, the public exhibition space was used
as the location for the observers who sat in
pairs in front of the glass partition opposite
114
Zoologica: New York Zoological Society
[34: 13
the communicating grill between the two
cages.
Because the activity of the gorillas varied
considerably with the time of day, the num-
ber of visitors, the daily routine and other
variables, it was decided to distribute the
30-minute observation periods over the whole
working day of the animal. However, since
the gorillas were allowed to be in the same
cage with one another only in the early
morning, there was a greater concentration
of sessions between 9 :00 and 10 :00 A.M. The
total number of observations made during
these hours was 10 one-half hour periods.
Sixty-eight sessions were devoted to taking
notes on behavior of the animals in isolation.
During these 68 periods, it was possible to
get data on both Oka and Makoko so that
each animal was studied an equal number
of times. Every hour between 10:00 A.M.
and 6:00 P.M. was covered. In addition to
these systematic observations which started
in July and lasted through the middle of
August, 1948, each experimenter observed
the gorillas several hundred times while
passing through the Primate House to and
from other areas in the Park. Any deviant
behavior or peculiar activities were noted
and added to the record.
VI. Results and Discussion.
Two factors limit the analysis of the data.
In the first place, quantitative analysis af-
fords very little insight into the problem at
hand. The purpose of this study was to obtain
information which would serve as a starting
point for further investigation of the gorillas
at the Bronx Zoo, particularly as they become
physiologically more mature and show active
sex and social behavior. In the second place,
it was thought desirable to point to both the
similarities and differences between our
gorillas and those at the San Diego Zoo
described by Carpenter.
In this comparison, there are many diffi-
culties and dangers arising from two sources.
In the first place, the gorillas belong to dif-
ferent sub-groups, ours being lowland and
Carpenter’s mountain specimens. Other va-
riables in this category include age and sex
differences between the two sets of observed
gorillas. In the second place, it is necessary
to stress the differences in the environmental
setting in which the San Diego and Bronx
gorillas carried out their daily activity. At
San Diego, the two male gorillas were housed
in outside cages equipped with tree trunks
for climbing and various devices which could
be manipulated by the animals, for instance
logs, swings, ropes, tires, etc. In addition,
the experimenter could insert objects into the
gorillas’ surroundings and study the effect
of such introductions. At the Bronx Zoo, the
separation of the gorillas from the public
was much more rigorous and the cages much
more bare. In interpreting the comparative
findings, the obvious individual and environ-
mental differences must be kept in mind.
However, despite these limitations, the com-
parison of the two groups should be of value,
if only to emphasize the danger of generaliz-
ing from any set of observations.
With the restrictions specified above, the
data in Table II represent the basic observa-
tions made upon the two gorillas at the Bronx
Zoo. Where information comparable to that
obtained by us was derivable from Car-
penter’s San Diego observations, it has been
included in the Table. The discussion of the
data in Table II will follow the general cate-
gories outlined above.
A. Posture and. Locomotion.
Posture and general locomotion seem fair-
ly well established as invariant gorilla pat-
terns. Both Carpenter’s mountain and our
lowland gorillas exhibited the same type and
frequency of gross motor activity. Walking,
running, standing and sitting were charac-
teristically alike for Oka, Makoko and the
San Diego pair. Differences were noted in the
frequency of observed sleep and in swinging
by the hands. Both of these differences may
be the resultant of variable environmental
and observational procedures. It was not
feasible for us to observe night behavior,
and swinging was made difficult for Oka and
Makoko by the absence of a place suitable
for that kind of activity. The complete ab-
sence of nest-building in the Bronx pair is
also related to the lack of adequate materials.
Both Carpenter and Yerkes report that it
was a fairly common behavioral pattern in
their subjects.
The observations on handedness in the
Bronx gorillas are not comparable to other
studies since this item was not listed by Car-
penter. Oka was observed to make differen-
tial use of her hands on 313 occasions and
Makoko 184 times. In both animals, the right
hand was more frequently employed regard-
less of the nature of the activity. The fre-
quency of use of this hand was 54% as com-
pared with the report by Yerkes (6) who
found that Congo used her right hand some
66% of the time. In Congo’s case the left foot
was preferred to the hand, whereas in our
gorillas there was relatively little pedal
manipulation.
B. Eating, Drinking and Elimination.
In the presence of an abundance of food,
Oka and Makoko both showed a form of be-
havior somewhat akin to the hoarding of rats
and lower mammals. The gorillas would
sweep the food into a heap with either the
hands or feet. The heaped food was then ex-
amined, tossed around or eaten. This behav-
ior is not mentioned in any other report on
the gorilla and may well be the unique result
of the absence of manipulatible material in
the cages in the Bronx Zoo. Placing of the
longer-stalked fruits and vegetables on the
heads of the gorillas was a frequent after-
math of the in-gathering of the material.
1949]
115
Riess, Ross, Lyerly & Birch: Behavior of Captive Lowland Gorillas
An interesting aspect of the behavior of
the isolated animals was a relatively infre-
quent passage of food from Oka to Makoko
through the bai-s of the intercommunicating
grill between the two cages. The initiation
of this activity usually came from Oka and
was noted particularly on the several occa-
sions when Makoko had been deprived of his
usual rations because of diarrhea or other
health considerations. In such circumstances,
Oka was observed forcing potatoes and cab-
bage through the double grill. Makoko did
not seem to be particularly interested in the
inserted foods. This behavior was seen on
three occasions.
The drinking of water is a form of be-
havior common both to our animals and to
those studied by Carpenter. According to
Yerkes, Congo drank but little water. The
manner of drinking in the Bronx animals
was to bend over the fountain and suck the
water into the mouth. Although there was no
opportunity to study comparative satisfac-
tion from milk and water, it is the opinion
of the authors that the milk-egg liquid was
preferred.
Regurgitation of the milk-egg mixture was
almost invariable. Following the feeding the
animals would typically squat on their
haunches, lean forward, and regurgitate some
if not all of the milk. The gorillas then would
examine the liquid manually and eventually
bend all the way and lick up the regurgitated
material. The time interval after ingestion
varied somewhat but was in the neighbor-
hood of 45 seconds. Some chemical changes
took place during the brief digestive stay
for the milk was usually curdled. This pat-
tern is seen not only in the gorillas at the
Bronx Zoo but also in the chimpanzees.
Whether or not this is a primate characteris-
tic and analagous to the similar behavior of
the pre-socialized human infant or whether
it is unique to the subjects in captivity re-
mains a matter for further research.
Voided fecal material was commonly
handled both by Oka and Makoko and fre-
quently was used to throw at the keepers,
the observers or the public. Handling was
casual and seemingly tactually motivated, for
the feces were not examined nor used except
for throwing. Since the keepers made heroic
efforts to keep the cages clean, the oppor-
tunities for greater concentration on feces
were limited. Carpenter makes no mention
of this type of behavior. Urination was a
casual affair and no localization of territory
for this or for defecation was noted.
C. Self-oriented Activity.
Self-manipulation of parts of the body was
a common form of activity in both Oka and
Makoko. The parts of the body selected for
handling or fingering were not consistent,
with the exception of the lack of attention to
or focus on the external genitalia. The major
phase of activity during which handling was
observed was while the gorillas were lying
on their backs or stomachs, when parts of the
body such as the lips, ears, eyebrows and nose
would be held. The absence of genital ma-
nipulation may well be the result of the small
size of the external genitalia of the gorilla,
as described by Carpenter and Goss.
Manipulation of objects in the environ-
ment is a frequent finding wherever gorillas
have been studied. Carpenter and Yerkes
mention this behavior pattern and it was
noticeable in the animals at the Bronx Zoo.
The female, Oka, showed some tendency to
manipulate and examine with greater fre-
quency than her cage mate. In the absence
of a variety of objects to examine, the fre-
quency of occurrence of this activity is all
the more remarkable.
The attitude of the gorillas toward the
water fountain has already been described.
An observable difference was noted in the
behavior toward drinking water and that
emanating from the pressure hoses used in
cage cleaning. As indicated above, in the dis-
cussion of daily routine, water was used to
separate the gorillas and to urge them toward
the shift cages. The initial reaction to the
stream from the hoses was retreat and ex-
citement. However, once wet, the animals
would face into the water and jump up and
down. The keepers reported that the animals
would on occasion approach more closely to
the nozzle of the hose when thoroughly wet.
No shaking of the body after the bath was
seen during the periods of observation.
Self-grooming does not seem to be a domi-
nant activity in the lives of either the San
Diego or New York gorillas. To what extent
the absence of this form of self-manipulation
is a function of the cleanliness of the en-
vironment and the animal is not established
by our observations but, as will be noted
later, grooming as a pattern of behavior
is markedly less present in the gorillas under
study than in other primates at the Zoo.
D. Inter-individual Behavior.
The data in this section were obtained
during those periods when the animals were
together in the same cage. To the extent that
the opportunity for such interaction was
limited, the enhancement of activity during
the periods of joint occupancy of the cage
may be a function of the limitation of time
during which the two animals could interact.
Both in Carpenter’s study and in ours, the
major forms of inter-individual activity were
running, chasing and wrestling. These be-
havior patterns were well marked and almost
stereotypical in appearance. Chasing was
especially vigorous when the cage floor was
wet and the gorillas spent much time sliding
in a pronograde posture from one end of the
cage to the other. Initiation of this activity
was fairly evenly divided between Oka and
Makoko. Wrestling, too, was not started con-
sistently by either male or female. A domi-
nance pattern was not apparent.
The sequence of individual motor acts in
116
Zoologica: Neiv York Zoological Society
[34: 13
TABLE II.
Comparative Behavior of Oka, Makoko and San Diego Gorillas *
Behavioral Classification
A. Posture and Locomotion
Walking, pronograde
Walking, upright
Running, pronograde
Running, upright
Standing, pronograde
Standing, upright
Sitting
Climbing
Swinging by hands
Hanging
Sliding
Left-handedness
Right-handedness
Lying down
Sleeping
Nest Building
B. Eating, Drinking and Elimination
Gathering food in heaps
Sharing food with cage mate
Drinking water
Drinking milk
Regurgitation of milk
Handling of feces
Attention to urination
C. Self-oriented activity (Play?)
Self-manipulation (non-genital)
Manipulation of genitalia
Manipulation of objects
Manipulation of food (non-eating)
Attitude toward stream of water
Self-grooming
D. Inter-individual behavior
Chasing
Wrestling
Grooming
Inspection and manipulation of genitalia
Presenting
Mounting
Pelvic thrusts
Chest thumping
Vocalizing
Dominance
E. Observer-oriented behavior
Throwing of feces
Throwing of non-fecal material
Attentional responses
Vocalization
* Key to symbols :
O— Never observed or reported
H — Little in frequency or amount
H — | — Some or fairly frequent
Frequency of Occurrence in
San Diego
Oka
Makoko
+ + +
+ + +
+ + +
4"
+
+ +
+ + +
+ + +
+ + +
ND
4"
4"
+ + +
+ + +
+ + +
+
4-
4"
+ + +
+++
+ + +
+ +
+
+
+ +
+
+
+
4~
4-
+ + +
+ +
+ +
ND
46%
46%
ND
54%
54%
+ + +
+ + +
+ + +
+ +
+
4"
+
NP
NP
ND
+ +
+
0
+
4*
+ + +
+ +
+++
+ + +
+ +
+ +
ND
+ +
++
ND
+
+
ND
+
+
+ +
+ +
+ +
0
0
0
+ + +
+ +
+
ND
+ +
+ +
positive + +
+
+
+ +
+
+
+ + +
+ + +
+ + +
+ + +
+ + +
+ + +
+ +
■+■
+
0
0
4-
0
0
0
0
0
0
0
0
+
+ +
+
+ +
+ +
ND
ND
+
+
+
ND
+
+.+
ND
4-
+
positive ++
negative -f+
negative +-
+ +
1
ND
ND
+ -j — | — Great deal, very frequent
ND— No data reported
NP— Not possible in the environment
the wrestling behavior was free of pattern-
ing and seemed to consist of random grap-
pling at the anatomical point nearest to the
initiator of the behavior. When the actions
of either animal seemed to approach the point
at which roughness would ensue, the animal
at the moment on the receiving end would
detach itself and a period of resting would
follow. Of all the behavior noted in this
study, wrestling was the most dramatic and
illustrative of the great strength of the
gorillas. As a rule, there were few vocaliza-
tions during the bouts.
Social grooming, certainly, does not seem
to be as predominant in the behavior of the
gorillas as in the case of other representa-
tives of the great apes or other primates.
Carpenter likewise observed little of this sup-
1949]
Riess, Ross, Lyerly & Birch: Behavior of Captive Lowland Gorillas
117
posedly socially oriented activity. It is pos-
sible that in the four animals for which data
are available, the age and sex differences
were not sufficiently well established to facili-
tate the appearance of this type of social
interaction. It may also be possible that
there is a real species difference in such be-
havior.
Genital manipulation and exploration in
the paired situation was not frequently ob-
served. It was seen only three times in Oka
and Makoko and was not reported for the
San Diego pair. In the Bronx specimens, the
initiator was always the male. This may be
a reflection of the relatively greater maturity
of Makoko. So, too, in our gorillas, mounting
and pelvic thrusts as precursors of mating
behavior were almost completely absent dur-
ing the periods of observation in New York.
Carpenter saw none of this at San Diego, but
his animals were both males. The one in-
stance of pelvic thrusts by Makoko occurred
during a wrestling bout and was not repeated
nor invited by Oka.
Chest-thumping was much more prevalent
during the periods of paired activity than
when the animals were in their own cages.
The causal sequences leading up to the
thumping could not be determined for Oka
and Makoko although the observers were all
of the opinion that the behavior was socially
oriented and significant. Some writers have
suggested that thumping of the chest is a
sign of well-being and general euphoria.
Yerkes states that the behavior is a sign of
“impatience or other mild dissatisfaction.”
There was clear evidence of neither causal
sequence in our observations. The range of
situations during which the thumping was
observed varied so widely that no specific
factor can be assigned as the reason for its
existence. The only statement that can be
made from our data is that the male, Makoko,
engaged in chest-thumping more frequently
than did Oka and accompanied the beating
with vocalizations more frequently than his
companion.
In the absence of sound-recording devices,
description of vocalization is difficult. Fur-
thermore, the public space was somewhat
soundshielded by the glass partition sepa-
rating the animals from the observers. In
our experience the occurrence of this activity
was less than that mentioned by Carpenter.
E. Observer-oriented Behavior.
The types of audience-attentive behavior
observed in the Bronx Zoo consisted mainly
of the throwing of feces or food at the glass
plate between the cage and the visitors. The
same aggressive behavior was noted during
an attempt to get photographs of the ani-
mals. Visitors invariably tried to attract
attention from the gorillas by tapping on the
glass partition and by yelling. The effect of
such devices was negligible. This difference
between our data and Carpenter’s may well
be the result of the more restrictive condi-
tions of the gorillas’ environment in the New
York Zoo.
VII. Summary.
This report describes the behavior of two
pre-adolescent lowland gorillas in the New
York Zoological Park during the summer of
1948. Oka, then an eight-year-old female, and
Makoko, a ten-year-old male, were observed
in their regular living cages when alone and
when placed together.
A check-list was prepared and regular
half-hour periods of observation were sys-
tematically made. Significant behavioral
items were compared for the sessions when
the animals were alone and when they were
together. Comparisons were also made with
the data collected by Carpenter from two
male mountain gorillas in the San Diego
Zoo.
The descriptive material obtained during
the period of observation should serve as a
base line from which to note variations aris-
ing from the maturation of the two gorillas
in the years to come.
References.
1. Bingham, H. C. 1932. Gorillas in a native
habitat. Carnegie Inst. Wash. Publ., No. 426.
Pp. 66.
2. Carpenter, C. R. 1937. An observational
study of two captive mountain gorillas
(Gorilla beringei). Human Biol., 9, 175-196.
3. Goss, Leonard J. 1947. The external geni-
talia of the gorilla, Gorilla gorilla gorilla
(Savage & Wyman). Zoologica, 32, 97-99.
4. Leister, Claude W. Gorillas. New York
Zoological Society, Popular Series No. 4.
5. Valker, A. 1931. La vie du gorille au Gabon.
Bull. Mus. Nat. d’Histoire Nat., 3.
6. Yerkes, R. M. The mind of a gorilla, (a)
Genet. Psychol. Monogr. 1927, 2. 1-193. (b)
Genet. Psychol. Monogr., 1927, 2, 375-551.
(c) Comp. Psychol. Monogr., 1928, 5, 1-92.
118
Zoologica: New York Zoological Society
[34: 13: 1949]
EXPLANATION OF THE PLATES.
Plate I.
Fig. 1. Makoko, the male lowland gorilla in
the New York Zoological Park. Esti-
mated age, 11 years; weight, 408
pounds.
Plate II.
Fig. 2. Oka, the female lowland gorilla in the
New York Zoological Park, is still
friendly and gentle with her keeper at
the estimated age of 9 years. Her
weight is 282 pounds.
Fig. 3. Oka playing with her keeper.
ilESS. ROSS. LYERLY & BIRCH
PLATE I
FIG. 1.
THE BEHAVIOR OF TWO CAPTIVE SPECIMENS OF THE LOWLAND GORILLA,
GORILLA GORILLA GORILLA (SAVAGE & WYMAN).
| ESS. ROSS. LYERLY & BIRCH.
PLATE II.
FIG. 2.
FIG. 3.
THE BEHAVIOR OF TWO CAPTIVE SPECIMENS OF THE LOWLAND GORILLA.
GORILLA GORILLA GORILLA (SAVAGE & WYMAN).
Beebe : Migration of Papilionidae at Rancho Grande
119
14.
Migration of Papilionidae at Rancho Grande, North-central Venezuela.1
William Beebe.
Director, Department of Tropical Research, New York Zoological Society.
(Plate I; Text-figure 1).
[This is one of a series of papers resulting
from the 45th, 46th and 47th Expeditions of the
Department of Tropical Research of the New
York Zoological Society, made during 1945, 1946
and 1948, under the direction of Dr. William
Beebe, with headquarters at Rancho Grande in
the National Park of Aragua, Venezuela. The
expeditions were made possible through the
generous cooperation of the National Govern-
ment of Venezuela and of the Creole Petroleum
Corporation.
[The characteristics of the research area are
in brief as follows; Rancho Grande is located
in north-central Venezuela (10° 21' N. Lat., 67°
41' W. Long.), 80 kilometers west of Caracas,
at an elevation of 1,100 meters in the undis-
turbed montane rain forest which covers this
part of the Caribbean range of the Andes. The
migration flyway of Portachuelo Pass, which is
also the water-shed between the Caribbean and
Lake Valencia, is 200 meters from Rancho
Grande. Adjacent ecological zones include sea-
sonal forest, savanna, thorn woodland, cactus
scrub, the fresh-water lake of Valencia and
various marine littoral zones. The Rancho
Grande area is generally subtropical, being uni-
formly cool and damp throughout the year be-
cause of the prevalence of the mountain cloud
cap. The dry season extends from January into
April. The average humidity during the expe-
ditions, including parts of both wet and dry
seasons, was 92.4%; the average temperature
during the same period was 18° C; the average
annual rainfall over a five-year period was
174 cm. The flora is marked by an abundance
of mosses, ferns and epiphytes of many kinds,
as well as a few gigantic trees. For further de-
tails see Beebe and Crane, Zoologica, Vol. 32,
No. 5, 1947. Unless otherwise stated, the speci-
mens discussed in the present paper were taken
in the montane cloud forest zone, within a
radius of one kilometer of Rancho Grande.]
For an account of Portachuelo Pass, to-
gether with a general introduction to the
groups of migrating insects and migrating
factors see “Insect Migration at Rancho
Grande,” by William Beebe, Zoologica, 1949,
Vol. 34, No. 12, pp. 107-110.
In Volume 26 of Novitates Zoologicae,
W. J. Kaye has a paper entitled “A Geogra-
phical Table to show the Distribution of the
American Papilios.” Under the heading
“Venezuela, North,” (pp. 352-355), the au-
thor lists thirty-one species. In a letter Dr.
Rene Lichy of Caracas sends me a list of
thirty-one species of this family which he
1 Contribution No. 852, Department of Tropical Re-
search, New York Zoological Society.
has collected in northern Venezuela. A con-
tinuation of this coincidence is that each list
contains seven species not found in the other
list.
Both lists contain all the species which we
took migrating through Portachuelo Pass,
with the single exception of crassus which
Lichy does not mention.
In the limited width of twenty meters of
Portachuelo Pass, and allowing a height of
net reach of a maximum of five meters, we
captured seventeen species of Papilio. This
area may be considered, not unrealistically
nor unconservatively, as, at the most, a mil-
lionth of the extent of north Venezuela. Yet
within this relatively microscopic bit of
Andean air, we secured almost half the pa-
pilios so far recorded from the entire north-
ern part of the country. Thus, in the con-
sideration of this family of butterflies, we
are made to realize the wide-spread, impel-
ling, migrational force affecting this group
of insect life.
So much of this migration — its causes and
extent — is at present unknown, that every
verifiable fact is of value. Reviewing the
known distribution of the seventeen species
of Papilio migrants, we find that most of
them extend from Mexico to Paraguay, south
Brazil or Argentina. The distribution of the
subspecies, however, presents a very differ-
ent picture, and a very significant one in its
over-all pattern. In twelve out of the seven-
teen, the subspecific range is confined to
Colombia and Venezuela, with a few exten-
sions to adjacent territory. Thus we may
expect to find the northern point of origin
of the movement of these forms a relatively
short distance away.
The twelve subspecies of Papilio with lim-
ited distribution are as follows :
anchises osyris
anchisiades anchisiades
agesilaus agesilaus
areas areas
cleotas coroebus
erithalion zeuxis
lycophron hippomedon
paeon thrason
polyxenes americus
protesilaus archesilaus
sesostris tarquinius
torquatus orchamus
120
Zoologica: New York Zoological Society
[34: 14
The remaining five forms of Papilio with
wider distribution are :
belus varus
crassus
phaon
polydamus polydamus
thoas neacles
Let us take as an example Papilio agesilaus
agesilaus or, as I called it before identifica-
tion, Small Zebra Swallowtail. I recorded
seven individuals captured and fifty-eight
seen with certainty. Added to this number
were the many papilios glimpsed too briefly,
flying too high or too fast to be recognized as
to species, and also those which must have
passed during the hours of our absence from
the pass. It was heartbreaking to realize
what a minute fraction we could honestly
record by sight identification, yet there is
no other way, at present known, to glean
definite, general knowledge of this phase of
the lives of these splendid insects. Without
exception, all sight-named species were sub-
sequently confirmed or discarded by compari-
son with captured, definitely classified speci-
mens.
In this as in some other species, the small
number of records from the year 1946 and
their entire absence from 1945 by no means
indicates the absence of the species on migra-
tion, but only reflects our non-recognition of
the importance of the Portachuelo Pass
migration during the early years, and con-
sequent slight attention paid to this phenom-
enon. An ultimate summary of the relatively
few, disconnected observations made during
1945 and 1946 reveals a general movement
on a scale equal in magnitude and as all-em-
bracing of insect orders as we recorded dur-
ing 1948.
The mere recording of the capture of
seventeen species of Papilio on migration is
a worth-while fact, and when more and more
individuals are taken on suceeding days and
weeks the phenomenon is enhanced in in-
terest. In few or in large numbers the insects
continue to fly past, slowly or circling or
alighting out of reach. As in the case of
many other organisms, the time has come
when sight records must be used to supple-
ment specimens in net, envelope and cabinet.
Ornithologists in general and British ento-
mologists in particular have gone far in sight
identifications, while at the same time main-
taining as perfect accuracy as possible with
man’s fallible eyesight and only too human
brain.
At the end of our many months of collect-
ing and observation at the pass, I found, in
my Journal, a significant assemblage of
shorthand names of papilios. They were, of
course, essentially personal, stimulated by
mental comparisons with swallowtails fami-
liar to me elsewhere, or in the case of strange
tropical forms, by outstanding wing shapes,
size, patterns and colors.
Lepidopterists recognize three “natural
groups” into which papilios may be divided.
Text-fig. 1. Map showing location of Rancho
Grande, Portachuelo Pass and surrounding ter-
ritory. ,
These are based on various factors such as
larvae, pupae, microscopical imaginal dis-
tinctions and/or food-plants. Our migrant
species fit into these three groups as follows :
Aristolochia : sesostris, erithalion, an-
chises, areas, polydamus , belus and crassus.
Fluted: polyxenes, thoas, paeon, lycophron,
anchisiades, torquatus and cleotas. Kite:
phaon, agesilaus and protesilaus.
Consideration of this arrangement shows
no logical, technical or scientific agreement.
This is only to be expected in sight identifi-
cation, which can take no account of sexual
relationships, or parallelisms, or the superfi-
cial resemblance brought about by mimicry.
After final identification of the seventeen
species of Papilio migrants, I arranged
mounted specimens of all in a large insect
drawer, placed this upright on a chair in
good light and studied them from a distance
of ten meters. From this distance I made
the following key :
A — White and Black
a — Small: agesilaus agesilaus
b — Large : protesilaus archesilaus
B — Yellow and Black
a — Yellow-banded
Broad-band-plus-spot, small: tor-
quatus, orchamus, male
Broad-band, medium : lycophron
hippomedon, male
Narrow-band : paeon thrason
thoas neacles
1949]
Beebe: Migration of Papilionidae at Rancho Grande
121
b — Y ellow-spotted
Small: polyxenes americus
Medium: polydamus poly damus
Large: cleotas coroebus
C — Red and Black (Hindwing)
a — Cream-spot-forewing
sesostris tarquinius, female
erithalion zeuxis, female
anchises osyris, female
areas areas, female
torquatus orchamus, female
b — Green-spot-forewing
erithalion zeuxis, male
anchises osyris, male
areas areas, male
c — Black-forewing
anchisiades anchisiades
D — Green and Black
a — Green-forewing
sesostris tarquinius, male
b — Green-hindwing
phaon
E — Black (Dominantly)
belus varus
crassus
lycophron hippomedon, female
I compared this key with the names made
up on the spot in the field, and found a grati-
fying agreement in species recognition. The
differences were chiefly substitution for pat-
terns and colors of the names of northern
species suggesting resemblances, species
with which I had long been familiar in the
eastern United States. For example, age-
silaus agesilaus was “small ajax or zebra,”
polyxenes americus was “small asterias,”
and thoas neacles was “cresphontes-like,” etc.
I cite all this as in no way directly possible
or in the same detail applicable for use by
another observer, but merely to show a
framework upon which can successfully be
erected an observer’s sight key. Further
comments, in greater or less detail, will be
found under the treatment of many of the
species.
My special thanks go to Mr. Henry Flem-
ing, entomologist of our Department of Trop-
ical Research, for many direct observations
and for frequent corroboration of my own.
In addition I am beholden to him for looking
up distribution data and for painstaking
identification of all the species.
Papilio anchises osyris Felder.
Species Range: Colombia to Brazil, Bolivia
and Paraguay.
Subspecies Range : Venezuela.
Field Characters: Both male and female
indistinguishable in the field from erithalion
zeuxis. Therefore all specimens observed
and not taken are combined under the two
species. Compared with areas areas the male
lacks bright green forewing spot, and the
female has a decidedly larger, 4-celled fore-
wing cream spot.
Number and Sex: Both sexes taken; eight
males, five females.
Date: April 13 to July 29.
Condition: All taken were fresh.
Record of Captures: 1945 — July 15
(male), 18 (female). 1946 — April 13 (male,
km. 20). 1948 — April 29 (male); May 1
(female), 31 (female); June 6 (male and
female) ; July 17 (male, km. 30), 23 (male,
km. 15), 29 (female at Pass, 2 males, km.
35).
Combined Sight Records: anchises and
erithalion : (Total 62). 1946 — May 27 (4
females) ; June 29 (2 females). 1948 — May
26 (4 females) ; June 4 (11 females passed
in 10 minutes), 15 (4 females resting on
shrubs), 22 (5 seen) ; July 2 (12 females),
9 (14 males, 3 females), 29 (3 females).
Papilio anchisiades anchisiades Esper.
Field Name: Red-spot Black.
Species Range: Mexico to southern Brazil.
Subspecies Range: Colombia to Bolivia
and Para (Brazil).
Field Characters : Black with red on hind-
wing. Closest in field appearance to wholly
black lycophron hippomedon.
Number: 2 specimens taken, a male and a
female.
Notes: This black-forewing-red-hindwing
papilio came through the Pass with what
were taken and identified as female areas
areas and anchises osyris, all captured to-
gether.
Record of Captures : A male in frightfully
worn condition collected on April 13, 1945,
No. 45456. A second individual, a female,
taken at the Pass May 1, 1948, No. 48474.
Papilio agesilaus agesilaus Guerin.
Field Name: Small Black-and-white Zebra
Swallowtail.
Species Range: Mexico to Paraguay.
Subspecies Range: Magdalena Valley, Co-
lombia, to northern Venezuela.
Field Characters: Unmistakable resem-
blance to our northern ajax or zebra. The
only optically related species is protesilaus
arehesilaus, but the present species is much
smaller (forewing 43 mm. as compared with
53 mm.). The difference easily recognizable
when either species is close at hand or near
other butterflies.
Number: Total recorded 58. Seven taken
(48543, 48731, 481344, 481494).
Sex : Both sexes taken.
Date: From April 29 to July 26.
Frequency: Recorded on seventeen days.
Moderately but markedly gregarious. Two-
thirds of agesilaus were recorded in groups
of 4 to 8, while one-third appeared singly or
in twos.
Flight: Except when alarmed or fighting
against a strong head wind, the flight was
unhurried and wavering, always steadily
south.
Condition: With one exception all that I
saw hovering or resting at the Pass were in
122
Zoologica : New York Zoological Society
[34: 14
good condition, with both long slender tails
visible to the naked eye or through three-
power binoculars. No. 481344, taken July
26, 1948, at kilometer 16, was badly muti-
lated, with one tail missing. Dissection re-
vealed that this male had already mated.
Record: 1946 — May 26 (2 seen), 27 (2 at
Pass, 3 at km. 20), 30 (6 singly). 1948 —
April 29 (2) ; May 14 (1 taken), 21 (4 seen,
1 taken), 23 (1 taken), 24 (4), 26 (4), 31
(8) ; June 6 (1), 17 (2 taken), 19 (2) ; July
13 (4), 16 (2), 23 (1 taken km. 16), 24 (6),
26 (1 at Pass, 1 at km. 16) .
Papilio areas areas Cramer.
Field Name: Green-spot (male). Two-
celled-cream-spot (female).
Species Range: Mexico to Colombia, Vene-
zuela and the Guianas.
Subspecies Range: Venezuela and the
Guianas.
Field Characters : Closest to anchises and
erithalion, from which it differs in the bright
green forewing spot in the male, and the
smallex*, two-cell forewing spot in the female.
It is also close to the exceedingly rare tor-
quatus orchamus. From the male sesostris
tarquinius this species differs in the red on
the hindwing.
Number: Total recorded 277. Twenty-one
taken.
Sex: Both sexes taken. In 1946 only fe-
males were seen or taken. In 1948 females
were dominant from May 1 to June 17, and
males from July 6 to July 22.
Date: May 1 to July 22.
Frequency: Recorded on 20 days: 1 (15
times), 2 (4 times), 4 (once), 6 (twice), 7,
13, 14, 16, 18, 29 and 85. Decidedly gregari-
ous, occurring singly and in twos nineteen
times, comprising one-fourteenth of the total
number passing in larger numbers — from
4 to 85. As mentioned above, the sexes showed
a decided segregation. The flocks were us-
ually compact waves.
Flight: Rather low and fluttering.
Condition: Most of areas observed were
in fresh condition, decidedly unworn.
Extent of Migration: On four separate
days specimens of this species were taken
both at the Pass and at kilometer 20, well to
the south, and at kilometer 35, half way to
the coast to the north.
Record: 1945 — July 3 (3 taken), 16 (2
seen, 1 taken, Limon). 1946 — May 28 (27
seen, 2 taken), 29 (7 at Pass, 6 at km. 20),
June 4 (female), 22 (female) ; September 7
(84 in half an hour, 1 taken). 1948 — May 1
(female), 4 (6 females), 5 (male), 28 (fe-
male) 29 (female) ; June 6 (2 males, flock
of 18 females), 17 (16 females flying low),
22 (male and female), 28 (28 females) ;
July 6 (2 males), 9 (male caught and eaten
by bat falcon, 21 males seen), 14 (7 males),
15 (11 males, 3 taken), 16 (13 seen, four
fighting in midair), 19 (4 males, 1 taken
km. 15) , 22 (2 males, 1 taken km. 35) .
Papilio belus varus Kollar.
Species Range: Mexico to Bolivia and
Para (Brazil).
Subspecies Range : Guatemala to northern
Venezuela and Ecuador.
Dichromatic Female, form latinus Felder.
Field Name: Greenish-hindwing-band
Black.
Field Characters: Black, with a curved
band of large, yellow-green spots on hind
wing.
Number: Total recorded 19. Eight taken.
Sex: Females only taken.
Date : May 29 to August 4.
Frequency: Taken singly. Five once seen
together.
Condition: Fresh.
Record: 1946 — July 7 (1 seen km. 20).
1948 — May 29 (1 seen) ; July 4 (4 seen at
8:30 A.M.), 14 (1 taken), 16 (3 taken), 21
(2 taken, 3 seen), 26 (1 taken, 2 seen km.
16) ; August 4 (1 taken).
Dichromatic Female, form varus Kollar.
Field Name: Cream-spot-forewing Black.
Field Characters : Irregular splash of yel-
low in forewing; hindwing blue-black.
Number: Total recorded 18. Three taken.
Sex: Females only taken.
Date: May 30 to July 26.
Frequency: On two occasions, five were
seen together.
Condition : Fresh.
Record: 1948 — April 29 (1 taken); May
30 (5 seen); July 8 (3 seen), 9 (2 seen
alighted, 2 taken, 5 seen km. 31).
Papilio eleotas coroebus Felder, form dione
Rothschild and Jordan.
Field Name: Large Asterias Swallowtail.
Species Range: Costa Rica to Brazil.
Subspecies Range: North Colombia and
Venezuela.
Field Characters : Rather like a very large
Asterias, or polyxenes americus, with fore-
wing 67 mm. as compared with 40 mm. A
very distinct species.
Number: Total recorded 19. One taken.
Sex: The single specimen taken was a
female.
Date: May 26 to July 17.
Record: 1948 — May 10 (4 seen), 26 (2
seen) ; June 6 (female taken, 2 others flying
in company with three of the small polyxenes
americus ) ; July 10 (2 at Pass, 2 at km.15),
17 (3 at Pass, 3 at km. 30).
Papilio crassus Cramer, male foi’m lepidus
Felder.
Field Name: Black Philenor.
Species Range : Costa Rica south to Brazil.
Field Characters : Wholly black except for
concealed bluish-white anterior border of
hindwing.
Number: Total seen 23. One taken.
Record: A single male specimen of the
form lepidus taken on July 21, 1948, No.
481538. Twenty more, distinctly seen, passed
1949]
Beebe : M igration of Papilionidae at Rancho Grande
123
at the same time, all out of reach. On the fol-
lowing day, July 22, two more of these black
papilios were seen. No other record.
Papilio erithalion zeuxis Lucas.
Species Range: Costa Rica to Colombia
and northern Venezuela.
Subspecies Range: North Venezuela and
Colombia.
Field Characters: Both sexes indistin-
guishable from anchises osyris. Differs from
areas areas in male lacking forewing green
spot, and female with larger, 4-celled fore-
wing cream spot.
Number: Total number taken 14.
Sex: Both sexes taken.
Date: June 29 to July 24.
Condition : All freshly emerged.
Record of Captures: 1945 — July 3 (female
taken, Limon). 1946 — June 29 (female, km.
20) ; July 7, 8 and 10 (Each day 1 female
taken, km. 20). 1948 — July 9 (3 males, km.
31), 14 (male), 15 (male), 17 (2 females),
23 (female, km. 15), 24 (female).
For joint sight identification records with
anchises osyris, see under latter species.
Papilio lycophron hippomedon Felder.
Field Name: Male, Broad-band Medium
Turnus. Female, Black Troilus-like Swallow-
tail.
Species Range : Mexico south to Argentina
and Uruguay.
Subspecies Range : Colombia and northern
Venezuela.
Field Characters: Male can be confused
only with the very rare, smaller, yellow-spot
torquatns orchamus; female recalling a mel-
anistic troilus or phaon with black hind-
wings.
Number: Total recorded 20. Eight taken.
Sex: Both sexes taken.
Date: May 10 to July 20.
Record: 1946 — May 27 (2 at Pass, 2 km.
20, all males) . 1948 — May 10 (3 males seen) ,
21 (female taken), 23 (male taken), 24
(male taken), June 6 (2 females taken), 17
(2 males taken), 29 (4 males seen); July
10 (male seen), 20 (male taken).
Papilio paeon thrason Felder.
Field Name: Rare Cresphontes-like Swal-
lowtail.
Species Range: Mexico south to Argen-
tina and Uruguay.
Subspecies Range: North Colombia and
Venezuela.
Field Characters: Indistinguishable in the
field from thoas neacles, but as only a single
specimen of paeon thrason was taken, com-
pared with more than one hundred of thoas,
I am assuming that all Cresphontes-like
papilios observed were of the more abundant
species.
Record of Capture: A single male taken
on May 23, 1948, No. 481539, in extremely
torn and worn condition. It was captured
at the Pass at 12:30 P.M., the day being
warm and sunny with a Force 4 wind from
the south.
Papilio phaon Boisduval, aberration
metaphaon Butler.
Field Name: Philenor-like Swallowtail.
Species Range: Mexico to Ecuador and
Venezuela.
Field Characters: This is the only black
papilio with green on the hind wings.
Number: Total recorded 254. Seven taken.
Sex: Both sexes taken.
Date: April 13 to July 21.
Record: 1945 — May 24 (1 taken). 1946 —
(September 9, numbers of these black pa-
pilios with large green spot on the hind
wings were flying too high to catch. Several
alighted and allowed detailed study with
Number three glasses. Counted 228 and
missed many more.) 1948 — April 13 (male
taken km. 20), 16 (male taken), 27 (male
taken, km. 20), 29 (female taken at Pass) ;
July 21 (2 taken, 19 seen).
Papilio polydamus polydamus Linnaeus.
Field Name: Medium Asterias Swallow-
tail.
Species Range : South Atlantic states,
West Indies and south to Argentina.
Subspecies Range: Georgia south to
Buenos Aires.
Field Characters: Differs to the eye from
polyxenes americus in the field by the single
instead of double line of yellow spots across
all wings. Another distinction is the larger
size.
Number: Total recorded 177. Nine taken.
Sex : Both sexes taken.
Date: May 15 to July 26.
Frequency: Decidedly gregarious. One-
eighth passed singly or in a scattering up
to five individuals. Seven-eighths were ob-
served in flocks of ten to forty-eight.
Record: 1946 — May 27 (2 seen). 1948 —
May 15 (5 seen), 29 (female taken) ; June 6
(female taken), 22 (11), 30 (female taken,
48 passing, 2 seen at km. 21) . July 9 (female
taken), 10 (2 males taken, 16 seen. 2 taken
km. 31), 11 (14 flurry, 1 single), 14 (1 taken,
23 seen), 17 (10 seen km. 30), 18 (3), 19
(3), 22 (3 km. 35), 26 (27 seen).
Papilio polyxenes americus Kollar, form
melasina Rothschild and Jordan.
Field Name: Small Asterias Swallowtail.
Species Range: Canada south to Cuba and
Peru.
Subspecies Range: Colombia, Venezuela
and northern Peru.
Field Characters : Under a new name this
proved to be the same species as our northern
Asterias. The only other resembling migrant
butterfly was the markedly larger polydamus
polydamus.
Number: Total observed 34. Although ob-
served on ten occasions during two seasons,
only three specimens were taken.
124
Zoologica: New York Zoological Society
[34: 14
Date: March 25 to July 20.
Frequency: 1, 1, 1, 2, 2, 2, 3, 3, 3, 6, 6.
Note: Three perfect specimens on June 5,
after rain, clung to the extreme ends of large
leaves. The wings were flat and expanded,
with the fore edge straight across so that the
transverse band and spots were continuous.
Record: 1946 — March 25 (1 taken, km.
21) ; April 19 (1 taken, km. 21) ; May 28
(2 seen at Pass). 1948 — April 29 (3 seen) ;
May 4 (6 flying together), 28 (2 seen);
June 5 (3 seen), 6 (3 seen) ; July 19 (6 seen),
20 (1 taken at Pass) , 22 (6 seen at km. 35) .
Papilio protesilaus archesilaus Felder.
Field Name: Large Zebra Swallowtail.
Species Range: Mexico to Paraguay.
Subspecies Range : Colombia, northern
Venezuela and western Ecuador.
Field Characters: Larger than (forewing
53 mm. as compared with 43 mm.) but in
general similar to agesilaus. Size difference
quite apparent when near, but not when
flying high, away from other known butter-
flies. At least fifteen individuals were not
counted because of uncertain sight identifi-
cation.
Number: Total recorded 42. Three taken
(48543).
Sex: Males only were taken. The female
seems to be quite unknown.
Date: Recorded on migration from April
29 to July 19.
Frequency: The relative gregariousness
corresponds to that in agesilaus. More than
five-sixths were in 4 to 12 groups, and six
only seen as solitary or dual migrants.
Condition : All observed in detail appeared
fresh and perfect.
Additional Notes: The flurry of 12 large
zebras on May 30, were in a compact body,
and at a time when neblina and rain, while
light, were continuous enough to discourage
all other migrants. Yet these great swallow-
tails flew steadily at a height of about 12
feet, up to and through the Pass and down
into the fog on the south slope.
The actions of six which passed on May
26 were typical. All flew slowly and with
slightly wavering flight at 10 feet, until I
swooped futilely at them with the net when
all swerved sharply out and down, two pene-
trating the underbrush and working their
way separately through the Pass before ris-
ing into the free air again. Three were fol-
lowed with the glasses far down the south
slope.
Record: 1946 — May 27 (1 seen, 1 at km.
20), 29 (4 seen, 1 taken). 1948 — April 29
(4) ; May 1 (3), 4 (1 taken), 10 (1), 21 (5
seen, 1 taken), 26 (6), 30 (12) ; July 19 (1
at Pass, 1 at km. 15).
Papilio sesostrh tarqulnius Boisduval.
Field Name: Male, Green-spot Black.
Species Range : Mexico to Bolivia and cen-
tral Brazil.
Subspecies Range: Panama, Ecuador,
northern Venezuela.
Field Characters: Male to be confused
only with male areas areas, but wholly lacks
the hindwing red.
Number: Total recorded 39. Three taken.
Sex: Males only taken.
Date: April 30 to July 20.
Frequency: 1, 4, 5, 6, 23.
Condition: All freshly emerged.
Record: 1948 — April 30 (1 taken) ; June
10 (4 seen), 17 (6 seen), 22 (22 seen, 1
taken) ; July 20 (4 seen, 1 taken).
Papilio thoas neacles Rothschild and Jordan.
Field Name: Common Cresphontes-like
Swallowtail.
Species Range: Texas to Buenos Aires.
Subspecies Range: Nicaragua to Ecuador,
Venezuela, Trinidad and the lower Orinoco.
Field Characters: Cresphontes-like. Un-
identifiable, even at close range, from paeon
thrason, but only a single specimen of the
latter was taken in two years of collecting.
Number: Total recorded 105. Nine taken.
Sex: Both sexes taken.
Date : May 4 to September 8.
Frequency: Usually seen passing in small
groups, five to eight, maximum sixteen.
Strong flyers, difficult to capture, but occa-
sionally alighting, affording opportunity for
a good look.
Record: 1946 — May 4 (male taken, km.
20), 27 (16 seen); July 7 (female taken,
km. 20) ; September 8 (3 seen) . 1948 — April
29 (3 seen) , 30 (3, km. 26, headed for Pass) ;
May 1 (4), 10 (2), 11 (2), 15 (4 seen, 1
taken), 23 (8 seen, male taken), 26 (6), 31
(1 seen) ; June 6 (3 taken, male, 2 females.
Eggs protruding from females), 10 (5), 18
(8), 22 (6), 29 (4), 30 (5); July 2 (5
seen), 9 (2 at Pass, 6 km. 31), 13 (male
taken), 19 (3 seen, 1 taken km. 18).
Papilio torquatus or chamus Boisduval.
Field Name: Male, Small Yellow-Band-
and-Spot. Female, mimic of areas.
Species Range: Mexico to Bolivia, Brazil
and Paraguay.
Subspecies Range : Colombia and northern
Venezuela.
Field Characters : Male somewhat similar
to but smaller than male lycophron hippo-
medon; female very close to female of areas
areas. The large, separate, anterior yellow
spot on forewing of the male distinguishes
it from the solid band of lycophron.
Number: Two males were seen, and two
females taken.
Sex: Both sexes seen, female only taken.
Dates: May 1 to July 2.
Record: On May 1, 1948, I watched two
new papilios fighting in the Pass. One flew
down and alighted just out of reach, and the
other soon followed. I made a detailed de-
scription of them, recording them as yellow-
19491
Beebe: Migration of Papilionidae at Rancho Grande
125
banded-with-spot, tailless papilio. Not until
our return north were we able to identify
the insects by comparison with a male tor-
quatus taken at Caripito.
On May 26 and again on July 2, 1948, a
female was taken. These were badly rubbed
and torn, whereas the males I saw were
freshly emerged.
126
Zoologica: New York Zoological Society
[34: 14: 1949]
EXPLANATION OF THE PLATE.
Plate I.
Seventeen species of butterflies of the
genus Papilio taken as migrants at Porta-
chuelo Pass, Rancho Grande, north-central
Venezuela.
Fig. 1. sesostris tarquinius ( male).
Fig. 2. sesostris tarquinius (female)
Fig. 3. erithalion zeuxis (male).
Fig. 4. erithalion zeuxis (female).
Fig. 5. anchises osyris (male).
Fig. 6. anchises osyris (female).
Fig. 7. areas areas (male).
Fig. 8. areas areas (female).
Fig. 9. polydamus polydamus.
Fig. 10. belus varus form latinus.
Fig. 11. belus varus form varus.
Fig. 12. crassus form lepidus.
Fig. 13. polyxenes americus form melasma
(male) .
Fig. 14. polyxenes americus form melasina
(female).
Fig. 15. thoas neacles.
Fig. 16. paeon thrason.
Fig. 17. lycophron hippomedon (male).
Fig. 18. lycophron hippomedon (female).
Fig. 19. anchisiades anchisiades.
Fig. 20. torquatus orchamus (male).
Fig. 21. torquatus orchamus (female).
Fig. 22. cleotas coroebus form dione.
Fig. 23. phaon aberration metaphaon.
Fig. 24. agesilaus agesilaus.
Fig. 25. protesilaus archesilaus.
:ebe.
PLATE I.
MIGRATION OF PAPILIONIDAE AT RANCHO GRANDE,
NORTH-CENTRAL VENEZUELA.
Smith: Notes on Ergasilus Parasites
127
15.
Notes on Ergasilus Parasites from the New Brunswick, New Jersey,
Area, with a Check List of All Species and Hosts
East of the Mississippi River.
Roland F. Smith1.
Dept, of Zoology, Rutgers University,
New Brunswick, N. J.
Introduction.
The members of the genus Ergasilus rep-
resent the most undifferentiated of all the
copepod parasites and clearly show a pos-
sible line of evolution from the free-living
forms to the very specialized parasites that
are to be found in other families and genera.
Even in the genus itself one finds the tran-
sition taking place, and E. chautauquaensis
(which has never been found as a parasite)
to E. elegans, which is parasitic only after
the eggs begin to develop, and on to the other
forms where the females are parasitic after
they have become sexually mature.
The main characteristics of the genus are
its cyclops-like appearance; 2nd pair of an-
tennae enlarged and prehensile, 1st antennae
six-jointed; first thoracic segment and head
fused to form a large carapace; five pairs of
swimming legs; the first four biramose, the
fifth pair very degenerate and uniramose.
Egg sacs are similar to those in Cyclops. The
genus is typically fresh-water, though some
ergasilids are to be found in brackish and
marine waters. The type species of the genus
is a very common European species, E. sie-
boldi, established by Nordmann in 1832.
These parasites are generally found cling-
ing to the gill filaments, but one species, E.
megaceros, has been found in the nasal fos-
sae of the Fulton cat, Ictalurus anguilla, and
another, E. elongatus, has been found at-
tached to the bony gill rakers of the spoon-
bill cat, Polyodon spathula. At the present
time they have been found to infest all of
the major groups of fresh-water fishes and
it is likely that no species of fresh-water fish
is entirely free from the possibility of be-
coming a host. As far as is known, fishes are
the only hosts on which this genus has ever
been found. (See Mueller, 1936; Tidd &
Bangham, 1945; Wilson, 1911, 1916, 1925,
1932; Wright, 1844.)
The males of Ergasilus are free-swimming
i
1 The author wishes to express his appreciation to Mr.
Herbert Groat, who first called attention to the epidemic ;
Dr. R. F. Nierelli for assistance in making this paper ready
for publication ; and to Herbert Treuting and others who
gave assistance when it was needed.
throughout their lives. They are small and
easily overlooked in plankton samples and
consequently few of them have ever been de-
scribed. The chief distinguishing character-
istics in the males are the powerful maxilli-
peds, which are lacking in the females, and
the small, weak second antennae.
The females are free-swimming during
their early developmental stages and only at-
tach themselves after mating. It is generally
concluded (Wilson, 1911) that mating takes
place only once while the female is still free-
swimming. The sperms are stored in the
semen receptacle and fertilize the eggs as
they pass out into the ovi-sacs.
The breeding season apparently extends
throughout the summer months. The length
will vary from season to season, or from one
region to another, depending on favorable
water temperatures. In this area females
were observed with fully extended egg
strings on March 31. In the laboratory the in-
cubation period was found to be around
eight weeks, which is about the period of
time observed by Wilson (“eight to nine
weeks”). Henderson (1926) in her paper on
E. luciovercarium from Canada, stated that
it is likelv that the females carrv their egg
strings throughout the winter. This is not
tha case in Westons Mills Reservoir, but it
may be that in Canada, where the summer
season is considerablv shorter, the last batch
of eggs does not get the chance to develop
before cold water temperatures come and
consequently must be carried over until the
warmer temneratures of spring. In New
Brunswick the breeding season is over by
the middle of November.
Observation on Epidemic of Ergasilus on
Fishes in the Westons Mills Reservoir,
New Brunswick, N. J.
During the latter part of November, 1947,
fishermen began to notice large numbers of
fish dying in Westons Mills, a reservoir from
which New Brunswick obtains its water
supply. Local residents estimated that the
fish were dying by the “thousands,” and in-
deed the number of dying fish was so great
128
Zoologica: New York Zoological Society
[34: 15
as to attract large flocks of sea gulls from the
nearby Raritan River. The fish affected were
apparently only two species, calico bass, Po-
moxis sparoides, and bluegill, Lepomis ma-
crochirus.
Some of these fish were brought to the
Rutgers University Zoological Laboratory
where they were examined for possible para-
sitic infestations. No intestinal parasites
were found in excessive numbers nor did
there appear to be any injury of body tissues,
either external or internal. The gills ap-
peared to be covered with an unusually heavy
coating of mucus and microscopic examina-
tion revealed great numbers of copepod par-
asites of the genus Ergasilus. The blue color
of these organisms, the three knobs on the
inner edge of the second antennae towards
the distal end, and the fact that they were all
found in between the gill filaments seemed
to indicate conclusively that these were of
the species caeruleus. The hairs and spines on
the appendages, along with other general
morphological characteristics, were not quite
in accord with Wilson’s caeruleus, but there
seemed to be a considerable amount of varia-
tion in this genus, depending on geographic
location (Mueller, 1936). Wilson (1911)
considers caeruleus a parasite of the vege-
tative Centrarchidae, and gives the explana-
tion that the copepods on these fishes must
locate themselves between the gill filaments
to escape the discomfort and irritation to
their gills from bits of vegetation.
A number of trips were made to the reser-
voir to collect fish during this period. Oxygen
determinations were made at all depths and
at no time was the O2 concentration less than
10 ppm. Most of the fish were alive when cap-
tured. They could easily be spotted as they
swam feebly on their sides on the surface of
the pond. It was possible to come up along-
side these fish in a boat and pick them up by
hand. Occasionally a fish would sound on ap-
proach, but only to rise slowly to the surface
after a short interval. On the first of such
trips about a dozen fish were collected and
taken back to the laboratory. These were
placed in a well-aerated aquarium. At first
they seemed almost dead — all lying motion-
less on their sides, with only an occasional
movement of a fin. The next day, however,
they seemed fully recovered and all were
swimming about apparently quite normal. A
few of these examined at this time were
found to have a very heavy infestation.
In brief, the meager facts obtained on the
epidemic from Nov. 27 to Dec. 10, 1947, can
be summarized as follows :
1. Many calico bass and bluegills died during
this period. No official estimates could be
obtained, but laymen who observed the
phenomenon estimated the deaths to be in
the thousands.
2. One calico bass died for every ten blue-
gills. These were apparently the only fish
affected.
3. All fish that were infested by these para-
sites appeared to have a heavy coating of
mucus over the gills.
4. Estimated number of parasites on each
fish was 250-300.
5. Age group (as determined by scale read-
ings) was 1-2 years.
During the late fall of 1948 these observa-
tions were continued. A large fish trap was
constructed and set in deep water as soon as
ice covered the reservoir. All of the species
of the Centrarchidae as well as one yellow
perch were examined. Later in the year the
fish were obtained by seining and from fish-
ermen.
In this survey the large-mouth bass, Mi-
cropterus dolomieu, was found to be para-
sitized by a second species, E. centrarchida-
rum. This species is generally larger, broad-
er, and found on the outside of the gill fila-
ments, in contrast to caeruleus which is
found between the gill filaments. In addition,
there are no knobs on the inside surface of
the distal end of the second pair of antennae.
Again there were morphological differ-
ences from Wilson’s description of centrar-
chidarum and this phase of the work will be
discussed in a later paper. Both species had
the blue pigment but centrarchidarum was
never as deeply pigmented as caeruleus.
Strangely enough, both species had the same
hair and spine formula on their appendages.
This is: 1st exopod, 1-0; 0-1; II-5, endopod
0-0 ; 0-1 ; II-3 : 2nd exopod, 0-0 ; 0-1 ; 0-6, end-
opod, 0-1 ; 0-2 ; 1-4 : 3rd exopod, 0-0 ; 0-1 ; 0-6,
endopod, 0-1; 0-2; 1-4: 4th exopod, 0-0; 0-5,
endopod, 0-1; 0-2; 1-3.
In addition to the sampling of fish from
Westons Mills, two other bodies of water in
the New Brunswick area were samoled.
These included the lower section of the Dela-
ware-Raritan Canal and the small pond in
Johnson’s Park. The bluegills and calico bass
in Johnson’s Park were found to be free of
ergasilids but the canal proved as fruitful as
the reservoir. A summary of the fish ex-
amined is given in Table I.
Some interesting facts are foreshadowed
in Table I. Although the amount of sampling
from both bodies of water was not extensive
enough to reveal fully the actual condition,
nevertheless the methods by which these fish
were obtained and the period of time over
which the sampling was made certainly in-
dicate the trend that one might expect to find
if a more extensive sampling were to be
taken.
For example, only five large-mouth bass
were obtained — but under totally different
conditions and at various times. The fact
that all five had infestations does not indi-
cate that all the bass are nai'asitized. yet one
would expect to find a high percentage of the
bass serving as hosts to these parasites. On
the other hand, not once during this entire
period of research has there ever been found
a pumnkinseed infested with these copepods.
Certainly one can sav that for this species
the incidence of parasitism is very low.
Again, viewing the information in the
same light, one should expect the bluegills to
be parasitized about 50% of the time in
f
Cali
Bto
Pui
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Yel
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sf
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(
1949]
Smith: Notes on Ergasilus Parasites
129
TABLE I.
Result of the Samplings from Westons Mills and Delaware-Ra>’>tan Canal
from Nov. 30, 1948- July 30, 1949.
Name
Total No.
Percent of
Average
Length
Caught
parasitism
Length
Range
Calico bass*
5
100%
5.79"
4"— 7"
Bluegills
15
60%
5"
2"-7%"
Pumpkinseed
6
0%
4"
3"— 5"
Large-mouth bass
5
100%
6.4"
2%"-12'
Yellow perch
1
0%
7"
7"
* The highest infestation
noted was on one calico bass
5'”K" in length :
259 caeruleua were counted
on this fish.
Westons Mills and the Canal. No data on
calico bass could be obtained from the Canal
but from the available information it would
seem that these fish have become the most
heavily infested in Westons Mills, not only
in number of individuals but in number of
parasites per fish. This is significant, for al-
though the calico bass was found to be the
most heavily infested as well as the most
commonly parasitized fish, the epidemic of
1947 killed only one calico bass to ten blue-
gills. Unfortunately, here also, too little in-
formation is at hand. On the basis of the
present data, however, there seems to be two
logical explanations for this:
1. The calico bass are more resistant to the
infestation of these gill parasites than are
the bluegills and have been able to build up
a resistance.
2. All the bluegills left are those that have
built up a resistance to these parasites,
or those that were only slightly parasi-
tized.
Both fish are to be found in the same gen-
eral type of environment and both have simi-
lar food habits, so apparently these two fac-
tors may be discounted.
Pathogenesis.
There has been a great deal of controver-
sy over the extent of damage done by the
ergasilids, as well as exactly what kind is
done. Wilson (1911) states that “Living as
they do upon the fish’s gills, there can be but
little doubt that they feed upon blood.”
Halisch (1939) in his observations on E. sie-
boldi and E. minor on the gills of tench
states that extra-intestinal digestion is im-
portant and that much more tissue is de-
stroyed than is ingested by the parasitic co-
pepod. Blood may be taken in. Fungi may
grow in the lesions.
In contrast to these two observations, Hen-
derson (1926), in her work on E. lucioper-
carium, observed that “The gills may be
heavily attacked without affecting the health
of the fish. It is a harmless parasite, which,
while it only benefits the unbidden guests,
causes no lesions and consequently does no
injury to the host.” (If this is the case it
should not even be called a parasite, but a
commensal). She goes on to explain how, in
her opinion, it is impossible for the parasites
to injure the gill tissue, since the mouth
parts of these organisms are too weak to
pierce the gill tissue.
One might suspect that the claw-like, pre-
hensile, second pair of antennae of the para-
site is capable of at least some damage to the
gill lamellae, although the mouth parts may
be too small to cause any injury. However,
the observations in the present studies
showed no evidence of mechanical injury;
neither was there any evidence of digested
gill tissue or blood in the intestine of the
numerous copepods examined, even in fe-
males with ovaries full of developing eggs.
We agree with Henderson (1926) that
these parasites in all probability feed on the
excessive mucus produced under the condi-
tion of the parasitism, or the many minute
mucus or bits of organic debris and bacteria
in this viscid material. However, it is alto-
gether possible that the parasites may feed
on sera, straining out the corpuscles.
A close examination of the mouth parts
of these ergasilids will reveal the mandibles
and second maxillae heavily fringed with
setae. These would seem ideal for feeding on
mucus or bits of organic debris and bacteria
but hardly suitable for feeding on blood. Any
pathological conditions of the gills may cause
the mucus glands to secrete an excessive
amount of mucus (Nigrelli, 1949). This has
been especially apparent during infestations
of trematode gill parasites. This condition
has also been observed on many of the fish
that were infested with these copepods2. It
may be that an excess mucus secretion over
the surface of the gills may lower the efficien-
cy of the gills to absorb the dissolved oxygen
in the water. Under normal water conditions
when there is sufficient oxygen and all the
chemical and physiological factors are in
proper balance, this may not have any dele-
terious effects. However, if, for example, a
factor such as the CO2 concentration in the
water should be increased, it might be suffi-
cient to reduce the efficiency of the gills in
absorbing the dissolved oxygen and so cause
the fish to die of suffocation.
The highest death rate has been found
among the younger fishes and it is the
- It must be kept in mind that the heavy coating of
mucus which seems to accompany fish that are heavily
infested with this parasite does not mean too much in
itself. Fish that have been placed in preservative or that
have died from other causes may show the same condition.
Moreover, some fish that were heavily infested did not have
an excess of mucus covering the gills. In such cases these
fish exhibited no symptoms whatsoever and appeared per-
fectly normal. Apparently, there is a physiological balance
here that is very delicately adjusted and which can be
thrown out of balance only under certain specific conditions.
130
Zoologica: New York Zoological Society
[34: 15
TABLE II.
Check List of the Ergasilus Found East of the Mississippi River.
Names and Synonyms Hosts
Localities Studied
Remarks
E. caeruleus
Wilson
Synonyms:
Yellow perch
Trout perch
Wall-eyed pike
Gray pike, Blue pike
Rock bass, Calico bass
Lake Mendota
Trout Lake region
Lakes Erie & Michigan
Oneida Lake
Mississippi River, Iowa
E. confuscus
Bere
White bass
Warmouth bass
E. skryabini
Mueller
Crappie
Green sunfish
Blue-spotted sunfish
Pumpkinseed
Bluegill
Lake trout
Cisco, White fish
Sucker, Long-nosed gar
E. centrarchidarum
W right
Rock bass. Calico
bass, Large-mouth
bass, Small-mouth
bass, White bass,
Warmouth bass,
Green sunfish,
Bluegill, Crappie,
Pumpkinseed, Sunfish (?)
Wall-eyed pike, Gray pike,
Sauger, Silversides, Smelt,
Microgadus tomcod
Lake Erie
Lake Michigan
St. Lawrence River
Watershed
Lake Champlain
Black Lake, N. Y.
Lakes St. John &
Chibogamo, Quebec
St. Andrew’s Bay, N. B.
Lake Maxinkuckee, Ind.
Mississippi River, Iowa
Clewiston, Fla.
Woods Hole
E. chautauquaensis
Fellows
Lake Champlain
Lake Mendota
Fairport, Iowa
Has never been
found as a parasite,
but may be like
elegans
E. cotti
Kellicot
Rainbow darter
Sculpin — Cottus bairdii
Lake Erie
Westerville, Ohio
E. elegans
Wilson
Ameiurus sp.
Northern black bullhead
Channel catfish
Short-nosed gar
Long-nosed gar
Lake Okeechobee
Myakka River & Canals
Peace River, Fla.
Mississippi River, la.,
Parasitic only after
eggs begin develop-
ing
E. elongatus
Wilson
Spoonbill cat
Mississippi River, Iowa
and Illinois
E. funduli
Kr0yer
Fundulus ocellaris
New Orleans
Salt and brackish
water
E. labracis
Kr0yer
Striped bass
Woods Hole
Baltimore
Marine
E. lanceolatus
Wilson
Gizzard shad
Cumberland River, Ky.
E. lizae
Kr0yer
Common killifish
Gulf killifish
Broad killifish
Striped mullet
White mullet
Englewood, Fla.
New Orleans
Salt and brackish
waters
E. luciopercarum
Henderson
Pike perches
Lake St. John
Lake Chibogamo, Que.
E. manicatus
Wilson
Silversides
Smelt
Two-spined stickleback
Top minnow
( Gambusia holbrooki )
Jordanella floridae
Englewood
St. Andrew’s Bay, N. B.
Woods Hole
Along Atlantic Coast
Marine
1949]
Smith: Notes on Ergasilus Parasites
131
Names and Synonyms Hosts
Localities Sttidied
Remarks
E. megaceros
W ilson
Synonym :
Fulton cat
Fall fish
Oneida Lake, N. Y.
Mississippi River, Iowa
Found in the nasal
fossae and
spiracles
E. fragilis
Mueller
E. mugilis
Vogt
Striped mullet
Beaufort, N. C.
Marine
E. nigratus
Wilson
Large-mouth bass
Mississippi River, Iowa
E. osburni
Tidd & Bangham
Burbot
North Central States
E. versicolor
Wilson
Synonym :
E. celestis
Mueller
Fulton cat
Red-mouthed buffalo fish
Channel cat
Common brown bullhead
Mudcat
Skip jack
\ ellow cat
Eel cat
Mississippi River, Iowa
Florida
Lake Erie
Lake Maxinkuckee
younger fishes that would require the great-
est amount of oxygen (in proportion to the
gill area), due to their greater metabolic
activity. At the same time it is the younger
fisnes that are more susceptible to attacks
from disease and parasites since they are
using all their available energy toward
growth. Wilson states that it is the young
nsh that are most heavily parasitized and
this is probably true — especially during an
epidemic. However, larger fish may also be
quite heavily parasitized. (See Table I).
'the ergasihds on the large-mouth bass
were never very abundant — never more than
thirty on any individual. Although Wilson
(1916) mentions fish fatalities from the
ergasilids he does not mention which species
of copepod causes death, nor which species
of fisn are killed. In checking all the litera-
ture on centrarchidarum, never were their
numbers found to be as great as for caeru-
leus. Therefore, one wonders if caendeus is
not the only one that may appear in such
numbers as to bring about the death of a fish.
This leads us to speculate on how many
ergasilids must be present on a fish to cause
death. It would seem that numbers that lead
to the death of a fish at one time, appear to
have no effect at another. However, during
the epidemic of 1947 none of the dead fish
had less than an estimated 250 copepods.
Whether fewer parasites can bring about
the death of a fish remains to be determined
by further study.
It is apparent from the literature that E.
centrarchidarum is the most widespread par-
asite and probably the best known. It has
been found in all the main regions studied,
including the marine habitat, but it has not
been found on as many hosts (16) as has
caeruleus (19), nor on as great a variety.
Caeruleus has not been found in all the
areas that have been studied and so far has
proved to be an exclusively fresh-water par-
asite. Additional research may also reveal
that caeruleus is more widespread than cen-
trarchidarum. It is unfortunate that most
of the workers in this field have failed to
make clear whether caeruleus has always
been found between the gill filaments of its
host. If it is found between the gill filaments
of such clear-water fishes as the lake trout,
cisco and white fish, certainly Wilson’s ex-
planation for their being found between the
gill filaments is not substantiated.
E. versicolor is apparently third in abun-
dance and seems to be more specific in its par-
asitism.
Many of the Ergasilus have only been
found on one or a few hosts, but it is still too
early to state definitely that they are spe-
cific for only those hosts.
Conclusions.
1. It seems quite likely that Ergasilus may in-
directly cause death to fish, although it
probably seldom causes extensive damage
to any given fish population.
2. Young fish (1-2 years) are apparently the
only ones on which these parasites may prove
fatal. Although infestations have been found
to be equally high on all age groups, more
young fish appear to be heavily infested than
older ones.
3. Ponds where the fish are overcrowded or that
are small in area seem to be the only places
where infestations become high enough to
cause death.
4. Ergasilids do not feed on blood, but prob-
ably on mucus or bits of organic matter
found in the viscid material.
5. Death of fish is probably due to a delicate
physiological imbalance affecting the diffu-
sion of oxygen through the gill tissues.
132
Zoologica: New York Zoological Society
[34: 15: 1949]
6. E. caeruleus can always be distinguished
from E. ceritrarchidarum by the three knobs
on the second pair of antennae, and by the
fact that it is always found in between the
gill filaments. Variations among these cope-
pods are exceedingly great.
7. Although Wilson considers E. caeruleus a
parasite of the plant-eating Centrarchidae,
it is also found on species that are quite
carnivorous and not generally found among
aquatic vegetation. It undoubtedly is found
most commonly on fishes that typically in-
habit an environment of aquatic vegetation,
such as the bluegill and calico bass.
8. Though ceritrarchidarum and caeruleus have
often been found on the same fish (Wilson,
1911, 16), they were never observed together
in the New Brunswick area nor was cen-
trarchidarum ever observed on either blue-
gills or calico bass in this area. Neither was
observed on any but the above species of fish.
9. It may be that certain factors, chemical,
physical, physiological and environmental,
or a combination of these, determine which
hosts may be parasitized.
10. No reason could be found to explain why
the pumpkinseed, Lepomis gibbosus, is free
from these copepod parasites in this region.
Summary.
An epizootic in Westons Mills Reservoir
which caused the death of many bluegills
and calico bass was investigated.
A copepod parasite, Ergasilus caeruleus,
was believed to be the indirect cause of the
death of these fish. Apparently, metabolic
wastes from the copepod or irritation to the
gill tissues causes an excessive secretion of
mucus. This mucus may lower the efficiency
of the gills, so that when certain other fac-
tors are not in proper balance the fish will
suffocate.
Large-mouth bass were found to be in-
fested with another species E. centrarchida-
rum.
A brief survey of two other bodies of
water in the New Brunswick area revealed
that the incidence of parasitism from these
ergasilids is quite high for bluegills, calico
bass and large-mouth bass, but does not seem
to be present on the pumpkinseed.
References.
Halisch, J.
1939. Anatomie und Biologie von Ergasilus
minor. Zeitschr. Parasitenk., 11 (2/3) :
284-330.
Henderson, Jean T.
1926. Description of a Copepod Gill Parasite
of Pike Perches in Lakes of Northern
Quebec, Including an Account of the
Free-Swimming Male and Some De-
velopmental Stages. Cou.tr. Canadian
Biol. & Fish., N. S. 3 (7) : 235-246.
Mueller, J. F.
1936. Notes on Some Parasitic Copepods and
a Mite, Chiefly from Florida Fresh
Water Fishes. The American Midland
Naturalist, 17 (5) : 807-815.
Nigrelli, Ross F.
1949. Notes from Lectures on Fish Diseases.
Tidd, W. M. & Bangham, R. V.
1945. A New Species of Parasitic Copepod,
Ergasilus osburni, from the Burbot.
Trans. Amer. Micr. Soc., 44 (3) : 225-
227.
Wilson, C. B.
1911. North American Copepods Belonging
to the Family Ergasilidae. Proc. U. S.
Nat. Mus., 39: 263-400.
1916. Copepod Parasites of Fresh- Water
Fishes, and Their Economic Relations
to Mussel Glochidia. Bull. U. S. Fish-
eries, 34: 331-374.
1924. New North American Parasitic Cope-
pods, New Hosts, and Notes on Cope-
pod Nomenclature. Proc. U. S. Nat.
Mus., 64 : 1-22.
1932. Copepods of the Woods Hole Region,
Massachusetts. Bull. U. S. Nat. Mus.,
No. 158: 1-623.
Wright, R. R.
1884. Notes on American Parasitic Cope-
poda. Proc. Canad. Inst., N. S. 1:243-
254.
Aronson: Reproductive Behavior in Tilapia macrocephala
133
16.
An Analysis of Reproductive Behavior in the Mouthbreeding Cichlid
Fish, Tilapia macrocephala (Bleeker).1,2
Lester r. Aronson.
Department of Animal Behavior, The American Museum of Natural History.
(Plates I-III; Text-figures 1-10).
Introduction.
Teleosts of the family Cichlidae are noted
for their elaborate patterns of courtship,
mating and parental care, and for the read-
iness with which they breed in the restricted
confines of the small aquarium. It is largely
because of these attributes that cichlids have
become the subjects of several extensive in-
vestigations of fish behavior. Outstanding
among these studies are those of Breder
(1934) on the blue acara, Aequidens lati-
frons ; Noble and Curtis (1939), Peters
(1941) and Seitz (1942) on the jewel fish,
Hemichromis bimaculatus ; Peters (1937) on
the small Egyptian mouthbreeder, Haplo-
chromis multicolor; and Seitz (1940) on a
closely related mouthbreeder, Astatotilapia
strigigena.
These students have investigated topics
such as schooling, sex recognition, courtship,
territory, social dominance, spawning, pa-
rental care, the stimuli causing the release
of various innate responses and many other
related items of behavior. In these studies,
mating behavior has been described qualita-
tively and in varying degrees of detail. Al-
though the reports in most cases have been
based on a number of observed spawnings,
the results are given in a generalized or “av-
eraged” form and the only suggestion of va-
riability in behavior is found in such broad
phrases as “this usually happens,” or “the
typical mode of behavior is.” Moreover, the
“averaging” is often achieved by means of
subjective impressions rather than in terms
of a calibrated or objectively weighted eval-
uation of behavioral characteristics.
Variability is a fundamental characteristic
of biological phenomena, a characteristic
1 The experiments herein reported were supported by
a grant from the Committee for Research in Problems of
Sex. National Research Council.
2 Mrs. Magda Schonwetter assisted in many of the ob-
servations. Drs. Frank A. Beach and Charles M. Breder
made numerous helpful suggestions on the conduct of the
observations and experiments. Dr. Myron Gordon helped
solve the feeding problem. Dr. T. C. Schneirla, Mr. James
W. Atz, Mrs. Marie Holz-Tucker and Mr. Christopher W.
Coates read the manuscript and made innumerable con-
structive criticisms. The author gratefully acknowledges
his indebtedness to these people and those past and present
associates of the Department of Animal Behavior who in
ways too numerous to mention made possible the comple-
tion of this study.
which always merits careful consideration
in studies of animal behavior. It is the writ-
er’s belief that the study of teleost behavior
cannot extend very far beyond the present
descriptive stages unless and until methods
of a more quantitative nature are employed.
Students of mammalian psychology, and par-
ticularly of rodent behavior, have made ex-
cellent progress by utilizing quantitative
procedures. The present study afforded an
opportunity for testing the applicability of
comparable techniques in the study of the
reproductive behavior of fish.
The present report is concerned with the
average behavior and the range of varia-
bility under constant aquarium conditions
of a type which can be readily duplicated. In
other investigations now in progress, the
mating activities of brain-operated and hor-
mone-treated animals will be compared to
the norms obtained in the present report.
Nothing appears to be known concerning
the mating behavior of Tilapia macrocephala
in the wild state, and the present report is
hardly intended as a substitute for such an
investigation. Nevertheless, wherever the be-
havior of fishes has been studied both under
field conditions and in captivity, agreement
has been fairly good, as for example in the
Centrarchidae. It is anticipated that the
over-all picture obtained in this study should
prove to be essentially similar to conditions
prevailing in the natural state, and that dif-
ferences if any would be expected only in
some of the lesser details.
Literature.
Information concerning the breeding hab-
its of Tilapia macrocephala and of related
species belonging to the same genus has been
furnished for the most part by aquarium
hobbyists and through cursory observations
by field naturalists. It is realized that because
aquarists’ reports often fall below generally
recognized standards of scientific accuracy,
as might be expected considerable confusion
exists in the literature concerning certain
aspects of the breeding patterns of Tilapia.
Some of these difficulties may no doubt be
attributed to an improper identification of
134
Zoologica: New York Zoological Society
[34: 16
the species in question, since aquarists some-
times trust the knowledge and dependability
of fish dealers for the identity of their sub-
jects. Nevertheless it is possible to obtain
from this literature a rough picture of the
reproductive habits of the genus Tilapia. For
these reasons the inclusion of numerous
aquarists’ accounts is considered expedient.
What may be offeied herein is by no means
intended to be a comprehensive review of
the extensive popular literature.
Brief descriptions of the breeding habits
of Tilapia macrocephala can be found in the
aquarium texts of Stoye (1935), Arnold and
Ahl (1936) and Innes (1944). In addit.on,
reports on the spawning of Tilapia lieude-
loti (which according to Boulenger (1915)
may be a variety of Tilapia macrocephala)
have been presented by b>reaer and
Schoenfeld (1934). These accounts tell us
briefly that (1) a nest is built by the mating
pair; (2; the female deposits the eggs in the
nest; (o) tne male fertilizes the eggs as soon
as tney are deposited; (4) shortly thereafter
the male taxes the eggs into his mouth; and
to; tne eggs natcn anu develop in tne mouth
of the maie. now long the eggs are retained
in the maie’s buccai pouch is not indicated,
but Stoye (1935; reports two cases where
eggs were carried 24 to 29 days respectively.
Stoye considers these periods abnormally
long as a result of excessive disturbances.
With four probable exceptions, all the
species of Tilapia whose spawning habits
have been reported are mouthbreeders. The
four exceptional species remove their larvae
to sand pits in typical cichlid fashion. These
non-mouthbreeding species are (1) Tilapia
guinasana (Rolon, 1938, 1939), (2) Tilapia
spaarmami (ney, 1945, 1941; Anon., 194S)J,
(3) Tilapia melanopleura (Svenssor, 1933;
Bertram, Borley and Trewavas, 1942; and
(4) Tilapia zillii (Bade, 1923; Stoye, 1935;
Arnold and Ahl, 1936; Bertram, Borley and
Trewavas, 1942). However, Liebman (1933)
describes Tilapia zillii as a mouthbreeder.
Incubation of the eggs is accomplished by the
females of Tilapia flavomarginata (Pella-
grin, 1906) i, Tilapia gaMlaea (Pellegrin,
1903, 1905), Tilapia martini (Boulenger,
1906), Tilapia microcephala 5 (Junghans,
1918) and Tilapia mossambica* * 4 5 6 * 8 (Bade, 1923;
Dietz, 1926; Roloff, 1937; Peters, 1937a,
1939; Seleuthner, 1941; Hey, 1947). The
same appears to be true for Tilapia squami-
pinnis, Tilapia lidole, and Tilapia shirana
(Bertram, Borley and Trewavas, 1942).
The male is credited with the care of the
eggs in Tilapia dolloi (Asch, 1939), Tilapia
heudeloti (Breder, 1934; Schoenfeld, 1934),
8 Also recorded in "Report No. 1 (1944) Inland Fish-
eries Dept., Union of South Africa”— 1946.
4 Designated by Boulenger (1911) as Tilapia andersonii.
5 Tilapia microcephala — Tilapia heudeloti. According
to Boulenger (1916, p. 178) Tilapia macrocephala, and
Tilapia multifaeciata “may ultimately have to be regarded
as varieties of T. heudeloti. I am unable to find characters
by which to separate them sharply.’’
8 Tilapia mossambica = Tilapia nataleneis.
Tilapia macrocephala (Stoye, 1935; Innes
1944), Tilapia microcephala (Schreitmiiller,
1920) and Tilapia simonis 7 (Lortet, 1875;
1883). However, there is some disagreement
on this point since both the male and female
are believed to incubate the eggs in Tilapia
simonis (Pellegrin, 1903; Liebman, 1933),
Tilapia galilaea (Liebman, 1933), Tilapia
microcephala (Locke, 1932), Tilapia nilotica
(Boulenger, 1901) and Tilapia zillii (Lieb-
man, 1933). Bodenheimer (1927) claims that
females alone incubate the eggs of Tilapia
simonis, and Arnold and Ahl (1936) claim
the same for Tilapia dolloi.
Irvine (1947) states that the male or pos-
sibly both sexes of Tilapia discolor and Tila-
pia heudeloti incubate the eggs, but con-
trary to the findings of Boulenger, Irvine
relegates this function to the female in Tila-
pia nilotica. Liebman (1933) believes that
it is quite general in Palestine cichlids for
both parents to incubate the eggs, but the
number of females performing this function
is higher than the number of males so doing.
The length of the incubatory period has
been reported for only a few species. Arnold
and Ahl (1936) say about 14 days for
Tilapia dolloi; Schreitmiiller (1920) gives 4
to 6 days for Tilapia microcephala; Roloff
(1937) reports 21 days for Tilapia mossam-
bica; Bade (1923) offers a value of 15 days
while Dietz (1926) and Seleuthner (1941)
both give 13 days as the incubatory period
of this species.
The retrieving into the female’s mouth of
newly released young has been reported for
Tilapia dolloi (Arnold and Ahl, 1936), Tila-
pia macrocephala (Stoye, 1935), and Tilapia
mossambica (Roloff, 1937; Seleuthner, 1941)
while in Tilapia microcephala the male is
credited with that activity (Schreitmiiller,
1920).
Nest making by these mouthbreeders has
received some general attention. On a num-
ber of occasions, Lortet (1883) witnessed
the female Tilapia simonis lay approximately
200 eggs m a small excavation which she had
hollowed out and cleaned in the silt among
the reeds. Nest building by both the male
and the female Tilapia nilotica was observed
in the field by C. L. Boulenger (1908 ) . Roloff
(1937) describes the nest of Tilapia mossam-
bica as being 20 cm. in diameter. Seleuthner
(1941) reports a nest for this species which
was 25 cm. in diameter and reached a depth
of 4 cm. in the middle, while Hey (1947)
pictures it as a “small saucer-shaped depres-
sion.” Bertram, Borley and Trewavas (1942)
describe the nest of Tilapia squamipinnis as
a circular depression.
Other mouthbreeding cichlids are listed
by Peters (1937) as belonging to the genera
Astatotilapia, Ectodus, Geophogus, Hap-
l Placed in the genus Tilapia by Boulenger (1899) but
now referred to a new genus Tristramella by Trewavas
(1942). Lortet (1876) called this fish Chromis pater-
familias.
1949]
Aronson: Reproductive Behavior in Tilapia macrocephala
135
locliromis, Pelniatochromis and TropheusA
It is to be noted that at least some of these
genera also contain non-mouthbreeding spe-
cies, suggesting a multiple origin of this
habit even within the cichlid family. This
problem has been considered in some detail
by Breder (1933) and Myers (1939).
The small Egyptian mouthbreeding cich-
lid, Haplochromis multicolor, and a closely
related form, Astatotilapia strigigena, have
been the most popular and intensively studied
of all the mouthbreeding fish. In addition to
the scientific investigations previously men-
tioned, more than 30 accounts of the spawn-
ings of these two fish have appeared in the
last three decades, the majority of them in
the Wochenschrift fur Aquarien-und Terra-
rienkunde. These accounts, which are rela-
tively consistent in their general implica-
tions, demonstrate that the spawning be-
havior of these species differs considerably
from that of the various species of Tilapia
described above. For this reason what is
known about the Haplochromis and Astato-
tilapia mating patterns is summarized brief-
ly for the purposes of comparison.
In these species the male does practically
all of the nest building. Upon the completion
of the nest, the female starts the oviposition
by depositing between four and ten eggs in
tne nest. Tne maie immediately fertilizes
the eggs after which they are picked up by
the female. This cycle is then repeated as
the female lays a second batch of eggs. Be-
tween five and ten such cycles have been re-
ported by various authors as comprising a
spawning. The eggs are carried for 9 to 20
days, after which the young are released.
However, the young are taken back into the
female’s mouth at night and at other times
when disturbed. Such a retrieving of the
young has been the subject of a special in-
vestigation by Peters (1937).
Material and Methods.
Tilapia macrocephala (Bleeker) is native
to West Africa, particularly in the region
of the Gold Coast. Boulenger (1915) de-
scribes the species as coming from the Gold
Coast, Ashantee and Lagos. Many of his
specimens were taken from the Ancobra
river and Secconda lagoon in the Gold Coast,
and from the Lagos lagoon. According to
Arnold and Ahl (1936) the fish is found in
the brackish lagoons of the coast and the
swampy deltas of rivers.
The individuals utilized in this study were
selected from a laboratory-bred stock which
had been maintained for a number of years
prior to the start of the present research.8 9
Males were chosen for the brightness of their
yellow operculum which is a secondary sex
8 An older listing of mouthbreeding cichlids given by
Pellegrin (1903) includes the genera Geophaaua. Aeara,
Chaetobranch.ua, Tilapia, Paratilapia, Pelmatochromis,
Ectodu8 and Tropheus.
9 I am greatly indebted to Miss Ethelwyn Trewavas of
the British Museum for kindly checking and verifying the
taxonomic identity of the fish as Tilapia macrocephala.
character (PI. I, Fig. 1). The females (PI.
I, Fig. 2) were selected on the basis of the
complementary sex character, namely a deep
red spot in the center of the gill cover.10 * *
These dimorphic color patterns appear at
sexual maturity and disappear after castra-
tion (Aronson, in manuscript) .
Pairs were established by random selec-
tion and were placed in 54-liter aquaria, 60
cm. X 30 cm. X 30 cm. each containing
roughly 36 liters of water. The side and rear
walls of these tanks were painted pale blue
to minimize any possible disturbing influ-
ences from neighboring tanks, and also to
facilitate the ability of the investigator to
follow the activities of the fish. The tanks
were located in a greenhouse the temperature
of which was maintained throughout the
year at 26° C. with a positive and negative
variation of approximately 3° C. To furnish
hide-outs for tne fish which at the same time
would not obstruct the observer’s view, a
mat of floating plants was placed in every
tank. Cabomba was extensively used for this
purpose, but Sagittana subulata was found
to be somewhat more suitable and was used
whenever available.11 The fish were fed
mostly a dehydrated preparation consisting
of dried shrimp, oatmeal, beef liver, lettuce
ana spinacn. At times tms was supplemented
by live tubifex worms. Occasionally the fish
nibbled at the stonewort Niteila, and this
was piaced in the tanks when available. The
tanks were aerated continuously, and the
water was changed whenever it became ex-
cessively murky. This was approximately
once a month. Tap water brought to the
proper temperature was used in washing
the tanks and for replacement.
In order to avoid injury to the fish due to
excessive nipping which often occurred after
spawning, a transparent glass partition was
placed in the aquarium, separating the male
from the female as soon as observations of
oviposition were completed. As the individ-
ual carrying the eggs (generally the male)
eats little or nothing during the incubatory
period, brooding fish were not fed during
this interval. By the time that the young
were released from the male’s mouth, the
females often were prepared to spawn again.
However preliminary observations indicated
that when such spawnings occurred males
sometimes behaved abnormally, due appar-
ently to the protracted period of inanition.
To avoid this difficulty an arbitrary rule
was established to the effect that males were
separated by a transparent glass partition
from females for one week after they had
released their young or had swallowed their
eggs. This interval allowed the males to feed
and regain their strength. While thus iso-
10 Examination of this spot by Aronson and Holz-Tucker
(in manuscript) has revealed that it is in actuality a semi-
transparent window through which the underlying red gill
can be seen.
11 The author wishes to express his appreciation to Dr.
Myron Gordon of the New York Zoological Society for
furnishing all of the sagittaria used.
136
[34: 16
Zoologica: New York Zoological Society
lated, females often spawned alone but ob-
servations showed that a considerable
amount of courtship took place through the
glass partition.
To prevent the parents from eating their
young after they were released, large masses
of the stonewort Nitella were placed in all
tanks where young were being incubated.
The stonewort was distributed equally on
both sides of the partition since the newly
released fry could easily swim through the
cracks at the intersection of the partition
and the glass walls of the tank. Disturbances
caused the young to swim into the fine inter-
stices of the Nitella where they would not
be followed by their cannibalistic parents.12
The criteria employed to indicate the ap-
proach of oviposition were (1) persistent
nest building, mostly by the female, (2)
heightened courtship activity, and (3) pro-
truding genital tubes. When these signs were
observed, continuous records were taken of
the courtship and mating activities of the
pair up to the time of spawning and for
one-half hour thereafter. Attempts were
made to secure continuous pre-spawning
records for three hours. However, this goal
was attained in only a small number of cases
with the result that the records vary from
just a few minutes before spawning up to
the full three-hour span. A serious difficulty
was encountered here in that the activities
of many promising pairs were observed
continuously for many hours up to a whole
day without the fish ever ovipositing.
After the present experiment had been
terminated, behavior during the interspawn-
ing interval was studied, using different
pairs of Tilapia. The experimental condi-
tions were the same as before with the fol-
lowing minor exceptions : (1) no plants were
used but instead inside aquarium filters
served as hide-outs; (2) the water in the
tanks was never changed; (3) the males
12 This was the author’s first experience with the main-
tenance of tropical fish. Since then, several innovations
have been developed. Inside aquarium charcoal filters are
now placed in every tank. These keep the water clean
and eliminate the need for chancing it. Plants are not
used since they grow poorly in Tilapia aquaria. The food
formulae have been modified as follows: (1) Wet mash:
2V> lbs. liver ; % lb. chopped lettuce and spinach : Va lb.
dried ground shrimp (mostly shell) : % lb. dried and ground
refined shrimp (mostly muscle) : Pablum (or other pre-
cooked infant cereal) —enough to make thick paste (ap-
proximately 3 lbs.) ; 1 pinch salt. The liver is chopped,
about 1 cup of water added, and the mixture is then
liquefied in a blending machine. All ingredients are mixed
together with sufficient Pablum to make a paste. The
food is further solidified and preserved by packing into
jars and immersing them in boiling water for about 10
minutes. (2) Dry food: 12 lbs. dried shrimp (mostly
shell) : 12 lbs. dried refined shrimp (mostly muscle) ; 10
lbs. liver ; 6 lbs. chopped lettuce ; 6 lbs. chopped spinach ;
28 lbs. Pablum ; 2 level teaspoons salt. The ground spinach
and lettuce are mixed with the Pablum and cooked for 15
minutes. The liver is cut into slices and boiled for 15
minutes in a minimum amount of water and then chopped.
All ingredients are mixed together and the resulting paste
spread about % inch thick on trays. When almost com-
pletely dry, the food mixture is ground and sifted through
screens of several coarsenesses.
The sexes are no longer separated after the spawning.
If the fry are to be saved they are forcibly removed from
the parents’ mouths on the tenth day post-oviposition,
and are placed in small aquaria. At this age the young do
very well without further parental care, and thus, losses
through cannibalism are easily avoided.
were never separated from the females. The
actual spawnings of these pairs were not
witnessed, all ovipositions being recorded
as having occurred at the time the eggs were
discovered in the male’s mouth. All pairs
were checked twice daily for eggs. A 15-
minute record of the behavior of a given
pair was taken 5 or 6 days after the spawning
and again on the 15th or 16th day. The in-
terval between successive spawnings varies
from 8 days up to almost a year with a mode
of 15 days (Aronson, 1945). Approximately
two-thirds of the intervals are less than 29
days. Thus the 5- or 6-day score serves as
an intermediate record for the shorter inter-
spawning intervals, while the 15- or 16-day
score serves in the same capacity for the
longer intervals. Obviously some of the 15-
day records could not be taken because of
intervening ovipositions. Many of these ob-
servations served, moreover, as behavior
records for varying days before spawning.
Qualitative Description
of Reproductive Behavior.
In order to furnish the reader with the
proper background for the quantitative in-
vestigation, it is appropriate to present first
a general description of mating activities.
This account does not take into consideration
the question of the range of variability and
any exceptional items of behavior. Details
concerning many of the generalizations
made here will be considered in the next
section.
Certain of the behavioral patterns which
increase in their frequency of occurrence
prior to spawning and which lead up to the
acts of oviposition and fertilization gener-
ally are classified as courtship activity. Such
behavior appears to express the level of sex-
ual excitability of the given individual. In
accordance with the views of Huxley (1914,
1938), Howard (1929) and Marshall (1936),
it is assumed that courtship tends to hold
the pair together, and through mutual stim-
ulation may lead to a well synchronized
spawning. In the terms of Schneirla’s (1946)
discussion, such relationship may be thought
of as involving trophallactic processes, and
the temporal aspects of these interactivities
are of significance from the standpoint of
adaptive function. Tilapia eggs (PI. I, Fig.
2; PI. II, Figs. 3, 4), as well as those of other
oviparous teleosts, are shed in a flaccid state,
but rapidly become hard and turgid upon
entering the water. That is, they “water
harden” (Breder, 1943). Hence, to insure
fertilization the male must deposit his sperm
over the eggs within a very short time after
they are laid. An adequate synchronization
of the pair’s reproductive processes thus
appears to be critically important for effec-
tive species survival.
There follows a description of the early
courtship behavior of Tilapia macrocephala :
(1) The male and female approach each
other and suddenly dip their heads; or one
member of the pair lowers its head. This
1949]
Aronson: Reproductive Behavior in Tilapia macrocephala
137
behavior has been termed “head-nodding.”
(2) When one member of the pair approaches
the other, spreads its opercula and expands
its buccal pouch, we have called this act a
“throat-puff.” (3) The male or female ceases
swimming movements and the trunk muscu-
lature appears to quiver for a fraction of
a second. We have named this a “body-
quiver.” (4) When one member of the pair
slaps the other with its tail, this has been
called a “tail-slap.” Included in this category
were the frequent cases where tail-slapping
motions were quite distinct, but where ac-
tual contact with the partner was not made.
Closely associated with the courtship acts,
but displayed as well in many pairs through-
out the interspawning interval, is a mode of
behavior which we have termed “nipping.”
This occurs when a fish swims after its part-
ner, and •‘-hen with a sudden dart nips or bites
the body of its mate. Nipping also occurred
at times without a previous chase. Sometimes
the male and female may nip each other
simultaneously and occasionally they may
even lock jaws. Frequently observed cases in
which the pursuing fish darts ahead but
misses the lleeing opponent also have been
included under the general heading of nip-
ping.
in addition to its association with court-
ship ana spawning, nipping behavior appears
to be related to the establishment of social
hierarchies and the formation of territories.
These further relations of nipping have not
as yet been investigated.
The above-mentioned patterns of behavior
usually appear as quite distinct, but occa-
sionally they tend to merge into one another,
so that discretion on the part of the observer
is often called for in assigning a particular
courtship act to its proper category. Often-
times two or more courtship acts may be
displayed in rapid succession, a frequent
combination being a head-nod, throat-puff
and body-quiver. Another commonly occur-
ring combination is the throat-puff and tail-
slap. i l
Headers acquainted with the courtship be-
havior of other cichlid fishes will readily
recognize the resemblance of the Tilapia pat-
tern with those of other cichlids. Reactions
such as the body-quiver, the throat-puff and
the tail-slap in some form seem to be preva-
lent throughout the family.
Nest-building is first observed after in-
tensive courtship has been in progress for
several hours or days. Most of this activity
is conducted by the female who begins scoop-
ing up mouthfuls of gravel from scattered
locations in the bottom of the tank. Soon the
excavating is confined to one location, and
the construction of a nest begins. Often two
or more nests are constructed prior to the
spawning, and sometimes nests are built and
then destroyed during the construction of an
adjoining nest. The nests are most often
round or slightly oval. If the gravel sub-
stratum of the aquarium is not too thick,
the fish dig down to the slate bottom of the
tank. If, however, the depth of the gravel
is more than 2 or 3 cm., the nests do not
reach the slate.
In our study the length of time taken to
complete a nest varied considerably from as
little as one-half hour up to what appeared
to be several days. In the latter case, the nest-
building activity occurred in spurts, followed
by periods of quiescence. The rapid builders
generally worked continuously until the nest
was completed. A small amount of nest-build-
ing was accomplished by sweeping move-
ments of the tail and pectoral fins. However,
this has been interpreted as incidental to
swimming and balancing movements and not
directly related to nest building.
After the nest is more or less completed,
nest-building decreases considerably and is
supplanted to some extent by nest-cleaning
(PI. I, Fig. 1) in which the female, and occa-
sionally the male, pick continuously at the
bottom of the nest. Nest-building and nest-
cleaning are always interspersed among var-
ious phases of courtship responses.
As the nest takes form, the genital tubes
of the male and female become more promi-
nent. At this time, the male begins to swim
slowly over the nest, rubbing his genital
tube over the bottom. We have called this
“passing-nest.” Later when the female com-
pletes the nest, she likewise “passes-nest.”
Thus the pair circle around and around, rub-
bing their genital tubes over the nest. This
behavior is often interrupted by periods of
courtship, nest-building and nest-cleaning
activity. As soon as a fish starts passing-nest,
the genital tubes become fully erected, but
if this activity ceases for a time, the tubes
generally recede somewhat. This suggests
that mechanical stimulation is one factor
causing the erection of the genital tube.
Since fish not on the verge of spawning are
sometimes seen with partially extended geni-
tal tubes, other stimuli seem to be involved
in the partial erection of the genital tubes.
Courtship activity and hormones are sug-
gested as possible factors.
After the passing-nest behavior of the
male and female has been in progress for
some time, the female stops in the nest dur-
ing a “pass-nest,” and her body musculature
quivers for a second or two. This has been
called a “spawning-quiver.” Males also ex-
hibit spawning-quivers, but in the male these
responses generally are less distinct and are
seen less frequently.
Spawning-quivers were the final pre-
spawning acts and indicated the imminence
of the oviposition. During one of these
quivers a batch of approximately 10 to 20
eggs is extruded by the female in what we
have termed an “oviposition movement”
(PI. I, Fig. 2). The female then swims just
past the nest. She is followed by the male
who passes-nest, usually rubbing his genital
tube over the newly laid eggs and sometimes
exhibiting a spawning-quiver. This com-
plementary act has been termed a “fertili-
zation movement” (PI. II, Fig. 3). Sperm
138
Zoologica: New York Zoological Society
[34: 16
apparently are emitted at this time, although
no male products were apparent in our ob-
servations. The female then repeats her ovi-
position movement which is followed closely
by a second fertilization movement of the
male. After two to four such egg-laying cy-
cles, the female swims rapidly from the nest
for a distance of 15 to 30 cm., then faces the
nest. Meanwhile the male swims in the vicin-
ity of the nest for a minute or so, then rapidly
picks up the eggs with his mouth (PI. II,
Fig. 4; PI. Ill, Fig. 5).
In our investigations there were a few
exceptional cases where the male did not
pick up the eggs. Then the female nipped
and tail-slapped the male violently, exhibited
some courtship behavior, and finally after
10 to 20 minutes of this activity she picked
up the eggs and carried them in her mouth
(PI. Ill, Fig. 6). We found that at times,
because of the unequal sizes of the male and
female, all the eggs could not fit into the
male’s mouth, the female would pick up the
remaining eggs, but not until 10 to 20 min-
utes had elapsed.
Post-spawning activity consists for the
most part in poking around the nest, first by
the male, and later by the female as well. If
any of the eggs are missed when the orig-
inal spawn was picked up, they are almost
always recovered during this poking activity.
After several minutes, this poking be-
havior sometimes gives way to extensive
nipping and mouthing in which one member
of the pair, generally the female, soon dom-
inates and the other retreats into hiding.
The eggs hatch in 5 days and are carried
from 2 to 15 days further, during which
time the embryos continue to develop. The
young are released abruptly and most of
them are sufficiently developed at this time
to suggest that further parental care would
not be advantageous to them. Parental care
appears to end suddenly with the release of
the young. In fact, parents sometimes eat
their newly liberated offspring. Never did
we see the young swim back into the parental
mouth as has been described by some authors
for this and other Tilapia species, and which
is such a striking characteristic of the small
Egyptian mouthbreeders, Haplochromis
multicolor (Peters, 1937).
With this brief description of the mating
pattern we turn now to an analysis of the
actual counts made of the frequency of occur-
rence in relation to the time of spawning of
many of the behavioral acts described above.
Analysis of the Mating Pattern.
For the purpose of analyzing the data,
records were organized in the following
manner. For each observed spawning, the
time of appearance of the first batch of eggs
was designated as the zero minute. The 15-
minute period just prior to the zero minute
was called the first pre-spawning interval.
The period 15 minutes to 30 minutes prior
to the zero minute was named the second
pre-spawning interval. Twelve pre-spawning
intervals were similarly measured. Again
starting from the zero minute, the 15-min-
ute interval which followed was called the
first post-spawming interval, and a second
post-spawning interval was likewise meas-
ured. The number of times that the various
courtship and mating activities (tail-slaps,
passing-nest, etc.) were recorded during
each 15-minute pre- and post-spawning in-
terval was determined for both the males
and the females for all observed spawnings.
With data assembled in this manner, a series
of distributions was obtained (one of each
behavior pattern of both the male and female
for each interval) . Almost all of these were
strongly skewed to the right. The arithme-
tical mean obviously is a poor representation
of the central tendency of a markedly skewed
distribution. Medians are generally more
suitable, but a better method of treating
such data is to employ a transformation. In
many cases by use of the transformation X
= vi, binomial distributions were ootained
which could be treated as normal curves.13
These were checked by plotting cumulative
distributions on arithmetic probability pa-
per. However some of the distributions were
not normalized following the above trans-
formation, but approximated closely the
Poisson series. This was particularly true
with infrequently occurring items, where
the highest frequency was zero and where
the mean was considerably smaller than one.
Theoretical Poisson distributions were cal-
culated from Pearson’s (1914) tables and
the goodness of fit of the actual distributions
was tested by the chi-square method.
Still other distributions did not approxi-
mate either the normal or Poisson series. As
will be noted later, these were not subjected
to further statistical analysis.
For the normal distributions, the means,
range, theoretical range (M ± 3<0, standard
deviation and standard error of the mean
were calculated. These were plotted graphi-
cally in a time sequence, using the method
of comparing ranges and means developed
by Dice and Leraas (1936) as modified by
Simpson and Roe (1939). Since the length
of the pre- and post-spawning records varied
inadvertently for each spawning, the calcu-
lated means for each interval are based upon
a varying number of spawnings. Simpson
and Roe (1939) point out that the method
of Dice and Leraas is less reliable when the
frequencies and standard errors of the mean
vary greatly. Therefore, in critical cases
where the graphic method was suspected of
being inaccurate, P values were calculated.
The solid lines in Text-figs. 1-6, 8 and 9, in-
dicate females; the broken lines males. The
heavy vertical lines designate actual ranges
of the distributions. The adjacent light verti-
cal lines indicate theoretical ranges (M ±
3<0. The large dots represent the means,
while the short horizontal lines above and
IS The writer wishes to acknowledge the aid given by
Dr. Charles P. Winsor in suggesting the use of this
transformation.
1949]
Aronson: Reproductive Behavior in Tilapia macrocephala
139
below the means indicate the range of M
± 2<*m. When these ranges overlap, it may
generally be assumed that the differences
between the means are not significant. Con-
versely, if M ± do not overlap, the dif-
ferences between the means are significant.
The limitation of this method has already
been noted.
For the Poisson series, the theoretical
ranges were considered to run from zero to
that value of the variate having a relative
frequency of .003 or less. Means were com-
pared by the method described by Snedecor
(1946). Because of the asymmetrical nature
of the Poisson distribution the graphical
method of comparing means described above
cannot be used. Hence, in the following
graphs, the range of M ± 2<fm are not indi-
cated for the Poisson distributions.
Where the distributions did not conform
reasonably well to either a normal or Pois-
son series, only the means and actual ranges
are presented on the graphs.
The 15-minute records on the 5th or 6th
post-spawning day and on the 15th or 16th
day were treated in a similar manner. Since
a number of pairs spawned again within two
weeks after these observations were made, it
was possible to use some of these data as
records of behavior on the 2nd, 5th, 6th, 9th
and 12th pre-spawning days. Because of the
small number of cases, only means and
ranges are indicated graphically.
Throat-Puffs. As seen from the graph
in Text-fig. 1, the females (solid lines) ex-
hibited this behavior very rarely on the sev-
eral days they were observed before the
spawning. One female throat-puffed just once
on the 9th pre-spawning day. However, by
three hours before spawning, the throat-
puffing frequency had reached a rather high
level, which was maintained with little fluc-
tuation right up to the spawning. Imme-
diately after the egg laying, throat-puffing
activity increased sharply. To be sure that
this rise was not due to chance fluctuation,
the means of the first pre- and post-spawn-
ing intervals were compared and were found
to differ significantly (P<.01).
The males showed the throat-puffing be-
havior much less frequently than the females
(Text-fig. 1, broken lines). On the several
days the pairs were observed prior to the
spawning, no throat-puffing by the males was
seen. At three hours before the egg laying, a
low frequency of throat-puffing was recorded,
and this level was maintained up to the
spawning. When these data were treated by
utilizing the transformation X = V x as al-
ready described, the frequencies of male
throat-puffs were found to be distributed in
a Poisson fashion with zero the highest fre-
quency, an indication that the mean frequen-
cies were less than one. This raised the ques-
tion whether the males of just a few pairs
were responsible for the bulk of the throat-
puffing activity. A partial answer to this
question was obtained by selecting the 25
spawnings in which continuous records for
the first hour before spawning were avail-
able. It was found that during this hour,
64.0% of the males exhibited no throat-puff-
ing at all. This contrasts with the figure of
only 4.0% for the female. Similarly, in the
seven pairs where continuous records for
the first two pre-spawning hours were avail-
able, 57.1% of the males did not throat-nuff.
From this we may conclude that a consider-
NUMBER OF SPAWNINGS OBSERVED (TOTAL FREQUENCY)
15 15 14 15 13 16 18 21 31 39 52 66 60 51
35 36
DAYS BEFORE SPAWNING
(ONE 15 MIN. INTERVAL
ON EACH OF ABOVE DAYS)
15 MIN. INTERVALS BEFORE SPAWNING
15 MIN. INTERVALS
AFTER SPAWNING
DAYS AFTER SPAWNING
(ONE 15 MIN. INTERVAL
ON EACH OF ABOVE DAYS)
Text-fig. 1. Fluctuation in male and female throat-puffing behavior before and after
spawning.
140
Zoologica: New York Zoological Society
[34: 16
NUMBER OF SPAWNINGS OBSERVED (TOTAL FREQUENCY)
DAYS BEFORE SPAWNING
(ONE 15 MIN. INTERVAL
ON EACH OF ABOVE DAYS)
15 MIN. INTERVALS BEFORE SPAWNING
15 MIN INTERVALS
AFTER SPAWNING
DAYS AFTER SPAWNING
(ONE 15 MIN. INTERVAL
ON EACH OF ABOVE DAYS)
Text-fig. 2. Fluctuation in male and female body-quivering behavior before and after
spawning.
able number of males exhibited little or no
throat-puffing behavior prior to the spawn-
ing.
Immediately after the egg laying, throat-
puffing by the male was no longer observed.
Since the mouths of the males were now
filled to capacity with eggs, it seems better
to say that after the spawning throat-puffing
could not readily be identified. By the 5th or
6th post-spawning day, many of the males
were no longer carrying eggs, and now the
throat-puffing behavior had reached the pre-
spawning level.
Body-Quivers. Only an occasional body-
quiver was exhibited by the males and fe-
males on the several days they were observed
prior to the spawning (Text-fig. 2), but by
three hours before oviposition the body-
quivers (solid lines) were very frequent oc-
currences in the female. They remained at
this relatively constant level until the spawn-
ing, after which there was an abrupt rise.
The means of female body-quivers for the
first pre- and post-spawning intervals were
compared, and the latter were found to be
significantly higher (P = .021). At five days
after spawning the females’ body-quivering
had dropped far below the immediate pre-
spawning level, and remained the same dur-
ing the observation period on the 15th or
16th day.
The body-quiver frequency of the males at
NUMBER OF SPAWNINGS OBSERVED (TOTAL FREQUENCY)
Text-fig. 3, Fluctuation in male and female tail-slapping behavior before and after
spawning.
1949]
Aronson: Reproductive Behavior in Tilapia macrocephala
141
NUMBER OF SPAWNINGS OBSERVED ( TOTAL FREQUENCY)
DAYS BEFORE SPAWNING
(ONE IS MIN. INTERVAL
ON EACH OF ABOVE DAYS)
IS MIN. INTERVALS BEFORE SPAWNING
15 MIN. INTERVALS DAYS AFTER SPAWNING
AFTER SPAWNING (ONE 15 MIN. INTERVAL
ON EACH OF ABOVE DAYS)
Text-fig. 4. Fluctuation in male and female head-nodding behavior before and after
spawning.
three hours before spawning was consider-
ably less than the females’ and again these
data were best treated as Poisson distribu-
tions. The behavior remained at this level
until the fifth pre-spawning interval when
it started to slope off, reaching a minimum
at the first pre-spawning interval. However,
when the male body-quivers of the fifth and
first pre-spawning intervals were compared,
this slope appears not to be significant
(P>.10). On the 5th or 6th post-spawning
day, and on the 15th or 16th post-spawning
day, the body-quiver frequency of the males
was very close to that of the females.
Of the 25 spawnings in which continuous
records were available for one hour before
the spawning, 100% of the females and
80.0% of the males exhibited body-quivering
at least once. In the seven ovipositions in
which continuous two-hour pre-spawning
records were taken, 100% of the males gave
body-quivers at least once. Thus while this
courtship pattern is exhibited more frequent-
ly by the females, practically all males show
some body-quivering activity prior to the
egg laying.
Tail-Slaps. With the transformation pre-
viously described, the data of both the male
and female were found to be distributed in
a Poisson fashion, excepting the first two
post-spawning intervals of the female which
were normally distributed (Text-fig. 3). On
the several days prior to the spawning occa-
sional tail-slapping by the female was ob-
served, but at three hours before the egg
laying, the occurrence of this behavior had
increased considerably. This level was main-
tained until the spawning, when there was
another significant rise (P<.01) during the
first post-spawning period.
No tail-slapping by the male was observed
on the several days prior to the spawning,
and during the three-hour pre-spawning in-
terval, the frequency of tail-slaps remained
low with relatively little fluctuation. Ap-
proximately this same frequency was ob-
served during all the post-spawning obser-
vation periods.
Of the 25 pairs for which continuous rec-
ords for the first hour were obtained, 100%
of the females were recorded as tail-slapping
at least once, but only 48% of the males. In
the seven spawnings with two-hour continu-
ous pre-spawning records, 71.4% of the
males tail-slapped at least once.
Head-Nods. Following the transforma-
tion, head-nodding data for the female was
characterized by a large number of zero fre-
quencies and a small number of rather high
frequencies. These did not fit Poisson dis-
tributions. At times, head-nodding was not
clear cut and easy to recognize, and it is
possible that a considerable amount of head-
nodding passed unrecognized. Before the
spawning, head-nodding activity was quite
high (Text-fig. 4), at least for some of the
142
Zoologica: New York Zoological Society
[34: 16
DAYS BEFORE SPAWNING
(ONE 15 MIN. INTERVAL
ON EACH OF ABOVE DAYS)
15 MIN INTERVALS BEFORE SPAWNING
15 MIN INTERVALS
AFTER SPAWNING
35 36
DAYS AFTER SPAWNING
(ONE 15 MIN. INTERVAL
ON EACH OF ABOVE DAYS)
Text-fig.5. Fluctuation in male and female nipping behavior before and after spawning.
females, and there was a still further rise
after the egg laying.
The data for male behavior fit Poisson dis-
tributions quite closely. The low mean values
indicate that this behavior occurred rather
infrequently, and little fluctuation was no-
ticeable befoi'e or after the spawning.
Of the 25 ovipositions from which con-
tinuous records were taken for the first pre-
spawning hour, 84.0% of the females and
only 20.0% of the males exhibited head-
nodding at least once. Similarly, of the seven
pairs where two-hour records were available,
100% of the females and 28.6% of the males
head-nodded at least once. We may conclude
that head-nodding is a typical female activity
and that a small fraction of the males head-
nod occasionally.
Nips. Nipping data of both the male and
female were treated as Poisson distributions.
Both sexes displayed some nipping behavior
on the several days they were observed be-
fore the spawning (Text-fig. 5). During the
three-hour pre-spawning observation period,
approximately the same amount of nipping
was shown by both the males and females.
After the egg laying there was a significant
rise (P<.01) in the nipping frequency of
the females. The rise in female nipping dur-
ing the ninth pre-spawning interval may be
significant (P = .05), but it was mostly due
to a marked spurt of activity of a single
female. 1 1 \
Analysis of the 25 spawnings where com-
plete records for the first hour before spawn-
ing were taken showed that 68.0% of the
females and only 28.0% of the males exhib-
ited nipping behavior at least once. Where
two-hour continuous records were available,
100% of the females, and 57.1% of the males
engaged in some nipping activity. This sug-
gests that practically all of the females and
at least half of the males do some nipping
before spawning.
Nest-building Acts. On the several days
before spawning, nest-building by either the
male or female was not observed (Text-fig.
6), but by the third hour before oviposition,
female nest-building activity had reached
a rather high frequency. Since the presence
of a nest and the occurrence of nest-building
behavior was one of the more important
criteria used to determine the imminence of
spawning, and hence to ascertain the ap-
propriateness of starting the observation,
these data are likely to be somewhat biased
in favor of early nest-builders. Actually at
three hours before spawning, the average
nest-building activity of the female may not
be as high as that indicated by the data.
The drop indicated in the seventh pre-
spawning interval appears not to be signifi-
cant if the nest-building values of the fifth
and seventh intervals are compared (P =
.13). On the other hand, there is a noticeable
downward slope between the fifth and first
pre-spawning intervals, and when these two
intervals are compared, the difference was
found to be highly significant (P<.01). It
is clear that female nest-building behavior
drops off as the time for the laying of the
eggs approaches, and it is gradually super-
seded first by nest-cleaning behavior (which
is clearly distinguishable from nest-build-
ing), and secondly by nest-passing activity,
which, as we shall see in the next section, is
increasing as the nest-building frequency
is declining.
Following the oviposition episode, nest-
building activity dropped to a very low fig-
1949]
Aronson: Reproductive Behavior in Tilapia macrocephala
143
NUMBER OF SPAWNINGS OBSERVED (TOTAL FREQUENCY)
(ONE 15 MIN. INTERVAL AFTER SPAWNING (ONE 15 MIN. INTERVAL
ON EACH OF ABOVE DAYS) ON EACH OF ABOVE DAYS)
Text-fig. 6. Fluctuation in male and female nest-building behavior before and after
spawning.
ure. On the 5th or 6th day nest-building by
females was not observed, and on the 15th or
16th day only one nest-building act was
observed during the 15-minute observation
interval by one female out of thirty-six.
The data for the male was characterized
by high frequencies of zero values and low
frequencies of high values which neverthe-
less did not fit Poisson curves even after the
aforementioned transformation. In contrast
to the extensive nest-building activity of the
female, that of the male was quite limited.
Similar to the female, there is a downward
slope in activity between the fourth and first
pre-spawning intervals. However, the de-
cline is not very pronounced and its statisti-
cal validity could not be readilv ascertained.
After the spawning the males no longer
engaged in nest-building except for a single
male which on the 15th post-spawning day
nest-built four times during the observation
interval. This male was paired with the one
female, which was also observed to build a
nest during the 15-day post-spawning in-
terval. Two nests were present in the tank
and it is apparent that this pair was ap-
proaching another spawning cycle.
Turning again to the 25 soawnings with
continuous records for the first pre-spawning
hour, it was found that 100% of the females
and 72% of the malps engaged in nest-build-
ing at least once. Of the seven pairs w’th
continuous two-hour nre-spawning records,
71.4% of the males did some nest-building.
It is probable that only a small percentage
of males do not engage in any nest-building
prior to the spawning.
Fifteen nests built by ten pairs were
measured shortly after the spawnings. In
each case the fish were first carefully re-
moved without damaging the nests. Since
in many cases the nests were oval, two diam-
eters were taken, namely the short diameter,
and at right angles to this the long diameter.
The points used in these measurements are
indicated in Text-fig. 7. The average short
diameter was 11.8 cm., the average long
diameter 13.2 cm., and the average depth
2.6 cm. The female fish (which as shown
above are primarily responsible for the con-
struction of the nests) varied from 10.7 gr.
to 19.2 gr. with an average of 15.1 gr. The
males were slightly heavier, weighing on
the average 18.0 gr. There was no indication
from these limited data of a correlation be-
tween size of fish and size of nest.
Passing-nest. The earliest nest-passing
by the fema’e was recorded for the eleventh
pre-spawning interval, two and one-half
hours before the egg laying (Text-fig. 8).
Following the previously mentioned trans-
formation, the data for this interval fit a
Poisson distribution. The same is true for
the records of the 6th, 8th. 9th and 10th
intervals. The data for the 2nd to 5th and
the 7th intervals consisted of a series of zero
or verv low frQouencies and a smaller group
of relatively high values, vaguely suggesting
144
Zoologica: New York Zoological Society
f34: 16
Text-fig. 7. Diagrammatic section through typical Tilapia nest showing points used for
nest measurements.
bimodal curves. The data for nest-passing
for the first interval were normally distrib-
uted. These data indicate a gradually rising
frequency of nest-passing as the spawning
approached, with a sudden spurt of activity
during the second and first intervals. After
the egg laying, nest-passing activity of the
female dropped to almost zero and none was
recorded on the 5th or 6th and 15th or 16th
days.
The nest-passing data of the male were
normally distributed for the first interval.
The records for the remaining pre-spawning
intervals were highly skewed to the right
with highest frequencies zero, which, how-
ever, did not fit Poisson series. While no
nest-passing was recorded for the males on
the several days prior to the spawning, a
substantial amount of nest-passing was ob-
served by three hours before the egg laying.
This level of activity remained fairly con-
stant until the second interval when it started
to rise precipitously. However, during the
first pre-spawning interval, the nest-passing
activity of the female surpassed that of the
male for the first time (P<.01). After the
spawning, the frequency dropped to almost
zero and nest-passing was not observed on
the two post-spawning observation days.
Observations indicated that male and fe-
male nest-passing were not entirely inde-
pendent of each other, and calculation of the
(ONE 15 MIN. INTERVAL AFTER SPAWNING (ONE 15 MIN. INTERVAL
ON EACH OF ABOVE DAYS) ON EACH OF ABOVE DAYS)
Text-fig. 8. Fluctuation in male and female nest-passing behavior before and after
spawning.
1949]
Aronson: Reproductive Behavior in Tilapia macrocephala
145
NUMBER OF SPAWNINGS OBSERVED (TOTAL FREQUENCY)
15 IS 14 15 13 16 18 21 31 39 52 66 60 51
ON EACH OF ABOVE DAYS)
Text-fig. 9. Fluctuation in male and female spawning-quiver behavior before and after
spawning.
coefficient of correlation for the first pre-
spawning interval yielded an r of + .63. This
was transformed to Z = + .74 which is a
highly significant correlation (P<.01). The
nest-passing data for the remaining pre-
spawning intervals appear to be comparably
correlated, but the data do not readily lend
themselves to this type of statistical treat-
ment. Prior to the spawning, all of the males
and females exhibited some nest-passing
activity.
Spawning-quivers. This behavior was not
observed during the observation periods on
the several days before the spawning (Text-
fig. 9). Female spawning-quivers were first
seen during the 8th pre-spawning interval,
1 % to 2 hours before the egg laying. Their
frequency gradually increased and reached
a peak during the first pre-spawning inter-
val. There was a marked drop to almost zero
after the egg laying, and on the 5th or 6th
days and 15th or 16th days none were seen.
A few male spawning-quivers were in
evidence during the 12th pre-spawning in-
terval, and a low level of this behavior was
maintained until the second interval, 20 to
15 minutes before the egg laying, when there
occurred an abrupt rise in frequency which
terminated during the fii'st interval. During
the first post-spawning interval, a very few
spawning-quivers were recorded, and none
were seen thereafter. While a few of the
males exhibited spawning-quivers long be-
fore the females, the peak of spawning-
quiver activity of the females during the
first pre-spawning interval was considerably
higher than that of the males. However, the
data did not permit further statistical anal-
ysis.
Selecting the 25 spawnings for which con-
tinuous records for the first hour before
oviposition were available, it was observed
that 96.0% of the females and 72.0% of the
males showed at least one spawning-quiver
during this hour. Similarly, in the seven
spawnings for which two-hour continuous
records were available, 100% of the females
and 71.4% of the males were recorded as
performing at least one spawning-quiver
during these two hours. It thus appears that
while all the females showed this behavior,
in about a fourth of the males spawning-
quivers could not readily be distinguished
from nest-passing behavior. Since all of the
females exhibited spawning-quivers and be-
cause of the sharp peak in the frequency of
occurrence of this activity just before the
spawning, this behavior can also be employed
as an indicator of the approaching oviposi-
tion.
Oviposition and Fertilization. The be-
havioral patterns considered thus far were
recorded in terms of the number of times
that the acts occurred during a short interval
of time (i.e., 15 minutes), and the relative
infrequency of some of this behavior ac-
counts in part for the marked skewness of
the distribution curves. On the other hand,
the oviposition data which follow, and the
data concerning the reactions of the male
and female to the eggs and young, are based
upon the total frequency of the behavior dur-
ing a given spawning, and as might be an-
ticipated, these data approximated more
closely binomial distributions which could
be treated as normal curves.
A nest-passing act by the female during
which eggs were oviposited was counted as
a single oviposition movement. A nest-pass-
ing act by the male when eggs were present
in the nest was recorded as a fertilization
movement. Actual contact with the eggs was
not considered essential as a criterion for a
fertilization movement, although in most
146
Zoologica: New York Zoological Society
[34: 16
instances the male rubbed his genital tube
over some of the freshly laid eggs.
In 76 observed spawnings, the mean num-
ber of oviposition movements by the female
was 3.41 ± .13 with a standard deviation
of 1.1 movements. The mean number of male
fertilization movements was 3.46 ± .15 with
a standard deviation of 1.3. The difference
between the means is .05 ± .2 which indicates
clearly that the number of oviposition move-
ments of the female does not differ signifi-
cantly from the number of fertilization
movements of the male. Finally, there is a
significant positive correlation (r = + .48,
z = + .52, P calculated from <.01) be-
o z
tween these two activities, indicating that
the number of times the males fertilize the
eggs is partly related to, and probably de-
pendent upon the number of oviposition acts
of the female.
Parental Behavior. In a total of 76 ob-
served spawnings, the male alone picked up
the eggs in 62 cases (81.8%), the female
picked up the eggs in 6 cases (7.9%) while
both male and female participated in this
activity in 8 cases (10.5%).
The time after the beginning of oviposi-
tion for the male to start picking up eggs
varied from 20" to 2'10" with a mean of 1'3"
± 3" and a standard deviation of 23". For the
female this interval varied from 3'18" to
11T4" with a mean of 7'59" ± 1'22" and a
standard deviation of 3'17". The difference
between the means of these two distribu-
tions is obviously significant, and from these
data we may conclude that the male starts
to pick up the eggs as soon as the ovinosi-
tion has terminated, while the female allows
several minutes to elapse before she will
collect any of the eggs still available. Here
then is an apportioning mechanism which
results in the observed fact that the male
usually incubates the eggs, and the female
does so on infrequent occasions.
Eggs remain in the nest available to the
female under two circumstances. First, if the
male’s mouth is of insufficient size to con-
tain all of the eggs, a few may be left over
in the nest. This was the situation in case
1 (Table I) where a small male was paired
with a lar^e female. It was quite clear to the
observer that in this instance not all of the
eggs could fit into the male’s mouth. Sec-
ondly, eggs would be available to the female
when the male behaved atypically and did
not touch the eggs. In three of these cases
males had released broods seven to twelve
days previous to the spawnings, and this
may be a contributing factor causing the
lack of response of the males to the eggs.* 1 2 3 * * * * * * * * * * 14
In most instances where the eggs remained
in the nest for any length of time, the females
would chase, nip and court the males. In a
few cases, the latter retaliated and violent
14 On the other hand, recent observations by Aronson
and Holz-Tncker (unpublished) reveal that males in the
process of incubatine young may on occasion fertilize and
pick up a new batch of eggs.
TABLE I.
Time from the Beginning of Oviposition for
Eggs to Be Picked Up. Eight Cases Where
Both Male and Female Engaged in This
Activity.
Male
Case No. Start Finish
Female
Start Finish
1 25"— 50'
2 ll'OO"— 11'30'
3 3'00" — 4'00'
4 4'05" — 4'30'
5 2'05" — 5'00'
6 4'10" — 5'10'
7 6'24" — 10'30'
8 3'50" — 4'00'
7'15" —
6'50" — lO'lO"
2'30" — 4'00"
4'15" — 4 '30"
4'30" — 5'00"
4'1 0" — 4'55"
5'50" — 6'36"
1'25" — 3'30"
fighting ensued; as a result the nests were
destroyed and the eggs scattered. In cases
3, 6, and 7 (Table I), as soon as the female
began to pick up the eggs, the males fol-
lowed suit and both gathered up the eggs
simultaneously. The typical pattern when
eggs are left in the nest may be summarized
as follows:
(1) Immediately after the eggs are ovi-
posited and inseminated, there is
often a period of extreme quiescence
lasting a minute or two.
(2) This is followed by a period in which
the female appears to be inhibited
from approaching or touching the
eggs, but at the same time she seems
to be excited by the eggs, resulting in
active nipping, chasing and courting
of the male who sometimes responds
similarly.
(3) After several minutes the inhibitory
action of the eggs begins to diminish;
the female now approaches the nest,
pokes around the eggs, and eventually
picks them up. It was at this time
that several of the recalcitrant males
listed in Table I also approached the
nest and in a few cases started to pick
up eggs ahead of the female.
It is suggested that in cases 2 to 8
(Table I), chasing, nipping and courting
by the female, and also her poking around
the nest, sufficiently stimulated the male to
pick up the eggs, thereby completing the pat-
tern.
Once started, the length of time it took
for males to gather up the spawn varied from
2" to 1'45" with a mean of 13" ± 2" and a
standard deviation of 16". The high varia-
bility noted here is a result of two excep-
tional cases, one where the male took 1'7" and
in the other 1'45". In the remaining 60
spawnings, the time was less than 46". On
the other hand, six females took from 35"
to 3’ with a mean of 2'6" ± 22" and a stand-
ard deviation of 51.7". Thus we see that not
only does the female wait longer before
starting to pick up the eggs, but once started
she performs this task at a significantly
1949]
Aronson: Reproductive Behavior in Tilapia macrocephala
147
slower rate. In most cases, the males gath-
ered up the eggs rapidly and then kept pok-
ing around the nest for some time. Thus any
scattered eggs were quickly recovered. Some
of the females, on the other hand, would
pick up part of the eggs, swim away from
the nest, return and pick up more eggs, swim
away again, and so forth.
The egg-gathering records for the female
were necessarily limited by the behavior of
the males as noted above. It was therefore
considered appropriate to use for compar-
ison data from other experiments. Aronson
and Holz-Tucker (unpublished data) ob-
served the spawning of an isolated female
that could see another female in an adja-
cent tank. The ovipositing female took 24' to
start gathering up the eggs and the process
itself took 1'25" to complete. Similarly, we
observed the spawning of a completely iso-
lated female. This female did not start to
pick up the eggs for 13'5". She took 1'15" to
gather up most of the spawn, but left six
eggs which she did not pick up for another
eight minutes. A large number of normal
females were paired with males suffering
various types of brain lesions (Aronson, in
manuscript). In 27 spawnings, these females
took on the average 12'2" to start picking
up eggs and an average of 1'15" to complete
the job. Hence these data support our orig-
inal conclusions. However, it is likely that
in our first observations, the mean time for
the six females to start picking up the eggs
is somewhat low, while the time it took to
complete the pi’ocess may be a little too high.
It is of interest to note that in a few spawn-
ings the females seemed unable to carry all
of the eggs that they themselves had laid.
Both the male and female are capable of
successfully incubating the eggs. The per-
centage of spawnings in which young were
recovered at the termination of the incuba-
tory period is shown in Table II. Where the
spawnings were not witnessed, the slightly
higher score made by the males may be ac-
counted for by a possible failure to record
a few cases where the spawn was swallowed
immediately after the oviposition, and before
it was observed. The data for the third set
of observations are taken from a second ex-
periment, (Aronson, 1945) . These spawnings
were also not witnessed. In this experiment,
aquarium conditions were considerably im-
proved by the use of aquarium filters, thus
avoiding any changes of water. The young
were forcibly ejected from the parental
mouth on or about the tenth day after spawn-
ing and were counted immediately, thus
largely eliminating the possibility of losses
through cannibalism.
Even with these improved techniques, only
40% of the males released viable fry. Two
factors account for this low yield of young
by the males. First is the failure of the eggs
to be properly fertilized, or death of the em-
bryos, with subsequent disintegration of the
eggs. A second factor is swallowing the
spawn. The relative importance of these two
factors will now be considered.
If freshly laid unfertilized eggs are placed
in a jar of Tiiapia-conditioned water which
is kept at approximately 26° C., very few of
the eggs will show any gross signs of degen-
eration before 24 hours. Starting with the
second day, however, some of the eggs will
have decomposed, and in all cases few if any
intact eggs remain after the tenth day. As
to the variation in the length of the incuba-
tion period, it will be seen in columns 6 and
7 of Table III that females may carry unfer-
tilized eggs for as long as ten days, after
which time it may be assumed that all have
decomposed. Note particularly that in almost
80% 15 of these cases, the dead eggs were
retained longer than one day, and it is highly
probable that in many of these cases the
eggs were carried until they were quite de-
generate. It was not unusual to examine the
contents of a male’s or female’s mouth and
is Since the presence of incubating eggs was checked
only twice daily, spawn swallowed shortly after oviposition
mi-ht have been overlooked. Hence this figure may be a
little too high.
TABLE II.
Per Cent, of Spawnings in which Young Were Recovered.
No. of spawn-
ings in which
males incubated
eggs.
No. of males
releasing
young.
% of males
releasing
young.
No. of spawn-
ings in which
female incu-
bated eggs.
No. of
females
releasing
young.
% of
females
releasing
young.
Spawnings
witnessed.
68
22
32.4
14
3
21.4
Spawnings
not
witnessed.
86
33
38.8
2
0
0.0
Spawnings
not
witnessed,
2nd experi-
ment.
70
28
40.0
4
4
100.0
Variation in Length of Incubatory Period.
148
Zoologica: New York Zoological Society
[34: 16
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Aronson: Reproductive Behavior in Tilapia macrocephala
149
find that the fish had been carrying a mass
of badly decomposed eggs, or a mixture of
decaying eggs and viable embryos. From
the appearance of the eggs it was frequently
apparent that the fish had been carrying the
dead eggs for many days. In columns 2 and
3 are listed the durations of the incubatory
intervals for males carrying fertilized eggs.
It will be noted that in 17.7% of the cases,
the spawn was swallowed within 24 hours.
The indirect evidence cited above leads to
to the conclusion that these eggs were swal-
lowed because of some failure of the male’s
incubatory mechanism, whereby the male
failed to discriminate between eggs and food.
On the other hand, those egg masses which
were retained in the mouth for a number of
days were only swallowed when they had be-
come extensively decomposed. It should be
noted in passing that decomposed eggs are
never found in the tanks, and it is assumed
that they are swallowed rather than spat out.
The stomach contents of several males were
examined shortly after the egg layings, while
the males were carrying eggs. In two of these
cases a few eggs were also found in the
stomachs.
Columns 4 and 5 show that in 54.6% of
the spawnings in which eggs are picked up
by the females, they were swallowed within
24 hours and in most cases within the first
hour after spawning. Although these data
are limited, they indicate that the female’s
incubatory mechanism is not as dependable
as the male’s, and that the female fails to
distinguish eggs from food much more fre-
quently than does the male.
The length of incubation by the male in
cases where young are recovered is shown
in columns 8 and 9 of Table III. These data
fit closely a normal curve, and from them we
have determined a mean incubatory time of
13.8 ± .27 days with a standard deviation
of ± 2.6 days. This would give us a theoret-
ical range of 6 days to 22 days. The few
cases in which the female successfully reared
young fall well within this range.
Thus far, only the presence or absence of
eggs and developing embryos have been con-
sidered. Now, the relative sizes of the spawn
and brood will be examined. A new group
of pairs was established, and on the day of
or day after oviposition, the spawn was
ejected from the male’s mouth and was
counted. This count may be taken to repre-
sent fairly accurately the number of eggs
laid by the female, since, in most instances,
all of the eggs are picked up and few if any
are swallowed. Eighty females whose mean
weight was 7.15 ± .38 gr. deposited an av-
erage of 49.7 ± 1.96 eggs.
In a second group of 31 pairs in which the
average weight of the females was only
slightly less (5.6 ± .38 gr. ), the males were
allowed to incubate the eggs and the fry were
counted soon after their release. Here it was
found that the average brood size was only
23.9 ± 2.9 young. It was thought at first that
this smaller brood size might be attributed
to the lesser weights of our second group.
To examine this hypothesis the body weights
of the females that had just oviposited were
compared with number of eggs in the spawn.
A low order positive correlation was found,
which was probably significant (r = +.23 or
z = + .236 ; P calculated from — — = .05) . A
o z
similar comparison of the weights of fe-
males (determined immediately after ovi-
position) with the size of the brood that was
eventually recovered after being incubated
by the male partner did not yield a significant
correlation (r = + .10 or Z = + .10 ; P calcu-
2
lated from — — = .6). When two regression
o z
lines are plotted (calculated by the method
of least squares), one for the rise in num-
ber of eggs oviposited as body weight in-
creases, and a second for the change in num-
ber of young recovered as body weight
increases (Text-fig. 10), the relationship
involved becomes clearer. From these re-
gression lines in Text-fig. 10, it can readily
be seen that for females of the same body
weight, the number of young successfully
incubated is considerably smaller than the
number of eggs laid. This loss can best be
accounted for by the failure of some of the
eggs to be fertilized and by the death of
some of the embryos. Since brooding fish
have never been observed to spit out decom-
posed eggs or embryos, and since such ma-
terial has rarely been observed on the gravel
substrata of the aquaria, it is assumed that
the incubating fish somehow manages to sort
out and swallow this dead matter.
These data also indicate that while larger
females tend to lay a greater quantity of eggs
than smaller females, the number of fry
successfully brooded by the males remains
constant regardless of the weights of the
females and hence of the magnitude of the
spawn. Therefore the mortality of eggs and
embryos must be directly proportional to the
size of the female and hence to the number
of eggs laid. Since the larger females were
in most cases older, this difference might
be based upon an aging factor. It is also con-
ceivable that such increased mortality was
due to overcrowding in the male’s mouth
during incubation.
It is an interesting fact that incubating
Tilapia generally carry some gravel inter-
mingled with eggs. Of 63 fish examined on
the day or day after the egg laying, 95.2%
were carrying one or more pieces of gravel.
Generally between 25 and 50 pieces (com-
mercial grade No. 2) were found along with
the eggs, and occasionally the count went
well over 100. The possible significance of
this fact is not known at present. It is not
clear whether or not the gravel is picked up
accidently along with the eggs, and whether
this behavior bears some relation to the sur-
vival of the embryos. For example, it is pos-
sible that since the eggs and gravel are con-
tinuously churned around in the mouth, the
150
Zoologica: New York Zoological Society
[34: 16
o
Ui
<E
Text-fig. 10 Regression lines showing relation of body weights of fe-
males to number of eggs laid during each spawning and relation of body
weights of females that spawned, to number of young recovered imme-
diately after their release by the incubating males.
latter might serve to rub off fungi or ecto-
parasites from the developing fish.
It is important to note that there is con-
siderable variability in the number of days
that given parents may incubate their young.
It is therefore to be expected that at the
time of release, the broods carried longest
will be the ones furthest advanced in devel-
opment. This, generally speaking, is found
to be true. Thus, fry released in less than ten
days still have a large yolk sac and their
swimming activity is sporadic, whereas
young incubated for longer intervals show
little or no trace of the yolk sac, also their
swimming ability is developed to the stage
where they are well able to elude their ene-
mies if reasonable shelter is provided. A com-
plicating factor is that, as can be seen from
the few samples in Table IV, there is a con-
siderable difference in the rate at which the
fry grow within the parental mouths. Thus
the average size of a given brood incubated
for 22 days was barely larger than another
one incubated only 15 days. Similarly, a given
brood retained in the mouth for only 11 days
reached the same average size as another
brood incubated for 16 days. It is possible the
number of young in the brood may somehow
be related to growth rate; however, our lim-
ited data on this point in Table IV do not
suggest such a relationship. It is also of in-
terest to note that the variation within the
brood was quite low, the average coefficient
of variation (V) for nine broods being 3.7.
This state of affairs is in striking contrast
to the great variability (V = 15 ± 1.60)
which resulted when a brood was kept to-
gether in a stock tank from the time of re-
lease to maturity (Aronson and Holz-Tucker,
in manuscript).
Discussion.
In most vertebrates the characteristic
mating behavior patterns of the two sexes
are distinctly different. Thus in the rat, the
reproductive habits of which have been
analyzed most intensively, the typical pattern
of the estrous female, lordosis, is only occa-
sionally exhibited by the male (Beach, 1938,
1945). Similarly, the typical male pattern
of ear - wriggling, mounting and pelvic
thrusts is seldom seen in the female (Long
and Evans, 1922; Hemmingsen, 1933; Beach,
1938).
The sex difference in behavior generally
is very clear although relative rather than
absolute. Under special conditions males
may be induced to exhibit female-like be-
havior, and the reverse can also be accom-
plished (Beach, 1941). The conditions pro-
ducing such results often are very special in
nature. Thus for example, the well known
fact that estrous cows frequently exhibit
male-hke mounting behavior may very well
result from the almost universal custom of
segregating the cows from the bulls. Simi-
larly Beach and Rasquin (1942) explain in
part the high incidence of masculine behav-
ior in their female rats as the result of re-
peatedly testing two females together. These
authors are also aware of the possibility that
the females of their particular colony may
have been more active in a masculine direc-
tion than are females from most other stocks.
However, we are concerned with the fact that
disparity of behavior between sexes is gen-
eral among the vertebrates.
A survey of the literature indicates that
in reptiles a behavioral dichotomy of the
sexes appears to be the rule, and the writer’s
extensive investigations of the sexual be-
1949]
151
Aronson: Reproductive Behavior in Tilapia macrocephala
TABLE IV.
Relation Between Average Size1 of Young in
Brood and the Number of Days the Brood Was Incubated.
No. of days young
were carried.
No. of fry
in brood.
Average length
of fry1 (mm.).
Coefficient of
variation.
10
13
9.2 ± .08
2.9 ± ,572
11
19
10.6 ± .10
4.0 ±: .65
11
3
10.4
11
8
9.8 ± .09
2.8 ± .70
12
44
10.8 ± .06
3.9 ± .41
14
18
9.4 ± .11
5.1 ± .85
15
82
11.2 ± .04
3.3 ± .26
16
58
10.6 ± .06
4.4 ± .40
16
38
10.6 ± .06
3.7 ± .43
22
58
11.5 ± .05
3.4 ± .31
1 Length from tip of mouth to end of tail fin.
2 Standard error of the coefficient of variation.
havior of the tailless amphibia have shown
that in the Anura, male and female sexual
behavior are quite specific with only occa-
sional evidence of bisexual behavior (Noble
and Aronson, 1942; Aronson, 1943, 1943a,
1944).
In many birds these distinctions are less
clear. Thus in the pigeon, billing and bowing
are common to both sexes (Whitman, 1919),
and while it is usual for the female to squat
and for the male to mount, copulation not
infrequently occurs with the positions re-
versed (Carpenter, 1933).
While all vertebrates appear to possess the
neuromuscular and hormonal mechanisms
capable of eliciting most elements of the mat-
ing pattern of the opposite sex (Beach, 1942,
194 < ), morphological differences, particu-
larly in the genitalia, hormones and other
genetic factors, greatly limit the incidence,
completeness and effectiveness of such be-
havior. Thus in the majority of vertebrates,
behavior patterns characteristic of the male
or female are readily distinguished.
In contrast to this typical vertebrate con-
dition, TiLapia appears to represent an ex-
treme condition. None of the patterns of
reproductive behavior investigated are en-
tirely characteristic of either sex. Qualita-
tively, male and female courtship and spawn-
ing behavior are exactly alike. Even in the
acts of oviposition and fertilization, the
overt motor patterns are the same in both
sexes. Both fish swim slowly over the nest
and rub their genital tubes on the substra-
tum. The one observable difference occurs
when eggs extrude through the genital aper-
ture of the female, while the male’s genital
tube releases sperm, which, however, are in-
visible to the naked eye. It is only when the
frequencies of the various reproductive acts
are considered that behavioral differences
between the sexes become apparent. It is true
even so that in Tilapia sex differences in be-
havior depend in some cases on the time in-
terval before the spawning. Thus, as we have
found, the females exhibit much more court-
ship and do most of the nest-building. Males
do more nest-passing than the females at one
to two hours before the spawning, but at 15
minutes prior to oviposition we find this
l-elationship clearly reversed. After the
spawning, both qualitative similarity and
quantitative dichotomy are still in evidence.
Thus males wait on the average only 1.3'
before they start to pick up eggs; whereas
females require on an average 7'59". Also,
males pick up the eggs much faster than
the females, and are less prone to swallow
tfieir eggs.
In some of the patterns, as for example
head-nodding, the quantitative difference
between male and female frequency of the
act is sufficiently large that such behavior
could possibly be called a female pattern.
However, our data have shown that in 25
pairs where continuous records for the first
hour were available, almost one-third of the
males exhibited some head-nodding. It is
highly probable that if the entire span of the
pre-spawning sex behavior could be observed,
an even greater percentage of the males
would be found to perform a minimal amount
of this behavior. Bisexual or homosexual ac-
tivity has generally been thought of as a
recognizable intrusion of the characteristics
of behavior in one sex to a greater or lesser
extent into the behavior patterns character-
istic of the opposite sex. Such partial obser-
vations of sex dichotomy are known to occur
in a limited portion of the population or
under special circumstances such as segre-
gation. Thus we are justified in considering
bisexual or homosexual behavior a rather
restricted phenomonon in most vertebrates.
It follows that in the case of Tilapia none of
the patterns should be relegated to one par-
ticular sex as is generally done in the higher
vertebrates.
One might hypothesize that this situation
in Tilapia represents a primitive condition
in the evolution of reproductive behavior
patterns. This, however, is doubtful since
cichlids are a highly specialized family of
152
Zoological New York Zoological Society
[34: 16
teleosts, and on the other hand clearly recog-
nizable, sexually divergent mating patterns
are in evidence in some of the anatomically
more primitive hshes. While our study of the
described condition concerns Tilapia, it is
apparent from the literature that qualitative
similarities and quantitative differences
such as we find between male and female
sexual behavior in this species will be found
to a greater or lesser extent in all cichlid
fishes, and may well be true of several other
families of fish.
Rather than being a primitive condition,
we might view these behavior patterns of
Tilapia as adjustments (in an evolutionary
sense) to a specialized mode of reproduction
in which the similarity of the sex behavior
patterns has a considerable adaptive value.
For the post-spawning parental behavior this
point is fairly evident. If both sexes are cap-
able of rearing the young, there is less like-
lihood of lost or wasted spawn. If we look
upon the action of courtship as a mutually
stimulating and a synchronizing mechanism
as well as one which keeps the pair together,
one might expect the sexes to develop com-
parable mechanisms to accomplish the same
outcome when not limited by morphological
dissimilarities.
Not all behavioral disparities between the
sexes in Tilapia are readily understandable.
On several occasions, males exhibited con-
siderably more courtship activity than the
females of given pairs. In no case did such
excess lead to a spawning. One observation
showed a male in a stock tank courting at a
very high frequency as he swam around the
enclosure. In the same aquarium a female
was engaged in building a nest. Actually she
did not court in relation to this sexually ac-
tive male, but rather her activities had to
do with two other males in the territory.
The significance of excessive courtship by
males is not clear. It is possible that it repre-
sents the equivalent of bisexual behavior,
that is, of males behaving like females.
In this study we have found it convenient
to group certain activities such as the throat-
puff, body-quiver, tail-slap and head-nod
under the category of courtship, as distinct
from subsequent items of the reproductive
series, namely nest-building, nest-passing,
spawning-quivers, oviposition and fertiliza-
tion. However, no sharp line of demarcation
is implied between these. If we follow the
functional definition of courtship as previ-
ously stated (page 136), one cannot alto-
gether exclude the latter group of patterns
from the courtship category. However, a
separation on the basis of functional or adap-
tive significance seems to be in order. Thus
courtship behavior is mainly concerned with
the formation and maintenance of the pairs
while the latter activities have most to do
with the immediate preparation for spawn-
ing, as for example the building of the nest
and the physiological preparation for ovi-
position and fertilization. Also there are
indications of an organic separation. It is of
interest to note in this connection that in our
observations on the several days before
spawning, most of what we are terming
courtship activities wei'e seen at one time or
another, out the acts of nest-buiiding, pass-
mg-nest and spawning-quivers were never
recorded. Thus in general reproductive be-
havior tends to arise and function in group
iashion.
The quantitative records show that all of
the courtship responses of the female in-
creased in frequency directly after spawn-
ing. The same was true for nipping. Two
factors appear to be responsible for this in-
creased activity. The first is a physiological
change consequent to oviposition, and the
second is the presence of eggs. While we have
not attempted to analyze the relative influ-
ence of these two factors, several observa-
tions are of interest here. First, the observed
heightened courtship activity generally lasts
several hours and subsides gradually. Sec-
ondly, the activity continues long after the
eggs have been removed to the male’s mouth.
The freshly laid eggs might possibly release
some type of chemical stimulus, but the evi-
dence for this is not forthcoming. Moreover,
if newly oviposited eggs are presented to
males and females that have not spawned
recently, such eggs are generally eaten with-
in a short time, and they do not stimulate
either courtship or nipping. The effect of this
heightened activity is not apparent in most
of the spawnings, but in the few cases where
the males are slow in picking up the eggs,
the courting seems to attract the male to the
nest and stimulates him to gather up the
spawn.
It has long been recognized that certain
external morphological characteristics of an
animal, together with specific modes of be-
havior, may act as exciting stimuli to other
members of the species (and sometimes to
members of another species) for the media-
tion of specific behavioral responses. Lorenz
(1935, 1937) has developed this concept as a
cornerstone of his theory of instinctive be-
havior. The stimulus or group of related
stimuli bringing forth a reaction are called
“releasers,” the responding individual is
designated as the “companion.” Mutual in-
stinctive responses of companions are sharp-
ly separated from learning processes al-
though some modifications of the former are
recognized. Furthermore, according to this
view the release of every unconditioned re-
action is considered to be dependent on a
special central nervous mechanism which is
called the “innate releasing schema” (Lo-
renz, 1935) or “innate releasing mechanism”
(Tinbergen, 1939, 1948).
These hypotheses have become quite popu-
lar on the Continent. In this country they
have received some consideration by students
of bird behavior, but they are largely out of
tune with the findings and interpretations of
a large segment of the American experi-
mental psychologists (Lashley, 1938) who
in general have paid little attention to the
1949]
Aronson: Reproductive Behavior in Tilapia macrocephala
153
Lorenz movement. To say that a special “in-
nate releasing mechanism” exists for every
unconditioned reaction implies an extreme
localization of function within the brain, a
claim that is without special support in this
country. Here the more popular view is that
most responses are capable of being elicited
by a broad array of well separated stimuli
(.Beach, 1942, 1947) and are not exclusively
dependent upon any single stimulus or group
of stimuli. Moreover, there is here a growing
tendency to think of innate and learned fac-
tors as closely interlocked in their influence
on behavior (Schneirla, 1941, 1946) with the
view that in the higher vertebrates at least,
purely innate behavior patterns as entities
may be simply matters of a convenient term-
inology doubtfully related to reality. Lack
(1940) has criticized Lorenz’s view as being
too simple. He points out that in many cases
the designated releasers may not be the sole
characters that bring forth the response.
Rand (1941) has been to date Lorenz’s sever-
est critic. According to Rand, the releasing
characters are by far too limited, and the
releasers and responses are mostly unidenti-
fied. The reality of releasers has accordingly
not been demonstrated but remains presump-
tive. Actually the experimental analytical
approach to behavior is not only untried by
Lorenz, but its validity is denied. Finally,
according to Rand, there is in Lorenz’s treat-
ment a negativistic approach which denies
the possibility of ever being able to elucidate
the fundamentals of behavior.
Tinbergen (1939) has modified Lorenz’s
hypotheses in several respects. First, releas-
ers are called “signals” or later “sign stimu-
li” and are subdivided into releasing stimuli
and directing stimuli. More important, Tin-
bergen recognizes a closer relation than does
Lorenz between innate responses released
and modifiable factors such as learning, en-
docrine reactions and neural processes (sum-
mation, conditioning and “higher mental
processes”) . Most important is Tinbergen’s
recognition of the validity of the experi-
mental approach, and his attempts, mostly
by means of artifacts and models, to demon-
strate releasers in this manner. Even so, it
must be emphasized that Tinbergen sees
releasers as very specific and limited mor-
phological and behavioral characters which
during the unfolding of a complex pattern
of response will hold to a relatively rigid
sequence.
Seitz, a follower of Lorenz and Tinbergen,
has analyzed the behavior of two related
cichlid fishes, namely a small Egyptian
mouthbreeder, Astatotilapia strigigena
(1940) and the jewel fish, Hemichromis
bimaculatus (1942) in terms of the releaser
concept. Seitz recognizes whole series of very
specific releasers which call forth specific
responses and which lead in an orderly man-
ner to the spawning. These he has summa-
rized in schematic form (1940, p. 82; 1942,
p. 100). Thus, in Astatotilapia, the presence
of a female releases a change to mating color-
ation in the male, and this change in its turn
releases a slight but not significant color
change in the female. The presence of the
female also releases a mode of behavior
caned by Beitz an introductory presentation
wiiicn in turn brings forth a passive response
in the female. This in turn releases a com-
plex of movement and color change called
oy Seitz a “Fegebalz” (lit., sweeping court-
snip dance). This Fegebalz of the male
releases a following reaction on the part of
the female, which in its turn releases circular
swimming in the male around the spawning
site. The circular swimming then releases
a strong following reaction of the female to
the spawning site which in turn brings forth
a response whereby the male slips under the
female. This releases circling movements in
the female which in turn release the same
movements in the male. The circling move-
ments of the male call forth additional cir-
cling movements by the female. These release
the oviposition movements and the latter re-
lease the fertilization movements of the male.
Our experiments were not designed to test
the releaser concept and this discussion is
not intended as a critique thereof. However,
we were interested in learning to what extent
our data would or would not support the re-
leaser hypothesis or fit into that pattern of
thought.
The significant correlation between male
and female nest-passing behavior appeared
most likely to fit in witn this concept if we
were to assume that nest-passing of male
and female released a like behavior in the
opposite sex. However, we had on record any
number of cases where the females were very
quiescent, exhibiting little or no courtship
or pre-spawning behavior of any kind, and
yet the males nest-passed consistently. Of
course, the nest itself might be a releaser
of nest-passing, but this would contradict
a large portion of our data where nest-build-
ing by the female and the presence of a well-
formed nest was not followed by nest-passing
on the part of the male. Similarly in the
spawning of the completely isolated female
previously referred to, the order of magni-
tude of nest-passing behavior was well with-
in the range of variability of our control
pairs. Yet there was nothing in that situation
which could be considered a releaser. In an
attempt to follow the lead of Seitz, we could
possibly view the various courtship patterns
previously described as releasers. For ex-
ample, the approach-throat-puff of the fe-
male might be construed as a releaser of
similar behavior by the male which in turn
might be thought of as releasing female
nest-building behavior. This may be espe-
cially so since an approach-throat-puff by
a female was often followed by a similar pat-
tern in the male, and soon thereafter the
female turned to the construction of the nest.
However, no consistent pattern of this type
was in evidence. Female throat-puffs were
also followed by almost any of the other
courtship patterns or by no particular re-
154
Zoologica: New York Zoological Society
f34: 16
sponses of the male. Again, female nest-
building was sometimes preceded by the
throat-puffs but often by head-nods, tail-
slaps or body-quivers. It is recognized that
in general observation, that is in “just
watching” these fish, one could easily gain
the impression that certain acts are in effect
releasers, and others a response to these
releasers. However, when observational tech-
nique involves an orderly and complete
quantification of response according to con-
dition of occurrence, the data do not support
such an interpretation.
We are inclined to view the courtship and
pre-spawning items of behavior together
with the territory and nest as having a gen-
eral stimulatory effect upon the other
member of the pair which would tend to
raise the level of sexual excitability in the
latter. Or, to put the matter in another way,
the given conditions may serve to lower the
threshold for the elicitation of various
courtship and pre-spawning patterns. Here
the particular response obtained would de-
pend upon a whole complex of factors includ-
ing the neural threshold, the immediate
topographic relation to the partner, the
territory, the nest and other environmental
conditions, as well as the internal physiologi-
cal balance of the individual at the moment.
In this sort of system, no specific releasing
stimuli may be properly postulated. For
example, a series of weak or only partially
eltective tail-slaps by the female might bring
forth a response in the male similar to one
very effective approach-throat-puff. More-
over, as the general level of excitability of
both members of the pair increased, there
would be a gradual shift in the statistical
probability of the elicitation of a given type
of response. In other words, throat-puffing
during the early stages of the pre-spawning
history of a pair might bring forth addi-
tional throat-puffing or other phases of the
courtship, while later, such behavior might
elicit return to the nest or nest-passing. As
spawning approached, nest-passing behavior
of one member of the pair was often followed
by like behavior of the other member of the
pair, but this was often interrupted by some
of the early phases of courtship such as
head-nodding and tail-slapping. In many of
the records, interruptions of the smooth flow
of passing-nest and spawning-quivers were
noted within minutes of the actual oviposi-
tion. While these data do not altogether con-
tradict the releaser concept, it is believed
that these findings can be more satisfactorily
understood by adhering to a considerably
more generalized interpretation of the com-
plexity and effectiveness of the stimuli than
the “releaser concept” implies.
Seitz (1940) and Tinbergen (1948) in
their discussion of releasers refer to the
“rule of heterogeneous summation” which
states that the release of a given behavior
pattern may result from the summation of
several different stimuli. Tinbergen also
emphasizes that “high internal motivation
may cause the reactor to respond to all
objects offering the minimum adequate ex-
ternal stimulation.” Finally, Tinbergen ob-
serves that some releasers have a general
excitatory influence, rather than to direct
the reactor’s response. If these three prin-
ciples noted here are sufficiently expanded,
some of the major objections to the releaser
concept are thus overcome, and except for the
sharp lines drawn between instinct and
learning processes, we begin to arrive at a
common ground for the understanding of the
nature of sexual behavior.
Summary and Conclusion.
Qualitative descriptions and quantitative
measurements of the patterns of reproduc-
tive behavior of the African mouthbreeding
cichlid fish, Tilapia macrocephala (Bleekerj,
have been presented. These patterns have
been grouped into three categories. The first,
namely courtship, includes head-nods, ap-
proach-throat-puffs, body-quivers and tail-
slaps. Most of the females exhibited these
courtship items during the observation
periods, and at a relatively high frequency.
The males performed these courtship acts
at a considerably lower frequency. A high
percentage of males showed some tail-slaps
and body-quivers, and it is believed that if
the entire spans of the pre-spawning activity
of the pairs could have been observed, all of
the males would have performed these court-
ship patterns. On the other hand, it appears
that a measurable portion of males do not
head-nod or approach-throat-puff prior to
the spawning.
It is hypothesized that courtship behavior
is an expression of the level of excitability
of the individual. It may be thought of as a
trophallactic process which through mutual
stimulation serves to regulate the behavioral
activities and physiological processes of the
male and female so that well synchronized
spawnings result.
Nipping, which is closely related to court-
ship and which also appears to be mutually
stimulating, was performed equally by the
male and female before the spawning, but
nipping on the part of the female rises
sharply directly after oviposition. Similar
post-spawning increases on the part of the
female were noted for all of the courtship
patterns. It is suggested that the physiologi-
cal changes following oviposition plus the
presence of eggs are the factors responsible
for this heightened activity. During the
inter-spawning interval, a low level of court-
ship is in evidence, especially on the part of
the females.
The second group of reproductive patterns
includes those acts which are concerned with
the immediate preparation for spawning.
Included here are nest-building, nest-pass-
ing, spawning-quivers, oviposition move-
ments and the act of fertilization. Consider-
ably more nest-building is exhibited by the
1949]
Aronson: Reproductive Behavior in Tilapia macrocephala
155
female than by the male, but it is likely that
all males do some nest-building before every
spawning. With passing-nest and spawning-
quivers the frequency is somewhat higher
for the males an hour or so before spawning,
but at 15 minutes before spawning this rela-
tionship is clearly reversed, with the females
at the height of their nest-passing and
spawning-quivers. A significant correlation
between male and female nest-passing during
the first pre-spawning interval suggests that
this behavior is mutually stimulating. The
mean number of oviposition movements of
the female did not differ significantly from
the mean number of fertilization acts of the
male. Moreover these behavior patterns are
highly correlated, suggesting that the num-
ber of times the male fertilizes the eggs is
partly related to and probably dependent
upon the number of oviposition movements
of the female. In contrast with the courtship
patterns, behavioral items in the present
category were not observed during the inter-
spawning interval.
The third category of reproductive acts
are those associated with the care of eggs
and young. Males start picking up eggs on an
average of 1'3" from the beginning of ovipo-
sition. Females, if given the opportunity,
took on the average 7’59". This is the appor-
tioning mechanism whereby males usually
incubate the eggs, and females do so only
on infrequent or special occasions. Similar
quantitative differences were found in other
phases of the parental pattern. Thus females
gather up the spawn more slowly and are
more prone to swallow the eggs.
A low order positive correlation was found
between the size of the female and the num-
ber of eggs laid during a given spawning.
Since brood size shows no correlation with
the size of the female, it is concluded that a
greater mortality occurs in the larger
broods. Incubating fish generally carry some
gravel intermingled with the spawn, but it
was not clear whether this bore any relation
to the survival of the embryos.
In the maioritv of vertebrates there are
distinct qualitative differences between the
patterns of reproductive behavior of the
male and female. While both seres have the
neuromuscular mechanism capable of elicit-
ing both the male and female patterns, bi-
sexual or homosexual behavior is limited
and p-cuorsllv appears under special condi-
tions. Tilavia are exceptional in this respect
insofar as there are no distinct oualitative
differences between male and female in their
sexual sctivit'es. TTowever. there are marked
quantitative differences in all of the natterns.
Several nrevious investigators have ana-
Ivzed cichl’d matin? behavior in terms of
I-overiz’s releaser concent Tt is felt that even
in tVio evnanded and modified form nresented
bv Tinbor<rDm th’’s concent is sti’l too re-
stricted to form an adennate basis for the
analysis of Tilapia reproductive behavior.
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158
Zoologica : New York Zoological Society
[34: 16: 1949]
EXPLANATION OF THE PLATES.
Plate I.
Fig. 1. Male cleaning nest. X .5
Fig. 2. Oviposition. The male is behind the
female, waiting for her to move along
so that he can pass over and fertilize
the eggs. X .5
Plate II.
Fig. 3. The male is fertilizing the eggs while
the female is circling the nest. By the
time fertilization was completed the
female was directly behind the male,
ready to lay a second round of eggs. X
.5
Fig. 4. Male picking up the eggs. All of the
eggs were gathered up in less than one
minute. X .5
Plate III.
Fig. 5. Male carrying eggs. X .7
(Photo, by S. C. Dunton, N. Y. Zool. Soc.).
Fig. 6. In special circumstances the female
may carry the spawn. An egg can be
seen at the tip of the open mouth of
the female. X .5
Addendum.
When this report was in page proof an article by Alfred Seitz (1948) — Verglei-
chende Verhaltensstudien an Buntbarschen (Cichlidae). — Zeitschrift fur Tier-
psychologie, 6 (22) : 202-235, was received from Germany. Here Seitz analyzes
fighting and courtship behavior in two cichlid species, Tilapia heudeloti and Tilapia
natalensis, in accordance with the theory of instinctive movements of Konrad
Lorenz. On page 134 of the present paper we have noted the very close similarities
of T. heudeloti and T. macrocephala; they may in fact be varieties or subspecies.
However, the pictures of T. heudeloti presented by Seitz, the descriptions of the ex-
ternal morphology, particularly coloration, as well as the descriptions of court-
ship and fighting behavior, all suggest that he was dealing with a very different
fish. It is not possible at this time to comment further on Seitz’s paper, nor do we
wish to venture any opinions concerning the complex problems of cichlid taxonomy,
except to suggest to the reader who may wish to compare Seitz’s paper with the
present report that the T. heudeloti of Seitz and our T. macrocephala are perhaps
very different species. — L.R.A.
ARONSON.
PLATE I.
FIG. 1.
FIG. 2.
AN ANALYSIS OF THE REPRODUCTIVE BEHAVIOR OF THE
MOUTHBREEDING CICHLID FISH. TILAPIA MACROCEPHALA (BLEEKER).
ARONSON.
PLATE II.
FIG. 3.
FIG. 4.
AN ANALYSIS OF THE REPRODUCTIVE BEHAVIOR OF THE
MOUTHBREEDING CICHLID FISH, TILAPIA MACROCEPHALA (BLEEKER).
ARONSON
PLATE III
FIG. 6.
AN ANALYSIS OF THE REPRODUCTIVE BEHAVIOR OF THE
MOUTHBREEDING CICHLID FISH. TILAPIA MACROCEPHALA (BLEEKER)
Crane: Salticid Spiders: Analysis of Display
159
17.
Comparative Biology of Salticid Spiders at Rancho Grande, Venezuela.
Part IV. An Analysis of Display.1
Jocelyn Crane. '
Research Zoologist, Department of Tropical Research,
New York Zoological Society.
(Plate I; Text-figures 1-9).
[This is one of a series of papers resulting
from the 45th, 46th and 47th Expeditions of the
Department of Tropical Research of the New
York Zoological Society, made during 1945, 1946
and 1948 under the direction of Dr. William
Beebe, with headquarters at Rancho Grande in
the National Park of Aragua, Venezuela. The
expeditions were made possible through the gen-
erous cooperation of the National Government
of Venezuela and of the Creole Petroleum Cor-
poration.
[The characteristics of the research area are
in brief as follows: Rancho Grande is located
in north-central Venezuela (10° 21' N. Lat.,
67° 41' W. Long.), 80 kilometers west of Cara-
cas, at an elevation of 1,100 meters in the un-
disturbed montane rain forest which covers this
part of the Caribbean range of the Andes. Ad-
jacent ecological zones include seasonal forest,
savanna, thorn woodland, cactus scrub, the
fresh-water lake of Valencia and various marine
littoral zones. The Rancho Grande area is gen-
erally subtropical, being uniformly cool and
damp throughout the year because of the preva-
lence of the mountain cloud cap. The dry season
extends from January into April. The average
humidity during the expeditions, including parts
of both dry and wet seasons, was 92.4%; the
average temperature during the same period
was 18° C.; the average annual rainfall over a
five-year period was 174 cm. The flora is marked
by an abundance of mosses, ferns and epiphytes
of many kinds, as well as a few gigantic trees.
For further details, see Beebe and Crane,
Zoologica, Vol. 32, No. 5, 1947. Unless otherwise
stated, the specimens discussed in the present
paper were taken in the montane cloud forest
zone, within a radius of one kilometer of Rancho
Grande.]
I. Introduction 159
II. Purpose 160
III. Materials and Methods 160
IV. Survey of Salticid Display 161
A. Form and Variety 161
B. Historical Review 170
C. Bases for Disagreements 172
D. Prognosis of Evolutionary Pattern 173
V. Factdrs in Display 175
A. Factors of the Internal Releasing Mechanism 176
1. Age 176
2. Fluctuating Epigamic Rhythm 176
1 Contribution No. 858, Department of Tropical Research,
New York Zoological Society.
This paper was awarded Honorable Mention in the A.
Cressy Morrison Prize Competition of the New York
Academy of Sciences for 1949.
3. Hunger and Thirst 179
4. Fatigue and Overstimulation 180
5. Attention 180
B. Factors of the External Releasing and Direct-
ing Mechanisms 180
1. Physical Environment 180
2. Sensory Elements and Sign Stimuli 181
a. Tactile Perceptions 181
b. Chemoperception 181
i. Chemotaxis 181
ii. Distance Chemoperception 182
c. Vision 184
i. Vision as a Primary Stimulus 184
ii. Motion 185
iii. Distance 187
iv. Size 189
v. Form 190
vi. Pattern, Intensity and Color 192
VI. Innate Releasing and Directive Mechanisms.... 199
A. Courtship 199
B. Threat Display 200
C. Comparison and Comment 200
VII. Behavior Related to Display 202
A. Displacement or Substitute Behavior 202
B. Dominance 202
C. Sociality 202
D. Territory 203
VIII. Functions of Display 203
A. Courtship 203
B. Threat 204
IX. Evolutionary Aspects of Display 205
A. Hypothetical Phylogeny 206
B. Origins of Display Motions 206
C. Relation of Secondary Sexual Characters to
Display 208
D. Sexual Dimorphism and Display 209
E. Climate and Display 209
F. Displays as Specific Barriers 210
X. Summary 211
XI. References 213
I. INTRODUCTION.
This is the fourth of a series of papers
dealing with the salticid spiders of Rancho
Grande, Venezuela. Part I (Crane, 1948.1)
discussed the taxonomy and life histories of
three species of Corythalia, Part II (1948.2)
described methods of study and Part III
(1949) dealt with systematics and behavior
in eight new species of various groups of
genera, as a basis for this and subsequent
sections. The present paper on display, while
an independent unit, will be followed by sec-
tions on comparative post-embryological
development and on evolutionary trends,
which must, to some extent, be anticipated
in the present paper.
My deep appreciation goes to Dr. William
Beebe for his continued advice, encourage-
ment and patience during the progress of
this study. Special thanks are due Miss
160
Zoologica: New York Zoological Society
[34 : 17
Louise A. Moore and Mr. Douglas G. Boyden
for their work on the drawings and dia-
grams, respectively.
II. PURPOSE.
The purpose of this paper is two-fold:
first, to analyze the innate releasing mech-
anism of display behavior in salticids; sec-
ond, through a comparison of display and
related behavior in widely scattered genera,
to shed light on evolutionary trends within
the family.
III. MATERIAL AND METHODS.
The observational and experimental ma-
terial in this paper has been drawn primarily
from about fifteen species of salticid spiders
which are common around Rancho Grande,
Venezuela. They were carefully selected
from among some thirty-five species in the
same locality, in which display behavior
was to some degree observed, on the follow-
ing bases:
1. They represent a wide divergence of
salticid forms, belonging to seven relatively
well-defined groups of genera. These groups,
although they have so far defied satisfac-
tory divisions in keys to include all their
borderline genera, are still rather generally
recognized as composed of naturally related
forms. For convenience, Petrunkevitch’s
(.1928, 1939) practice of terming the groups
“subfamilies” is followed, although their
unequal values and the usual difficulties of
salticid systematics are fully recognized.
Since the tracing of a pattern of relation-
ships is one of the purposes of this paper,
groupings must be attempted. The genera
chosen are relatively non-controversial, as
are, in their broad outlines, the subfamilies
to which they are referred ; borderline cases
have purposely been omitted. The selected
forms are fairly typical members of the
following subfamilies designated by Pet-
runkevitch: Marpissinae, Synagelinae, Den-
dryphantinae, Magoninae, Hyllinae and
Plexippinae. The Lyssomanes group, how-
ever, in spite of its aberrant characters
will be given only subfamilial status. These
terms will be used throughout, in order to
simplify correlation by future workers, in
spite of questionable validity or priority in
any of the old, type genera. Each subfamily
corresponds to a group in Chickering’s 1946
arrangement, based on Panamanian genera,
except that in the present paper the plex-
ippinids and hyllinids will be kept separate,
largely in deference to basic behavior dif-
ferences. From these limitations, it will be
seen that the study cannot be expected to
help in practical fashion in the determina-
tion of fine points of relationship among
puzzling genera; rather, it is hoped only to
clarify some of the broad outlines of the
family’s organization.
2. The second criterion of selection has
been that each subfamily have a northern as
well as neotropical distribution. This has
a double advantage, since some comparable
display behavior has in each case been al-
ready recorded and future comparisons will
be facilitated.
3. In three genera, Sassacus, Phiale and
Corythalia, intrageneric behavior patterns
could be compared.
The thirteen species listed in Table I
form the paper’s foundation, in the sense
that most of the observations and experi-
ments were performed on them. Ten of them
were described as new in Parts I and III of
this study (1948.1, 1949) ; the remaining
three are well known in the tropics. For
further comparison, a few casual references
will be made to the following Rancho Grande
species which are systematically undeter-
mined: Magoninae: Hypaeus sp., Dep’t. of
Tropical Research Cat. No. 45138; Hyllinae:
Phiale sp., Cat. No. 481408; Plexippinae:
Eustiromastix sp., Cat. No. 45110; Plexip-
pinae : a genus, probably new, near Capidava,
Cat. No. 4586.
In Table II an effort has been made to
correlate the, relevant observations made
upon northern species of the same subfam-
ilies by previous investigators. Complete
spider display references to 1939 are given
in Bonnet’s bibliography (1945, p. 718).
Kaston (1936) and Bristowe (1941) give
excellent selected references on the subject.
The general methods of observation and
experiment at Rancho Grande have already
been recorded (Part II, 1948.2). Special
information regarding particular experi-
ments will be presented in the body of this
paper. See also PI. I, Fig. 2.
It remains to remark on the usage of
several terms. The phrase “epigamic display”
has been advocated by a number of behav-
iorists to replace “courtship,” a word which
has an undesirable historical burden of an-
thropomorphism. In the study of salticids, the
choice is complicated by the wide range of
distinctness of male-female, female-male
and inter-male display; a general term
seems necessary to cover the entire field.
Again, “aposematic display” has been pro-
posed to designate inter-male encounters;
in the literature, however, this term is used
so widely to cover displays against potential
predators — a type of display which is not
known to occur in salticids — that a more
restricted name appears desirable. There-
fore, for present purposes, the following
terminology has been adopted: Epigamic
display refers to special behavior of either
sex which is normally used in any sexual
situation, including encounters between
males. Courtship applies to those forms of
epigamic display which normally precede
copulation. Threat display is used for all
inter-male display, whether or not it is vis-
ually distinct from courtship, and whether
or not occasional contact fighting occurs.
“Ornamentation” is used as a convenient
term to cover special colors, patterns, tufts,
brushes, etc., which, conspicuous to human
eyes, are specializations possibly connected
1949] Crane: Salticid Spiders: Analysis of Display 161
TABLE I.
Subfamily acc. to
Petrunkevitch (1939)
Group acc. to
Chickering (1946)
Name
Lyssomaninae
Marpissinae
Synagelinae
Dendryphantinae
Dendryphantinae
Dendryphantinae
Magoninae
Hyllinae
Hyllinae
Plexippinae
Plexippinae
Plexippinae
Plexippinae
Lyssomaninae
Marpissa Group
Zuniga Group
Metaphidippus Group
Metaphidippus Group
Metaphidippus Group
Amycus Group
Phiale Group
Phiale Group
Phiale Group
Phiale Group
Phiale Group
Phiale Group
Lyssomanes bradyspilus Crane, 1949
Menemerus bivittatus (Dufour, 1831)
Semorina brachychelyne Crane, 1949
Ashtabula furcillata Crane, 1949
Sassacus flavicinctus Crane, 1949
Sassacus ocellatus Crane, 1949
Mago dentichelis Crane, 1949
Phiale dybowskii (Taczanowski, 1871)
Phiale flammea Crane, 1949
Plexippus paykullii (Audouin, 1827)
Corythalia chalcea Crane, 1948.1
Corythalia fulgipedia Crane, 1948.1
Corythalia xanthopa Crane, 1948.1
I with epigamic displays. It and similar words
are put in quotation marks when it seems
i wise to emphasize their non-anthropomor-
! phic use. The quotations will simultaneously
act as reminders, as advocated by Korzybski
(1948), of the every-day, inexact connota-
tions of the terms; another of his safety
j devices, the frequent use of etc., is employed,
not to disguise ignorance, but to emphasize
the incomplete state of our knowledge.
The terms sign stimulus, releaser, director,
innate releasing mechanism, stimulus con-
figuration and heterogeneous summation are
used with the meanings attached to them in
Tinbergen’s general discussion, in English,
of social releasers (1948).
Throughout the present paper, the terms
reaction and response are used interchange-
ably to indicate overt display behavior dis-
cernible to the human observer.
Kaston’s (1948) nomenclature is followed
in all references to species of salticids oc-
curring in the United States.
IV. SURVEY OF SALTICID DISPLAY.
A. Form and Variety.
The general habits and life histories of
all salticids are similar and typical of many
hunting spiders; they spin no snares, but
stalk and leap upon their prey, which they
eat promptly; the male copulates in the dor-
sal position; the female guards the egg co-
coon until the emergence of the spiderlings.
In the development of their eyes, however,
salticids are unique even among long-sighted
hunting spiders. In company with the latter,
notably the lycosids, and correlated with
the acute vision, the salticids have evolved
a courtship display which is primarily vis-
ual. This is in contrast to the behavior of
short-sighted forms, such as the web-spin-
ning epeirids, in which courtship depends
on senses other than vision.
As in all groups, there is considerable
intra-family variation in details of behav-
ior, particularly in the form and care of
the cocoons, in jumping methods, in pre-
ferred ecological niches and in physiolog-
ical tolerances of various kinds. A most
important series of differences is based upon
an apparently unreported correlation be-
tween visual acuity and chemical dependence
on the one hand, and method of locomotion
on the other. Because of its basic connection
with display behavior, this subject will be
considered in detail further on (p. 173).
The outstanding directions of development
within the family are shown in the very
remarkable variety of epigamic display
patterns and in the apparently correlated
ornamentation of the spiders. All salticids,
before inserting the sperm-charged palps
into the epigynum of the female, perform
some sort of preliminary motions, the form
of which, under natural conditions, is a fixed
part of the behavior pattern for the species.
These movements range from brief and
simple elevations of the first legs to com-
plicated and prolonged sequences involving
most of the other appendages.
In some salticids there are clear visual
distinctions between courtship and inter-
male threat display, which often employ not
only distinctive motions but also different
appendages. In each kind of epigamic dis-
play, when complete, there are always dis-
tinguishable two major stages. Specific
display differences, as well as distinctions
between courtship and threat, always lie
principally in Stage I. Stage II, which im-
mediately precedes either copulation or true
fighting as the case may be, is very similar
throughout the family; it consists in extend-
ing the first legs forward, parallel to each
other and slightly elevated. (PI. I, Fig. 1).
Although salticid display is dominantly
visual, the role of other senses is proving
of considerable secondary importance. UnT.
foi’tunately, because of the limitations of
human sense organs and the lack of appro-
priate instruments, specific display descrip-
tions must still be largely confined to the
reporting of gross visual components. For
instance, possible changes in chemical stim-
uli, which may be emitted in sequence during
display, are still necessarily overlooked.
Table II presents correlated data on the
displays recorded within the subfamilies
162
Zoological Neiv York Zoological Society
[34: 17
TABLE II.
Comparative Display Data in Seven Subfamilies of Salticid Spiders.
Name
R3tP«
Display References
Courtship Posture (Stage I)
Marpissa undata
(De Geer)
Marpissa rumpfi.
Scop.
Menemerus bivittatus
(Dufour)
Hyctia nivoyi Luc.
Hyctia pikei Peckham
Salticus cingulatus Panz.
Salticus scenicus (Linn.)
Synageles Venator Luc.
Semorina brachychelyne
Crane
Gertschia noxiosa (Hentz)
Peckhamia picata (Hentz)
Marpissinae
Peckham, ’89, p. 43; fig.
(“Marptusa familiaris” )
(U.S.A.)
Bristowe, ’29, p. 330 352.
Bristowe, ’41, p. 480; fig.
(England)
Crane
(Unpublished observations)
(Venezuela)
Berland, ’27, p. 15; fig.
Bristowe, ’29, p. 329 ; fig.
Bristowe, ’41, p. 484; fig.
(Europe)
Kaston, ’48, p. 455; fig.
(U.S.A.)
Bristowe, ’29, p. 332.
(England)
Peckham, ’89, p. 39.
(“Epiblemum scenicum ”)
Gerhardt, ’21, p. 131.
(“ Epiblemum scenicum”)
Monterosso, ’24, p. 1.
Bristowe, ’29, p. 332.
Bristowe, ’41, p. 480, 499 ; fig.
(Europe & U.S.A.)
Synagelinae
Bristowe, ’41, p. 485.
(England)
Crane, ’49, p. 37.
(Venezuela)
Kaston, ’48, p. 451; fig.
(U.S.A.)
Peckham, ’89, p. 43.
Peckham, ’90, p. 121 ; fig.
(U.S.A.)
Carapace high.
1st & 2nd legs raised
stretched laterally.
Abdomen pendent.
slightll
Carapace high.
1st legs raised vertically, parallel.!
Abdomen raised.
hope
Carapace low.
1st legs raised slightly, stretchcjtions;
antero-laterally ; 2nd legs e:|torint
tended forward.
Abdomen level.
1st legs stretched forward @
45° ; raised & lowered.
Abdomen often bent sideways.
Dendryphantinae) .
Z ‘I
(c1
Carapace low.
1st legs raised vertically @
45°.
2nd legs extended forward.
Abdomen raised.
Z cl
Similar to S. scenicus.
ikecc
1st legs slightly raised, stretchy]
laterally. 1 to si
Chelicerae opened. j\;0 B
Palps stretched laterally.
1st legs stretched forward, parallel!
Abdomen wriggled or waved froirl
side to side, sometimes raised ~
little.
1st legs stretched forward, @ 90
Z, slightly raised.
Abdomen raised; tends to remain
high with increasing excitement.
1st legs braced laterally.
Abdomen raised vertically & swung |
from side to side.
Carapace high.
1st legs on ground, bent, convex sur- 1
face forward.
Abdomen raised vertically.
Spider sways from side to side.
1949]
Crane: Salticid Spiders: Analysis of Display
163
TABLE II (cont.)
Comparative Display Data in Seven Subfamilies of Salticid Spiders.
Principal Apparent
ireat Posture & Fighting Morphological Locomotion & Remarks
Adaptations to Display
Marpissinae (cont.)
Slight enlargement basal half
of 1st & 2nd legs.
$ vibrates palps.
aws opened; pressed against
those of opponent.
1st legs elongated, enlarged,
darkened.
Jusually $$ pay no attention to
each other; no distinctive mo-
tions ; rarely courtship to mir-
ror image in closed vial.
1st legs elongated, enlarged,
darkened.
Runs; jumps only on prey or
to cross gaps.
1st legs little used in walking.
$ vibrates pale palps.
1st legs elongated, enlarged,
darkened.
1st legs elongated, enlarged,
darkened.
Abdomen boldly striped.
1st legs not used in walking,
extended forward & raised
to clear ground.
Chelicerae elongated, en-
larged, darkened.
1st legs slightly elongated.
jike courtship, but jaws opened
wider and palps not stretched
to sides.
No mirror display. Crane obs.
U.S.A.)
Chelicerae elongated, en-
larged, darkened.
1st legs slightly elongated.
Jerky walk; hops when pur-
sued. (Crane obs. U.S.A.) .
Synagelinae (cont.)
«
1st legs enlarged.
apparently no inter-male dis-
play. No mirror display.
1st legs elongated, enlarged,
darkened.
Abdomen dark with 2 dorsal
white spots.
Runs; jumps only on prey or
to cross gaps.
1st legs stretched to front,
parallel ; do not touch
ground.
9 vibrates palps.
1st legs elongated, enlarged,
darkened.
Abdomen dark with dorsal
white bands.
Runs slowly, irregularly, like
ant.
(Emerton, 1909).
1st legs elongated, enlarged,
darkened.
Abdomen dark with dorsal
white bands.
164
Zoologica : New York Zoological Society
[34: 17 | iJi
TABLE II. (cont.)
Comparative Display Data in Seven Subfamilies of Salticid Spiders.
Name
Display References
Courtship Posture (Stage I)
It'."
Lyssomanes bradyspilus
Crane
Ashtabula furcillata
Crane
Hentzia mitrata (Hentz)
Icius elegans (Hentz)
Sassacus ocellatus Crane
Lyssomaninae
Crane, '49, p. 34; see also
Text-fig. 8, this paper.
(Venezuela)
Dendryphantinae
Crane, ’49, p. 41; fig.
(Venezuela)
Peckham, ’89, p. 49; fig.
(“Icius mitratus”)
(U.S.A.)
Peckham, 89, p. 46.
(“ Dendryphantes elegans”)
(U.S.A.)
Crane, ’49, p. 45.
(Venezuela)
Carapace high, occasionally bobbcj
No legs raised; 1st 3 prs. bracl
forward.
Palps sometimes tap ground.
Abdomen pendent, twitched.
Prolonged posing in this positic
as —
Rate of retinal motions within A!
are accelerated.
Carapace high.
1st legs stretched laterally, held
right A to body.
Abdomen twisted to side.
Carapace high. ■
1st legs stretched laterally, wave*
up & down.
Palps jerk; later are quiet.
Abdomen turned to side.
1st legs waved “in way that remind
one of a wind-mill”. Later, r«
volves on tip-toe.
Carapace scarcely elevated.
1st legs stretched up & out at righ
A to each other.
Chelicerae closed.
Palps vibrated occasionally.
Abdomen sometimes twisted slightl
to side, held motionless.
Sassacus flavicinctus
Crane
Metaphidippus protervus
(Walck.) and/or
M. galathea (Walck.)
Paraphidippus marginatus
(Walck.)
Crane, ’49, p. 41.
(Venezuela)
Peckham, ’89, p. 45; fig.
( “Dendryphantes capitatus ”)
(U.S.A.)
Peckham, ’89, p. 50; fig.
(“ Philaeus militaris")
(U.S.A.)
Carapace moderately high.
1st legs stretched up at wide A t(
each other.
Chelicerae stretched sideways, bu
closed.
Palps stretched sideways.
Abdomen trailed inconspicuously
from side to side.
Later : Carapace & abdomen slowly
rocked from side to side.
Carapace low.
1st legs stretched forward, close tc
ground, slightly curved with tips
turned up.
Palps given circular movement.
Later, lies on side, legs still extend-
ed.
1st legs raised, curved toward each
other, tips nearly meeting.
Palps moved up and down.
1949]
Crane : Salticid Spiders: Analysis of Display
165
TABLE II (cont.)
Comparative Display Data in Seven Subfamilies of Salticid Spiders.
I Threat Posture & Fighting
1
Principal Apparent
Morphological
Adaptations to Display
Locomotion & Remarks
Lyssomaninae (cont.)
Threat display rarely induced;
posture as in courtship, but no
bobbing, no abdominal twitch-
ing, no acceleration of retinal
motions.
Fighting & mirror display ab-
sent.
Distal portion of black retina
contrasts with green sur-
roundings.
Chelicerae elongated but not
specially displayed.
Runs in short spurts. Jumps
only in final pounce on prey.
Palps pendent, pat ground
during pauses.
1st legs take part in locomo-
tion.
2 sags to side when watching
display; muscular activity
in eyes increase as in $.
Dendryphantinae (cont.)
Apparently no inter-male dis-
play. No mirror display.
1st legs elongated, enlarged,
darkened, fringed.
Abdomen with lateral white
stripe bounded by irides-
cence.
Runs; jumps only on prey or
to cross gaps.
1st legs unused in progress,
held forward, they & palps
tap ground; waved in air
in pauses.
2 vibrates palps.
Same as courtship.
Chases rivals away.
1st legs elongated, fringed.
Abdomen with lateral white
stripe bounded by darker.
2 “indifferent”; sometimes at-
tacks $.
Stage I: Similar to courtship,
except chelicerae partly un-
sheathed, & no distinct ab-
dominal twisting.
Stage II: Actual fighting, 1st
legs raised vertically; chel-
icerae wide open; palps ex-
tended laterally. Clinching
frequent, with occasional in-
jury; no deaths seen.
Stage I: Like courtship except
no rocking.
Stage II : (Rare) . Chelicerae un-
sheathed, extended forward ;
1st legs raised vertically;
clinching; no injuries seen. ’
Occasional mirror display.
Carapace with frontal tufts.
1st legs elongated; tibia
fringed, black-spotted.
Both sexes covered with iri-
descent scales.
1st legs elongated, enlarged,
darkened, with white scales.
Chelicerae elongate, enlarged
darkened.
Clypeus with white band.
Abdomen with large, sub-
distal, white-barred black
spots.
1st legs elongated, enlarged,
darkened.
Chelicerae elongate, en-
larged, darkened.
Clypeus with yellow band.
2 irritable, prone to attack;
later raises abdomen or
turns it sideways.
Runs; jumps only to catch
prey & cross gaps.
1st legs held forward, scarce-
ly touch ground; these &
palps wave in air & tap
ground, during pauses.
2 once did weak mutual dis-
play.
Runs; jumps occasionally
when pursued, as well as to
catch prey & cross gaps.
Palpates ground & waves 1st
legs less than S. ocellatus.
Distinct from courtship. 1st legs
raised. Sparring, chasing &
clinching without injury.
Distinct from courtship.
1st legs raised.
Palps vibrated.
Abdomen dragged to either side.
Chasing and clinching without
injury.
1st legs elongated, enlarged.
White markings on dark
palps & clypeus.
1st legs elongated, enlarged.
Chelicerae elongated.
Runs, but hops freely when
pursued, as well as to catch
prey to cross gaps.
Palps often jerk.
1st legs not waved. Take
some part in walking.
(Crane obs. U.S.A.)
$$ guard immature 22 until
molt.
166 Zoologica: New York Zoological Society [34: 17
TABLE II. (cont.)
Comparative Display Data in Seven Subfamilies of Salticid Spiders.
Name
Phidippus audax (Hentz)
Phidippus clarus
Keyserling
Phidippus purpuratus
Keyserling
Phidippus whitmanii
Peckham
Display References
Dendryphantinae (cont.)
Peckham, ’89, p. 45.
(“ Phidippus morsitans”)
Kaston, ’36, p. 120; fig.
(U.S.A.)
Kaston, ’36, p. 118; fig.
(U.S.A.)
Kaston, ’36, p. 121; fig.
(U.S.A.)
Peckham, ’89, p. 44.
(“Phidippus rufus ”)
(U.S.A.)
Courtship Posture (Stage I)
Carapace moderately high.
1st legs raised in 2 jerks, stretched
obliquely out at Z of 45° to
ground; later waved alternately.
Palps raised and lowered rapidly
when close to $.
Carapace moderately high.
1st legs stretched out, the femur
obliquely up, other segments
paralleling ground.
Palps sometimes widely spread.
Abdomen now & then moved from
side to side.
Carapace high.
1st legs stretched out, held higher
than in clarus & audax ; waved.
Palps held wide apart, parallel with
each other, occasionally drummed
on ground.
Abdomen dropped.
Carapace high.
1st legs stretched forward & up,
crossed at tips.
Palps held wide apart, parallel with
2nd legs.
Abdomen dropped.
Sways during advance.
Hyllinae
Evarcha falcata Bl. Bristowe, ’29, p. 333.
Bristowe, ’41, p. 480.
(England)
Homann, ’28, p. 249; fig.
(“E. blanchardi”)
(Germany)
Philaeus chrysops Poda Berland, ’14, p. 116.
Thomas, ’29, p. 267.
Bonnet, ’33, p. 139; fig.
(France)
Some waving of 1st legs & twitching
of abdomen, but tends to leap on
9 with almost no display.
Carapace high.
1st legs raised & waved up & down.
Palps vibrated.
Abdomen slightly elevated.
Courtship sometimes almost absent.
Leaps on 2 suddenly.
Phiale fiammea Crane Crane, ’49, p. 48.
(Venezuela)
Carapace high.
1st legs raised at 45° Z to each other
& ground.
Palps vibrated irregularly.
Abdomen lowered.
Carapace rocked from side to side.
Later, sinks low, 1st legs stretched
to front.
Crane: Salticid Spiders: Analysis of Display
167
1949]
TABLE II (cont.)
Comparative Display Data in Seven Subfamilies of Salticid Spiders.
Threat Posture & Fighting
I
Principal Apparent
Morphological
Adaptations to Display
Locomotion & Remarks
J
Dendryphantxnae (cont.)
Like courtship, until opponents
close; then chelicerae & fangs
opened wide; some 9 fighting.
1st legs elongated, enlarged,
with white fringes and
scales.
Palps with white scales.
Both sexes savage, prone to
eat mate.
As in P. audax.
1st legs elongated, enlarged,
lightly fringed.
Palps with white band.
1st legs elongated, enlarged,
heavily fringed.
Clypeus with white fringe.
1st legs elongated, enlarged, Hopping well developed,
heavily fringed.
Hyllinae (cont.)
1st legs elongated, enlarged,
darkened ; palps pale, white-
haired.
Differs from courtship in quiet
palps & absence of low stage.
At close quarters, chelicerae
open, 1st legs spread widely,
often touch opponent’s; no in-
juries.
Displays freely to mirror.
1st legs elongated, darkened
with light tufts & scales.
1st legs elongated, darkened.
Clypeus & palps with huffy
yellow bands against black.
Excellent visual acuity; hops
skilfully when pursued.
Berland & Thomas saw fre-
quent leg-waving when
spiders were alone in field
and in clean boxes; Bonnet
saw waving only before 99.
Basically a runner although
jumps readily over gaps;
1st legs little used in walk-
ing, habitually waved dur-
ing pauses; true also of 99
& young.
9 raises 1st legs & vibrates
palps during display.
168 Zoologica: New York Zoological Society [34: 17
TABLE II. (cont.)
Comparative Display Data in Seven Subfamilies of Salticid Spiders.
Name
Display References Courtship Posture ( Stage I )
Plexippus paykullii Aud.
Plexippinae
Crane (unpublished obs.) ;
PI. I, fig. 1, this paper.
(Venezuela)
Eustiromastix sp.
Crane (unpublished obs.).
(Venezuela)
Carapace very high.
1st legs stretched forward, up & out
at varying wide &, higher wit!
excitement; no waving.
Palps quiet.
Abdomen pendent.
Prolonged posing in display posi-
tion.
Carapace high.
1st legs raised antero-laterally @
Z of 45° to ground; sometimes
jerked, & raised higher.
Palps vibrate.
Abdomen horizontal.
Some posing in display position.
Saitis barbipes Sim.
Corythalia xanthopa
Crane
Berland, ’23, p. 206.
Berland, ’27, p. 16; fig.
(France)
Crane, ’48.1, p. 35; figs, (also,
see footnote, this paper, p.
183).
(Venezuela)
3rd legs wave.
Carapace moderately high.
Palps hanging, motionless.
Abdomen slightly below horizontal.
Stage la: Side-to-side rocking, all
feet on ground.
Stage lb: 1st legs stretched for-
ward, straight, parallel @ Z of
45° to ground.
No posing in display position.
Corythalia chalcea Crane Crane, ’48.1, p. 21; figs.
(Venezuela)
Carapace moderately high.
Palps hanging, motionless.
Abdomen about horizontal.
3rd legs stretched out, waved up &
down in unison above the hori-
zontal.
No posing in display position.
Corythalia fulgipedia
Crane
Crane, ’48.1, p. 28; figs.
(Venezuela)
Carapace progressively lowered.
Palps jerked in unison.
Abdomen about horizontal.
1st, 2nd, & 3rd legs stretched far out
to sides, the 3rd legs being slight-
ly intermittently raised & vi-
brated up & down in unison with
body.
No posing in display position.
1949] Crane: Salticid Spiders : Analysis of Display 169
TABLE II (cont.)
Comparative Display Data in Seven Subfamilies of Salticid Spiders.
Principal Apparent
Threat Posture & Fighting Morphological Locomotion & Remarks
Adaptations to Display
Plexippinae (cont.)
No inter-male displays seen. 1st legs elongated, enlarged,
Mirror response not tested. darkened.
White clypeus band against
black.
Hops often even during ordi-
nary progress.
9 completely passive.
Scarcely developed ; rarely, brief
threat similar to early court-
ship but abdomen lowered &
palps quiet.
Occasional brief mirror display,
sometimes ending in attack.
Same as courtship.
Completely distinct from court-
ship.
Carapace very high.
Palps motionless, flexed, their
yellow scales continuing band
of clypeus.
Abdomen pendent.
2nd, 3rd & 4th legs off ground,
raised successively higher.
Prolonged posing. No fighting.
Displays freely to mirror.
Distinct from courtship, although
same legs used.
Differs in waving 3rd legs below
horizontal, often in an arch;
prolonged posing.
Palps flexed & stiff.
No fighting.
Displays freely to mirror.
Distinct from courtship, although
similar in very first phase, &
same legs used throughout.
Whereas in courtship carapace
sinks lower, in threat it rises
higher. At peak, all legs are
drawn close in & stretch up,
2nd tarsi leave ground & 3rd
legs are raised slightly; body
rocks from side to side ; posing
with 3rd legs arched frequent.
Palps jerked at beginning, later
flexed & stiff.
No fighting.
Displays freely to mirror.
1st legs slightly longer, thick-
er, blacker than others.
3rd legs elongated, com-
pressed, fringed.
Palps & clypeus with yellow
bands.
2nd, 3rd & 4th legs fringed,
compressed, with iridescent
patches; 3rd & 4th legs
elongated.
1st, 2nd & 3rd legs fringed,
compressed, with iridescent
patches ; 3rd legs elongated.
1st, 2nd & 3rd legs fringed,
compressed, with iridescent
patches strongly developed ;
3rd legs elongated.
Palps with white patches.
Hops often even during ordi-
nary progress.
1st legs take active part in
walking.
Palps quiet.
9 extremely savage both be-
fore & after mating; often
kills $.
Performs display motions
even when alone in clean
boxes.
Ordinary progress consists
largely of hopping.
1st legs take active part in
walking.
Palps quiet.
9 not aggressive ; when watch-
ing display, sits high with
braced legs until receptive,
then crouches.
Locomotion as in xanthopa.
9 sometimes aggressive, may
do some form of reciprocal
display.
Locomotion as in xanthopa.
9 more aggressive than most
salticids, although not seen
to injure $; when stimu-
lated performs rough, re-
ciprocal display.
170 Zoologica: New York Zoological Society [34:17
TABLE II. (cont.)
Comparative Display Data in Seven Subfamilies of Salticid Spi-ders.
Name
Display References
Courtship Posture (Stage I)
Magoninae
Maevia vittata (Hentz)
Peckham, ’89, p. 54; fig.
(“ Astia vittata ”)
(Wisconsin, U.S.A.)
Painter, ’13, p. 625.
(Connecticut, U.S.A.)
Displays of the 2 forms differ.
Gray form : Carapace high ; 1st legs
raised, waved, palps stretched
sideways; abdomen down; later,
whole body low, 1st & 2nd legs
forward, tips touching; palp^
forward. (Peckham, Wise.).
Low stage precedes high stage, i
(Painter, Conn.).
Black, tufted form: Carapace very
high ; 1st legs raised & waved, or
held high in pose.
(Peckham & Painter).
Mago dentichelis Crane
Crane, ’49, p. 48; fig.
(Venezuela)
Carapace slightly raised.
1st legs stretched sideways &
slightly forward ; waved alter-
nately up and down or held in
pose.
2nd tarsi touch ground, far out &
slightly forward.
Abdomen sometimes vibrated.
Ballus depressus Walk.
Bristowe, ’31, p. 1409; fig.
Bristowe, ’41, p. 484; fig.
(England)
All legs on ground, drawn in, sways
body from side to side.
under consideration in this paper. Text-fig.
1 and Table III will give some idea of the
variety of display motions throughout the
family. All displays unite at least several
of the listed movements, and many involve
all of the major headings except that of
retinal motion, which so far appears to op-
erate as part of the signal mechanism only
in Lyssomanes.
No less varied are the forms of “orna-
mentation” in the family; these occur too
often on displayed appendages to be ex-
plained only by chance mutations, metabolic
processes, other behavior patterns, etc. They
include many specializations of size, shape,
hair concentration, scalation and pigmenta-
tion, several forms usually occurring in a
single spider. For example, the first legs,
which always enter at least into Stage II
courtships, are often elongated, thickened
and much darker than the others; in ad-
dition, the palps, which frequently vibrate
during display, in many species show patches
of shiny white scales. Again, in Corythalia
xanthopa, the fourth legs, elevated only in
threat display, are prominantly fringed;
in the related C. chalcea, in which they never
leave the ground, fringes are absent. Many
other examples occur in the literature, of
which the Peckham’s original series are
among the best (1889, 1890). The striking
point is that any appendage specially ex-
hibited in epigamic display usually shows
some differentiation which is to human eyes
conspicuous.
However, two other obvious points must
be kept in mind. First, not all parts active
in display are so ornamented; an example
is the brown abdomen of Semorina, relieved
only with small, pale inconstant spots. Sec-
ond, not all parts conspicuously ornamented
(again, to human eyes) play any demon-
strable part whatsoever in display. Examples
are the bright scarlet abdomen of Phiale
flammea and the striking carapace bands
of various Phiale and Corythalia.
B. Historical Review.
The history of the study of salticid display
reflects the changing ideas of evolutionists.
The Peckhams pioneered both in the record-
ing and explanation of epigamic display and
of the apparently correlated ornamentation
(1889, 1890). According to them, the phe-
nomena were due to direct Darwinian sexual
selection; the females, consciously or un-
consciously, were affected by the perform-
ances, colors and beauty of the males, select-
ing the most handsome or at least the most
striking. Male battles were initiated both for
simple possession of a mate and as sham con-
tests for the females’ edification. The Peck-
1949]
Crane: Salticid Spiders: Analysis of Display
171
TABLE II. (cont.)
Comparative Display Data in Seven Subfamilies of Salticid Spiders.
Threat Posture & Fighting
Principal Apparent
Morphological
Adaptations to Display
Locomotion & Remarks
Magoninae (cont.)
Wave 1st legs at each other but
“quarrels . . . harmless”
(Peckham).
Displays to mirror.
Tufted form is black anteri-
orly, has cephalic tufts.
Peckham & Painter disagree
on selection value of di-
morphism.
99 irritable; fight each other.
Stage I: Similar to courtship.
Stage II: 1st legs waved higher,
almost meet overhead; when
lowered, 1st & 2nd tarsi rub
together. Rarely, chelicerae
opened and knocked repeatedly
against opponent’s; no inju-
ries.
Displays to mirror.
1st & 2nd legs enlarged,
blackened.
5 other Venezuelan magoni-
nids use all legs actively in
walking, hop freely during
ordinary locomotion.
All include rubbing of 1st &
2nd tarsi in display.
9 usually extends 1st legs dur-
ing courtship.
1st legs enlarged, partly
blackened.
9 vibrates light palps.
hams disagreed strongly with Wallace (1878,
1889), who attributed both performance and
ornamentation to the greater vigor of the
male, especially during the breeding season.
Montgomery (1910) in general held Wal-
lace’s views; furthermore, he thought the
displays originated in self-defense move-
ments, which were combined with external
evidence of physiological excitement. Ber-
land (1923, 1927) attributed the behavior al-
together to the exceptional physiological act-
ivity of the breeding season and its correlated
excitement.
Bristowe (1941 et ante ) has held the view
that courtship in spiders in general has two
fairly distinct functions, namely recognition
and stimulation. Recognition he holds to be
particularly important, since in spiders the
female is not only carnivorous but often much
larger than the male. Inter-male display and
fighting he attributes essentially to mistaken
identity: “It, would appear from what has
been said above that the battles, which are
preceded by normal courtship reactions, are
the outcome of frustrated instinct, and are
modifications of courting procedure brought
about as a result of the different reception
experienced at the hands of another male to
that of a female.” (1929, p. 352) . He believes
that display arose from the groping and
chemotaxing motions of primitive spiders,
which have been modified by signs of sexual
excitement, such as twitching of the abdo-
men. He points out that a male which pro-
duces a striking pigment spot easily visible
to the female during his approach has a bet-
ter chance to survive than has an individual
less easily identified.
Savory agrees with Bristowe, except that
he considers the division between “recogni-
tion”— for which he prefers the term “real-
ization”— and stimulation invalid. He gives
the following summary of his views on spider
courtship: “If we may venture to summarize
in a few words the results of so complex an
activity as courtship, we may say that court-
ship is a chain of related instinctive actions,
in which the reproductive urge suppresses
the normal habits of self-protection and self-
nourishment, and is accompanied internally
by the physiological changes necessary to
make the subsequent union possible.” (1928,
p. 221).
Gerhardt, who has recorded numerous
spider pairings, has been most interested in
copulation methods, showing that the various
positions assumed by the male shed light on
phylogeny. In salticids the copulation posi-
tion is invariably dorsal and similar through-
out the family. He has made few observations
on the courtship phases of reproduction.
There has been as great discrepancy in
172
Zoologica: New York Zoological Society
[34: 17
views regarding the senses involved in salti-
cid display, as in those concerning its origin
and functions. The Peckhams (1894) con-
cluded that sight was essential, both for
recognition and as stimulus for display;
Petrunkevitch (1910) agreed. Montgomery
(1910) apparently saw a salticid court after
merely touching a female. Berland (1914,
1923, 1927) thought smell must be involved
in locating the female though sight was
needed to stimulate courtship. Bristowe and
Locket (1926) and Bristowe (1929, 1941)
thought that both sight and chemotaxis (re-
ferred to as “smell” in the earlier papers)
were involved. Savory (1928, p. 215), after
speaking of inter-male courtships, of the
courting of immature individuals and those
of other species, sometimes through glass,
says in regard to spider courtship in general :
“It is clear, therefore, that the stimulus
which initiates the male’s performance is
vague, rather than definite and specific. It
may act upon the sense of sight, of smell, or
of touch, but the appearance or the scent of
the female does not seem to be readily dis-
tinguishable from that of the male.”
The most recent group of observers have
concluded, along with the Peckhams and
Petrunkevitch, that sight alone among salti-
cids is the only adequate and necessary stim-
ulus for salticid display; these workers in-
clude Homann (1928), Bonnet (1933), Heil
(1936) and Kaston (1936). The latter gives
a detailed review of previous work, followed
by a report and discussion of observations
and experiments of his own ; these were con-
ducted on thomisids, pisaurids, lycosids and
salticids. His salticid examples are distrib-
uted among four Connecticut species of the
genus Phidippus (Dendryphantinae) . His
conclusions, which include his results with
that genus, are as follows (p. 152) : “On the
basis of a large number of observations and
experiments with the males of 19 species
from 4 families of vagabond spiders, it is
pointed out that the senses involved in court-
ship may vary with the species. There is no
evidence that a sense of smell is used in sex
recognition by any spiders. At least this
sense plays no part in initiating courtship
activity in the male. There is no evidence
that Attid males can ‘recognize’ the females
by any sense other than sight. At any rate, it
appears that the visual stimulus is the only
one that suffices to incite courship in this
family.”
C. Bases for Disagreements.
The disagreements and uncertainties
among previous investigators appear to have
been due principally to the following causes :
1. Although all observers have realized
that display stimuli among the various fam-
ilies differed, and were roughly dependent on
the development of vision, yet it does not
seem to have been clear that there are large
differences within the family, at least in
salticids, in the relative importance of the
senses. On these sensory differences depend
many differences in behavior.
2. Systematic attention does not seem to
have been paid to the fluctuating physiolog-
ical states of individual spiders.
3. The early experiments, as usual
throughout the history of biological science,
were not properly organized to ensure the
testing of only one stimulus or characteristic
at a time.
4. In later work, the compound natures of
such concepts as “visual stimuli” do not
seem to have been taken fully into account.
5. The recent concepts of innate releasing
and directive mechanisms, as developed by
Lorenz, Tinbergen, and their co-workers, do
not appear to have been applied to spiders.
6. Certain motions involving sensory per-
ception, such as waving of the first legs, have
perhaps occasionally been mistaken for epi-
gamic display.
TABLE III.
Principal Motions Reported in Salticid
Display Literature.
1. General manoeuvers, involving spider as a
whole.
a. Tacking.
b. Direct approach.
c. Leaping.
d. Sidling.
e. Semi-circling and circling.
f. Posing.
g. Crawling.
h. Elevation of carapace.
i. Rocking motions.
j. Bouncing motions.
2. Retinal motions within antero-median eyes.
3. Palp motions.
a. Vibrations, jerks, rotations; synchro-
nized or alternate.
b. Flexion or extension in special atti-
tudes.
c. Palpation of female or opponent.
4. Chelicerae motions.
a. Lateral extension of basal segment.
b. Extension of distal segment.
c. Clinching.
d. Biting.
5. First leg motions.
a. Elevation and extension forward, up or
out, and to any intermediate degree.
b. Vibrations, waves, jerks, rotations;
synchronized or alternate.
6. Second leg motions.
a. Extension forward, to rub second tar-
sus against first.
b. Elevation to form portion of fan-type
display.
7. Third and fourth leg motions : Elevation and
extension to various degrees, sometimes
forming part of fan-type display involving-
all legs.
8. Abdominal motions.
a. Depression, when carapace is raised.
b. Twitching or vibration; vertical or
horizontal.
c. Twisting or bending to side.
d. Elevation.
Crane: Salticid Spiders: Analysis of Display
17:
1949]
Text-fig. 1. Examples of display motions in salticid spiders. A-F, Subfamily Plexippinae;
G, Magoninae; H, Synagelinae; I, Dendryphantinae. A, Corythalia xanthopa, threat;
B, C, same, courtship; D, C. chalcea, threat (dotted lines indicate peak position of legs
during courtship); E, C. fulgipedia, threat; F, same, courtship; G, Mago dentichelis,
courtship; H. Gertschia noxiosa, courtship (from Connecticut, U. S. A.; similar to that
of the Venezuelan genus Semorina) ; I, Ashtabula furcillata, courtship. H, after Kaston,
1948; others reprinted from Parts I and III.
7. The great range of development of
threat display within this single family does
not appear to have been recognized, since
the days of the Peckhams’ scattered remarks
on the subject.
D. Prognosis of Evolutionary Pattern.
In order to bring into perspective the ma-
terial in the following pages, it is necessary
to anticipate some of the conclusions reached
not only in this paper but in future sections
of the series on post-embryology and general
evolutionary trends.
1. Correlation of Jumping with Sensory
Development. The genera studied at Rancho
Grande showed great variety in the fre-
quency of jumping in the course of normal,
unexcited progress. Taking into considera-
tion their behavior in other fields, notably
display, it seems certain that this variety is
based on differences in the dependence on
various senses. In some forms more depend-
ence is placed on chemical senses and less on
vision. Evidence is presented later showing
that these chemical senses include both
chemotaxis and a distance chemoperception
appearing very similar in its operation to the
vertebrate sense of smell. In other forms,
vision seems both more acute and more far-
sighted, while the chemical senses are rele-
gated to relative unimportance. Experiments
have not yet been performed at Rancho
Grande on variations in visual acuity, al-
though several experimenters, notably Ho-
mann (1928) and Heil (1936) , present mod-
ern data on visual perceptions in species of
the subfamilies Marpissinse and Hyllinae.
A few minutes’ observation of unconfined
salticids of selected genera will, I think, give
convincing empiric support of the hypothesis.
For example, in Ashtabula the progress is “a
scurrying run, the first legs held flat and
low, straight in front of the body; both they
and the palps palpate the surface almost con-
stantly during progress. During pauses the
first legs are usully elevated, and they and
the palps jerked rapidly up and down.” (Part
III, 1949, p. 41). These “runners,” as salti-
cids of similar habits will be called, never
resort to hopping or jumping except in cross-
ing gaps in the surface and in the final stages
of prey capture. In contrast to other kinds,
they are more easily observed in glass-cov-
ered dishes than when permitted to run free-
ly on a table (cf. Part II, p. 143) . Their com-
paratively poor vision and their dependence
on chemotaxis, or near-chemotaxis, make
their exploration of a new environment a
highly “restless” and “nervous”-appearing
proceeding, as they race to and fro.
In strong contrast is the behavior of an
almost completely vision-dependent salticid,
such as a Corythalia. When allowed to drop
on an unfamiliar surface, he tends to pivot
almost where he is placed while he looks all
around ; finally he moves off, without haste or
174
Zoologica: New York Zoological Society
[34: 17
205 and 175) for explanation; cf. Table II. Non-Venezuelan genera are enclosed in
parentheses; those not observed alive by the author are followed by a question mark.
The characteristics (“foreleg lifters” etc.) under each subfamily indicate the most
generally present display distinction; it usually is not found in every genus, and occurs
sporadically in other subfamilies as well; cf. Lorenz (1941) on the phylogeny of display
in ducks.
visible agitation, in a series of short runs and
hops, interspersed with pauses for further
observation. His palps, usually motionless,
are held quite clear of the ground ; his first
legs are used as much in walking as all the
others. To the human observer, in an anthro-
pomorphic moment, the contrast in “poise”
between a chemotaxis-dependent and a
strongly vision-dependent salticid is ludi-
crous. When startled or pursued he hops al-
most altogether. In the future, salticids pro-
gressing in the Corythalia fashion will be
called “hoppers.”
Phiale is a good example of an interme-
diate stage, in which hopping is moderate
and eyesight obviously keen. Considerable
dependence appears to be placed on distance
chemoperception, however, although not on
chemotaxis. The specially sensitive first legs
and palps are often carried up, in normal
progress, and during the pauses they are
waved up and down, without touching the
ground; it seems certain that they are re-
ceiving sense impressions which, by defini-
tion, are not chemotactic.
Characteristics of locomotion, where
known, are included in Table II.
2. Preliminary View of Evolutionary Pat-
tern. It is generally agreed that the salticids
are more specialized than the lycosids, par-
ticularly in regard to the development of
vision. In this characteristic, indeed, they
lead all other families. It seems likely, there-
fore, that the salticids in which vision is
19491
Crane: Salticid Spiders: Analysis of Display
175
best developed are those farthest from the
primitive stock. This hypothesis is supported
by evidence from a number of other fields,
. although the pattern of specialization, as
shown even in these few present-day genera,
is far from a simple “family tree.” As in
all other groups, salticids having many prim-
itive characteristics may be extremely spe-
cialized in a few directions.
Briefly summarized, the hypothetical pat-
tern may be stated as follows :
The evidence in regard to salticid evolu-
tion— based on post-embryology, external
morphology, ornamentation, locomotion,
general habits and epigamic display — all
points to a radiative type of development.
Many forms with primitive characteristics
still survive. Each subfamily so far studied
in any detail includes genera ranging
through at least two stages (Text-fig. 2).
The first, most primitive stage includes
species with low carapaces (Text-fig. 3), low
visual acuity, high dependence on chemo-
tactic stimuli, locomotion of the running
type, courtship simple and threat display
absent.
The second stage is characterized by in-
termediate carapaces, visual acuity, hopping
proficiency and courtship development; by
depending on distance chemoperception
rather than on chemotaxis; and, especially,
by the presence of threat display which is
based on the mistaking of males for females,
and is undifferentiated from courtship except
that it often ends in true fighting.
The third and most specialized stage is rep-
resented by species with high carapaces, ex-
treme visual acuity, chemical perceptions of
minimized importance, locomotion consisting
principally of hops, courtship often complex
but sometimes secondarily simplified, and
threat display separate from courtship and
altogether ritualistic.
I realize keenly the extremely small num-
ber of forms investigated at Rancho Grande
in relation to the large size of the family.
However, nublished records, in so far as they
can be analyzed from modern points-of-view,
and my own sporadic observations in other
localities, appear to lend support to the hy-
pothesis. It seems that the basic framework
will prove to have validty.
V. FACTORS IN DISPLAY.
The releasing and directive mechanisms
of salticid display have proved much more
varied and their interrelationships more
complex than was expected. Although all epi-
gamic display is fundamentally visual in this
family, chemoperception is important in cer-
tain phases ; simple adulthood is not the only
physiological essential for performing or re-
sponding to display; vision itself must be
divided into perceptions of motion, distance,
size, form, intensity, pattern and color for
an approach to understanding display
stimuli.
In the following pages an attempt is made
to isolate and evalute these factors, giving
I* INSTAR 2* INSTAR AOULT 3
3
12)
Text-fig. 3. Change of carapace proportions
with growth in representative genera of salti-
cids. Left column, 1st instar; center column, 2nd
instar; right column, adult male. A, Menemerus
bivittatus; B, Semorinv megachelyne; C, Sassa-
cus flavicinctus ; D, Mago dentichelis; E, Phiale
flammea; F, Corythalia chalcea.
in each case a summary of the observations
and experimental results upon which the con-
clusions are based. Because of space limita-
tions, it is impossible to publish full details
of field and laboratory data; however, de-
scriptions of especially illustrative observa-
tions and experiments are quoted now and
then directly from the original notes. Labora-
tory procedures have already been described
(Part II, 1948.2). Reference to Text-fig. 6
will help clarify the relationships of display
elements during the course of this analysis ;
in the subsequent section on innate releasing
mechanisms, beginning on p. 199, these
factors are viewed as closely integrated,
mutually dependent series of stimuli activat-
ing, controlling and directing both courtship
and threat.
A. Factors Of The Internal Releasing
Mechanism.
1. Age. In general, no sexual interest ap-
pears between the sexes until both are adult.
Exceptions occur when a mature male in a
particularly responsive state briefly courts
a pre-adult female of similar appearance to
the adult; such displays never pass beyond
Stage I. A number of workers have reported
behavior of this kind and I have seen it in
various genera, always of the hopper type,
especially in Corythalia. Very rarely juvenile
individuals perform abortive displays; ex-
amples will be given below.
176
Zoologica: Neiv York Zoological Society
[84: 17
The Peckhams (1889, p. 50) recorded the
interesting case of Philaeus militaris (—
Paraphidippus marginatus ) in which the
adult males actually kept guard over indivi-
dual preadult females until they molted. The
nearest approach to this seen at Rancho
Grande was the behavior of a single male
C. fulgipedia. He was captured clinging to
the outside of a cocoon, within which a female
was molting to the adult instar. He did not
even wait for her to dry and harden, but
raced through the briefest display as she
emerged, and mated. She was still soft and
weak, and gave no resistance or apparent
response at all. The eggs appeared on sched-
ule, some two months later, and were fertile.
This was an exception to the general behavior
pattern of the genus.
Similarly, matui'e males rarely display to
juvenile males except in reinforced experi-
mental situations.
During experiments on display sign
stimuli with C. xanthopa and C. chalcea,
several cases of display behavior were noted
in immature individuals of both sexes. They
were exceptions, however, to the rule. Once
a preadult (6th instar) male chalcea leapt
repeatedly on a dead, mounted adult male,
when the latter was appropriately moved. He
then gave a medium complete threat display,
through the fan stage with stiffened palps,
following this with a Stage I courtship, and
ending with a clumsy attempt to mate, ap-
proaching atypically from the rear. When a
normal adult male was presented to him,
without a rest period, the young one at once
gave a complete threat display; when the
adult responded in kind, the other retreated.
When a preadult female was presented to the
young one, she attacked him, whereupon he
retreated, turned and courted briefly, Stage I.
This sequence with the female happened four
times, she charging him each time.
A single case of display in a 5th (pre-pre-
adult) instar chalcea was recorded. This in-
dividual did a moderately complete threat
display (through the arch phase) to two
different adult males, punctuating them with
retreats. One of the adults responded with
threat. There was no courting or other re-
action to females.
Preadult females of both xanthopa and
chalcea gave incipient threat displays on a
number of occasions, to adult displaying
males, alternating the rudimentary fan-pose
with the usual abortive attacks and retreats.
In each case, it was clearly rudimentary
threat, not courtship. These motions were
never made in the presence of potential prey
or predators, even of similar size.
Display is usually delayed, however, even
after the final molt has taken place. Cory-
thalia males normally do not display to each
other or to a female until at least two days
thereafter. One exceptional xanthopa did his
first threat displays on the third and sixth
days, but did not court at all until the seventh
and then only a rudimentary Stage I; full
threat and courtship were not attained until
the tenth day. Although freshly molted fe-
males are both attractive and receptive, they
normally do not emerge from their cocoons
for two or three days. Judging by the re-
sponses of males, female xanthopa do not
attain their maximum attractiveness until
about the fifth day.
The age at which males and females lose
interest in display varies with the species.
After two months in the adult stage, un-
mated C. fulgipedia males usually cease to
initiate display in threat or courtship, and
do not respond to threats of other males.
However, one example, reared from the egg,
three months after the final molt briefly
courted an aged female who had completed
molting more than six months before. Un-
mated males of C. xanthopa are in their adult
prime less than a month; for several weeks
thereafter they may respond to stimulus
situations which are both complete and re-
inforced, but never to the extent of carrying
a courtship beyond Stage I. An exception
was a 32-day-adult male which mated with
a 33-day-adult female, after a courtship of
13 minutes; the usual courting period for
the species is three to five minutes. Their
healthy brood was reared through several
instars.
Unmated females of fulgipedia remain at-
tractive to males and will accept them at
least four months after the final molt, even
though they have, at the usual time (about
two months after molting), started laying
the customary successive clutches of eggs.
Females of xanthopa start losing both their
responsiveness to display and their attrac-
tiveness after about three weeks, although
the eggs in this species are not laid until
about the fortieth day after molting.
No Rancho Grande female, in spite of re-
peated attempts on a number of well-dis-
tributed genera, was ever found to mate
more than once, when more than a few hours
had intervened; usually, also, she lost most
of her attractiveness. However, there are a
number of recorded northern observations
of repeated salticid matings by both males
and females (e.g., Peckham, 1889, p. 38).
Rancho Grande males, as in the north, copu-
lated readily at least several times in Cory-
thalia, Phiale and Menemerus ; other genera
were not investigated, nor were such aspects
as the duration of fertility and the effect of
copulation on subsequent display initiation
and duration.
2. Fluctuating Epigamic Rhythm. Not only
is the period of display and acceptance at
Rancho Grande limited to moderately young
adults of both sexes and to unmated females,
but there is also a definite fluctuation in both
sexes from day to day in sexual aggressive-
ness and receptivity. Such fluctuations are
of course well known, and are in fact the
rule in higher vertebrates. Hints of it have
also been detected in fiddler crabs (Crane,
1941, p. 153 ff.) , but it does not seem to have
been much investigated in invertebrates.
1949]
Crane: Salticid Spiders: Analysis of Display
177
In all salticids under special observation
at Rancho Grande, it was noted repeatedly
that young, vigorous adults of either sex
varied daily in behavior. Those which were
particularly lively in display or responsive-
ness on one day might, on the next, be alto-
gether unresponsive to sexual situations,
although their general health and activity
seemed unimpaired. These individuals had
not been allowed to mate on the preceding
day, nor had there been a break in the feed-
ing routine, or noticeable changes in other
conditions such as moisture or general
weather. A day or more later, they might
recover all their previous epigamic energy.
This phenomenon was specially investi-
gated in more than twenty adult Corythalia
i xanthopa males of varying ages. About half
of them passed the final molt in the labora-
; tory, so that their exact length of adulthood
| was known. None of this group was allowed
to mate. All were kept under conditions as
similar as possible, although all twenty were
not observed simultaneously, but in small,
overlapping groups. All were tested for va-
rious periods up to 43 days with stimuli of
such proven efficacy that they came to be
termed “standard stimuli” for the species.
They consisted of a particular, normal male,
dried and mounted in semi-threat stance
(see p. 169), a mirror image, young and
older adult females, and live normal males.
In the case of low-threshold males, various
unnatural or incomplete stimuli were also
presented. The mounted specimens and mir-
ror were each presented in a series of up to
a dozen successive jerking approaches to the
tested male, after his attention had been
caught by the stimulus. The quickness of
response (if any) was taken into account
when determining his sexual tone, because,
obviously, a stimulus which drew a response
only on repetition was reinforced by sum-
mation. Again, the “startle” element of re-
peated presentations of the same stimulus
in quick succession, rather than continu-
ously, was important in drawing responses
from low-tone individuals; hence all hand-
controlled stimuli, for the sake of uniformity,
were presented in this fashion: the mount
or mirror was lifted high in the air for an
instant after each approach to the spider.
The working of the law of heterogeneous
summation (Tinbergen, 1948, p. 35) was
minimized by always working with a group
in every test period ; each individual could
thus be tested with one stimulus followed
by a rest, yet with a minimum wastage of
experimental time. The same general rules
were followed in all salticid testing (see
also, Part II, p. 143) .
In the course of the work, the fluctuating
internal drive appeared divisible into four
major degrees of sexual tone, with the fol-
lowing characteristics :
A-Tone. Internal drive strong, that is, male
with low epigamic threshold. Only minimal
stimulus needed for maximal response, which
follows promptly and appropriately to all
normal stimuli and to many deficient stim-
ulus situations as well. Display follows
through swiftly into Stage II in courting
receptive young females; old females courted
with considerable persistence. Responses
continued during a prolonged test-period. No
“vacuum responses,” however, have ever
INDIVIDUAL TONE SUCCESSIVE DAYS 0? OBSERVATION
1. MOLTED TO A
ADULT IN
LAD. B
C
D
2. MOLTED TO a
ADULT IN n
LAB. B
C
D
3. ADULT A
WHEN
caught. a
c
D
4. ADULT A
WHEN r.
CAUGHT.
C
D
5. aDULT ' A
WHEN
caught b
c
D
Text-fig. 4. Daily fluctuation of epigamic threshold in five male Corythalia xanthopa .
Spiders of A tone are most responsive to epigamic stimuli, those of D tone unresponsive.
The tone was determined by the individual's response to three standard stimuli : his own
mirror image, a particular dead male mounted in a semi-threat position, and a young,
living female. See p. 177. Blanks in the graphs indicate days when observations were
not made.
H N n '4*««OC'COO»OrHCJ
H N to ^ IO tO C* CD O) O i
N N W N CNJ W CM N CJ tO i
::=2
S*
LV*1
' L
178
Zoologica: New York Zoological Society
been obseiwed, in this or any other salticid;
that is, no display is performed without
some external stimulation.
B-Tone. Internal drive moderate, the epi-
gamic threshold being moderately low. Maxi-
mal stimulus needed for normal response;
either threat or courtship or both may be
elicited, but only through summation, though
display is eventually complete. Courtship of
a young female may be energetically per-
formed throughout its entire course, but
older females attract little or no attention.
All courtships are likely to die out in the mid-
dle, when the male, instead of following up
the female during her normal, periodic re-
treats, simply circles around her previous
resting place, palpating the ground in the
manner of less specialized, runner-type sal-
ticids. Again, the male sometimes gives
atypical responses, leaping briefly at the
threat stimulus before performing threat
display, or even courting it; equally briefly
and rarely, he may threaten a female. He
may or may not l'espond to one or two slightly
incomplete stimulus situations. Invariably
ceases to respond in a test period after rela-
tively few stimuli have been presented.
C-Tone. Internal drive weak, the epigamic
threshold being moderately high. Maximal
stimulus needed for minimal response. Male
responds either to normal threat or normal
courtship stimuli, but not both; summative
threat stimuli more often draw a response
even than young females. No courtship is
ever followed through to completion, and the
palpating of the female’s resting place is pro-
nounced, the male spending much of his at-
tention on one of her former positions, even
though she is at the moment sitting in full
view, and facing him, only an inch away.
Leaping at threat stimuli is frequent and,
unlike the case in B-tone spiders, is rarely
followed by the normal response. Incomplete
stimulus situations attract no attention, ex-
cept, sometimes, brief notice followed by es-
cape behavior. The attention of C-tone
spiders is difficult to attract and hold, they
often move about in “restless” fashion, and
always stop responding quickly, often after
only one or two stimuli have been presented.
D-Tone. Internal drive imperceptible, the
the epigamic threshold being very high. Male
does not respond with display to stimuli of
any kind, although other daily activities,
such as feeding and moving about, are un-
imnaired.
The above four states naturally merge in-
to one another, and in recording the daily
changes of tone, it was found convenient to
recognize plus and minus degrees. For ex-
ample, a spider which did not display, but
nevertheless showed enough interest to fol-
low a jerked mount or a young female with
with his eyes ( through the twisting of his
carapace), was counted D-plus.
Typical examples of daily fluctuations are
shown in Text-fig. 4. Text-fig. 5 diagrams
the degree of influence on each other of
spiders of various tones.
[34: 17
Text-fig. 5. Diagram showing mutual influence
of salticids of various epigamic tones. For ex-
ample, even an A-tone spider never draws a
display response from one of D tone, although
he himself displays readily toward individuals
of any tone and either sex; B- or C-tone spiders
may respond to the displays of an individual
of higher tone, but rarely initiate it themselves.
The boundaries between tones should not be re-
garded as hard and fast lines. See pp. 176-179.
In salticids, courtships are rarely completed in
the field, or threat displays energetically prose-
cuted, except by spiders of A tone, that is, of the
lowest epigamic threshold. This is in contrast
to the condition in some vertebrates, where the
behavior of a correspondingly low-threshold in-
dividual is not regarded as typical (cf. Part II,
1948.2, p. 143, and Tinbergen, 1948, p. 39).
The one result that emerged clearly from
the records was the complete lack of a defi-
nite rhythm in xanthopa. In general, how-
ever, individuals in their prime remained
from one to four days in A-tone, then
dropped within 24 hours to C or D; they re-
mained at these high-threshold levels from
one to three days, and then climbed back
to B-plus or A in about 48 hours. Some spi-
ders attained A-tone only once in the course
of their four weeks of prime, the rest of their
top display levels being B’s, with prolonged
C and D stretches between. A rarely active
male, caught adult, had three A-periods of
three, four and one dav each, during a pe-
riod of two weeks, with only one day of D
intervening; the subsequent portion of his
active display prime tapered off in irreg-
ularly alternating B’s and C’s of one to three
days each.
Just-molted males start at D, paying no
attention whatever to epigamic stimuli dur-
ing at least the first two days; thev then
climb, gradually or abruptly, to their first
B or A period, which may come at any time
from the third day to the tenth. After about
the 28th day, display responses taper off,
fluctuating gently between C’s and D’s. Often
they flow along for four or five days on about
a C-minus level, responding, for example, one
day to a single, summative threat stimulus
situation, and the next with a series of leaps
only to the same stimulus, but with the ad-
dition of a few abortive rocks to a young
female.
That the above phenomena are not the re-
sults of laboratory conditions is proved by
the fact that a number of males were tested
1949]
Crane: Salticid Spiders: Analysis of Display
179
in the field, before capture, and their be-
havior was found to correspond to that im-
mediately afterwards in the laboratory;
also, these and other males were taken, as
adults, in all four phases, some young, judg-
ing by their future plotted curves, some old.
Their behavior curves differed in no wise
from those of males reared through at least
one molt in the laboratory, except that in
general the wild-caught adults tended to hit
and hold A-tone oftener and longer.
Although males of other genera and spe-
cies were not tested systematically, there is
no doubt but that they do have periods simi-
lar to those of xanthopa. Females, too, show
noticeable behavior fluctuations, both in gen-
eral aggressiveness and in reciprocal dis-
play, although their periods were not studied.
One C. chalcea, for example, was especially
prone during several days to do a particular
type of jiggling in mutual display; I never
saw it in another of her species, and it soon
dropped from her usual behavior, reappear-
ing only sporadically. Although she was ex-
posed to the display of a number of males
during and after this time, she was not
allowed to mate for another two weeks, when
the process went off perfectly normally.
3. Hunger and Thirst. Hunger, as might
be expected, was an important factor in the
behavior of females, since a hungry female
was usually far more aggressive and needed
far longer courting than one which was
well-fed. However, in no genus except Eu-
stiromastix have I ever seen a female try
seriously to kill a male, although occasionally
she will leap at him. Moderate hunger in
males lowers their epigamic tone only
Text-fig. 6. Factors of the innate releasing mechanism in salticid display. The circles
are represented as loosely interlocking to indicate the interdependence of the three
groups of factors. Similarly, the spokes are not continued to the circumference in order
to show the mutual influence of the sections. Finally, the spokes are represented as
broken lines, to emphasize the variability and instability of their relative importance;
this varies not only throughout the family, but even in the same spider under different
conditions. For example, among the Synagelinae, the chemical factors are more im-
portant than among the Plexippinae, while the same sections are of still different
relative value in the end-forms of the Dendryphantinae. The sizes of the sections, there-
fore, are merely a guide to their average apparent importance within the family. The
term “X-factor(s)” is included in each circle to emphasize the incomplete state of our
knowledge. “Distance” is omitted from among the sign stimuli since it seems to be more
of a compound factor in display than do the others, based largely on apparent size, on
chemical stimuli or their absence, and on physiological conditions; perception of dis-
tance and depth does exist however (see Homann, 1928; Heil, 1936). Tactile factors are
omitted since they do not release display in salticids, although they are important in
the final display stages. The sign stimuli circle applies to courtship; for threat display,
the chemical factors should be eliminated.
180
Zoological New York Zoological Society
r 34 : 17
slightly — from B-minus to C-plus, for ex-
ample. In all the experimental work, no spi-
ders were tested without having been fed
at most 48 hours previously.
Thirst cannot of course properly be sepa-
rated from the external factor of humidity,
although spiders do drink water as needed.
Salticids that for some days had been kept
without special moisture in the laboratory
went into silk shelters, if they did not die
first, became semi-torpid and did not react
to display stimuli ; this behavior is doubtless
equivalent to aestivation in the field.
4. Fatigue and Overstimulation. These
two factors have not been properly distin-
guished. Actual physical fatigue, however,
sometimes appeared clearly involved during
experiments. Often initially restless indi-
viduals, even of hopper-type genera, were
tested which raced about the table before
settling down to attending to stimuli; they
often had even to be pursued to the floor;
afterwards, even when otherwise of A- or
B-tone, they always reacted to stimuli for
a shorter length of time than those which
responded without preliminary activity. On
the other hand, overstimulation appears to
be a better term for the cessation of reaction
that takes place in spiders at the peak of
their internal drive, when they have had
practically no exercise but have been pre-
sented with a succession of stimuli. In every
individual the moment is reached, as in other
animals, when no further response is drawn,
no matter what the stimulus. The length of
reaction time is highly variable ; C. xanthopa
is, however, as typical as any : A-tone males
may be tested for around half an hour with
only minimum pauses between stimuli (about
three minutes). When a large group is used
(the customary experimental procedure) re-
sponses may spread over three or four hours,
the long rotation period allowing individual
rests of 10 to 30 minutes between stimuli
presentations. An individual which is tested
in the morning to the point of complete
non-response is usually again responsive by
mid-afternoon; full recovery (regardless,
that is, of internal drive changes) is the rule
by the next day. Spiders with low internal
drive, e.g., of C-tone, have extremely short
response periods, usually totaling only five or
ten minutes; often there is only a single
response, regardless of length of rest be-
tween stimuli.
5. Attention. This factor is quite distinct
from the others, and a sine quo non of dis-
play. Although spiders with strong drive
undeniably are more easy to stimulate with
display situations than weak-drive individ-
uals, all vary irrespective of sexual tone in
the amount of time needed to attract their
attention to a stimulus. Sometimes they are
so restless when first put on the table that
their only motive appears to be to get some-
where else; some minutes pass before they
seem suddenly to “wake up” with an almost
literal jerk and “notice” the stimulus, which
may have been equally close in front of them
many times before. This applies to both
threat and courtship stimuli. The same is
true of a spider engaged with one display
stimulus, or with food, when another and,
at the moment stronger, situation is pre-
sented.
B. Factors of the External Releasing
and Directive Mechanism.
1. Physical Environment. The effect of the
physical environment has not been precisely
analyzed in regard to exact tolerances and
requirements of temperature, humidity, light
and altitude. However, the following facts
have been determined:
Corythalia xanthopa in the Rancho Grande
laboratory displayed readily at temperatures
between 65° and 80° Fahr., the extremes
tested. As has been noted by others, responses
were somewhat accelerated at the higher
temperatures. The humidity varied from 50
to 95 per cent. The same species displayed
at Weston photometer readings as low as 0.8
foot candles; below that there was no re-
sponse. During regular experimental ses-
sions, reflected daylight from the laboratory
table ranged from about 25 to 75 foot candles,
Weston; these readings were comparable to
those obtained in xanthopa’ s natural habitat
during periods of high activity. Menemerus,
with apparently poorer vision, seemed to
need considerably more light, but no accurate
measurements were taken.
Corythalia fulgipedia, C. chalcea, Eusti-
romastix and Phiale flammea, although cap-
tured at 3,500 feet, all displayed at sea level
in New York, having been brought there
alive in the early instars. They displayed
upon reaching maturity regardless of the
month, of the highly variable and unnatural
extremes of temperature and humidity in
which they had been reared, and of the lower
altitude.
Displaying individuals of most species
could be found at Rancho Grande from Feb-
ruary to September, the period of our res-
idences, which includes most of the dry sea-
son and half of the wet. However, as in other
groups, most species had a clear peak of
breeding adults in May, June and July, the
early part of the rainy season.
Both in Venezuela and in New York, spi-
ders displayed under extremely unnatural
conditions, with no time whatever needed
for adjustment, short of their being literally
frozen, deeply chloroformed, dried out or
far overheated. That is, if the spider were
not injured physically beyond recovery, if
he remained capable of primary activities —
moving freely, catching prey and spinning
a shelter — he could and did display when
in a period of moderate or strong drive and
when properly stimulated. All salticids ob-
served displayed as readily in a glass
container or on a table top as in their own
forest environment, and in every case the
pattern of display behavior in captivity was
identical with that recorded in the field. The
only exception was that, rarely, captive
1949]
Crane: Saltieid Spiders: Analysis of Display
181
Corythalia males, in close quarters under
hot lights, occasionally injured each other
in actual fights; ordinarily, aggressive be-
havior was confined to highly ritualized
threat display.
In the field, salticids invariably disap-
peared under leaves and therefore presum-
ably did not display, both during every
shower beyond a drizzle and during all hours
of intense heat, drought or baking sunlight.
Display during the night in the laboratory
was successfully stimulated in A- or B-tone
Corythalia and Phiale which were forced
out of their shelters and provided with ade-
quate light. Hence an innate diui'nal rhythm
does not appear to exist in these forms.
In brief, the only external physical require-
ments for display appear to be tolerable con-
ditions of temperature and humidity, along
with adequate light (sun, photoflood or
mazda) . In Corythalia at least, this illumina-
tion is less than that sufficient for comfort-
able reading by a human being.
2. Sensory Elements and Sign Stimuli.
The sign stimuli (or perceptual signs) con-
sidered here are those external signals which
are involved in releasing or directing epi-
gamic display in salticids. They are unex-
pectedly complex, especially in courtship, in
which they customarily involve compound
tactile, chemoperceptual and visual stimuli.
Of them all, only two or three combinations
of a few visual stimulus-elements can be
termed primary releasers, since no saltieid
displayed at Rancho Grande unless the an-
tero-median eyes were photopically stimu-
lated.
Reference to Table II and Text-fig. 2 will
be helpful in maintaining orientation in
regard to the various displays and the pre-
sumed degree of specialization of species
discussed.
a. Tactile Perceptions. Although in many
web-spinning spiders tactile perceptions are
of great importance in courtship, in salticids
their role is confined to advanced Stage II,
where the stimuli are probably mingled with
chemotactic stimuli. They will be discussed
together on p. 182.
b. Chemoperception. For many years a
possible “sense of smell,” as earlier ob-
servers called it, in spiders has been the
subject of extended experiment and con-
siderable controversy. In their studies of
courtship in various spiders, modern work-
ers, particularly Kaston (1936) and Bris-
towe (1941), have drawn necessary and
precise distinctions between contact chem-
operception and distance chemoperception.
In contact chemoperception, conveniently
called chemotaxis, the spider must actually
touch the female herself, her silk or exuviae,
or at least her trail, in order to receive a
sensory chemical impression. This sense,
both authors agree, often enters into spider
courtship, although it is not always essential
for the stimulation of display, and, in their
experiments, was not necessary in salticids.
The Rancho Grande results were in agree-
ment with these conclusions.
On the other hand, in courtship stimuli
involving distance chemoperception, a spider
would have to react to volatile, airborne par-
ticles given off by a female, her silk or exu-
viae, or her trail. Such stimuli have not been
found by these authors to take any part in
stimulating courtship display in any spiders
which they have studied, nor do they con-
sider that evidence has yet been brought
forward which necessitates the operation of
such a sense in spider courtship. Here the
Rancho Grande results differ; it seems nec-
essary to postulate the use as a secondary
sign stimulus in courtship of a sense analo-
gous to smell.
The two aspects of chemoperception — con-
tact and distance — will now be considered
in reference to the Rancho Grande salticids.
i. Chemotaxis. In some other groups of
spiders, all with vision less well developed
than in salticids, chemotaxis is a primary
releaser of courtship, with or without sight
of the female. In salticids this is certainly
not the case. Kaston, reexamining the sit-
uation in 1936 (pp. 129-130), concluded that
it was not essential in two species of Phidip-
pus, and Bristowe (1941 et ante ) agreed.
Similarly, at Rancho Grande, when chem-
otaxis was positively eliminated from the
stimulus situation, in every genus tested
A-tone males displayed without chemotaxis.
Contrariwise, no individual ever displayed
on the stimulus of chemotaxis alone, although
excitement over chemotactic stimuli was
often shown. The tested forms were the fol-
lowing: Menemerus, Semorina brachy-
chelyne, Ashtabula, Sassacus (2 spp.), Phi-
ale (2 spp.), Plexippus and Corythalia
(3 spp.).
However, in all of the numerous observa-
tions and experiments, variations were very
obvious in the minor exercise of chem-
otaxis. These were present on all levels —
genex'ic, specific and individual. When chem-
otaxis was allowed, the initiation of display
in all the runner genera ( Menemerus , Sem-
orina, Ashtabula) was definitely expedited,
although that of the hopper and intermediate
forms (e. g., Corythalia, Phiale) was appar-
ently not affected in the case of A-tone
spiders. However, many individuals in all
genera, when of B-tone, initiated display
only when chemotaxis was included in the
stimulus situation.
Salticids of all conditions, except those of
D-tone, and in all genera, but particularly
among the runners, often paid great atten-
tion to the recent resting places of females,
tapping them with palps and forelegs and re-
volving round and round the area. All B-tone
or tiring spiders tended to break off display
and palpate at length her just-vacated rest-
ing spot even as the aroused female sat and
watched only an inch or two away; some-
times, absorbed in these palpations, a male
allowed a prospective mate to wander com-
pletely out of sight; this reaction was noted
182
Zoologica: New York Zoological Society
[34: 17 *
most frequently in the hopper genus Cory-
thalia.
In the runners there was more of a tend-
ency to follow a trail, by intermittent palpa-
tion of the ground, over which the female had
passed. In Sassacus ocellatus, Phiale and
Corythalia, there were no tendencies to trail-
following chemotaxis at all; once a female
had attracted his attention, a male often
followed her zigzag or curving retreat by
taking the most efficient short-cuts, whether
or not display had been initiated.
The latter half of Stage II is so similar
and well known throughout the family that
it will not be specially treated here. It begins
with the male climbing over the female, pat-
ting her carapace with palps and first legs,
and follows through to the twisting of the
abdomen and successive insertion of the
palps into the epigynum, usually after some
palpation of its surface. Here it will only be
pointed out that chemotactic as well as purely
tactile perceptions are probably involved.
Chemotactic perceptions are also not nec-
essarily eliminated from the stimuli which
continue to lower the female’s threshold
during the same period. Her final resistance
is probably broken down by the stimulating
action of the patting motions ; however, since
her chemotactic receptors appear to be
scattered over her entire body, as well as
concentrated on the appendages (Kaston,
1936 & ref.), they very likely are stimulated
at this time.
Except in the advanced stage noted above,
chemotaxis certainly plays no important
part in the acceptance of a male by a female.
Since she usually retreats a number of times
during normal courtship, she infrequently
crosses a male’s trail during display in the
field. It is true that some individual males
have a tendency to circle the female com-
pletely during display. Also, females of Phi-
ale and Menemerus have often been seen
to palpate a male’s recent resting place, while
he is still displaying. Nevertheless, on open
table tops in more than 20 experimental
situations and in uncounted casual observa-
tions, involving all the experimental genera,
males were accepted without the females
once crossing their trail.
I can find no evidence whatever that chem-
otaxis plays any part in inter-male display ;
I have never seen a male palpate the surface
or show apparent excitement of any kind
when placed on the spot recently vacated
by another male, even when both have been
or are displaying to each other. However, a
male Menemerus, a typical runner, could be
induced to display to his mirror image only
by corking him in a clean glass vial and
holding the mirror against the glass end.
After a few minutes, display sometimes took
place in A-tone individuals, each presum-
ably activated by his own strongly reinforced
chemical stimuli. This experiment was per-
formed on six different males; the display
was indistinguishable from Stage I of
courtship and was always of very brief
duration.
The occasional males in various genera
that enter upon Stage II of threat, when the
first legs of each opponent touch, practically
always finish the encounter at this point
with no damage done to either. The fangs
are very rarely unsheathed during these
moments. Chemotaxis, or, rather, the ab-
sence of the female chemotactic stimulus,
may be involved here.
ii. Distance Chemoperception. The appar-
ent part played in display by a sense analo-
gous to olfaction will now be discussed. The
majority of previous experiments concerned
with it have dealt with the reactions of spi-
ders to essential oils, chiefly of an apparently
irritating nature. It has been pointed out
several times (e.g., Kaston, 1936, p. 146)
that sensitivity to these stimuli does not nec-
essarily indicate possession of a true olfac-
tory sense. Such a sense has, however, been
proved to occur in a number of terrestrial
invertebrates, including moths, beetles and
bees. The only aspect directly concerned in
the present display study is the part, if any,
taken by a sense analogous to odor percep-
tion.
If such a sense exists, the necessary af-
fectors do not, of course, necessarily con-
sist of volatile particles; they need only be
airborne and, presumably, invisible to human
eyes. It is regrettable but natural that we
human beings, as visually dominated organ-
isms, should tend to be impressed with and
to study chiefly visually or at least aurally
dominated displays in animals. Moths and
other nocturnal creatures, both vertebrate
and invertebi'ate, may, for all we know,
give off in epigamic display regular sym-
phonies of smells in a rhythmic succession.
More difficult to imagine, but perhaps easier
to check, is the following possibility : Females
of certain animals, at the peak of physio-
logical readiness, may actually radiate cer-
tain wavelengths invisible to us. These may
be either infrared or ultraviolet, using
those terms in their broadest sense. An alter-
native possibility is that these frequencies
may be reflected rather than radiated. The
emission, of whichever type, would doubtless
be under hormonal control, and Would have
some signal values for males. It would not
necessarily of course be perceived visually.
In that connection may be mentioned the old,
recurrent speculations on possible wave-per-
ceptive functions of insect antennae. The
claims of some investigators that blind-folded
human beings respond muscularly to red light
is another related topic (See Birren, 1938,
ref.).
Toward the proof or disproof of the pos-
sibility of such emissions, not one jot of ex-
perimental proof is offered here. The subject
is mentioned only because of its potential
connection with salticid display, and in the
hope of stimulating investigation. If found
valid, it would explain one or more aspects of
1949]
Crane: Salticid Spiders: Analysis of Display
183
animal display which at present are inexpli-
cable. The proving of the perception of spe-
cial frequencies would solve puzzles in certain
fiddler crabs and birds, for example. In each
of these cases, males appear instantly to dis-
tinguish receptive females of their own kind
from non-receptive ones in some fashion
which does not appear to depend either on her
behavior or on her emission of any recognized
sensory signal (see Beebe, 1928, p. 64; Crane,
1941, p. 157 ; and Armstrong, 1947, p. 340
& ref.).
It is true that the operation of scent has
not yet been eliminated under experimental
conditions in the above instances. Since the
published work on crabs, however, many field
examples have been seen where the particu-
larly stimulating female crab was several
yards to leeward in a stiff wind; the poor
olfactory development in birds is well known.
According to Kettlewell (1946), however,
moths have not yet been proved ever to ap-
proach females with the wind, the contrary
evidence of earlier experiments having been
due to incomplete observations.
In view of its highly controversial aspect,
therefore, this present speculation may be on
the order of postulating improbable sunken
continents in order to explain an inconven-
ient distribution of fauna, flora or culture.
Nevertheless, it is set down in the hope that
adequate instruments will soon be developed
capable of testing the possibility. Proofs of
the use of supersonic signals by bats (Hart-
ridge, 1945) and the perception of polariza-
tion of light by bees (von Frisch, 1949;
Thorpe, 1949) are suggestive recent results
of research in unusual directions. In our own
future researches and experiments we cer-
tainly intend to take such possibilities into
account.
In regard to salticid spiders, however, it
seems that the phenomena described below
can be most simply and adequately explained
by postulating the action of typical, airborne,
volatile, chemical particles acting upon ap-
propriate receptors. They will be so consid-
ered in this paper.
The work at Rancho Grande seems to
point conclusively to the existence of such a
chemical sense. It appears likely that it is
merely a further development of chemotaxis,
in that the affectors, and perhaps the recep-
tors, are basically similar or identical. This
sense plays an important secondary role in
courtship.
The evidence for this conclusion is based
chiefly on the following observation : Young,
living, unmated, adult females at Rancho
Grande were in general markedly more suc-
cessful in drawing courtship display from
males than were females in other categories.
Their superior attraction was noted in
more than 25 species, in fact, in all material
plentiful enough to yield comparative obser-
vations ; these included all the Rancho Grande
forms discussed in this paper. The Peckhams
(1894, p. 251) attributed a similar observa-
tion to the fact that females about to lay
eggs were so heavy that they tended to move
about less, and that it was the lack of move-
ment which made them relatively unattrac-
tive. In the basic experiments at Rancho
Grande, this factor was eliminated by chloro-
forming the subjects. They were then pre-
sented alternately as motionless, or uni-
formly moving stimuli on L-mounts. Chemo-
taxis was ruled out as previously described
(Part II, pp. 143 and 144). The males were
allowed only front views on a plane surface,
to avoid the visual variables of large vs. small
abdomens, or fresh vs. rubbed patterns. The
effects of summation were guarded against,
as usual. For experimental purposes, a fe-
male was counted as “young” when she had
been adult for less than two weeks, “old”
when more than four.
The general results, in this and other gen-
era, were as follows: A-tone males, when
given a choice, almost always devoted their
attention to a young female, although an old
or mated individual was usually courted ener-
getically if no other was available. B-tone and
C-tone males, as well as all near-senile males,
showed no courtship display response at all
to any but young females. A-tone males often
courted a young female which was chloro-
formed, placed on a mount and held motion-
less ; males of the same tone rarely took any
notice of an old or mated individual similarly
presented. Dead, thoroughly dried females of
all ages either drew no reaction when mo-
tionless, or, when appropriately moved, oc-
casionally drew confused threat display.
Finally, although certain paper models of C.
xanthopa males successfully drew threat dis-
play, models of females were never success-
ful.
Special test situations, involving C.
xanthopa 2 and Phiale, gave the following re-
sults which are pertinent:
The abdomen of a young adult female
xanthopa had been painted black for another
experiment on the tenth day after her molt.
After repeated intermittent chloroforming,
she died about five o’clock one afternoon. At
that time, when crumpled, motionless and of
atypical coloring, she readily drew display
from several males. Chemotaxis had, as usual,
been eliminated from the test situation. By
seven o’clock the next morning, although she
was not yet stiff, she had lost attraction for
three A-tone males to the extent that, al-
though some attention was paid her when
she was jerkily moved, no display whatever
was drawn. The normally colored abdomen of
a chloroformed female, adult for fourteen
days, was then cut off from the cephalothorax
- The published description of C. xanthopa display (Part
I, p. 36) erroneously states that rocking precedes threat
as well as courtship ; further observation showed that this
occurred only in small, closed dishes involving strong
reinforcement of chemical stimuli. Normally, rocking does
not occur in threat display. See Table II, this paper, for
synopsis of threat and courtship distinctions. Also, the
threat display figured (l.c.. Text-fig. 14 C) represents a
moderate phase; in the extreme form the 4th legs are
held even higher, well above the carapace, and the second
tarsi leave the ground.
184
Zoologica: New York Zoological Society
[34: 17
and placed on the mount 10 mm. from the
dead female. Two males now displayed
promptly to the black corpse, following
through to attempted mating with it, disre-
garding completely the normally colored, de-
tached abdomen.
The detached abdomen was then rubbed
over a dried, mounted male, which had here-
tofore always di’awn prompt threat display.
It drew brief, confused display, with both
threat and courting elements, from a male
which had threatened it a few moments be-
fore, and threatened standard stimuli im-
mediately after (without a rest period).
During the test, however, he followed the
confused display with a leap at the mount and
finished with an abortive attempt to mate.
On another occasion, a male xanthopa was
threatening a dried male mount. Another
freshly detached abdomen of a young female
was hidden under a tiny paper tent, 5 mm.
to the left of the displaying male which in
turn was 50 mm. from the mount. The male
stopped threat display, went into a courtship
stance and rocked. When the mount was re-
moved, there was no further reaction.
A xanthopa male paid no attention to a
scrap of black cloth alone on a mount, or to
the same cloth with a young female under-
neath it, with or without motion of the
mount. Then a large, dried Eustiromastix
male was given a yellow clypeal band similar
to that of xanthopa, and placed on another
mount. This stimulus drew prompt threat
display when the mount was moved. The
piece of cloth was then placed in a heap un-
der the Eustiromastix. Threat again fol-
lowed. However, when the chloroformed fe-
male was once more hidden beneath the cloth
and the whole placed under the Eustiroma-
stix, the test male paid swift attention with-
out display, then vibrated the palps, took a
courtship stance, and, finally, after one or
two rocks, leapt on top of the Eustiromastix
and tried to mate with it. This involved such
a violent attempt to twist the stiff abdomen
that the whole structure toppled over. With
a changed sequence of stimuli, the same
series was presented to two other males, with
similar results, except that the courtship re-
sponse died out before mating attempts.
Young females of both Corythalia and
Phiale, when painted to resemble males, drew
unmixed courtship, not threat, although old
painted females drew mixed display (p. 194) .
When female epigyna of Menemerus,
Phiale and Corythalia, in a total of six indi-
viduals, were sealed with paraffin, no diminu-
tion in attraction was noted; the attractive
substances are probably secreted rather gen-
erally by body and appendages; the experi-
ments of Kaston (1936) and others on non-
salticids support this idea.
It should be mentioned incidentally that
young adult females have a stronger
odor to human noses than either males
or older females, at least in Menemerus
and all three species of Corythalia. In a
series of tests on Corythalia, the odor of
young females was the only one percep-
tible to two of five observers; the other
three people in every case considered the
young female’s odor strongest, but indistin-
guishable in quality from that of males or
other females. The latter seemed stronger
than the males’. No generic or specific dif-
ferences were detected, during casual tests
with various genera. The terms used in at-
tempts to describe the salticid odors included
“spicy,” “pungent,” and “faintly antiseptic.”
Further incidental data, by no means to be
regarded as actual evidence for distance
chemoperception, include the different meth-
ods of using palps and first legs in the various
genera (Table II). This seems to be in ac-
cord with the various degrees of reliance on
contact and distance chemoperception.
The chemoreceptors (including those of
chemotaxis and the distance chemorecep-
tors, whether or not they are distinct) , have
been shown to be generally distributed on the
spider, but concentrated near the tips of the
palpi and first legs (Kaston, 1936 and ref.).
With this view the Rancho Grande experi-
ments agree. Females without palps or with-
out first legs readily accepted males in Mene-
merus and Corythalia. When both palps and
first legs were removed, reaction time was
slowed in Menemerus but not in Corythalia.
Males similarly handicapped displayed right
through to mating attempts. When use of the
first legs was the essential part of display,
spiders without them would suddenly leap at
the female after a prolonged period of atten-
tion. This response was specially noted in
Menemerus. Reaction time was decidedly
slowed in handicapped males in initiating
courtship, but not in responding to any of the
threat stimuli, including the mirror and
painted mounts. It was much slower in muti-
lated runners of the genus Menemerus than
in the hopper genus Corythalia.
In summary, the following conclusions
appear valid. At Rancho Grande, distance
(airborne) chemoperception is a positive
factor in courtship, but not in threat display.
A strong, invisible stimulus is given off by
young females, serves as a secondary sign
stimulus for releasing courtship and is sur-
passed in importance only by the male’s abil-
ity to use his antero-median eyes. It appears
to have little directive value. When courtship
and threat are distinct, the addition of air-
borne chemical stimuli from a young female
to a stimulus configuration tends to change
threat to courtship.
b. Vision.
i. Vision as a primary stimulus to display.
Use of the AME (antero-median eyes) by
the male is the only single, sine qua non of
display. The investigators who have experi-
mented with the reactions of blinded male
salticids agree that they will not display
when the eyes are completely covered, no
matter what other stimuli (e.g., contact and
airborne chemical stimuli) are present.
Apparent excitement eaused by other stimuli
Crane: Salticid Spiders: Analysis of Display
185
1949]
has sometimes, however, been observed.
The species previously tested are well dis-
tributed among a number of subfamilies:
Dendryphantes elegans, Saitis pulex, Phidip-
pus rufus and Astia vittata (Peckham, 1894,
p. 248) ; Evarcha blancardi (Homann, 1928,
p. 254) ; Aelurillus v-insignatus (Bristowe,
1929, p. 343) ; Phidippus clarus, P. audax
(Kaston, 1936, p. 131). Homann confined
some of his blinding experiments to the
AME, finding that display failed to occur
as completely as when spiders were totally
blinded.
The present experiments support and ex-
tend these previous conclusions. Two males
each of Corythalia xanthopa, C. chalcea and
Menemerus bivatattus, all of A-tone, were
used. In one of each pair all eyes except the
AME were painted, as described in Part II
(1948.2, p. 144) ; in the other, only the AME
were covered. After the spiders had appar-
ently recovered from the chloroform (the
minimum time allowance was one hour) ,
standard test stimuli (p. 177) were presented
on three successive days, unless a positive
response was obtained earlier. Regardless of
all-negative responses, the paint was then
removed and, in tests counted as successful,
positive responses obtained to at least one
of the standard stimuli within one hour.
In each species, the males with the AME
uncovered performed complete display (both
threat and courtship, in the case of Cory-
thalia) within the allotted period, although
their reaction time in general appeared .a
little slowed. In contrast, those with the
AME painted never displayed at all. The
latter did, however, show some awareness
of moving objects, including females, by
shifts in position and slight “following” with
the eyes at distances of half an inch or less.
This note is typical: “X64 male sat quiet,
although he ran and jumped when prodded.
Took no notice of X34 female, as she watched
him only one-half inch away; although she
actually brushed against him several times,
even his palps hung quiet. Later he moved
off, crossing her trail and very recent resting
place without pausing. Finally, when she
jumped past him, he did turn toward her;
the stimulus, however, could have been a vi-
bx-ation of her jump, rather than either a
chemical or visual stimulus. Even in the same
vial with her, with all her chemical stimuli
reinforced, there was no reaction.”
The most interesting result, from the view-
point of phylogeny, was that Menemerus,
which appears to depend more on chemotaxis
and distance chemoperception and less on
vision than Corythalia, was decidedly less
handicapped by the loss of the AME. He
moved around much more actively, and pal-
pated the females’ resting places in apparent
excitement. No attempt was made by either
genus to pursue prey without the use of the
AME.
The AME of a single young female C.
xanthopa were covered. Just before the op-
eration she had allowed a male to reach an
advanced Stage II, when the pair was pur-
posely separated. After painting, she refused
him and four other actively displaying males
which subsequently displayed to her, two or
three in succession, on four successive days.
One male, on the third day, reached Stage II
repeatedly, . . but at the least touch, or
before, she moved off. Male was very per-
sistent. She did not turn to keep him in view;
her palps hung motionless. Since he circled
her time after time in displaying, as usual,
she crossed his trail frequently during her
retreats. She never once assumed the raised-
carapace stance so many xanthopa females
temporarily adopt when attention is gained.
He persisted for twenty minutes, then gave
up. I cleaned her eyes, introduced the same
male within half an hour, and they were
mating five minutes later.” (Field lab. note.)
No other female blinding experiments
were completed through the final stage of
obtaining positive mating responses after
the AME were uncovered. However, all the
indirect evidence — in the observations of
others and in my own — points to the usual
necessity for the female to see the male be-
fore allowing mating to take place.
In summary, use of the AME, and of the
AME only, is a prime requisite in salticid
courtship.
ii. Motion. It seems well established that
at least some salticids will not only leap on
motionless prey, but will display to a motion-
less female (e.g., Peckham, 1894, pp. 243-
248; Heil, 1936, p. 10). Nevertheless, the
present study showed that movement is the
most important single element in the visual
sign stimulus pattern, in both courtship and
threat display, in all the salticids tested. Al-
though display was occasionally obtained by
a motionless spider, the courtship response
always occurred under the following condi-
tions: first, the stimulus was a female; sec-
ond, the displaying male was of A-tone with
exceptionally low threshold; third, the mo-
tionlessness of the stimulus was the only
unusual factor in the situation. On other
experimental occasions in which the above
conditions were fulfilled, however, only nega-
tive responses were drawn. Table IV gives
test data and results of pertinent experi-
ments.
In many (certainly more than twenty)
unrecorded, casual observations, a normal
spider, which was sitting quietly when the
male to be tested was introduced, would
not draw a display until it moved, even
though the two spiders were close together
and the test male, judging by his actions —
his turning, or moving toward the stimulus
— had certainly become aware of the stim-
ulus.
In more than 300 tests on 10 species, in
which the stimulus was in some way un-
natural, no positive response was ever ob-
tained until the stimulus was moved. Many
times the slightest, brief, manual twitching
of the cardboard mount was enough of an
186
Zoologica: New York Zoological Society
[34: 17
TABLE IV.
Responses of Males to Motionless Females.
Series A. Conditions. Male of A-tone; normal female chloroformed, placed on cardboard
mount in approximately natural position; chemotaxis, but not distance chemoperception, elimi-
nated (i.e., fresh sheet of paper on open table used for each test) ; same female in experimental
session drew display from same male both before test (followed by rest) and immediately after.
Exposure of each male to stimulus consisted in allowing him to drop near, or run toward, female,
chivvying him gently with brush if necessary, to ensure her being in his direct line of vision.
Even in negative responses, attention was often gained, even extending to feeling of the female
with palps and legs, although display was not released.
Species.
No. of
Individuals.
No. of
Tests.
Negative
Responses.
Positive
Responses.
Menemerus
bivittatus
5
7
6
1
Semorina
brachychelync
i
2
2
o
Sassacus
flavicinctus
2
2
2
0
Phiale
dybowskii
.
1
5
0
5
Phiale
flammea
2
3
0
3
Corythalia
chalcea
i
1
0
1
Corythalia
xanthopa
4
12
11
1
(delayed)
Total
16
32
21
11
Series B. Chemotaxis permitted. Conditions as in A, but males permitted to cross repeatedly
trails of the now motionless females.
Species.
No. of
Individuals.
No. of
Tests.
Negative
Responses.
Positive
Responses.
Menemerus
bivittatus
4
9
8
1
Sassacus
flavicinctus
2
2
2
0
Corythalia
xanthopa
4
8
8
0
Series C. Elimination of Distance Chemoperception and Motion. Conditions as in A, but female
isolated in glass vial. No tests completed, since males did not display even when the stimulus
female regained consciousness and moved naturally. However, in spite of the imperfect conclu-
sions, it is worth recording that not one positive response was drawn in a total of more than 20
tests involving the same species listed in Series A. The enclosing of tested males, rather than
stimulus females, would not have given significant results, since the effect of possible self-stimu-
lation through crossing of their own tracks or responding to their own distance chemical stimuli,
would not be eliminated. Similarly, mirror responses automatically involved sight of the males’
own motions (however slight).
Series D. Alteration of Female Appearance. Conditions as in A, but female painted or upside-
down. More than 25 tests, using stimuli which gave positive responses when moved, were all nega-
tive when motionless. See pp. 191 and 193 for tested species; all were given motion response
tests in the same session. No stimulus which was unsuccessful when moved drew display when
motionless.
Series E. Use of Models. No motionless model, even when successful in motion, drew a positive
response. See p. 190. Test individuals were given motion-response tests in the same session.
1949]
Crane: Salticid Spiders: Analysis of Display
187
addition to the stimulus situation to draw
a prompt display.
This manual jerking of the mount of a
chloroformed or fresh dead female was as
effective as the female’s own motions in
drawing display. Her further motion was
not essential, once the male had initiated
display ; it is, however, customary in normal
courtship for even A-tone females to turn
and twist and, especially, to retreat a few
inches at least once or twice during courtship.
Also, the finer motions of females during
male display almost certainly have value. In
various species — in C. xantliopa, for example
—females early in courtship brace them-
selves high in a position similar to a pre-
threat stance in males. Just before or after
the beginning of the male’s Stage II, they
crouch low. Again, in very many species, the
females vibrate the palps rapidly during the
early stages of courtship. Finally, in a few
species, notably C. fulgipedia and Sassacus
ocellatus among Rancho Grande examples,
excited females often make weak copies of
the males’ motions, with occasional individ-
ual quirks of their own. None of these female
motions ever proved to be in the least neces-
sary either to release or to direct male dis-
play. There seems no question, however, but
that they have minor directive value, and
the female’s eventual, crouching quiescence,
as in other animal groups, certainly acts as
a positive factor in permitting the continua-
tion of Stage II. Also, it is this crucial point
which in intermediate genera largely deter-
mines the continuation of display as Stage
II courtship or as contact threat or actual
fighting. In final summary, however, the
function of female posture and motion
changes is minor: in no genus which I have
observed closely does the release of Stage
II depend, in lock-and-key fashion, on any
motion, or cessation of motion, in the female.
From the female’s viewpoint, the sight of
a male making appropriate courting motions
is essential for acceptance. Exactly what con-
stitutes, for each species “appropriate mo-
tion’’ is, however a completely unsolved ques-
tion of obvious evolutionary interest. Since
no method has yet been devised of testing
this aspect, only the following general re-
marks may be made.
Little or no notice is taken of motionless
males, although occasionally an A-tone fe-
male will approach or even touch one, without
any attempt to injure or eat him (cf. be-
havior toward a partly blinded male, p. 185) .
No male was ever allowed to mate without
complete courtship for the species, although
sometimes the whole display was raced
through in less than a minute, with no repe-
titions of stages, where both individuals
were of high tone. In the occasional instances
where males, because of low tone or a con-
fusing experimental factor, omitted display
and leapt directly on the female, she in-
variably drove him off or escaped.
Many males of high tone court females of
similar-appearing or closely related species.
These females are often attentive for long
periods and make no effort to attack or re-
treat. It is usually the female that with-
draws, often before Stage II is reached. It
has yet to be determined how much her with-
drawal depends on the, for her, “incorrect”
pattern of male motion, and how much on the
unsatisfactory character of other elements in
the stimulus configuration.
In the release of inter-male threat display,
motion is usually essential. Once a C. xan-
thopa, of especially high A-tone, did per-
form threat to a chloroformed male; this,
however, was the only exception in more
than 25 trials on different species. Those
males which react to their mirror images
will very rarely display unless the mirror
is moved jerkily forward.
One more point should be emphasized in
regard to motion in general. The positive
responses to motionless stimuli, of whatever
nature, usually occur when the test spider
is dropped suddenly near the stimulus. It
seems likely that the visual effect to the
spider may be similar to that obtained when
the stimulus is moving — roughly analogous
to a human passenger’s confusion when one
of two parallel trains starts to move.
To sum up: Male display is sometimes
released in the presence of motionless fe-
males. In these cases, however, the males
are of extremely high epigamic tone (A-
plus), the females of unaltered appearance,
and the general external conditions propi-
tious. Therefore, although movement of the
stimulus is not a primary releaser for court-
ship, it is an important secondary releaser,
and, doubtless, a director as well. Appro-
priate motion of the male is essential for
acceptance by females. The response to mo-
tion does not appear to vary with the degree
of specialization within the family.
iii. Distance. The distance at which sal-
ticids perceive their prey and mates, and
at which they start display, has been meas-
ured for a variety of species (e.g. Peckham,
1894, p. 242; Homann, 1928, p. 247 ff.). In
this study it is pertinent to add the follow-
ing remarks and conclusions.
Corythalia xanthopa, which averages
around 4.3 mm. in length, is typical of mod-
erately small species at an advanced stage
of visual evolution. Males will come to a
state of attention and approach a female on
the same level, from a maximum distance
of ten inches, but the usual limit of attention
(as distinct from display) is not more than
six inches for courtship, much less for threat.
Neither courtship nor threat usually takes
place at more than three inches, and usually
at 1.5 to two inches. At the resumption of an
interrupted series, however, it may start
at 3.5 to four inches. Stage II, in courtship
or threat, starts at less than one inch, usually
at about half an inch. C. chalcea and fulgi-
pedia, which have apparently equal visual
dependence with xanthopa, are both some-
what larger than the latter; in correlation
188
Zoologica: New York Zoological Society
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1949]
Crane: Salticid Spiders: Analysis of Display
189
TABLE VI.
Spectral Reflectance of Paints Used in Salticid Experiments.
Conditions: Ultraviolet (UV) reflectance judged optically from comparative inspection of
negative images of paint samples; lens, Leitz 50 mm. Summar; film, Eastman Super XX; filter,
Wratten No. 18A; exposure, 1 sec. @ F 3.5; illumination, light from north sky, partly cloudy,
noon, June, latitude 10° 21' north. Spectral reflectance readings from curves furnished by Elec-
trical Testing Laboratories, N. Y., made from same samples on Hardy spectrophotometer; re-
flectance factor in terms of magnesium oxide.
|
Reflectance Factor
Wave-
length
(m/x)
UV
White
(Devoe)
very
strong
Blue
(Devoe)
strong
Green
(Devoe)
weak
Yellow-
green
(Devoe)
very
weak
Yellow
(Devoe)
very
weak
Yellow
(Flo-
quii)
very
weak
Orange
(Devoe)
very
weak
Red
(Devoe)
weak
Red
(Flo-
quil)
very
weak
Red-&-
white
(Devoe)
(Pink)
strong
Black
(Devoe)
very
weak
400
.715
.432
.080
.067
.115
.067
.050
.080
.030
.450
.025
410
.725
.450
.082
.070
.120
.067
.055
.087
.032
.462
.032
420
.735
.480
.090
.075
.130
.067
.057
.092
.033
.477
.035
430
.750
.515
.095
.082
.145
.067
.058
.095
.035
.492
.035
440
.755
.535
.110
.090
.165
.068
.060
.098
.035
.512
.035
450
.765
.550
.135
.107
.197
.070
.063
.013
.035
.522
.035
460
.770
.545
.180
.145
.250
.070
.065
.012
.037
.520
.035
470
.775
.505
.255
.215
.330
.075
.067
.095
.038
.518
.035
480
.780
.440
.365
.345
.425
.078
.068
.087
.040
.513
.035
490
.782
.365
.475
.475
.520
.087
.072
.085
.043
.507
.036
500
.785
.280
.545
.565
.597
.105
.075
.077
.045
.498
.037
510
.785
.215
.552
.617
.640
.157
.077
.072
.052
.492
.037
520
.787
.160
.520
.637
.667
.265
.080
.068
.057
.488
.037
530
.787
.125
.470
.637
.680
.402
.083
.070
.060
.493
.037
540
.790
.097
.405
.620
.685
.515
.105
.070
.062
.502
.037
550
.790
.080
.335
.592
.692
.575
.160
.067
.067
.507
.038
560
.792
.067
.275
.560
.697
.615
.270
.068
.075
.510
.038
570
.795
.060
.220
.535
.700
.640
.420
.075
.080
.530
.038
580
.800
.057
.185
.525
.705
.657
.550
.100
.085
.580
.039
590
.805
.055
.167
.523
.713
.670
.625
.180
.115
.650
.040
600
.810
.057
.155
.525
.715
.677
.665
.335
.195
.715
.040
610
.812
.060
.147
.532
.725
.678
.688
.550
.310
.747
.040
620
.817
.062
.137
.550
.737
.678
.705
.575
.395
.755
.040
630
.820
.067
.132
.577
.747
.678
.718
.625
.450
.777
.040
640
.825
.080
.135
.615
.752
.677
.735
.657
.478
.787
.040
650
.830
.092
.145
.640
.763
.677
.750
.680
.493
.798
.040
660
.835
.115
.167
.657
.777
.678
.765
.700
.502
.807
.041
670
.845
.145
.195
.665
.790
.680
.780
.722
.510
.818
.041
680
.850
.190
.225
.657
.795
.683
.792
.740
.513
.827
.042
690
.855
.242
.280
.653
.805
.685
.802
.755
.522
.838
.042
700
.860
.316
.342
.663
.815
.688
.813
.767
.527
.843
.043
710
.863
.392
.407
.692
.820
.690
.820
.773
.532
.850
.043
720
.865
.465
.463
.720
.823
.693
.827
.787
.535
.853
.042
730
.867
.530
.515
.745
.827
.697
.832
.795
.538
.858
.042
740
.870
.577
.558
.770
.830
.698
.835
.780
.540
.860
.042
750
.872
.610
.595
.790
.834
.702
.840
.887
.542
.862
.042
they have slightly greater attention and dis-
play limits, thus illustrating a general prin-
ciple of the spatial limits of salticid re-
sponses.
In comparison with Corythalia, species
which still place a relatively high depend-
ence on chemotaxis and/or distance chem-
operception, appear relatively short-sighted.
Thus, Ashtabula and Semorina, although
similar in length to C. xanthopa, rarely give
evidence of first attention at more than four
inches, and usually much less. Display be-
gins even closer, and, as always, Stage II
starts closer than Stage I.
It must be kept in mind that two spiders
are seldom in the same plane in the field;
therefore, one customarily enjoys the ad-
vantage of an obliquely downward view on
the expansive dorsal surface of the other.
Once more the importance of the physio-
logical state should be emphasized: A-tone
males tend to start attention and display
responses at greater distances than others.
iv. Size. The display responses of salticid
males to stimuli of unusually large and small
size do not appear to have been previously
investigated. In this study C. xanthopa was
subjected to a series of experiments, the
190
Zoologica: New York Zoological Society
[34: 17
distinctness of its threat and courtship re-
sponses making it an especially interesting
species. Because of the importance of dis-
tance chemoperception in its courtship re-
leasing mechanism, males rather than fe-
males were usually selected as stimuli. The
results of the tests appear in Table V. In
casual observations on other species, primar-
ily concerned with inter-specific display, it
was always true that where two species had
display relations, there was never much dis-
crepancy in size.
Conclusions : First, Appropriate size is an
important secondary sign stimulus to display.
No male xanthopa will give any kind of a
positive display reaction to a spider with a
frontal area of more than five times, or less
than one-third, of his own. In other words, in
linear measurements, positive responses may
be given to stimuli measuring up to about
twice natural size and down to about one-
half.
Second. The responses within this range are
highly variable and signs of confusion are
frequent. The courtship response is often
given in these unusual size ranges to stim-
uli to which a threat response would seem
more appropriate.
Third. Addition of a clypeal band of yellow
paint to an otherwise black frontal view,
sometimes changed negative to positive re-
sponses. This yellow resembled, to human
vision, that of xanthopa males. Both brands
used had slight reflectance below about 520
mg (Table VI).
Fourth. As usual with unnatural stimuli,
responses were usually obtained only by de-
liberately reinforcing the stimuli. This was
accomplished by repeated presentations in
quick succession, and constant chivvying to
keep them moving in the direct front visual
field of the spider. Spiders below A-tone
gave consistently negative results.
Fifth. There was a tendency to respond to
large stimuli at longer range than the nor-
mal.
v. Form. The general shape of the spider
is another secondary sign stimulus for re-
leasing display. Heil (1936), working with
life-size pictures of Evarcha, performed
some experiments on this subject. He found
that the number of positive display responses
decreased rapidly with the simplification of
the drawing, particularly in the reduction
of lines representing the legs. The following
results, which support and extend those of
Heil, were obtained at Rancho Grande.
As in other factors, the importance of
form varies inversely with the normality of
the test situation, and with the drive of the
individual; an A-tone C. xanthopa of espe-
cially strong drive will respond (with threat)
to a jerked, life-size paper model of great
simplicity, to the jerked carapace of a dried
male, or to a jerked or naturally moving nor-
mal male with a high black paper “hat.” None
of these extremes is successful under other
conditions, or in combination with one an-
other. For example, none of them draws dis-
play from a male of mediocre drive, while a
legless paper model or a motionless carapace
is unsuccessful even with males of the high-
est tone.
Because of the variety of contributing fac-
tors, a tabular representation of the experi-
ments in this series would not give as true a
synopsis of the results as the following run-
ning account with selected and annotated
field notes. All of the experiments were per-
formed under the conditions described under
Table VI. In this series the tested species
were Corythalia xanthopa, C. chalcea and
Phiale flammea, all visually advanced species
in which chemotaxis and distance chemoper-
ception are of reduced importance.
Models. Cardboard models were tested on
male C. xanthopa. Drawings of successful
and unsuccessful models, made through a
binocular microscope, are shown in Text-fig.
7. “A” represents a careful drawing of a male
in threat position, for comparison, and was
not used as a model; such detail, would, of
course, have been impossible to include on a
Text-fig. 7. Examples of models used in experi-
ments on form-pattern-color perception in
Corythalia xanthopa. The stippled areas were
painted yellow, the dashed areas white, the
backgrounds light green; spectrophotometric
data in Table VI. A, detailed drawing of C. xan-
thopa in threat position; B-E, drawings of paste-
board models which drew threat responses, in
order of their success, from the most (B) to
the least (E): F-J, unsuccessful models. Note
that the only difference between successful and
unsuccessful in some cases was the presence or
absence of a yellow median area. See text.
1949]
Crane: Salticul Spiders: Analysis of Display
191
surface 5 mm. across. Similar models, but
relatively larger or smaller than those shown
were never successful. Even the unsuccessful
models (right column) drew some attention
from A-tone males. All were drawn in Hig-
gins India Ink on bits of pasteboard ; inter-
stices between “legs” were painted pale green
(see wave length analysis, p. 189 for this and
other colors mentioned hereafter), from the
same jar as the mounts and similar in reflec-
tance curve to the paper substratum. This
tint, used frequently as background in series
of experiments involving living stimuli, ap-
peared, from the tested spiders’ behavior, to
provide for them excellent contrast.
The most successful model was B, the least
successful in the positive series, E. The best
of them, however, drew only delayed display.
It will be noted that all successful models had
the following characteristics : a roughly
squared or rounded central portion, painted
matte black and yellow, with a greater or
lesser series of lateral extensions. The “eyed”
model (F) was never successful, nor were
models similar to or identical with the suc-
cessful one except for black or white in place
of yellow in the central region. Plain black
quadrilaterals — i.e., without “legs,” but cov-
ering the same area as B and D — were always
unsuccessful, even when furnished with a
median yellow spot or bar. Also unsuccessful
were all more extreme models, including
equilateral and isosceles triangles; the latter
were tried both vertically and horizontally,
and all were furnished with a median yellow
area. Finally, no small, median portion, rep-
resenting the carapace only, ever drew a
response.
Form Experiments with Specimens.
General Shape. The following experiments
also show the necessity for a general shape
approaching the normal. A young adult fe-
male C. chalcea was lightly chloroformed,
placed on an L-mount and covered with a bit
of flimsy black silk (chiffon), which scarcely
enlarged her, but effectively concealed her
shape. When she was jerked before a high
tone male, no display was drawn. However,
when the whole was surmounted by a large,
dried Eustiromastix sp. (a male, plain dark
brown with a painted yellow clypeus), the
chalcea promptly courted, following through
to attempted mating — not with the hidden
chalcea, but with the Eustiromastix. Single,
fresh abdomens, of both males and young
females, were presented to males of xan-
thopa, chalcea and P. flammea; none ever
drew display, or even attention, fi'om a total
of about 25 tests. Detached fresh legs, sin-
gly or in groups, were equally ineffective.
Importance of Legs. A legless, dried male
xanthopa cephalothorax was very rarely suc-
cessful in drawing threat display. Similarly,
a legless and palpless young female chalcea
(but with abdomen intact) drew delayed
display in only three of nine tested males.
However, none of these positive responses
was complete : one male broke off display and
jumped, several times in succession, at the
female; another, after a delayed Stage I,
merely climbed onto the cardboard mount
beside her, palpating its surface; a third
reached Stage II, but did not follow through
to twisting the abdomen, his display dying
out in palpation of her body and the sur-
rounding mount.
A single real leg was then laid flat at each
side of the female, without drawing a re-
sponse; but when two pairs were used, de-
layed but complete courtship followed.
When a sling of stiff black cotton, shaped
like a broad, inverted W, was substituted for
real legs, brief and abortive display followed.
When a second sling was added, so that the
front view showed lateral stripes of alternat-
ing light and dark, prompt, complete court-
ship ensued. When the second sling was re-
moved, there was once more no response,
which however followed promptly upon the
second sling’s replacement.
Throughout all the experiments with xan-
thopa, it proved axiomatic that mounted
dried or chloroformed males with the legs
stretched out at the sides drew threat dis-
play from males of mediocre drive faster
than those in a huddled position.
Upsidedown females of xanthopa, chalcea
and P. flammea were tested. These almost
never drew display, the rare exceptions being
from males of the very highest tone. The dis-
play never started until abnormally close to
the stimulus and Stage I was exceptionally
brief ; occasionally a male would approach a
female without display, palpating her for a
time before wandering off. No actual mating
attempts, which were always preceded by
some display, were ever successful, the males
groping about in evident confusion. Upside-
down males, on the other hand, never di'ew
threat display or, in fact, more than passing
attention.
Importance of Carapace Height. Compared
with the legs, the height of the carapace is of
very minor importance. Its elevation off the
ground has no perceptible significance, judg-
ing from the prompt response of male xan-
thopa to the various mounts and of other
forms, including Menemerus, Sassacus, C.
chalcea and Phiale, to various dried or chloro-
formed mounted examples of their own
species, none of which was ever presented
balanced in life-like fashion on the legs. It
must be remembered, however, that in nor-
mal display one of the most usual factors is
the elevation of the carapace high on the
legs.
Black and white paper “hats” of various
sizes and shapes were fastened above the
eyes to males and females of C. chalcea and
P. flammea with rubber cement. So long as
other conditions approached the normal, ap-
propriate display was delayed little or not at
all, and a female chalcea readily accepted a
black-hatted male. However, in the case of
hatted females, the males did not carry dis-
play through to actual mating attempts, in
192
Zoologica: New York Zoological Society
[34: 17
this small set of “hat” tests which totalled
about a dozen. The highest addition tried
measured twice the height of the clypeus-
plus-AME-diameter.
Conclusions. First, shape is of secondary
significance in the release and direction of
display.
Second, deviations in shape which do not
hinder display release nevertheless often
prevent mating.
Third, the primary shape requisite for re-
leasing and directing either courtship or
threat in Corythalia and PhiaXe are a roughly
quadrilateral figure, broader than high, the
lateral portions of which show some sugges-
tion of dark and light horizontal stripes.
Fourth, the vertical dimension is less im-
portant than the horizontal as a factor in
display.
Fifth, when the shape of a stimulus de-
viates from the normal range of variation,
the tested spider must be of high tone and
the other factors in the display situation
must in general be normal, in order to draw
a display response.
vi. Pattern, Intensity and Color. The parts
played by pattern, intensity and color in
courtship and threat displays are, as in other
groups, exceedingly difficult to determine and
to distinguish ; in the present study only the
surface of the question has been scratched.
It should be pointed out that the distinction
made here between “shape” and “pattern” is
obviously artificial, although, in the preced-
ing section on shape the emphasis was on the
general form of the figure, rather than on
the details; however, the stimulus value of
the legs certainly belongs as much under
“pattern” as under “shape.”
The Peckhams (1887, p. 403) were the
pioneers, as usual, in investigating color per-
ception in spiders. Working altogether with
lycosids, they approached theproblemthrough
a choice method, the lid of the cage being
covered with glass squares differing in col-
or. Since neither intensity nor wavelength
was controlled, their results can have no final
significance for modern workers. Neverthe-
less, it is very suggestive that the vast major-
ity of Lycosa chose “red,” since these spiders
are largely negatively phototropic ; very
likely the hunting spiders in general, like
the majority of insects, will prove to be rela-
tively insensitive to the red end of the
spectrum.
The Peckhams performed another series
of experiments, painting female salticids
partially or completely “blue.” Following
painting, the previously displaying males
paid them much less attention, either not dis-
playing or delaying the response, although
it was resumed promptly when normal fe-
males were introduced. A number of uncon-
trolled factors were of course involved in
this series (1894, p. 249).
Kaestner (1949) has just published a pre-
liminary report indicating that color percep-
tion does occur in the European genus
Evarcha.
Apart from the question of actual color
perception in salticids, the minor role played
by color, or at least by lightness and dark-
ness, in courtship is shown by Maevia vittata,
the North American salticid with dimorphic
males. Painter (1913) seems to have been the
most recent worker on the subject. The di-
morphism is controlled by the presence or
absence of a small sex chromosome, and in
the population studied the two forms were
about equally abundant. The principal dis-
tinctions consist of general color — black in
one, gray with orange palps in the other —
and the presence of a pair of tufts on the
carapace in one, absent in the other. In spite
of considerable individual variation, the two
forms are quite distinct visually; in addi-
tion, the display patterns show differences.
Nevertheless, both were readily accepted by
females, as were dark males with their tufts
missing. According to the Peckhams (1889,
p. 54), who worked in a different part of
the United States, the darker, tufted form
was more aggressive, and was preferred by
the females; also, differences in display of
the gray form from that described by the
Peckhams were reported by Painter. In spite
of the fact that in neither study were con-
trolled experiments performed, which guard-
ed against summation, etc., it still seems
likely that a most interesting series of be-
havior differences has evolved in this wide-
spread species. Here is certainly splendid
material for worthwhile work.
The disagreements between the Peckhams
and Painter do not affect the conclusion that
degree of lightness or darkness in males is
not, in that species, of primary importance in
gaining acceptance by the female.
In the present experiments, two groups of
colors were used, the Floquil “Flo-paque”
series and Devoe opaque water colors. Since
spectrographic analysis could not be con-
ducted in the field, library cards were painted
with each hue and combination used; later
these were analysed spectrophotometrically
(Table VI), along with fresh samples from
new jars. The latter check was employed to
determine the degree of changes in the sam-
ples since their field use; these differences
proved insignificant. Although the paints are
far from a theoretical monochromatism, most
of the hues do show steeply ascending curves
near the regions of highest reflectance.
During the field experiments, the paints
and the yellow of C. xanthopa (cut in two,
without exposure to chemicals, just before
the photograph was made) were photo-
graphed through Wratten Filter No. 18A,
which screens out virtually all light except
the ultraviolet; the spider’s yellow areas did
not perceptibly affect the negative according
to observations through a binocular micro-
scope. (Spider photographic data: lens, Leitz
90 mm. Ektar; exposure, 13 sec. @ F12.7;
other data as in Table VI) . In view of these
1949]
Crane: Salticid Spiders: Analysis of Display
193
negative results, it is apparent that ultra-
violet is not a factor in the effect of xanthopa
yellow as a sign stimulus; they do not of
course, preclude the possibility of the spider’s
visual sensitivity to ultraviolet wave-lengths.
So far the yellow of the spiders has not been
further analyzed : dried specimens appear to
be too changed to give trustworthy results.
It will now be shown that the various
characteristic “display ornaments” are in
themselves of minor importance in the sam-
ples studied as display sign stimuli, except in
certain inter-male relationships. C. xanthopa
was the subject of most of the experiments;
C. chalcea, C. fulgipedia, Menemerus bivit-
tatus, Phiale flammea and P. dybowskii were
used to a lesser extent; a few illustrations
were taken from Semorina, Ashtabula and
Sassacus.
The Role of Female Pattern, Intensity
and Color in Stimulating Male Display. The
female’s pattern and color equipment may be
divided between that of the frontal and the
dorsal (particularly abdominal) regions. It
must be kept always in mind, however, that
in practice the male’s first view of the female
is often obliquely downward, so that both
regions are simultaneously visible. In the ex-
periments below, the two regions were kept
separate, the stimuli being presented on a
plane surface in direct frontal or rear views.
Frontal pattern. (Text-fig. 8). In females,
this may be divided functionally into two
well-marked classes; first, differentiation of
frontal markings, including those of clypeus
and palps as a unit, and, second, distinctions
concerning the palps alone. The sexual dif-
ference of frontal markings is noticeable in
many salticids, but ranges through all de-
grees and is often absent. The following ex-
amples are from the present series: In C.
xanthopa and Phiale spp., the females are
black-faced while the males have strong yel-
low or white clypeal bands which continue,
when the palps are bent, onto certain palpal
segments. In contrast, in C. chalcea, C. fulgi-
pedia and Eustiromastix sp., the females are
frontally moderately or very hairy, chiefly
gray and white, while the males are almost
completely black and naked. In Menemerus
bivittatus, both sexes have the hairs pale,
but the female more so than the male, where
they are confined to a white clypeal band.
Finally, in Mago acutidens,the frontal region
is black and naked in both sexes. It should be
remembered that the legs, particularly the
first, are often much darker in males than
in females.
Experiments were made with C. xanthopa
and P. flammea, both having sexual dimor-
phism in this respect, to determine the effect
on the male of alteration of the clypeus.
When normally black-faced young female
xanthopa were given yellow clypei and palpi,
resembling those of the male, the courtship
display was always drawn as promptly as
ever with no hint of threat or confusion.
Chemotaxis, but not motion or distance chem-
Text-fig. 8. Examples of frontal sexual di-
morphism and types of “ornamentation” in
representative salticid genera. A, Lyssomanes
bradyspilus, female; B, C, D, same, male, show-
ing three positions of ocular color “change”;
E, Menemerus bivittatus, male; F, same, female;
in the male, one-half of the white clypeal band
has been shaved off to show narrowness of true
clypeus; G, Phiale flammea, male; H, same fe-
male; I, Mago dentichelis, male; J, same, fe-
male. All figures drawn to same scale; note how
in Mago the individual female depicted happens
to be smaller than the male ; this often occurs in
salticids, although the carapace size of the aver-
age female in a given species is always some-
what larger than that of the average male.
The adaptive value of the “ornamentation”
is by no means equal throughout the family. In
Phiale, the clypeal band of the male is a releaser
for threat display; in Menemerus there seems
normally to be no inter-male display at all,
although a similar clypeal band is well devel-
oped, nor is it of apparent importance in court-
ship; Mago, lacking all clypeal “ornamenta-
tion”, but having a plain, high, black carapace
and clypeus, has both courtship and threat well
developed and moderately well differentiated.
Note the prevalence of pale palps in the female;
these are usually vibrated during display, but
do not, in the several tested genera, function
as either primary or secondary releasers of
male courtship. Sexual dimorphism varies from
negligible (e.g., Mago ) to extreme (e.g., Mene-
merus) ; see also Part I, 1948.1, text-figs. 12-15
inch, for examples of the latitude of dimorphism
within a single genus, Corythalia.
194
Zoologica : New York Zoological Society
r 34 : 17
ical stimuli, was excluded. (Test-material: 2
females, 6 males, 9 tests).
Four weeks later, one of the same females,
still unmated, but now more than “middle-
aged” for that species, was given a new coat
of paint and presented to a fresh series of
males. The results were quite different:
(Test-material: 1 female, 7 males, of which
6 were of A-tone and 1 of B-tone; 10 tests.)
Fii'st round: 2 threat, 3 courtship, 2 unde-
cided ; second round (3 only tested) : 1 threat
(last time this individual was uncertain), 1
courtship (last time this individual also
courted), 1 courtship (last time this individ-
ual was uncertain). Total of responses: 3
threats, 5 delayed courtships, 2 confused re-
sponses (including that given by the low-tone
individual) . The threat reactions were all
promptly given, while courtship followed a
period of uncertainty characterized by at-
tention, “following” with the eyes, and in-
termittently dropping the abdomen (i.e., al-
ternating in courting and threat position).
It seems apparent, in conjunction with other
experiments, that the waning airborne chem-
ical stimuli of the female were here respon-
sible for the inclusion of threat and uncer-
tainty in the results.
Two male xanthopa were now altered; in
one the yellow scale-hairs of clypeus and
palps were painted black; in the other the
palps were detached and the clypeal scales
scraped off with a razor. The results (5 males
tested, 5 tests) consisted of 2 very rudimen-
tary brief courtships (1 single rock each), 1
abortive, brief, courtship-plus-threat and 2
responses which were negative except for
brief attention. When a mirror was pre-
sented to the palpless, shaved male, as he
stood on the same spot where a young fe-
male had just been sitting, he gave prompt
attention, coui’tship stance and a few rocks,
then retreated briefly, and ended by leaping
at the mirror, but above the image level. The
same response was repeated on fresh paper,
but since no rest time was given in this in-
stance, summation may well have been in-
volved ; unfortunately because of an accident
to the shaved male, and lack of time for repe-
tition, the testing was not resumed.
When similar experiments were conducted
on Phiale, the results were as follows :
When a normally black-faced young P. dy-
boivskii female was given white spots on the
palps and, later, a white clypeal band similar
to those of the males, repeated tests always
drew complete courtship, not threat.
When an old P. flammea female was simi-
larly treated, two males courted relatively
promptly so long as she was motionless, but
approached her, for mating, from the side,
out of sight of the abnormal, white palps and
never got properly oriented toward the
abdomen.
When the female mount was moved during
the presentation, the male first gave threat,
changing to a prolonged, side-anproach court-
ship only when he was unusually close to her.
When her paint was washed off, both males
gave prompt, normal, complete courtships.
The entire frontal aspect of a young chloro-
formed C. chalcea female, excepting the eyes,
was painted white. As she sat motionless on
a mount, an A-tone male was introduced with
the following results : “He soon vibrated
palps in the air, making semi-circles until he
came close to the mount. Then he palpated
and further vibrated the palps, circling off
on the surrounding paper and returning to
mount time and again. However, there was
no trace of display, and no attempt to mate.
It was exactly like the behavior of a male
introduced to the spot where a female had
recently been sitting or moving about. Fi-
nally, after more than five minutes, he went
away. When he was brought back, to a dis-
tance of about two inches, the female was
jiggled infinitesimally. He displayed at once,
with good courtship. The first mating at-
tempts in Stage II were from her unpainted
rear. Then he displayed Stage I again and
approached with a typical Stage II from the
front.” (Field lab. note).
One typical feature of the female frontal
appearance in many species is the rapid vi-
bration of contrastingly pale palps at a cer-
tain stage of courtship; it occurs usually in
the earlier phases, during her first apparent
awareness of the male’s Stage I. Sometimes,
as in Semorina, Ashtabula, Phiale and Mago,
contrast to the otherwise dark front (in-
cluding or excluding legs), is attained prin-
cipally through pale integument; sometimes,
as in Menemerus, the effect is accentuated by
long white fringes. In species such as C.
chalcea, where vibration of the palps by the
female is of casual and infrequent occur-
rence, they are not notably lightened or “or-
namented.” When present, the paleness and
“ornamentation” usually extends to most
segments, unlike special spotting with scale-
hairs of certain segments which is so typical
of some males, as in C. xanthopa and Phiale.
In order to determine their importance as
display sign stimuli, the female’s palps were
removed from examples of Menemenis and
Phiale. In every case the males courted the
mutilated females as readily as normal forms.
( Menemerus : 1 female, 8 males, 10 tests;
P. dybozvskii: 1 female, 1 male, 2 tests.)
Palps were not removed fi'om any of the other
species, but males of all genera mentioned
in the preceding paragraph always displayed
as readily to a chloroformed female when
her mount was slightly twitched as to a nor-
mally moving individual; the separate mo-
tion of contrasting palps is certainly not
even a secondary sign stimulus.
In most femaies, as in males, the eyes are
rimmed more or less conspicuously with red-
dish, yellowish or white scales. Since males
display as readily to clumsily blinded fe-
males, in which all trace of the scales is
covered, as to normal ones, these scalesseem
to have no primary or secondary display
value in modern times.
19491
Crane: Salticid Spiders: Analysis of Display
195
Dorsal pattern: Dorsal carapace markings
are almost always absent in the species
studied, except in C. fidgipedia and Phiale,
where they are highly variable and practi-
cally identical with those of the male; the
white submarginal carapace bands charac-
teristic of many species do not appear to be
displayed in any of the ones I have observed
and, indeed, short of a theoretical lateral dis-
play, involving flattening of the legs, it is
difficult to see how they would ever be plainly
visible.
The females of C. xanthopa have a naked
black carapace, but a prominent abdominal
pattern, consisting of a pair of broad, longi-
tudinal, yellow stripes on a black ground.
Males "frequently display to the rear view
of normal females before they have seen her
dark front view at all. Blackening of the
female abdomen produced no change in reac-
tion time (1 female, 2 males, 6 tests). Then
two young females were selected, of similar
age and condition ; the abdomen of one was
blackened ; they were then placed, facing
away from the center, about eight inches
apart on the ten-inch line of the concentric
test circles (Part II, 1948.2, p. 145), this
being about the limit of perception-response
in this species. Six males in A-state were
then placed in succession in the circle’s cen-
ter, facing exactly between the two females.
In every case the male became first aware
(as judged by his shifting position to look
toward her, and the adoption of preliminary
courtship stance) of the female whose mount
was twitched first, no matter how slightly.
His progress toward her was invariably in-
terrupted by a movement from the other
female, regardless of her color. By the time
the male’arrived within actual courting dis-
tance (several inches away), he would either
court first one, then the other, impartially,
or simply sit, restlessly, between them in
apparent confusion. In all, four of the six
males courted somewhat, and two sat quiet
at courting distance and eventually retreat-
ed. Of the courters, two eventually stopped
and ran between the females out of range,
one retreated, and the fourth’s test was
broken up when the normal female came
out of the chloroform and moved off. At that
point the male made no attempt to follow,
but concentrated at once on the black female
nearby, now not being twitched, and followed
through into advanced Stage II before they
were forcibly separated. A second round of
tests, using the same males, brought similar
results, although a given male did not always
behave in the second test as in the first. The
overwhelming effect in all tests was uncer-
tainty and conflict. The only possible dif-
ference noted in the response to the two
females was a very slight tendency to notice
the normal female first at the greatest dis-
tance (10 inches), w'hen both were twitched
simultaneously.
Thus, although the female’s pattern cer-
tainly has no value as a releasing stimulus,
it seems likely that in the long run the two
light stripes may have some slight directive
value in the field. Under those natural con-
ditions, three-dimensional vision is of course
the rule. Therefore, the striking contrasts
may make the attraction of the male’s at-
tention from farther than 10 inches, beyond
the range of distance chemoperception,
easier, among the dead leaves and shadows.
To human vision, both males and females of
this species are conspicuous in their normal
habitat. However, I have never been able to
gather the least evidence of this possible
function, although it has been kept con-
stantly in mind in field observations. In re-
gard to C. fulgipedia females, which exhibit
an abdominal pattern and color stinking
enough with their strong black and white,
there are no distinctions from either im-
mature specimens of almost equal size or
from those of adult males; therefore the
adaptive nature of abdominal pattern in any
fashion, except that perhaps of disruptive
coloration, is even more unlikely.
In female chalcea, any adaptation to dis-
play use is the most questionable of all; its
dull bronzes, pale gold and black bandings
are not only indistinguishable without a mi-
croscope from those of the preadult female,
but they merge exceedingly well into dead-
leaf or tree-trunk backgrounds, and when
the spider moves it is even less conspicuous
than an all-black form. If the color and pat-
tern have any adaptive significance, it
would appear to be as disruptive coloration
of a protective nature; certainly it cannot
help in attracting the attention of the male.
A further point is that chalcea and fulgipedia
females, although of strikingly different ap-
pearance, each draws display from males of
the alternate species even when the males
have only a posterior view.
When the abdomens of xanthopa females
were painted solidly yellow to resemble those
of males, there was no confusion: as when
the clypeus and palps were painted, the male
courted from the rear if the female was a
young adult; as in the case of the altered
front view, he became confused with old
painted females.
In Phiale, the quite constant flaming rusty
red of many species in the male is variable
in females, ranging from almost male vivid-
ness to nearly black. The white spots and
bars, though also variable, are nevertheless
always present and, to human eyes, conspic-
uous, especially posteriorly. However, when
the female abdomen was painted uniformly
black in P. flammea, three males courted her
promptly from the rear at close range. Never-
theless, in this genus pursuit is a funda-
mental, normal part of the courtship, and
as often as not the male is displaying for
a time to the female’s posterior view. It
seems therefore that while female white pos-
terior markings have no releasing value,
they, like the xanthopa stripes, are probably
directive in the sense that they facilitate
following.
196
Zoologica: New York Zoological Society
f 34 : 17
In perhaps a majority of genera, the ab-
dominal pattern appears completely undis-
tinguished, except as probable examples of
protective coloration. In Sassacus ocellatus,
one of the species in which the female abdo-
men is both striking and closely similar to
that of the male — i.e. iridescent green with
posterior black-and-white spots— it is note-
worthy that abortive female display was
once observed (Part III, 1949, p. 46).
There is, however, one female abdominal
marking which appears of importance in
courtship. This is the sub-basal pale band,
usually white, which is of such frequent oc-
currence in both sexes throughout the family.
It is found even more often in females than
in males. It is present in all of the adult fe-
males discussed in the present paper except
Lyssomanes, Semorina, Ashtabula and Cory-
thalia xanthopa. It always crosses the high-
est point of the abdomen and is preceded
only by the naked black region immediately
behind the pedicel.
In order to test its display value, abdomens
were variously covered with white, yellow,
black or red paint in females of C. xanthopa,
C. chalcea and P. flammea. In painted xan-
thopa, in which the abdomen is normally
striped, display proceeded though normal
mating, as described earlier. In the other two
species, which normally have sub-basal bands,
the males appeared to become confused in
the middle of Stage II, when the time came
for twisting the abdomen. The following in-
stance is typical: A chalcea female was
painted completely white above, except for
the thoracic slope of the carapace, which was
inadvertently left black. Three males in
seven tests courted normally until advanced
Stage II, whereupon all of them tried to twist
the carapace, not the abdomen, and made
fumbling efforts to insert the palp too far for-
ward. All of them gave up and moved off.
When black paint was applied to this area
also, lest the behavior difference involve neg-
ative chemical or other stimuli from paint,
the same reaction followed. Finally, the base
of the abdomen was blackened, the crest and
all posterior to it remaining white. The two
males still in display condition now promptly
started mating after normal courtship, in
the typical position. Cases of similar con-
fusion occurred in Phiale : When the female
abdomen was completely blackened, the males
did not locate the epigynum; when a sub-
basal white band was added in the usual
position, courtship was completed without
delay. (1 female; 3 males; 3 tests). The con-
fusion following lateral approach in response
to legless females has already been discussed
(p. 191).
The following interim summary may now
be made in regard to the role of female pat-
tern and color in stimulating male display:
First. They are of the most minor impor-
tance, in comparison with other stimulus
situation components, having no releasing
functions and only rarely a directive value.
Second. Young adult females in the tested
species, when painted to resemble males, or
when the typical pattern is severely altered,
are nevertheless courted as females, so long
as the remainder of the stimulus configura-
tion remains normal; otherwise there is no
response.
Third. Old females so painted, or males
painted to resemble females, draw delayed or
confused responses, or, sometimes, complete
threat display.
Fourth. The correlation of palp ornamen-
tation and vibration in females, although
it may well increase visibility to and/or ex-
citement in the male, is certainly not even
of secondary releasing value; males court
palpless females of Menemerus and Phiale,
as well as chloroformed — and hence quiet-
palped — females of these and other genera,
promptly and completely.
Fifth. While the white posterior markings
or other patterns of some females may in-
crease visibility to pursuing males, this is
pure speculation; the markings are not nec-
essary to release display, nor are any of the
other tested female abdominal patterns.
Sixth. The white sub-basal abdominal band,
which occurs so frequently in the family,
proves to be a directing mechanism for cop-
ulation in Corythalia chalcea and Phiale
flammea, the only two species tested. There is
contributory evidence that this is also true
in other species.
The Role of Male Pattern and Color in
Gaining Acceptance by the Female: The fol-
lowing paragraphs summarize the experi-
ments in this series. The difficulty in assem-
bling data was in bringing treated males
back to full A-tone display condition after
the operation, and in having receptive fe-
males on hand at the right time.
Menemerus bivittatus. Five females all
paid prompt attention to the display of two
males, each with the white clypeal bands
and palpal spots scraped off, showing black
integument beneath. Though all the females
allowed a male to reach the part of Stage
II in which he was entirely out of range of
her AME, on top of her, mating was not
completed in any case. However, these fe-
males on the test days rejected even normal
males, and two of them proved to be already
fertilized.
Phiale flammea. Four females paid prompt
attention to two displaying males, each with
Text-fig. 9. Antero-dorsal view of a female
abdomen ( Phiale flammea). The pale band acts
as a director for turning the abdomen in the
final part of Stage II courtship display.
1949]
Crane: Salticid Spiders: Analysis of Display
197
the white clypeal bands and palpal spots
scraped off (as in Menemerus ) ; in two cases
the males, during prolonged (20 minute)
sessions, were allowed to reach Stage II;
there were no acceptances; the females ap-
peared very sluggish, bat normal, control
males were not then available for determin-
ing the females’ true tone.
Corythalia chalcea. A female accepted
promptly a male painted to resemble C. ful-
gipedia, having white pigment on the femur
and patella of the normally black palps. These
two species display to one other freely, but
never, in numerous observations, did dis-
play proceed beyond early Stage II.
C. fulgipedia. A female accepted promptly
a male with the white-spotted palps black-
ened to resemble those of C. chalcea (see
above) . Another female accepted a male with
the frontal aspect, including the legs, com-
pletely covered with white. It will be recalled
that in fulgipedia the iridescence and fringes
of the legs are displayed in courtship as
well as in threat. Complete blackening of
the legs did not affect the female’s reaction
in the least, nor did removal of the fringes.
In three other pairs the male was variously
painted with red, from a median red spot
above the AME through covering of the cly-
peus and palps to a complete coating of the
frontal aspects (excepting the eyes) includ-
ing again the legs. Once more acceptance
was complete in each case, and within the
normal acceptance time of the species (3 to
6 minutes).
C. xanthopa. Two females each accepted
a male with the yellow of palps and clypeus
changed to white, and another in which the
clypeus had been painted black and the palps
removed. When the latter male reached ad-
vanced Stage II, he could not of course pro-
ceed and eventually backed off the carapace ;
the female, however, had showed no signs of
restlessness. Another female accepted a male
with the yellow areas replaced with orange.
From these examples it appears that de-
viations in the color and pattern of the male
play a very minor part in acceptance by the
female. To summarize: In three species of
Corythalia, altered males were readily ac-
cepted by six individuals. In Menemerus
and Phiale, although mating was not com-
pleted by frontally altered males, they were
allowed to reach advanced Stage II ; in these
two genera the available females during the
tests appeared to be in non-receptive con-
dition.
The Role of Pattern and Color in Inter-
male displays. This aspect was investigated
rather fully in C. xanthopa and less so in C.
chalcea and P. flammea. The following re-
sults were obtained :
C. xanthopa. Males of A-tone responded
invariably with more or less prompt threat
display to the following moving stimuli in
which the yellow clypeal band and spotted
palps appear to be involved: conscious,
chloroformed or ftiounted-and-dried males
with yellow, orange or yellow-plus-white
paint covering the front yellow areas; legs
painted dark brown or black (covering iri-
descence) ; two-dimensional models of appro-
priate size and general shape so long as a
yellow or orange bar or spot appeared across
the middle ; dried females painted like males ;
unrelated species of large size with a yellow
clypeal band added ; preadult male xanthopa,
or other Corythalia species of similar sub-
xanthopa size, in which males were painted
like adult xanthopa males.
On the other hand, males responded with
incomplete courtship or did not respond at
all to the following stimuli : conscious, chloro-
formed or mounted-and-dried males with the
yellow of palps, clypeus and abdomen altered
to any of the following colors : red, blue, blue-
plus-white, red-plus-blue; six intensities of
black-plus-white which included values com-
parable, by Weston meter, to those of the yel-
low, yellow-plus-white and orange which
elicit threat display ; to receptive young adult
females of other species of Corythalia
painted like male xanthopa.
To the stimuli listed below, the reaction
was variable and included incomplete court-
ship, threat, no reaction and a confused
mixture of threat and courtship. The vari-
ability was not only between individuals,
but in the same individual on different days
or even at different parts of the same test
period. These borderline stimuli are the fol-
lowing : males, or the testee’s mirror image,
with the yellow parts painted black as in fe-
males, white, red-plus-white or green ; males
with the black legs altered to white, yellow
or red ; females, middle-aged or old, painted
to resemble males; a normal male Phiale
flammea, characterized frontally by strong
white markings on a black ground. When a
male xanthopa face was blackened and the
legs painted yellow, there was no reaction.
When, as occasionally happened, a blackened,
chloroformed male suddenly became con-
scious and threatened a normal subject, the
latter either appeared startled, leaping in the
air, and retreated, or, if he were courting,
stopped courting and, in several instances,
changed the courtship to threat display.
Male xanthopa in B-state responded to
very few unnatural stimuli, and the few re-
actions which did result were so erratic and
variable that no general listings would be
profitable. The only invariable reaction was
that orange or buff-faced males were threat-
ened as promptly as yellow-faced examples.
The solid abdominal yellow of xanthopa, as
seen from the rear, drew as prompt threat
display as a front view ; males with the abdo-
men striped black and yellow like females,
also drew threat display. Males with the
abdomens blackened, however, drew confused
responses.
Unfortunately, during the inter-male se-
ries of xanthopa tests, which took place early
in the salticid experimental period, the full
198
Zoologica: New York Zoological Society
[34: 17
importance of the amount of mount-jerking
in some species was not fully realized ( infra ) .
Nevertheless, in all the xanthopa tests, the
responses did not appear in general to be
dependent on the type of motion. It can be
stated without reservation that the stimuli
which drew the most variable, confused or
inappropriate reactions (i.e., anything ex-
cept threat) were almost always ineffective
unless strongly reinforced by constant jerk-
ing and repeated presentations: in other
words, the single twitch which in C. chalcea
and Phiale sometimes made the difference
between threat and courtship responses (see
below) was inadequate for the less successful
stimuli of the xanthopa series.
C. chalcea and C. fulgipedia. These two
species, in one of which the palps are black
and in the other spotted with white, display
to each other freely, each maintaining rigid-
ly its own threat pattern.
Under experimental conditions, the follow-
ing tests were made : Two chalcea males each
displayed promptly with threat to another
male with his normally black face painted
yellow (as in the smaller xanthopa, to which
species it will not display). They also dis-
played to a larger male Eustiromastix sp.,
dried and mounted, his normal dark brown
having a painted yellow clypeus. Again, a
chloroformed, mounted male chalcea with the
legs whitened, drew various responses from
a single A-tone male; these ranged from in-
complete courtship with a poorly oriented
side approach, to prompt threat, depending
on the motion of the mount: when the latter
was not moved, attention only resulted; a
single twitch was followed by courtship ; con-
tinued jerking drew threat. When the
same mount was presented to the same male
on one of his B-tone days, the incomplete
courtship responses occurred, with the pre-
viously successful single twitch, but no
threat could be drawn with any type of mo-
tion. A male chalcea' s legs were blackened,
to eliminate iridescence, and the now com-
pletely black indivdual mounted on a black
background, and jerked slightly before an
A-tone individual. Incomplete courtship was
eventually drawn, with no trace of threat.
From the above it appears that at least in
chalcea, change of the normal black color on
the mid-frontal region to white or yellow
does not affect threat response, while con-
fusion may result when the legs are painted
white; this complex would seem however to
be more concerned with shape than with pat-
tern and color in the sense used in this sec-
tion; the importance of type of motion is
once more emphasized; the response to the
mount on black seems to be connected with
visibility.
Phiale flammea. The white scales were
scraped from the palps and clypeus of two
males, so that they now had in frontal view
the wholly black appearance of females.
Neither male was chloroformed thereafter
and each was of A-tone. When placed to-
gether, both gave complete threat displays
starting at three inches, followed by brief
sparring, in six tests on two successive days.
The displays were never mixed with court-
ship, except that 'on one day the larger, less
aggressive male tended to start with a gen-
eralized display (high stance, first legs
high). However, when another male was
chloroformed, palps and clypeus blacked and
the whole mounted, the following occurred:
“A normal male took no notice when the
mount was not moved. When the mount was
given a single twitch, the testee hesitated,
adopted a high, generalized stance, then, at
one inch (unusually close) he changed into
a typical crawling courtship; he continued
to persistent attempts to mate, even includ-
ing complete twisting of the mount’s abdo-
men.” ( Field lab. note ) . On the second round,
when the mount was persistently jerked,
prompt threat response was drawn ; courtship
resulted once more when the movement was
restricted to a twitch. Here again, as in
chalcea, the significant difference was in the
amount and type of motion; and as in the
yellow of xanthopa, the median white frontal
markings were, under appropriate condi-
tions, badges of masculinity, (cf. also p. 194
for threat display toward females with whit-
ened palps, and the role played by distance
chemoperception, p. 183) .
When the first legs of a normal male were
painted white, incomplete threat was drawn
from three individuals.
When the flame-red abdomen of a dried
male was painted completely black, including
the white terminal spots, prompt threat dis-
play was drawn from three individuals; the
same results followed when the white cara-
pace stripe and submarginal band were also
removed. However, when the abdomen was
whitened, there was no display from any of
the three in a total of ten tests; one of them
stalked and leapt upon it once, apparently
treating it as prey, but dropped it promptly
(perhaps because it was stiff) . When it was
re-blackened, all three males once more
threatened promptly. Intensity, rather than
hue, seems to be the important factor here.
The abdominal red of Phiale shows no evi-
dence of being an aposematic hue. Small
frogs and lizards of several species stalked
and ate Phiale without hesitation or apparent
distaste. Furthermore, a Phiale, in all three
tests made, showed a decided fear-flight re-
sponse when dropped near a young Anolis;
this occurred several seconds before the liz-
ard appeared aware of the spider.
The results of inter-male display experi-
ments may be summarized as follows :
First. The light-colored contrasting cly-
peal and palp markings of male Phiale and C.
xanthopa have definite releasing value for
threat display, subject to superior control by
airborne chemical stimuli and motion. White
clypeal and palp markings in the naturally
black-fronted C. chalcea have no such value.
Second. Reflected light appearing in the
(to human eyes) yellow region of the spec-
1949]
Crane: Salticid Spiders: Analysis of Display
199
trum has threat releasing value in xanthopa,
irrespective of intensity, when occurring in
the palp-clypeal region. Other spectral re-
gions in general release incomplete or inap-
propriate responses.
Third. Reflected light in the red region,
added to the black frontal view, does not af-
fect threat display in C. chalcea.
Fourth. When the scarlet abdomen of
Phiale is covered with black, threat responses
are not affected.
Fifth. White paint applied to legs or abdo-
men of male chalcea or Phiale usually draws
an incomplete response, or none.
Sixth. The obliteration of iridescence in
Corythalia legs by black or brown paint does
not affect display responses.
General Summary of Value of Color-Pat-
tem-Intensity in Epigamic Display.
First. Pattern, color and intensity are of
minimum importance in the stimulus config-
uration of courtship, at least in Menemerus,
Phiale and Corythalia : Male or female “or-
namentation” is not a primary or secondary
releaser for either female acceptance or male
courtship display, respectively.
Second. It is probable, however, that in
both sexes, certain spots and patterns, par-
ticularly when correlated with motion (as in
spotted palps) have definite directive and/or
excitatory value, or at least function as vis-
ual aids. An example of an unquestionable
directive stimulus is the pale sub-basal ab-
dominal band which, at least in Phiale and
Corythalia functions during advanced Stage
II as a copulation guide for twisting the
abdomen.
Third. Clypeal and palp markings con-
trasting strongly in brightness with the
black integument and present only in the
male have definite releasing value in inter-
male display at least in two species, Phiale
flammea and Corythalia xanthopa. In two
other Corythalia, however, one with minor
markings and one with none, facial ornamen-
tation is not a threat releaser.
Fourth. There is some evidence that at
least Corythalia and Phiale have poor sensi-
tivity in the red region.
Fifth. The application of white paint to
extremities in Corythalia and Phiale some-
times affects display reactions, perhaps be-
cause of apparent form alteration.
Sixth. The obliteration of leg iridescence
in Corythalia has no perceptible affect.
The significance of deviate responses will
be discussed under displacement behavior
(p. 202).
VI. INNATE RELEASING AND
DIRECTIVE MECHANISMS.
From data given in the preceding sections,
the innate mechanisms in courtship and
threat may now be viewed as integrated pat-
terns. In all display, adequate physical con-
ditions, including light, humidity and tem-
perature, are prerequisites, as described
earlier; they, as well as the general good
health and nourishment of the spiders are
essential and will not be referred to again
in the following discussion. Courtship and
threat displays as a whole will be taken up
in turn, from the points of view of both sexes,
followed by a more general discussion.
A. Courtship.
1. Mechanisms of courtship display
in males.
a. Releasers in A-tone males of
hopper groups.
i. The stimulus must be within
range of the AME.
ii. It must fulfil certain rough
size-shape-distance require,
ments.
iii. One of the following factors
tors must be present.
(a) . Airborne chemical sti-
muli.
(b) . Generalized motion
(for Stage I) ; lack of
motion is customary for
advanced Stage II.
iv. The following often contri-
bute to the configuration, but
are not essential as releasers :
(a) . Special frontal and/or
abdominal patterns, of-
ten displayed with spe-
special motions, such as
the vibration of pale
palp.
(b) . Chemotaxis.
(c) . Cessation of motion, in-
cluding that of palps,
often accompanied by
crouching; this behav-
ior has releasing value
for Stage II, although
it is not essential ; close
proximity is of first im-
portance, with or with-
out crouching.
b. Directors of A-tone males of
hopper groups.
Groups iii and iv above are
probably partly and primar-
ily, respectively, directive in
function. A light, sub-basal
abdominal band is sometimes
a specific director for copu-
lation.
c. Releasers and directors of A-
tone males in runner groups.
These differ from the hopper
group requirements in the
greater importance of both
airborne and contact chem-
ical stimuli, as well as of
proximity. The visual stim-
uli of i and ii remain essen-
tial, however, as releasers;
200
Zoologica: New York Zoological Society
[34: 17
the visual stimuli of iv are
probably of less importance
as directors.
d. Releasers and directors of
males of lower tones.
Stimuli from all groups, act-
ing in conjunction when ne-
cessary. Alternatively, a
few stimuli may release and
direct courtship display when
strongly reinforced.
2. Mechanisms for courtship re-
sponse in females.
a. The stimulus must be within
range of the AME.
b. It must fulfil cex'tain motion
requirements.
c. Airborne chemical stimuli
are probably involved at close
range in Stage I, at least in
the runner groups.
d. Tactile (and probably chem-
otactic) stimuli are essen-
tial in Stage II.
e. No obvious secondary sexual
charactex-, including special
structures, “decorations” and
colors, is essential to succes-
ful mating in the genera
studied.
3. Course of mutual display in court-
ship.
In general, a system of progres-
sive, mutual stimulation exists
between the sexes, on the order of
that demonstrated in stickle-
backs (Tinbergen, 1948 et ante).
However, in the salticids, espe-
cially in the more specialized
forms, the situation is far less
clear-cut. Omitting, for the sake
of clax*ity, special exceptions and
qualifications, the usual course of
normal field or laboratory display
in salticids is presented in Table
VII.
4. Comments on various aspects of
courtship.
Stage II is never entered upon without at
least a rudimentary Stage I, even when it is
only a resumption of a display briefly broken
by the female’s temporary retreat during
Stage II, or even in the middle of copulation.
The x'ole played from the female viewpoint
by the size and shape of the male, as well as
by his motion-configuration, has so far un-
fortunately px'oved impossible to test. The
most pex'tinent data beax-ing on this are sup-
plied by observations on the behavior of fe-
males watching males other than those of
their own species in display ; in each case the
size and shape were similar to those of their
own males, and they always retx*eated before
allowing Stage II to commence; this subject
will be further discussed under species bar-
riers.
The female’s own occasional reciprocal dis-
play during Stage I and early Stage II is not
a vital factor in self-stimulation, at least in
Phiale and Corythalia, since it is not of reg-
ular occurrence; it is practically absent in
xanthopa, and only fairly common in the
other two Corythalia species; much of the
apparent display in Phiale females px-obably
should be interpreted rather as distance
chemoperception (“sniffing”) motions with
the front legs and palps.
B. Threat Display. The mechanism of
threat display differs fx’om that of courtship
in the absence of positive x’eleasing or di-
x'ective value of any chemical stimuli and of
the greater importance of special colors or
“decorations.” These badges combine with
non-female behavior — i.e., height and width
accentuation, plus increasing instead of de-
cx'easing activity — to release threat. The
various signals, in order of importance, are
general motion, motion configuration, a
fox-m-size element and, finally, any special
male pattex'n-and-color badge. In nature, all
normally act together as a configuration.
In runner genera, inter-male displays
px-actically never occur; when they do, they
are indistinguishable from courtship and
bx-eak off before actual contact, probably be-
cause of the absence of female chemical re-
leasers and/or directors. In intexunediate
genera, courtship and threat are usually
identical until the spiders are practically
touching. As in the runnex*s, inter-male dis-
plays in these gi'oups probably ax*e the re-
sult of a male treating another male as a
female. When the requisite close-range
signals — visual, chemical or both — are not
fox*thcoming, displacement behavior then
occux-s as special fighting or sparring. In
hopper genera, cases of mistaken identity
appear to occur only rarely, since threat and
courtship are usually distinct throughout
display.
C. Comparison and Comment on In-
nate Releasing Mechanisms in Courtship
and Threat. Before proceeding to a discus-
sion of the functions and evolution of display,
it may be helpful briefly to compare the prin-
cipal aspects of the innate x*eleasing mechan-
isms of display.
The perceptual sign stimuli for releasing
display are ovenvhelmingly visual in both
courtship and threat, involving motion, form
and size. Neither contact nor airborne
chemical stimuli alone will release display of
any form in the genei’a studied, no matter
how great the reinfoi’cement, nor how strong
the intex'nal drive. However, both forms of
chemical stimuli play an important differen-
tial role in display, their importance vax*y-
ing phylogenetically : in general, the pres-
ence of an airbonxe and/or contact chemical
signal is a positive differentiator for court-
ship, while a pattera-and-color signal in
some species plays a corresponding positive
role in threat. Absence of either one usual-
ly acts as a positive signal for the alternative
1949]
Crane: Salticid Spiders: Analysis of Display
201
type of display, other factors being equal.
Airborne chemoperception takes precedence
over a color-pattern badge in test situations
where both are present, since males court,
not threaten, young adult females painted
like males. Motion-configuration also super-
sedes color-and-pattern, since males change
courtship to threat when a previously quies-
cent, female-painted male starts threat dis-
play. Motion-configuration itself is a strong
differentiator of the two types of display,
particularly in the earliest and late stages;
in the middle portions of courtship (i.e., late
Stage I), the tendency of some females to
reciprocal display never confuses the male.
As in other groups of animals, deficiency
of one signal can often be compensated for
by increase in another in the same stimulus
situation. An example may be given of an
unnaturally painted spider, which stimulates
no response when running freely in the field.
When it is chloroformed or killed, and
mounted, so that it can be persistently mani-
pulated with appropriate jerks close to the
tested spider, it will frequently arouse a re-
sponse through reinforcement. The same
situation occurs when an old female is placed
with a male in a small, closed container;
with either contact or airborne chemical
stimuli reinforced, the male frequently dis-
plays, although he would not be sufficiently
stimulated to do so in the field.
Also as in other groups, maximal stimu-
lus is needed to arouse minimal response in
spiders of weak internal drive, while spiders
of strong internal drive give a normal re-
sponse to a minimal stimulus, often respond-
ing to fraction of the usual configuration.
As previously noted, summation readily oc-
curs in this family.
Highly stimulated spiders in unnatural
test situations, and spiders in states of mod-
erate and low internal drive, frequently con-
fuse threat and courtship reactions and be-
have inappropriately in other respects. These
actions, which may often be classed as typi-
cal displacement behavior, give provocative
clues to evolution ; they will be considered
later.
Wholly aside from test situations, how-
ever, the reaction of spiders in the field is
to complex, closely integrated, mutually de-
pendent configurational stimuli. Although
there are no simple, lock-and-key arrange-
ments, the more the stimulus situation de-
parts from the normal, the less likely is it
to release display.
In an over-all view, salticid display pre-
sents a complex combination of rigid and
fluid aspects. It is true that display patterns
are wholly fixed and instinctive. Learning
plays no discernible part in this field of ac-
tivity, since males reared in solitude per-
form either courtship or threat to perfection
on their first attempt. Again, there is no evi-
dence that imprinting ever takes place.
Finally, when two males of different species
display, each maintains rigidly his own dis-
TABLE VII.
Generalized Course of
Male.
Becomes aware of $; starts display, Stage I.
{Minimal releaser : several sight factors; air-
borne chemical stimuli also usually involved).
Approaches, in zigzags, or follows (if female
retreats), continuing or resuming display.
{Minimal releaser and director : above sight
factors, plus type of female motion or lack of it) .
Special $ signs, such as vibrating palps and
light abdominal spots probably have directive
value.
Speeds up display tempo. {Releasers and di-
rectors : reduced motion of female, plus chemical
stimuli. Self-stimulation is doubtless also a
factor) .
Enters Stage II.
{Releasers: primarily, proximity of female;
also involved, usually, her lack of motion, low
position, and, doubtless, reinforced chemical
stimuli) . Copulation follows unless female with-
draws. ( Director : sometimes a pale abdominal
cross-bar) .
Display in Salticids.
Female.
Retreats, or watches $, usually in braced, high
position, often vibrating palps. Rarely attacks.
( Minimal releaser and director: several sight
factors) .
Becomes completely attentive; sometimes gives
weak reciprocal display. {Minimal releaser:
summative effect of display motions).
Ceases motion and, usually, crouches low, legs
drawn in.
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Zoological New York Zoological Society
[34: 17
play pattern; no temporary modifications
were ever seen to occur; by contrast, imita-
tive behavior is frequent in vertebrates, as
in the cormorants which altered their flight
pattern to match that of their pelican com-
panions (Beebe, 1938, p. 106).
Nevertheless, in working with living sal-
ticids, both their own individuality and the
dynamic, unfinished, untidiness of display
mechanisms are constantly apparent. A given
spider’s behavior is scarcely more predictable
than that of a single electron. It depends, at
any given instant, on great numbers of fac-
tors, external and internal, all in various
stages of evolution, all related, and all them-
selves in a state of constant change. Wasted
energy, abortive displays and lost opportun-
ities are the rule. Altogether, the lumber-
ing, complex, display mechanisms do not ap-
pear efficient. The most that can be said of
them is that they work adequately enough,
often enough, to ensure the perpetuation of
the species. In the end, however, all the
billions of salticids, hopping at this moment
about the globe, are lively proof of their
success.
VII. BEHAVIOR RELATED TO DISPLAY.
The basic similarity of behavior trends
in all higher organisms — whether inverte-
brate or vertebrate — becomes increasingly
clear. Not only do animals agree in the ob-
vious, basic activities essential to all life, but
they prove also to be surprisingly similar,
among the end forms, in the more complex as-
pects of behavior. It has long been recog-
nized that when vision becomes the domin-
ant sense, elaborate, visually dominated
courtships often evolve; the principle ap-
plies in certain cephalopods, crabs, spiders
and insects as well as among fish, lizards and
birds. The importance and frequent dis-
tinctness of threat display in salticids has
been emphasized throughout this paper, and
is in accord with results of recent behavior
studies of birds and other vertebrates. Sim-
ilarly, it is increasingly evident that dom-
inance hierarchies, social and territorial be-
havior and displacement activities very of-
ten occur in higher invertebrates, though
usually in primitive form. Although these
latter aspects were studied only incidentally
in the Rancho Grande salticids, the follow-
ing observations appear to be worth record-
ing.
A. Displacement or Substitute Behav-
ior. Experimental work sometimes drew in-
appropriate responses which correspond
closely with typical displacement behavior in
birds and other groups. When a stimulus con-
figuration, while partly effective, was yet too
exaggerated or incomplete to draw an ap-
propriate display, the spider either retreated,
attacked or by-passed the stimulus, ex-
changed threat for courtship and courtship
for threat or regressed to a more primitive
level of display behavior. It is interesting,
however, that at no time were the substitute
activities wholly outside the sexual field.
That is, never did the spider stalk an insect,
pick up a discarded fly, spin a retreat, or even
resort to grooming — that substitute action
so common throughout much of the verte-
brate world.
This restriction of displacement behavior
appears to be an example of the rigidity typ-
ical of higher invertebrates. The compart-
ments of life are kept distinct even in the
midst of “mental” confusion, shock or frus-
tration. With such restrictions, a girl would
not reach for her lipstick when frightened
by a bomb, nor a startled bird fidget with the
makings of an off-season nest. To lapse into
anthropomorphism, a salticid, when his prey
escapes, never picks a fight with another
male, nor does a rejected suitor gorge on
extra flies.
B. Dominance. In salticids, little hint
has yet been found of long-term dominance
relationships. Day to day variation, as
shown in energetic courtship and threat or
apathy and retreat, is on a rhythmic, physio-
logical basis. It is true that some males
never develop as high tone levels as others,
the peaks of their rhythm curves being
lower; in this sense dominance may be said
to occur. I have never found, however, that
rank is altered by a series of successes or
failures in threat display or in actual com-
bat. In primitive groups, where inter-male
combat does not ordinarily exist, nothing ap-
proaching dominance relations was seen
(apart from the usual physiological fluctua-
tions, which determined the degree of court-
ship activity at a given time) . In groups of
young and old spiders placed together in a
cage, the small ones invariably showed a
healthy wariness of the large individuals ; in
view of the carnivorous character of the
spiders, however, this behavior certainly
needs no more explanation than the obvious
one of self-defense. (See also p. 203).
C. Sociality. C. xanthopa was one spe-
cies in which traces of social behavior oc-
curred. Along the leaf-strewn Water Trail,
the richest habitat for this species, three to
ten individuals were often found fairly close
together. These groups included individuals
of various ages and both sexes. They occur-
red in an area of about a square yard or two,
divided by 10 to 20 feet from the next group.
There was absolutely no discernible differ-
ence in the ecological characteristics of the
populous and barren stretches, and indeed
they shifted back and forth in quite irregular
fashion over a period of weeks. Intra-spe-
cific relations within a group appeared pure-
ly casual. Abortive courtships and threat
displays were brief and frequent. There was
a high degree of mutual toleration, but, ex-
cept for sexual situations, no inter-individual
associations. It is noteworthy that quite
small individuals, measuring less than half
the size of the adults, were included in these
groups, and apparently not stalked as food.
Often their attention was attracted by the
1949]
Crane: Salticid Spielers: Analysis of Display
203
displays of nearby adults, and at times they
appeared to watch attentively these encount-
ers of their elders, although they took no
part whatever. The same behavior was
noted in non-displaying adult males, and in
fat females, obviously soon to lay eggs. On
other occasions, these non-displaying mem-
bers of the group simply continued their pre-
vious occupations, paying no attention to the
others; they either maintained hour-long
lookouts from the top of a projecting twig,
stalked prey with indifferent success, hop-
ped slowly along a meandering course, or
simply moved occasionally out of the shifting
sunlight.
An example of more advanced sociality
was found in Semorina megachelyne. A large
silk shelter was taken containing a group of
14 individuals of various instars, including
several adult males and females. The young
ranged from the first through the preadult
stadia, the youngest being sheltered in a
subcocoon. The morphologically primitive
position of Semorina, compared with the ad-
vanced niche of Corythalia, indicates that
sociality is not dependent in this group on
phylogenetic specialization. Comparable
dissociation of these two factors is found in
other groups, including non-salticid spiders,
lepidopteran caterpillars and birds.
It is probably significant that in neither
of these examples of incipient sociality,
Semorina and Corythalia, is inter-male fight-
ing developed ; in the one case the males fol-
low the primitive pattern of mutual disre-
gard ; in the other, combat has been subli-
mated into threat. Never have I seen any
signs of group formation in the more pugna-
cious genera.
D. Territory. A territorial concept al-
most certainly exists in salticids, but work
has scarcely been started on this angle. In
captivity, all salticids generally returned to
the same shelter, if they made one, night af-
ter night, even occupying it through several
successive molts, so long as it was undis-
turbed. It seems probable that such a prac-
tice occurs also in the field. It is certainly a
likely beginning for a concept of territory, or
at least of home range. When siblings of
Phiale, Corythalia and Eustiromastix young
were reared in groups of three or four
through the early molts, they not only got on
peacefully at all times, but each always — on
the many occasions when they were observed
with this point in mind — returned to his own
shelter at night and during eedysis. One
group of three Phiale dybozvskii was reared
to adulthood in this fashion. Individual va-
riations in size and pattern made the identi-
fication of individuals easy after the second
instar.
Cannibalism is exceedingly rare in this
family, and when it occurs it is only under
extreme provocation of hunger or gross size
disparity. In the few cases where females
killed the males, they were eaten only twice,
both in Eustiromastix. After inter-male
battles, the losers were never eaten, even
when they had been disabled or killed.
On the other hand, there is no evidence yet
that salticids actually defend a territory,
even in the case of adult males. I am quite
sure that the young at least do not have one,
although the concept of home range seems ap-
plicable, at least in C. xanthopa. This term,
as defined by Burt (1943), signifies that
wandering of individuals is limited, although
the area is shared at least tolerantly with
others of the same species.
It is notable that although in the orb-
weavers the adult males wander, in Cory-
thalia xanthopa — the only species where field
marking has been initiated — the females did
the roaming. One female, recaptured after
34 days, was taken 186 feet from her original
location, while several marked males were
seen repeatedly during a single month within
two feet of their original positions. A similar
situation exists in fiddler crabs (Crane, 1941,
p. 160) and, of course in vertebrates (e.g.,
Evans, 1938; Lack, 1943).
When a strong adult male xanthopa in A-
tone was dropped close to a wild male who
had hitherto been undisturbed, the usual
threat display took place, but I saw no sign
whatever that the strange male was usually
driven off by the previous “tenant,” or
showed much perturbation. The response
was tested more than ten times. This result
is in strong contrast to the behavior of an
orb-weaver dropped into a strange web. The
retreat of a protagonist seemed to depend
only on his physiological condition, rather
than on any general reduced pugnacity or
sense of security when out of his own range.
The other intermediate and hopper group
salticids were in general strongly individual-
istic ; two or more adult males were rarelv
shaken from the same herb or bush, and it
may be that in these cases definite territori-
ies are maintained and defended. On the
other hand, possession or invasion of a terri-
torv is certainly not a prerequisite to display,
judging by the prompt reactions of A-tone
males dropped simultaneously on a strange
table-top. Altogether, development of ter-
ritoriality appears to be a very primitive
level.
VIII. FUNCTIONS OF DISPLAY.
A. Courtship.
The theories concerning the functions of
courtship display in spiders have already
been reviewed (p. 170) . The conclusions re-
sulting from the Rancho Grande studies are
as follows: First, courtship display is un-
questionably a necessary preliminary to mat-
ing, and not merely an outlet of excess energy
for males in breeding condition, as suggested
by Wallace and Berland. Second, sexual se-
lection in the original sense used by Darwin
and the Peckhams is not operative. Third,
as Savory points out, the concept of recogni-
tion as distinct from, and preceding stimu-
lation, does not appear to be necessary.
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[34: 17
Indeed, any concept of preliminary recog-
nition seems to be highly questionable, even
when recognition is regarded merely as a
kind of realization by the female that
a potential meal is not at hand. Rather,
it seems probable that it is through her
sexual stimulation that her feeding im-
pulses are inhibited. In this view, the two
processes are merely different results of
the same psycho-physical sequence, devel-
oping simultaneously and governed by the
same stimulus configuration. To distinguish
between recognition and stimulation in sal-
ticids seems as difficult as to differentiate
temporally between human fear and loss of
appetite when a charging bull interrupts a
picnic: in each case two emotions, or at least
sensations, are involved, one positive, one
negative, and bearing an inverse ratio to
each other.
As modern observers agree (e.g., Savory,
1928), the stimulation of the female is prob-
ably physically as well as psychologically ne-
cessary, since alterations appear to be made
in the epigynum itself in order to permit the
insertion of the palps.
The following summary may now be given
of the functions of salticid courtship, as they
appear from the Rancho Grande studies. It
is, in essence, a selection and elaboration of
certain earlier views, especially those of Sav-
ory and Bristowe. Most of it applies to ani-
mal courtship in general. The term “court-
ship,” as used throughout this paper, in-
cludes the responses of the female as well as
those of the male, and the production as well
as the reception of sign stimuli.
The primary functions of courtship in sal-
ticids, then, may be expressed as follows:
Courtship serves to bring to mutual atten-
tion and proximity two individuals of the
same species, opposite sex and requisite
physiological condition; simultaneously it in-
hibits their usual predator and self-protec-
tive behavior while stimulating each sex so
that copulation may take place. These are
the primary functions, of obvious and imme-
diate biological use in the life of the individ-
ual.
Secondary functions, which may be consid-
ered by-products of the above, are of impor-
tance in the life of the species. First, sexual
selection operates in the sense that males of
chronically mediocre drive — among which
are doubtless individuals of genetic weak-
ness— do not use sufficient energy and per-
sistence to win acceptance by females. Sec-
ond, courtship display acts as an effective iso-
lating mechanism between similar but well-
evolved species: Since crosses would, judg-
ing by analogy in other animals, often prove
sterile or unfit, the progressively exclusive
action of individual courtships must prevent
considerable germ cell wastage. The effec-
tiveness of display as an isolating mechan-
ism will be discussed in the section on evolu-
tionary aspects.
B. Threat.
Earlier views on the cause and function of
fighting and/or threat display among salticid
males were summarized on p. 170. To reca-
pitulate briefly the two extreme hypotheses,
the Peckhams accepted the unmodified Dar-
winian premise that inter-male fighting was
brought on by rivalry; as a result, the fe-
males mated with the strongest or bravest or
boldest, either through male conquest or fe-
male choice. Bristowe, on the other hand,
holds that inter-male display occurs when
males temporarily mistake each other for fe-
males; actual fighting sometimes results
through frustration.
Neither of these views explains satisfac-
torily the condition in many salticids, includ-
ing Corythalia. Against the Darwinian hy-
pothesis stand the facts that the winner of
a combat does not necessarily get the adja-
cent female, females certainly do not neces-
sarily “choose” the winners, the fights are
usually no more than slight psychological
sparring matches, at which females may or
may not be present, and the winners — that is.
the more aggressive — are certainly not ne-
cessarily the largest or brightest of the
males. I have seen a six or seven-legged
male, of small size, prevail over larger in-
dividuals on successive days; almost certain-
ly the epigamic physiological rhythm was in-
volved; but some individuals appear regu-
larly to be more dominant (in their A-tone
periods) than others of the same threshold.
(See also p. 202). In the species Bristowe
observed in England, courtship and threat
display were identical. For such as these, his
hypotheses of mistaken identity is completely
satisfactory. According to the hypothesis
of phylogeny presented in the present paper,
these represent an intermediate phase of de-
velopment of dependence on sight practically
superseding dependence on chemical stimuli.
In genera taken to represent more primitive
forms, such as Menemerus, Ashtabula and
Semorina, all far more dependent on chem-
ical stimuli, threat display is apparently non-
existent and mirror display never occurs (ex-
cept in self-stimulation in Menemerus, p.
182). Finally, in the most visually depend-
ent genera, the hopper groups, of which the
best ultimate examples are in Corythalia,
threat and courtship are completely divorced.
In C. xanthopa such an extreme is reached
that different appendages are used in the
two kinds of display, and true fighting has
never been known to occur except under ex-
tremes of crowded, hot laboratory conditions.
It seems that here there is a sublimation of
aggressive impulses, as in many birds, and
that a type of mutual stimulation takes place
which is useful in maintaining or increasing
sexual tone.
Just as in courtship display, . it must be
kept always in mind that male spiders are po-
tentially dangerous to one another, both be-
cause of their carnivorous habits and their
poison glands; this is rarely the case in other
1949]
Crane: Salticid Spiders: Analysis of Display
205
groups of animals. Where mistaken iden-
tity in salticids results in actual fighting,
with consequent frequent casualties, the fol-
lowing conclusions concerning its functions
appear to be valid. All of them are of the
secondary type, of importance to the species,
rather than to the individual.
1. The elimination through disability or
death of weaker males, thus strengthening
the strain through natural selection.
2. The prevention of weaker males mat-
ing as often as do stronger individuals.
3. The encouragement of excess males to
go elsewhere in search of mateless females,
and so ensuring the more even distribution
of the sexes.
These three points conform to Darwin’s
general hypothesis, excepting only that fe-
males do not deliberately choose superior
males. Although it is true, as Bristowe and
others maintain, that actual fighting, to the
point of inflicting injury or death, is rare
among spidei'S, it most certainly does occur,
under natural conditions at that, among some
salticids at Rancho Grande. For example, an
apparently new genus near Capidava (not
yet described] time and again fought to the
death in large display boxes and on open
table tops ; twice they were seen fighting be-
side trails in the forest; in one of the latter
cases a male was killed; in the other both
were injured.
In the majority of salticids, and especial-
ly in Corythalia, where threat display is
highly ritualized and actual fighting occurs
only rarely and atypicallv, the physical elim-
ination of males does not take place. In these
cases, the threat display undoubtedly repre-
sents an advance over the more wasteful
practice of actual fighting.
In these ritualized displays, the additional
function of maintaining emotional tension
is probably of importance to the species as a
whole, although it is not necessary to indi-
viduals. Perhaps males having frequent op-
portunitv for epigamic display maintain
A-tone for longer periods. Armstrong. 1 947.
discusses tone maintenance in birds and
gives excellent examples. No experiments
whatever appear to have been done on this
question in spiders, but its importance is be-
coming increasinglv recognized in other
groups. Exneriments at Rancho Grande
proved onlv that threat disnlav is not a ore-
requisite to successful mating, either from
the male or female pomt of view. Phiale
and two snecies of Corvthah'a all showed con-
clusively that virgin males disolaved oromnt-
Iv and completelv to virgin females and were
accepted by them, without any of the males
ever having seen another male, much less
practised or observed threat disolav. Each
of the individuals tested was reared through
at least three previous instars, and permit-
ted to molt to the adult, in complete solitude.
Precautions were taken, as in all displav ex-
periments. to eliminate the danger of chem-
ical stimuli remaining from previous spiders.
Conversely, virgin male Phiale, C. chalcea
and C. xanthopa performed threat display
perfectly to virgin males of their own species
without ever having laid eyes on or received
chemical stimuli from a female or another
male.
Nothing is known of territoriality in any
of the species except in xanthopa: here it ap-
pers to exist, but in rudimentary form (p.
203). This amount, however, might account
for the development and maintenance in
this species of a completely separate threat
display. But it seems certain, as said above,
that at least equally important to the species
is its function of mutual stimulus and of
keeping males to some pitch of excitement,
perhaps in a state of long-sustained A-tone
for dealing with wandering and scattered fe-
males. There is no question but that males
tend to display to each other more promptly
in the presence of a female: in photograph-
ing threat displays, one female was always
placed with the two male subjects, since fe-
males were repeatedly found to have this
decided catalytic effect. Often the males
spent more time displaying to each other
than to the female.
Another function of threat display doubt-
less is its usefulness as a “safety valve,” its
displacement behavior aspect, for spiders
already keyed to display pitch (p. 202) .
No true group displays, comparable to
those reported by the Peckhams (1889, p.
40) for several dendryphantinids, were seen
at Rancho Grande. They undoubtedly should
be regarded as rudimentary social affairs,
again with the double function of inhibiting
hunger and stimulating sex.
IX. EVOLUTIONARY ASPECTS OF DISPLAY.
The purpose of this section is to interpret
salticid display from an evolutionary point of
view. Although the dangers of top-heavy
generalizations are fully realized, it seems
advisable to organize the limited data avail-
bale. As Menninger puts it, “Classifications
must never be taken too seriously — but the
fear to use them has prevented much more
thinking.” (1945, p. 34] .
A. Hypothetical Philogeny.
While many display similarities obvious-
ly coincide with natural affinities, others re-
flect only an extensive parallelism among the
subfamilies. The salticids show a “bush
type” of evolution, with similar basic trends
in each branch.
The parallelism is well shown in the loco-
motive, sensory and displav differences that
have been discussed from time to time in this
and preceding papers (Parts II and III). At
Rancho Grande, Menem erus, Semorina and
Ashtabula are examples of the apparently
primitive “runner” group; Sassaeus and
Phiale are intermediate forms; and Cory-
thalia and Mago are advanced “hoppers.”
Each of these groups contains representa-
tives of two or more subfamilies, and each
^Ub Zooloyica: New York
subfamily usually includes genera in two or
more groups. Their various characteristics
will now be assembled.
The runners” never jump or hop during
horizontal progress except to cross a gap or
leap on prey. According to experiments, they
depend more on chemotaxis than do the other
groups, court less readily in its absence,
rarely or never respond to a mirror image
and are rather strongly affected by the loss
of their palps and first legs. The first legs are
often held in front of them, not helping in
locomotion, but barely clearing the ground,
while the palps may pat the ground lightly.
When in a strange place, they run to and
fro, palpating ceaselessly and appearing end-
lessly “restless” and “nervous.” Although
they stalk and leap on prey like the hoppers,
they nevertheless tackle large prey in pref-
erence to small. Menemerus chooses moths
and the larger Diptera instead of Droso-
phila; in captivity, when no choice is given,
they miss fruit flies repeatedly and eventual-
ly become thin. Large, hopper-group Cory-
thalia of similar size, in contrast, can live out
their lives on a fruit fly diet, catching them
with ease. Morphologically, the most obvi-
ous characteristics of the runners are low
carapaces and few, weak leg spines.
The intermediate group seems to be in the
middle of changing from moderate chemo-
tactic to nearly complete visual dependence
by way of distance chemoperception. They
are long-sighted, hop when hard-pressed,
pursue their mates via efficient short-cuts
and are relatively little affected in display
when deprived of pabs and first legs. Yet
they appear to gain chemical sense impres-
sions from a distance largely through these
members which, from the second instar, of-
ten wave in the air during normal explora-
tion. Phiale, when near mates or food, be-
fore display or stalking begins, is especially
prone to wave the first legs up and down. It
seems probable that they are “sniffing” the
air. Berland’s accounts of Philaeus chrysops
and others (1914, 1923, 1927), waving their
legs when completely alone in clean boxes,
sound as though these were intermediate-
group spiders. Philaeus, incidentally, is a
close relative of Phiale. Bonnet’s (1933)
Philaeus, studied in another part of France,
never waved the legs except in true display;
why these observations should differ so rad-
ically remains, for the present, a mystery.
Berland’s Saitis barbipes waved the third
legs at random, as well as in display.
Among salticids generally, it may be that
the female is “sniffing” the male when during
display she often vibrates the palps and first
legs; if so, the sense of distance chemoner-
ception may evolve through the female. Cer-
tainly in the runners, the female’s palps are
usually particularly active, whether or not
they quite touch the ground in true chemo-
taxis. It does not seem advisable to consider
this activity solely as a symptom of nervous
tension.
Zoological Society [34 : 17
The “hoppers” are the visually dominated
salticids ; chemotaxis means little or nothing
to them under natural conditions; distance
chemoperception dperates as a secondary re-
leaser for courtship display. Their sight is
magnificently developed, and their courtship
sign stimuli overwhelmingly visual. These
are the “poised” spiders; they sit quietly
when dropped on the table, look about them,
then hop away, at leisure and with frequent
pauses, interspersed with a measured walk.
All eight legs remain firmly on the ground .
except during a hop or display ; there is other-
wise no raising at all of the first legs, or
carrying of them clear of the ground. Cara-
paces are high and leg spines strong and
numerous.
An inspection of Table II and Text-figs.
2 and 6 will show that among runners court-
ship is simple and threat practically non-
existent. In intermediate groups, actual
fighting is frequent, but its early stages are
little or not at all differentiated from court-
ship. In the hoppers, true fighting is rare and
special threat display the rule.
The divisions appear to be based primarly
on the different degrees of visual dominance.
The eyes in each successive group seem to
take over more and more from the chemical
senses. And it is the dependence on the latter
which appears, both functionally and mor-
phologically, the closer to the pre-salticid
stock. But, as in various studies of animal
and human societies, the forms in midevolu-
tion seem most subject to combativeness.
They appear to be in the middle of changing
from one way of life to another, mistakes are
made, frustration results and combat ensues.
One vitally interesting clue to {he evolu-
tion of salticid display is given in the be-
havior of the highly specialized hopper, Cory-
thalia xanthopa. Senile, over-stimulated or
low-tone spiders often use primitive chemo-
taxis during courtship to {he frequent and
inappropriate exclusion of other sign stimuli
(p. 178) . These and other hoppers in a simi-
lar condition tend to leap at threat stimuli
rather than perform their characteristic dis-
plays. This behavior seems unquestionably
atavistic, regressive in a phylogenetic sense.
Similar examples among displacement behav-
ior (p. 202) consist in the replacement of
threat with physical attack in experimental
situations.
B. Origins op Display Motions.
i The biological principle of least effort, as
presented by Zipf (1949), proves to be of
considerable help in this attempt to under-
stand the development of salticid display.
The subprinciples of permutation (i.e., com-
bination) and multiple function seem espe-
cially applicable. Armstrong in particular
has implied their operation throughout his
discussion of bird display (1947, e.g. p. 61).
From this point of view, display evolved
through the use, with modification, of ac-
tions, senses and structures already serving
1949]
Crane: Salticid Spiders: Analysis of Display
207
less specialized purposes in the economy of
the organism. Only rarely would a display
action evolve especially to fill an epigamic
need.
In direct agreement with this general con-
cept is Bristowe’s view of the origin of sal-
ticid display. It appears to him that the
rituals grew from the groping and fending-
off motions of the primitive stock. Certainly,
it seems^ far more likely that display grew
'from gestures like these, which had a dif-
ferent original function, than that an en-
tire new series of motions arose for display
alone.
The acceleration and exaggeration of sim-
ple motions, so typical of display, are clearly
related to the excitement, to the nervous
agitation, connected with breeding activity.
The difference between this view and those
of Wallace and Berland is that, to them,
display appeared to have no function ; it was
simply an expression of high vitality or ex-
citement.
The “groping” motions certainly in-
cluded chemotaxic behavior, such as is used
by many runners in casual exploration as
well as in responding to the trails of females.
The similar behavior of very young runners,
just out of the cocoon, contributes largely
to this conclusion. Again, high-in-air leg
waving, found in numerous displays through-
out the family, seems to have an undoubted
origin in distance chemoperception of air-
borne stimuli.
Plausible origins of a number of other
common display gestures may be proposed.
Thus, the vibration of palps originated in
1 the “sniffing” motions so characteristic of
chemoperception, including both contact and
distance types. The vibration is perhaps also
useful in dissipating nervous tension dur-
ing courting excitement.
Twitching or lowering the abdomen is an-
other frequent display gesture which may
be due to permutation. Probably it was con-
cerned originally only in attaching silk to
the substratum, in the usual salticid fashion »
of ensuring a quick escape. An almost uni-
versal part of display is the habit of ap-
proaching a female in zigzags, which ob-
viously gives added time for necessary
stimulation. When zigzagging was first com-
bined with lowered spinnerets, it is easy to
see how a side-swinging abdomen could have
developed-: in the more primitive, long-ab-
domened Dendryphantinae, it seems a nat--
ural result analogous to that of a train
rounding a bend.
The elevation of the abdomen is charac-
teristic of all the ant-mimicking or pseudo-
scorpion-like salticids in which display is
known. It was also typical of normal prog-
ress in Semorina and related genera as early
as the second instar. Its ultimate origin is
not yet clear. Possibly the gesture started
as the end of the dropping motion, as a spi-
der hits the ground at the end of a silk cable.
Possibly, actual scorpion or pseudo-scorpion
mimicry was involved among larger for-
bears; these hypothetical spiders may well
have been large enough to win a mimic’s
protection from frogs and lizards (cf. Part
III, p. 37).
Again, it is possible that abdominal lift-
ing, quiverings, and even the swift rockings
of some displays are concerned in the emis-
sion and wafting of a chemical display sig-
nal. I have as yet no corroborative experi-
mental data; work with sealed genital
grooves and epigyna — a promising approach
— has not been adequate. The possibility is
mentioned here as a potential X-factor, an
“etc.” that may prove important!
In Lyssomanes the retinal motion within
the light-green, antero-median eyes seems to
play a definite role in courtship. The result-
ant color shifts increase in tempo with excite-
ment. They seem clearly to be a ritualization
of incidental effects resulting from the me-
chanics of vision. Although the same motions
occur throughout this and other large-eyed
families, they are rarely as noticeable because
of the dark pigment which usually surrounds
the retina. The morphological basis has been
discussed by Homann (1928, p. 235).
All of the display motions mentioned up
to this point can have evolved through per-
mutation of activities normally occurring
in the simplest epigamic situations as well
as in other fields — dropping from overhead,
fixing a silk drag-line for quick escape, grop-
ing forward, and so on. In this way these
postulated display origins parallel the epi-
gamic character of displacement reactions
(p. 202). Several possible exceptions will
now be discussed.
The display crawling motion may have
originated from the hunting rather than the
sexual field of behavior. In this phase, the
male lowers the carapace and creeps directly
forward. Usually it occurs during late Stage
I or early Stage II of courtship. It crops up
sporadically, with variations, in almost every
subfamily observed. It may have grown from
a displaced stalking motion, definitely out-
side the sexual field. It appears more prob-
able, however, that there is, a sort of innate,
anticipatory imitation: the crouch is ex-
tremely similar to that usually assumed
by the female during the latter stages of
courtship. As in many birds, this position
often acts as an important releaser for the
latter part of the male’s display. Its previous
assumption by salticid males, in combination
with a typically masculine approach and leg
stretching, perhaps has value in empath-
ically inducing the female to assume a simi-
lar position. Whatever the origin, this mo-
tion is one of the most remarkable in all sal-
ticid display.
Two widespread groups of display gestures
do not seem to be based on economical per-
mutation, on made-over motions. They are
the lateral leg displays and the stretchings
up to maximum height. It seems, rather, that
these salticids have evolved independently
the widely successful animal custom of ap-
208
Zoologica: New York Zoological Society
[34: 17
pearing larger than life in a crisis. It is ex-
ceedingly interesting that they never use the
technique in other departments of living;
never do they display before prey or hungry
enemies; in salticids, the “big bluff” is re-
served solely for an epigamic crisis.
Posing is a frequent occurrence in displays
of the size-increasing type, particularly in
threat and especially among the plexippinids.
The motionless state, amounting to a kind
of catatonia, often persists for many seconds
after the exciting object has been removed.
Similar behavior occurs in many animals
under various conditions; it has been dis-
cussed at length by Armstrong in connection
with bird display (1947). Its origin in sal-
ticids remains obscure.
This effect of seeming larger — whether
in connection with courtship or threat — is of
special interest in comparing displays within
the advanced genus Corythalia. Courtship
and threat display are distinct in the three
species studied. However, in xanthopa, court-
ship is simple and primitive, exhibiting only
the first two legs, while the remaining pairs,
all specialized, are reserved for the highly
developed threat display. In chcdcea and ful-
gipedia, on the other hand, the first legs are
used only in Stage II of courtship, while
more posterior legs take part in Stage I of
both courtship and threat. It seems that
xanthopa represents the more primitive
form, with the size-increasing threat fan
occurring first. Then, by permutation with
modification, the original threat function
was extended, in the other species, to form
a more complicated courtship. In support of
this hypothesis, xanthopa appears decidedly
more generalized than the other two species
in several structural and developmental char-
acters; these include eye proportions, spin-
ulation, course of color development and den-
tition of tarsal claws.
The displays of two advanced Dendryphan-
tinae give similarly interesting clues to their
relative evolutionary status. One species,
Sassacus ocellatus, shows a vestige of the
side-swing typical of the subfamily; in the
other, S. flavicinctus , no swinging occurs.
It is the latter form which is morphologically
more advanced.
The subject of vestigial behavior charac-
ters is, of course, one of the most contro-
versial. Which traits are to be considered
vestigial and which rudimentary? One
point, however, appears increasingly clear.
Any successful animal species includes in
its organization a hodgepodge of behavioral
relics in addition to the usual morphological
trash. When a dog revolves before lying
down on a grassless rug, he is performing a
well-worn example of a functional antique.
Similarly, Sassacus, in a moment of face-to-
face courting, swings his stubby abdomen
sideways, even though it and its ocellus are
well hidden behind the sturdy legs; surely
this example belongs equally in a behavior-
istic museum.
C. Relation of Secondary Sexual
Characters to Display.
The question of vestigial characters leads
directly to the problems of ornamentation,
where morphological and behavioral rela-
tions are even more involved. Some “orna-
ments,” including the fringes and iridescence
on the displayed legs of Corythalia, are prob-
ably recently evolved structures: they are
highly variable in individuals; their adap-
tive value is so slight that it has not been
proved experimentally to exist at all; they
develop (as do most epigamic characters)
only in the final instar; and they are con-
fined altogether to males. Yet their place in
the perceptual sign situation seems unques-
tionable; on a mathematical basis these re-
cent refinements would doubtless prove to
have definite adaptive value, at least to the
extent of making their possessor more con-
spicuous during display.
Next to these come such signs as the yel-
low or white clypeal bands of C. xanthopa
and Phiale males, which have decided, con-
temporary releasing value for threat display.
Finally, in a confusing mixture, comes the
mass of “decorations,” including vestiges,
characters linked genetically to selected
structures, patterns aiding in camouflage,
and mere byproducts of metabolism. Only
rarely can their origins be satisfactorily
discovered or inferred. They include the sex-
chromosome-controlled tufts of dark-phase
male Maevia vittata (Painter, 1913), the
scarlet-red of Phiale abdomens and the cir-
cles of colored scales around most salticid
eyes.. These circles occur as early as the sec-
ond instar and are irrespective of sex. Pos-
sibly, among primitive salticids, they had
adaptive value, making the large eyes or
frontal regions more conspicuous in display;
now they have no demonstrable function, and
their early appearance, far back in ontogeny,
indicates a vestigial, if not a purely meta-
bolic, character.
A related instance is the occurrence of
white scales on the legs of intermediate in-
stars in Corythalia, followed by their obsoles-
cence on the front of legs in adult males ; they
may well represent an old secondary or minor
display character which has been superseded
by iridescence; a similar explanation may
apply to the reduction of clypeus white in
adult male chalcea and fulgipedia.
The variable distal abdominal ocelli of
Sassacus ocellatus, mentioned above, seem to
belong rather definitely among the vestigial
character group.
All of these relics, again in agreement with
the principle of least effort, persist unless
it takes less “effort” for the organisms to
drop them than to carry them along
The old question as to the significance of
elongate chelicerae recurs. Bristowe has
pointed out that they are only rarely used
in courtship or fighting, and also are un-
questionably less efficient than those of the
usual size (1929, p. 339) . It seems to me that
19491
Crave: Salticid Spiders: Analysis of Display
209
they are enlarged, not to aid fighting, but
much as first legs are specialized, primarily
for an increase in conspicuous area. It ap-
pears, too, that secondarily their very ineffi-
ciency may be adaptive in the way that a
threat display is adaptive : the danger of mu-
tual injury is reduced without reduction of
the presumable toning value of aggressive
behavior.
It is noteworthy that these enlarged cheli-
cerae, in the genera with which I am fa-
miliar, occur in the middle groups: in the
Dendryphantinae, and in Salticus (morpho-
logically an advanced marpissinid) , in which
aggression reaching the contact stages is
most developed. Elongate jaws occur also in
the highly aberrant lyssomaninids, but I
have never been able to induce any threat
display in them whatsoever. Another point
is that in some genera, for example Sassacus
and Ashtabula, the length of the chelicerae
is highly variable among individuals, exactly
as in certain beetles (e.g., Beebe, 1947) with-
out any reference to their general tone or
health. No use as a specialized, female-hold-
ing tool, as is found in Pachygnatha, has
ever been observed in a salticid.
In salticids, as seems increasingly clear
in other groups, function appears basic,
structure follows after. The male salticid
raises his forelegs or middle legs, or jiggles
his palps ; elongation, thickening, blackening,
whitening, polishing or fringing may or may
not follow. A spider does not, as the older
naturalists naively implied, show off his dec-
orations; rather, his decorations evolved in
a conspicuous position. Every experiment
made in this study showed that motion —
function — was of more importance than mor-
phological detail. This view does rfot, of
course, rule out the obviously great7 recip-
rocal influence of behavior and morphology
during selection. Nevertheless, in any gen-
eral trend, it must be the function which
usually changes first, while vestigial tufts
and spots, seemingly placed to accent an ob-
solete display, persist indefinitely.
D. Sexual Dimorphism ani\ Display.
The correspondence in forms of sexual
dimorphism in both salticids and birds was
discussed at length by the Peckhams (1889,
1890). Here their general points will only
be summarized and remarks made concern-
ing the relationship of dimorphism to fe-
male behavior. As in birds, dimorphism is
of three principal types. First, and appar-
ently most primitive, neither sex is strikingly
marked, or greatly differentiated in second-
ary characteristics ; display is primitive and
the female remains completely passive (ex-
amples: Semorina, Mago) ; in the second
type, sexual dimorphism is strong, the fe-
male remaining dull and adolescent in color-
ing and her behavior passive (examples:
Sassacus flavicinctus, Plexippus paykullii,
Corythalia chalcea ) ; in the third, both sexes
are strikingly marked and similar, with the
female often acting aggressively or, in ad-
vanced groups, tending to reciprocal display
with the male (examples: Sassacus ocellatus,
Corythalia fulgipedia). However, there is
by no means a fixed correspondence between
female aggression and striking pattern: for
example, the most aggressive females found
at Rancho Grande were those of Eustiromas-
tix sp.; structurally, this species is strongly
dimorphic with the females dull and little
differentiated from the young; they always
killed the males shortly after mating if kept
together, and, in courting, the males had to
be excessively cautious. A similar situation
occurs in some dendryphantinids (cf. Peck-
ham, 1889) . As in birds, all degrees of di-
morphism, both of morphology and behavior,
can sometimes be found at low taxonomic
levels, including the genus. It seems certain
that hormones and neurohumors are as
deeply concerned in these characteristics as
in vertebrates.
E. Climate and Display.
Another interesting aspect of phylogeny
refers to a possible effect of climate on the
development of display. It seems likely that
there are fundamental differences in behav-
ior in the tropics and in the north, which may
well be responsible for some of the differ-
ences between the Rancho Grande observa-
tions and those in the temperate zone.
The differences in breeding season length
is usually considerable. At Rancho Grande
it is months long, while in the north the adult
males are often active only a few days or
several weeks. In the short-summered north
the relatively few salticids which adapted
themselves to the severe winters also had to
adjust in less obvious ways. Basically this
meant the more perfect coordination of the
breeding mechanisms of the two sexes; in
a short season there would be little time for
the vagaries of physiological rhythms or of
regressive behavior, both of which are so
evident at Rancho Grande. Also, in a climate
where many individuals of few species are
the rule, instead of vice versa, there would
be no economy in restricting a female to the
single insemination which appears typical
at Rancho Grande. Again, the same northern
conditions might encourage the development
of the almost communal displays described
by Peckham (1889, p. 40), and which I have
never had the fortune to see in the tropics.
It must be remarked, however, that com-
munal displays in birds reach a high tropical
development, as in birds of paradise and
cocks-of-the-rock.
Finally, in the brief northern summer
persistent courtships and protracted fights
apparently are usual, from accounts in the
literature. By contrast, in Venezuela court-
ship and threat displays are almost always
short; either they are unsuccessful, which
is usually the case, and quickly broken off,
or else they are consummated in a few min-
utes. This difference is probably also tied up
210
Zoologica : Neiv York Zoological Society
[34: 17
with the leisure of physiological rhythm in
the tropics, with the lack of a pressing hurry
to coordinate the mechanisms.
F. Displays as Specific Barriers.
Almost every student of salticid behavior
is familiar with the frequent occurrence of
display between different species, both in
courtship and in threat. At Rancho Grande,
it was found that in general any A-tone male
would initiate appropriate display before any
moving male or female that showed a few
sign stimuli roughly similar to those of its
own species. The females, presumably be-
cause of the chemical factors, were invari-
ably close relatives; the males, in inter-male
display, had to confoi-m in appearance only.
The heterosexual pairs of species which dis-
played regularly to each other, under uncon-
fined laboratory conditions, were the fol-
lowing: Corythalia chalcea and C. fvlgipe-
dia; Plexippus paykulli, female, and Eustiro-
mastix sp., male ; any two species of Phiale.
This weak selectivity would, on first sight,
seem to indicate that differences in display
have little value in erecting or maintaining
specific barriers. Closer study, however, re-
sults in the following observations.
First, display is only initiated; it is rarely
carried on beyond early Stage I. When it is
continued longer (in courtship only), by a
male of exceedingly high tone, the female
always bi’eaks away well before the end of
Stage II. Even in experimental situations
with chloroformed females, the male himself
was never, in more than a dozen trials, seen
actually to copulate; it seemed that in each
case a mechanical barrier was reached in the
epigynum. On the other hand, fertile eggs
were secured after copulation with chloro-
formed females in one pair of Eustiromastix
sp. and one of Corythalia chalcea, showing
that the drugged condition of the female
was not the final deterrent.
Second, when individuals of their own
species are introduced to a pair of mis-
matched displaying spiders, attention is
quite promptly turned to the appropriate
newcomer.
The above does not of course show that
inter-specific crosses may not occur in na-
ture; it only indicates that, if they do so,
they are probably unusual in the area stud-
ied, even though several pairs of closely re-
lated species occur, each having similar dis-
plays and occupying overlapping ecological
niches. These include Phiale dybowskii with
Phiale sp. ; the latter with P. flammea ; and
Corythalia fulgipedia with C. chalcea.
The particulate nature of display explains
its apparent inefficiency. It does not act as a
single unit, but rather as a series of strainers
of progressively finer mesh. An interspecific
display is interrupted when the constantly
changing stimulus configuration — releasing,
directive or both — becomes too weak, from
the viewpoint of one partner, to draw the
requisite response. This positive response
may, of course, appear active or passive, de-
pending both on the stage of display and on
the sex of the partner. The point at which
the display breaks off depends largely on the
physiological conditions of the two protag-
onists.
It was pointed out on p. 204 that display
may aid in the economy of germ cells by pre-
venting unproductive mesalliances. It seems
clear, however, that as a practical barrier
between established species it has relatively
little importance; the primary walls are
ecological and morphological. As in most
other animal groups, there is usually slight
overlapping of the microgeographical ranges,
of the ultimate ecological niches, in closely
related species; under natural conditions,
interspecific display must occur but rarely.
Also, even if such display runs its full
course, the specific differences of palps and
epigyna are probably in most cases effective
final barriers.
In the formation of species, display may
prove of more importance. As Mayr empha-
sizes (1947), the growth of geographic iso-
lation is doubtless the essential factor in
all species formation. Granting this prec-
edence, it seems likely that cumulative
slight differences in display, as in other be-
havior, often pave the way for final mor-
phological breeding isolation. No one who
has been struck, in other animal groups, by
display distinctions within a species or sub-
species in different parts of its range can
doubt that functional change tends to pre-
cede structural differentiation. To give ex-
amples from my own experience, a crab and
a bird may be mentioned. The waving rhythm
of the fiddler, Uca pugnax rapax (Smith),
varies from Porto Rico down through the
islands to Trinidad and British Guiana, and
on west through Venezuela to reach a peak
of complexity in Cartagena, Colombia. Simi-
larly, the choruses of chachalacas, Ortalis
ruficauda (Jard.), when heard in Tobago
and Caracas, sound as distinct as the calls
of ducks and turkeys in a barnyard. The ap-
parent display differences in Maevia vittata
in Wisconsin and Connecticut (Peckham,
1889, p. 53; Painter, 1913, p. 634) hint at
similar geographic distinctions, of obvious
evolutionary interest, among salticid spiders.
X. SUMMARY.
(See also sectional summaries on pp. 184,
185, 187, 190, 192 and 196-199).
1. A study was made of the epigamic dis-
play behavior of fifteen species of salticid
spiders distributed through seven subfam-
ilies. All observations and experiments were
made at Rancho Grande, Parque Nacional
de Aragua, Venezuela.
2. The factors comprising the innate re-
leasing mechanisms prove to be similar
throughout the family, but their relative
importance varies from genus to genus even
within the same subfamily, and,, in minor
sign stimuli, from species to species.
1949]
Crane: Salticid Spiders: Analysis of Display
211
3. The principal controlling factors of the
internal releasing mechanism appear to be
age, fluctuating epigamic rhythm, hunger,
thirst, fatigue, overstimulation and atten-
tion.
4. The epigamic rhythm plays an espe-
cially important part in daily responses. Only
spiders of the highest tone (i.e., lowest epi-
gamic threshold) usually carry courtship
to completion in either field or laboratory.
No regular periodicity, however, was ob-
served.
5. The controlling factors of the external
releasing and directive mechanisms are
divided into those of the physical enviro-
ment and of the configurational stimulus
situation (sign stimuli).
6. Temperature, humidity and light are
recognized as the principal environmental
variables. Display may be released through-
out a wide range of these factors.
7. Tactile, chemical and visual stimuli are
regarded as the major components of the
configurational stimulus situation.
8. Display cannot be released through
touch, although it plays an important role in
the final stages of courtship.
9. Neither is display released through
chemical stimuli alone although, in the pres-
ence of certain visual stimuli, they are im-
portant secondary releasers of courtship.
They appear to stimulate two overlapping
senses, contact chemoperception (chemo-
taxis) and distance or airborne chemoper-
ception (apparently akin to odor). The im-
portance of each type, both in comparison
with each other and with visual stimuli, de-
pends on the degree of visual dominance of
the genus; in the more primitive genera, al-
though they too are visually dominated,
greater dependence is placed on chemical
senses than in more advanced forms. Ab-
sence of chemical stimuli sometimes changes
courtship to threat display or actual fight-
ing.
10. Visual stimuli alone are sufficient to
release display. The principal factors of the
visual stimulus situation appear to be mo-
tion, distance, size, form, pattern, intensity
and color. The relative importance as well as
the characteristics of these factors varies
throughout the family, and no single one ever
proved to be an essential, primary releaser.
11. In general, motion, form and appar-
ent size are the most important factors, and
the sign stimuli among these categories in
each genus or species may vax*y within fairly
wide limits. Pattern, intensity (except for
visibility contrast) and color, on the other
hand, have little releasing value. Sometimes,
however, a special contrasting or colored
marking does act as a definite releaser or di-
rector for display. An example is the yellow
clypeal band of C. xanthopa, which, in com-
bination with an adequate supporting visual
configuration, is a releaser for inter-male
threat display. An undoubted director is the
white sub-basal abdominal band of many fe-
males, which acts as a copulation guide. Most
clypeal and palp max-kings, iridescent
patches, leg fringes, etc., have no high x*eleas-
ing value, since complete display readily
takes place in their absence without strength-
ening of other elements in the stimulus con-
figuration.
12. This appax-ent uselessness of epigamic
“decox-ations” does not, of course, mean that
they ax-e necessax'ily lacking in adaptive value
for display or that selection has not been
acting upon them toward that end. It is mex-e-
ly a fux'ther instance of the unequal value of
sign stimuli, and of the lack of a lock-key
i-eleasing formula: the effective configui'a-
tion is made up of many small bits ; togethex*,
they have cumulative x*eleasing, directive or
merely excitatoi’y value, but a number may
be missing from an individual situation
without noticeable effect. To only one dy-
namic compound element can the term
“primax’y x-eleaser” be applied : this is a unit
composed of any of sevex-al adequate visual
stimuli; very few x-eleasers and dix-ectox-s are
of even secondary importance.
13. Evidence is px-esented that color per-
ception occurs at least in the yellow x*egion ;
the two tested genex*a, Corythalia and Phiale,
appear to be insensitive to the i*ed end of the
spectrum.
14. All of the components — extex-nal and
internal — of the innate releasing and direc-
tive mechanism form a closely woven, mu-
tually dependent, dynamic whole. When one
impox-tant part of the normal configuration
is weak or absent, x'einforcement of another
pax't — including especially high tone in the
x-eceptor spider — can bring a complete re-
sponse : the concept of hetex-ogeneous sum-
mation is as important in salticids as in other
animal gi'oups.
15. Nevertheless, in the field coux*tships
ax'e usually incomplete when the stimulus
configuration is weak in any pax*ticular.
16. Although individual spiders, particu-
larly females, sometimes showed slight, tem-
pox-ary idiosyncx'asies, display pattex-ns are
fixed and instinctive. No evidence at all was
found of display learning, imprinting ox-
copying of other species’ pattex*ns.
17. Displacement behavior is confined to
actions which normally occur in the sexual
field.
18. Little evidence has been found of dom-
inance x-elationships in the vertebrate sense;
tempox'ary dominance seems to be due to
fluctuating epigamic rhythms.
19. Sociality and home range concepts
apply in some species on a px-imitive level; no
defended territories wei'e observed.
20. The functions of courtship in this
family are held to be stimulatox-y with a si-
multaneous blocking of hunger drives; the
concept of “x-ecognition” does not seem to be
necessary.
212
Zoologica : New York Zoological Sooiety
r 34 : 17
21. The explanations and functions of
threat display depend on the phylogenetic po-
sition of the genus in reference to chemical
dependence. In the apparently most primitive
groups, the males take little or no notice of
one another; there is neither fighting nor
threat display, nor even inter-male court-
ship; mirror display does not normally oc-
cur. In intermediate genera, with greater
visual dependence but strong reliance on air-
borne chemical stimuli, courtship and threat
are similar, usually identical in the first
stage, and appear to result from the mis-
taking of males for females, as suggested by
Bristowe. When the mistake is “discovered,”
fighting often results. In the genera with
the least dependence on chemical stimuli,
however, courtship and threat are distinct,
fighting does not occur and mirror display
is readily induced. In these forms, and to
some extent in the intermediate group,
threat display seems clearly to have a stimu-
latory function which is totally apart from
any direct competition for mates, and which
is not concerned in territorial defense.
22. A tentative hypothetical phylogeny is
presented, in which the studied genera and
some northern relations are placed accord-
ing to dependence on chemical stimuli, dis-
play criteria and method of locomotion. The
correlation with morphological characters, to
be specially treated in a subsequent paper,
are indicated. The evolutionary trend within
all the subfamilies appears to be similar in
various major characteristics.
23. The origins of most display move-
ments appear to be in accordance with the
principle of economical permutation, since
the majority probably developed from or-
dinary motions connected with daily activity.
24. Atavistic behavior was noted in which
low-tone, overstimulated or senile individ-
uals of advanced genera resorted to inappro-
priate aggression and chemotaxis; this was
strikingly similar to the normal behavior of
more primitive groups.
25. It is held that in salticids, the display
motions probably preceded most morpho-
logical secondary sexual characteristics, and
that the latter often persist vestigially after
they have ceased to function as part of the
display stimulus configuration.
26. In salticids, as in other groups, sexual
dimorphism may occur in all degrees within
a closely related group, even within the same
genus, and appears to hold little phylogenetic
significance. In species having both sexes
strongly and similarly marked, the females
are often more aggressive than usual and
tend to perform definite reciprocal or mutual
displays.
27. It is suggested that tropical salticids
may differ in behavior from northern forms
because of the prolonged breeding season,
which necessitates less close correlation of
breeding rhythms between the sexes.
28. Although it is agreed that geographic
isolation is the prime essential of species for-
mation, it appears that growing display dif-
ferences may be a strong secondary factor in
salticid speciation. As a barrier between spe-
cies already established, however, display
seems to be an effective, but usually super-
numerary, isolating mechanism.
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EXPLANATION OF THE PLATE.
Plate I.
Fig. 1. Plexippus paykullii. Courtship, Stage
II. Typical of Stage II courtship in the
Salticidae. In this family, specializa-
tion of display occurs principally in
the earlier phases.
Fig. 2. Observation table at Rancho Grande,
showing presentation of a dried spider
to a test salticid. The spider is mounted
on an L-shaped strip of pasteboard.
The table is covered with oilcloth ruled
in concentric circles for distance ob-
servations. When chemotaxis was to
be eliminated, fresh sheets of paper
were used in each test.
CRANE.
PLATE 1.
FIG. 1.
FIG. 2.
COMPARATIVE BIOLOGY OF SALTICID SPIDERS AT RANCHO GRANDE. VENEZUELA.
PART IV. AN ANALYSIS OF DISPLAY.
Tavolga: Differential Effects of Sex Hormones on Platyfish
215
18.
Differential Effects of Estradiol, Estradiol Benzoate and
Pregneninolone on Platypoecilus maculatus.
Margaret Cordsen Tavolga1.
New York University and The American Museum of Natural History.
(Plates I-V ; Text-figures 1-5).
Androgenic effects of estrogens have thus
far been described as affecting only secon-
dary sex organs in mammals (Allen, Hisaw
and Gardner, 1939; and Witschi, 1939). In
the present work, data are presented to show
gonodal stimulation and androgenic effects of
an estrogen in the platyfish.
The platyfish, Platypoecilus maculatus
Gunther, is a viviparous cyprinodont belong-
ing to the family Poeciliidae. It has been
used frequently for genetic studies (Bellamy,
1928 and 1933; Fraser and Gordon, 1929;
Gordon, 1927, 1931, 1937a, 1947a and b), for
studies of melanoma (Gordon, 1937b, 1948a;
Gordon and Flathman, 1943; Levine and
Gordon, 1946) , and for embryological studies
(Tavolga and Rugh, 1947), but until 1940 it
was not used as an experimental animal for
endocrinological work.
In 1941, Cohen, Gordon and Nigrelli re-
ported on the spontaneous development of
gonopodia in females of Platypoecilus, while
in 1940 and 1942 Grobstein worked out the
development of, and endocrine effects on,
gonopodium differentiation. Cohen in 1942
and 1946 did the first work on the effects of
sex hormones on the platyfish. At that time
he found that pregneninolone has an andro-
genic effect upon the gonads and anal fins of
the fish and that alpha estradiol benzoate has
feminizing effects on the male. Pregneninol-
one, in mammals, has been known as a prog-
estogen, although its effects have been dis-
credited in recent years as ineffective and
partially androgenic (Corner, 1942; Freed,
1942 and 1943). As above indicated, it has
been found since that time that on lower ver-
tebrates, such as fishes, the hormone has an
entirely androgenic effect. In experiments
preliminary to the present work, it was found
that the effects of free estradiol did not co-
incide with those produced by the benzoate
ester, and it was decided to continue and ex-
pand this aspect of the work as well as to
determine the effects of androgens on the
male and estrogens on the female. Pregnen-
inolone was used as an androgen with which
to compare the effects of alpha estradiol when
1 Submitted in partial fulfillment of the requirements
for the degree of Doctor of Philosophy at New York
University.
it was found that this substance did not ex-
hibit strictly estrogenic activity.
MacBryde et al. (1942) found differential
effects of estrogenic substances on the mam-
malian liver. It was decided, therefore, to
investigate the effects of the treatment upon
the liver of the experimental animals as a
possible source of information as to the
reason for the differential effects of the two
estrogens. With these purposes in mind the
present experiments were undertaken.
The author is indebted to Dr. Charles M.
Breder, Jr., of the American Museum of
Natural History and New York University
for his aid and criticism in the progress of
the work and in the preparation of the man-
uscript, to Dr. Myron Gordon of the New
York Aquarium of the New York Zoological
Society and New York University for his
help and criticism during the experiments
and for the stock of fishes2 with which the
work was carried out, and to Dr. Lester R.
Aronson of the Department of Animal Be-
havior of the American Museum of Natural
History for his generosity in supplying
equipment and laboratory space. Thanks are
due also to Dr. Robert Gaunt of Syracuse
University, who provided the initial stimulus
for the problem, and to Dr. Milan J. Kopac
of New York University and Dr. Irwin
Schwenk of Schering Corporation for obtain-
ing the initial supply of alpha estradiol ben-
zoate.
Materials and Methods.
The fishes used for these experiments were
platyfish, Platypoecilus maculatus, of the
New York Zoological Society’s Genetics Lab-
oratory Culture 180 (Gordon, 1948b), their
immediate offspring and those of several sub-
sequent inbred generations. In this strain the
males are the heterogametic sex (XY). The
Y chromosome carries the “spotted” gene
( Sp ), regularly transmitted from father
to son when a spotted male (X)+/(Y)sp is
mated to the recessive female (X)+/(X)+.
The effect of the Sp gene is such that
2 These animals were obtained from the Genetics Labora-
tory of the New York Aquarium, New York Zoological
Society. The work of this laboratory is supported by a
research grant from the National Cancer Institute of the
National Institute of Health, U. S. Public Health Service.
216 Zoologica : New York Zoological Society [34: 18
groups of macromelanophores are distrib-
uted over the major part of the body. The pig-
ment cells begin to show immediately after
the birth of the animals, affording a con-
venient method of identifying males and
females within a day or two after birth. The
females carry the recessive gene for macro-
melanophores (+ or sp ) and appear as gray.
In other strains of the platyfish which do not
possess this or a similar feature, sex identifi-
cation is delayed until such time as the fish
mature, when the anal fin of the male is
transformed into an “intromittent” organ,
the gonopodium. Experiments carried out on
such fish necessitate knowledge of the ratio
of males to females usually derived in a brood
from these fish and the use of statistical
methods in determining the deviation from
such a ratio when the work involves the use
of hormones which produce effects on pri-
mary and secondary sex organs. This type of
analysis is unnecessary with the strain used.
In seven generations only one crossover
occurred, and this was in the control group.
An animal in that group appeared to be a
normal female 21 mm. long, but on sectioning
she was found to possess normal testes for
a fish of that age (66 days). Fraser and
Gordon (1929) indicated that crossing over
of the sex chromosomes in the platyfish is
likely to occur at the rate of 1%.
The experimental individuals were kept in
two-gallon tanks, approximately eight imma-
ture specimens of both sexes to a tank. Plants
and gravel were removed at the beginning of
each experiment in order to secure more
uniform conditions. The room in which the
tanks were kept was maintained constantly
at a temperature of 80° F., plus or minus 3°.
The fish were fed daily on a fresh liver-
Pablum mixture (Gordon, 1943) in amounts
such that each tank received approximately
1.5 cc. per week. The method of introduction
of hormones received a considerable amount
of attention. The literature contains accounts
of hormone administration both by dropping
hormones into the water and by injection
(Berkowitz, 1937; Eversole, 1941; Grob-
stein, 1942a). It was felt that the animals
used in these experiments were too small to
receive injections, since at the beginning of
treatment they were only two to ten days old
and averaged only 8 mm. in length. There-
fore, dropping powdered hormone into the
tanks was attempted.
It was soon noted, however, that the ani-
mals learned early to distinguish between
food and hormone. At first they ingested it
freely, but after a day or two they were ob-
served to ignore it completely and the pow-
der settled to the bottom of the tank, where
it remained until removed. Thus the greater
part of the hormone was not utilized. The
possibility that it may have dissolved and
been absorbed was not overlooked, and experi-
ments designed to check this possibility will
be discussed later. However, since the solu-
bility of the hormones is known to be rela-
tively small, another method of more direct
administration was attempted, and was
found to give good results.
The powdered hormone was introduced in-
to the semisolid liver-Pablum paste and
thoroughly mixed. The food was given in
small lumps each day and the fish were ob-
served to pick at and ingest it freely each
time it was given for the duration of the ex-
periment. They consumed the amount given
in about twenty minutes and close observa-
tion showed that they did not reject any part
of the food, nor could any be seen at the bot-
tom of the tank when later observed. Because
of this it was assumed that the entire amount
was ingested, and with it all the hormone
which did not dissolve in the twenty-minute
period which was needed for ingestion. Al-
though no accurate measurement could be
made of the amount taken in by any individ-
ual fish, it was seen that all the fish had ac-
cess to the food and that there was little fight-
ing among them for it. Each animal, there-
fore, received approximately the same
amount of food and of the hormone. The hor-
mone in oil preparation was mixed into the
liver paste in the same manner, and though it
changed the consistency of the paste to a
slight degree, the preparation which was
chosen contained a high concentration of hor-
mone per cubic centimeter and little of the
solution was needed to secure the necessary
concentration in the food.
The hormones used were pregneninolone
(Pranone, Schering), alpha estradiol (Pro-
gynon DH, Schering), and alpha estradiol
benzoate (Progynon B, Schering). The first
two were prepared in tablet form. The initial
supply of estradiol benzoate was in powder
form and the remainder in solution in sesame
oil. The two forms of estradiol benzoate pro-
duced indistinguishable results and so are
considered together.
The amounts of hormone used were selec-
ted in an attempt to secure doses which
would be adequate to gain results and at the
same time avoid much of the toxicity which
was found to occur, especially from preg-
neninolone (see Text-fig. 1). Pregneninolone
was used in doses of 5, 2.5, 1.25 and 0.625 mg.
per 3 cc. of food. Estradiol and the crystalline
estradiol benzoate were used in doses of 0.5,
0.25, 0.13 and 0.06 mg. per 3 cc. of food. The
estradiol benzoate in oil was obtained in am-
pules containing 1000 rat units per cubic
centimeter of solution in sesame oil, corres-
ponding to 0.166 mg., and the doses given
per 3 cc. of food measured 0.166, 0.083, 0.0415
and 0.0275 mg. These dosages were obtained
by mixing 1, 1/4, etc. cc. of the solution
or the same fraction of tablets with 3 cc. of
food.
Control tanks containing littermates of
the experimental animals were run simultan-
eously with each experiment. In experiments
where oil solution of estradiol benzoate was
used, tanks of littermates which were fed
with a mixture of food and sesame oil alone
were also used.
The doses were given for various intervals,
1949]
Tavolga: Differential Effects of Sex Hormones on Platyfish
217
varying from ten days to seven weeks, and at
the end of each interval at least one male and
one female were removed from each experi-
mental tank and from the control tank. They
were immediately fixed in Bouin’s picro-for-
mol and preserved after fixation in 70% al-
cohol. Each fish was then prepared for fur-
ther study as follows : the anal fin or gono-
podium was removed, cleared and mounted
entire, and the belly of each animal was slit
to facilitate infiltration of solutions. The ani-
mal was decalcified in a nitric acid-phloroglu-
cinol mixture for a period of 24 to 48 hours,
depending on the size of the fish, placed in
several changes of 70% alcohol to remove the
acid, and dehydrated by means of Dioxan.
After infiltration in 60-62° paraffin, the an-
mals were cut at 10 n and the sections stained
with Harris’ haematoxylin and eosin, or with
acid fuchsin and aniline blue (modification of
Masson trichrome stain) .
The sections thus obtained were examined
to ascertain the condition of the ovary or
testis, and the general condition of the other
organs, and the anal fins were examined for
signs of structural changes which might in-
dicate a progressive shift toward maleness
or femaleness. The sizes of the gonads were
determined by measuring the widths of these
glands. It was found that while the lengths of
the gonads were roughly correlated with the
standard length (length from the tip of the
snout to the base of the caudal fin) of the
animals, the width of the gonad was also cor-
related with the development and apparent
activity of the gland as induced by hormone
treatment.
In order to determine size differences be-
tween the eggs of the treated animals and
those of the controls, one or more widely sep-
arated sections of each treated and control
animal was subjected to analysis. The sec-
tions selected were as widely separated as
possible in order to insure that no duplicate
measurements could be made on any egg. Ex-
cept for this consideration, the sections were
chosen at random. In each case, 100 eggs
were measured in order to obtain a good dis-
tribution and a sufficient and representative
sample. The means and standard errors of
the means were obtained and according to the
formulas given by Simpson and Roe (1939)
the significance values were calculated. These
results are given in Text-fig. 4 and Table
IV. In the case of the testes, the same method
of analysis was applied to primary sperma-
tocytes , secondary spermatocytes and sper-
matophores. In addition, an adult control
male was sectioned in order to afford a com-
parison between the testes of the treated
young animals and a normal adult testis. The
testis of this animal was analyzed in the
same manner.
Results.
Two hundred forty-six animals were used
experimentally. Each experimental group
contained 26 to 33 animals when finally pre-
pared and sectioned. Since with the dosages
used there was no statistical difference be-
tween those treated for different periods and
with different dosages, the descriptions
given will cover all periods of treatment and
all dosages, except where otherwise indi-
cated.
General Considerations.
As was stated above, the animals ate the
hormone-impregnated food freely and there
was observed to be no fighting among the
members of any tank for greater amounts of
food. Each pellet of food was eaten within
about twenty minutes of its introduction in-
to the tank, assuring almost complete intro-
duction of the hormone into the animals. It
was possible, however, that the hormone
might be entering the animals by way of ab-
sorption taking place from the amount of
hormone which dissolved into the water dur-
ing the time when the food was present in the
tank. In order to establish whether such dis-
solution took place, and whether, if it did,
the dissolved hormone remained in an active
state, two experiments were set up.
In the first of these, the water between
two tanks was circulated through glass wool
by a conventional air-lift filter in such a way
that the water passed from one tank to the
other without any undissolved particles pass-
ing in either direction. To the immature
fishes of the same strain which were placed
in the first tank was fed the same liver-Pab-
lum paste as was used for the main experi-
ments. The fishes in the second tank received
the same food without the hormone. The
water was transferred from one tank to the
other at the rate of about 240 cc. per hour,
with a complete turnover at the rate of once
in every 30 hours. At the end of a week, the
fish in the second tank, which received no hor-
mone in their food, had developed the same
effects as those in the first tank which were
fed directly, showing, first, that dissolved
hormones or their metabolic products af-
fected the animals ; second, that dissolution
had taken place in an amount great enough
to produce an effect on the animals; and
third, that the hormone was stable during
the time taken for the change of water from
one tank to the other.
Three weeks to a month after the termina-
tion of the main experiments, immature fish
were placed in the tanks used for these ex-
periments without changing the water or
washing the tanks. The animals showed no
effects of any sort and matured into normal
adults, showing that after this period of
time the hormone was no longer active.
During the main body of the experiments,
the effect of the hormones on the experi-
mental animals was first noticed on the males
which were treated with pregneninolone.
Within four to five days after treatment was
begun, when the animals were six to fifteen
days old, and measured 8 to 9 mm. in length,
each animal’s anal fin was modified into the
general form of a gonopodium. A few days
later, the anal fins of the females in the same
218
Zoologica: New York Zoological Society
[34: 18
tanks had also acquired this characteristic.
At about the same time (ten to twelve days) ,
the males in the estradiol tanks developed the
same type of modification, also followed in a
few days by the females. Note that the estra-
diol, while it had superficially the same effect,
was delayed in its action in comparison to
the pregneninolone. Those fish in the tanks
treated with estradiol benzoate, both males
and females, for the duration of experimen-
tation never developed any structures even
superficially resembling a gonopodium. That
the gonopodia developed by the androgen-
treated animals were not typical nor perfect
gonopodia with the characteristic hooks,
spines, serrae, etc., of the platyfish gonopo-
dium will be discussed later. It should be said
here, however, that they were modified in
the male direction sufficiently to be consid-
ered greatly affected by the hormone treat-
ment, and that the fins of the pregneninolone-
and estradiol-treated animals reacted in the
same general manner.
Pregneninolone in the amounts given had
a serious effect on the viability of the treated
animals (see Text-fig. 1) . Forty-eight percent.
(32) of these animals died before the term-
ination of the experiment and therefore
were not considered in the results given. It is
believed, however, that this death rate must
be a significant consideration in the general
effect of the hormone on the metabolism of
Text-fig. 1. Percentage survival of control and
treated animals over a period of eight weeks.
the animals, and should therefore be included
in the general results. The effect of estradiol
was similar, but again quantitatively less, as
only 24% (17) of the animals treated in this
manner succumbed. This number was sig-
nificant as compared to the death rate of the
control, where there was only 4% mortality,
but not as compared to the estradiol benzo-
ate-treated group, since 30% (20) of the
estradiol benzoate-treated animals also died.
The locomotor activity of the treated ani-
mals was not impaired in any way. Sexual
activity, normally absent at this stage of de-
velopment, appeared precociously, and the
tiny animals with miniature gonopodia were
seen vigorously following the females, in a
manner suggestive of precopulatory behavior
of adults, as early as one week after the be-
ginning of treatment, when they were but
two weeks of age. This type of activity con-
tinued until the end of the experimental pe-
riod. The females of the group, although they
exhibited the male type of behavior, showed
it to a lesser degree. Although they tended
to follow each other, they did so less often
and less vigorously. Females were not seen
following males. In the estradiol benzoate
tanks there was no evidence of male behavior
during the entii'e course of the experiment
on the part of either males or females.
Effect on Gonads.
Control Males.
All animals in the experiments were young
healthy specimens, ranging in size at the end
of the experimental period from 8 mm. for
those treated for one week to 22 mm. for
those treated for seven weeks. None of the
animals at the end of the experiments were
old enough to be normally sexually mature,
and they would not normally have become so
for about two months, as Platypoecilus macu-
latus matures at the age of about four to
six months under the laboratory conditions
maintained here. At the ages of one to seven
weeks, therefore, the testes were small com-
pact masses, fused but showing their pri- j
mary bilobed nature, their anterior ends ap-
pearing between two coils of the intestine at
approximately the same cross sectional level
as the caudal tip of the liver. They wei-e at-
tached to the dorsal peritoneum by a short
mesorchium and in a few cases were seen
dorsal to the intestines. See PI. I, Fig. 1. They
ranged in width from 0.08 mm. for the small-
est animals (7.0 mm.) to 0.35 mm. for the
larger ones (19.0-21.0 mm.). The younger
gonads could be seen to contain numerous
groups of cells (cysts) which could be only
poorly differentiated from the main mass of
tissue, except under the higher powers of
magnification at which they could be seen
to be spermatogonia. Between these cysts ex-
isted numerous connective tissue cells and
fibers, making up the stroma of the gland.
The sperm duct in these smaller animals was
poorly differentiated. In the larger speci-
mens, none of which was more than eight
19491
Tavolga: Differential Effects of Sex Hormones on Platyfi.sk
219
weeks old, the cysts were slightly better dif-
ferentiated. They could be seen to be separ-
ated from their surrounding stroma, which
was less distinct, and the smaller cysts had
now taken up a position relatively peripheral.
At this stage these cysts measured 20 to 33 a
in width. A few larger cysts, 36 to 46 a in
width, could be seen toward the center of the
gland. These, under higher magnification,
could be seen to be primary spermatocytes.
These cysts, when present, surrounded the
now well formed duct. These descriptions
confirm those of Wolf (1931) on the devel-
opment of the testis in Platypoecilus macu-
latus at this stage. In no case did the gonads
contain any cellular formations acceptable as
secondary spermatocytes, spermatids, ma-
ture sperm or spermatophores.
Males Treated with Alpha Estradiol Ben-
zoate.
The testes of these animals showed a
slightly retarded development as compared
with the controls (see PI. I, Fig. 2). Their
size range was equal to that of the normals
(0.10-0.34 mm.). The mean was 0.19 mm.
The two testes were slightly separated, indi-
cating an inhibition of their development
toward a fused gland. The gland in general
consisted of a number of peripheral cysts,
surrounding stroma cells which were abun-
dant and ducts which were slightly less well
formed than the normal. The cysts measured
11 to 21 a in width, but since the testes were
so small there were not enough of them to
justify a statistical analysis. A few larger
cysts were present, but they were less dis-
tinctly demarcated than those of the control.
There were no statistically significant dif-
ferences between the widths of the testes
of this group and those of the controls
(P = .05— -see Table II).
Males Treated with Pregneninolone.
Pregneninolone has been shown to have
androgenic activity in the guppy by Ever-
sole (1941) and in the platyfish by Cohen
(1946). It was expected, therefore, that it
would have a similar effect here. That ex-
pectation was justified. The testes of even
the smallest of the males thus treated showed
definite stimulation effects. These testes
were significantly larger than those of the
control (P = .001), measuring from 0.36
to 1.50 mm., with a mean of 0.81 mm. (see
Text-fig. 2 and Table II). In each case,
whether the animal was treated for a short
period or a long one, the results were the
same except for the general size of the
gland, which had sufficient time to grow
larger in the animals which were treated
for a longer period. The cellular effects, in
1.25 P
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t-
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£
2
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2
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<
2
CONTROL
ESTRADIOL BENZOATE
ESTRADIOL
PRESNEN/NOLONE
0.75
0.50
0.25 .
1
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10
1
15
doss midpoints (mm.)
STANDARD LENGTH
1
1
20
Text-fig. 2. Growth of control and hormone-treated male gonads as determined
by measurements of testis widths.
220
Zoologica: Nev> York Zoological Society
TABLE I.
Comparative Sizes of Gonads in Treated and Control Females.
[34: 18
Treatment
Number of
animals
Mean
length
Range
Mean gonad
width
Extremes
Control
13
13 mm.
8-19
0.42 mm.
0.11-1.40
Estradiol benzoate
13
17 mm.
8-24
0.31 mm.
0.11-0.61
Pregneninolone
12
16 mm.
13-23
0.45 mm.
0.30-0.85
Estradiol
16
16 mm.
13-23
0.45 mm.
0.20-0.88
every case, were the same. The effect was
to stimulate the testes to maturity far ahead
of the time at which it would ordinarily be
functional. Cysts of spermatogonia were
present, measuring 30 to 51 y in width, but
in all the animals of the group the sperma-
togenic process had gone far beyond the
stage of spermatogonia and primary sperm-
atocytes into secondary spermatocytes,
spermatids and spermatophores, the pres-
ence of which is the usual sign of a functional
gland (see PI. I, Fig. 3) . The groups of cells
had become differentiated into cysts of ma-
turing primary spermatocytes measuring
93 n plus or minus 2.1 (see Table III), sec-
ondary spermatocytes measuring 97 v- plus
or minus 1.5, or later stages, each cyst con-
taining only one stage of spermatogenesis,
as is found in mature fish. Many of the cysts
contained spermatids in the process of grow-
ing tails. Others contained nearly mature
spermatozoa with heads embedded in Ser-
toli cells, beginning the formation of the
typical ring of the spermatophore. Still
others were found (69 y plus or minus 1.5 m)
which possessed the completed ring form
of the spermatophore, containing mature
spermatozoa, tails inward, dark heads form-
ing a ring and massed together. In many
cases the released spermatophores were
found in the ducts, which is typical of the tes-
tis of the mature fish. In all the cases sperma-
togenesis was active in all its stages ; there
was an abundance of every stage from the
earliest spermatogonia to spermatophores.
In a general way the progress of spermato-
genesis was from the outer portion of the
gland inward toward the duct, and spermato-
gonial cysts were found mainly at the peri-
phery progressing through primary and sec-
ondary spermatocytes to spermatids and
spermatophores which were located near the
center of the gland and adjacent to the ducts.
Since the animals were not treated for more
than seven weeks, it is possible that the
maximum effects were not obtained. Exhaus-
tion effects in Lebistes, in which all sperma-
togenesis is in very late stages and no sperm-
atogonia are present (Eversole, 1941), were
not found. It is possible, therefore, that a
longer treatment would have produced glands
showing lack of germinal elements such as
those described by him.
The position of the glands was also dif-
ferent in the treated fish. In the controls at
this age they tended to be placed, as stated
before, between the coils of the intestine, and
only rarely were situated dorsal to this gen-
eral position. In the pregneninolone-treated
animals, however, the testes had grown so
large that they pushed dorsally. They often
occupied a position completely dorsal to that
of the normal gonad, while in all other cases
at least part of the gland projected above
the coils of the intestine in cross section.
Often they occupied the major part of the
body cavity.
The interstitial tissue had also changed.
In comparison to the size of the treated gland
it was very sparse, being seen as mere
threads between the cysts of maturing germ
cells. However, toward the center of the gland,
where it filled in spaces between the cysts
and the sperm duct, slightly more abundant
interstitial tissue was often seen. In the main
it appeared like connective tissue, often with
large oval cells. A few collecting tubules lined
with cuboidal epithelium could also be seen.
TABLE II.
Comparative Sizes of Gonads of Treated and Control Males.
Treatment
Number of
animals
Mean
length
Range
Mean gonad
width
Mean
W/SL*
t
P
Control
13
15.1 mm.
7.0-21.0
0.20
0.0122
Estradiol benzoate
12
16.0 mm.
8.5-21.5
0.19
0.0125
.17
.05
Pregneninolone
10
16.2 mm.
8.5-21.5
0.81
0.0518
8.9
.001
Estradiol
17
15.2 mm.
8.0-22.0
0.35
0.0209
2.4
.02
* W/8L equals the ratio of the gonad width divided by the standard length.
1919] T avoir; n : Differential Effects of Sex Hormones on Platyfish 221
TABLE III.
Sizes of Spermatogenetic Cysts of Treated and Control Males.
Treatment
Structure
of testis
Number
of
animals
Sample
Mean
diam.
(micra)
±
General effect
Control at
experimental
stage
Primary
sp. cytes
14
Few — insufficient
for significant
count
secondary
sp. cytes
14
None pi’esent
sp. phores
14
None present
Control mature
male
primary
sp. cytes
1
100
60
1.4
secondary
sp. cytes
1
100
73
1.3
sp. phores
1
100
49
.9
Estradiol
benzoate
primary
sp. cytes
12
Very few
secondary
sp. cytes
12
None present
sp. phores
12
None present
Pregneninolone
primary
sp. cytes
10
100
93
2.1
Enlarged over
mature control
secondary
sp. cytes
10
100
97
1.5
Enlarged over-
mature control
sp. phores
10
100
69
1.5
Enlarged over
mature control
Estradiol
primary
sp. cytes
2
100
56
1.4
Not significantly
smaller than
control
secondary
sp. cytes
2
100
56
1.1
Significantly
smaller than
control
sp. phores
2
100
42
1.1
Significantly
smaller than
control
Significance Values Calculated from Means in Table III.
Significance values are calculated as the difference between two
means divided by the standard error of the difference.
Primary spermatocytes
Estradiol
Pregneninolone
Control mature male
2.0f
13.0
Pregneninolone
14.8
Secondary spermatocytes
Estradiol
Pregneninolone
Control mature male
9.9
12.1
Pregneninolone
22.2
Spermatophores
Estradiol
Pregneninolone
Control mature male
5.0
11.2
Pregneninolone
14.6
* <7m equals standard error of the mean,
t These values are not to be considered significant.
222
Zoologica : New York Zoological Society
[34: 18
These observations corresponded very well
to Wolf’s (1931) description of the intersti-
tium of a young mature male.
Males Treated with Alpha Estradiol.
In mammals it has been found that alpha
estradiol is usually an estrogenic hormone
(Willier, 1939; Witschi, 1939), although in
some cases paradoxical effects have occurred
which have affected the secondary sex or-
gans. However, the gonads are not ordinar-
ily changed in these cases. When the present
experiments were in their earliest stages, it
was found that apparently this was not true
in the platyfish. Therefore further experi-
ments were carried out in order to determine
the effects of this substance. The animals in
this group fell into two sets, the difference
being one of size and depending not at all
on dosage or length of treatment. In all ani-
mals under 18 mm. in length, the testes pre-
sented a normal control picture. The testes
were small, compact, showed spermatogonia
and early spermatocyte stages, compared well
in size with those of the controls, except for
a very small increase, and generally showed
no significant effects. In all animals, however,
over 19 mm. in length, the developmental
picture was entirely changed. The gonads in
these cases were intensely stimulated organs,
showing all stages of spermatogenesis. Dis-
crete cysts of primary and secondary sperm-
atocytes, 56 plus or minus 1.4 y, and 56 plus
or minus 1.1 y, respectively, spermatogonia
25 to 30 g at the periphery, and normal
spermatophores of 42 plus or minus 1.1 g at
the center were present. Note that the sizes
of these cysts were significantly smaller than
those of the pregneninolone animals (see
Text-fig. 3 and Table III). The ducts also
were large and well formed, typical of the
ducts of a mature male, and they were fre-
quently filled with spermatophores. These
spermatophores, like those of the pregnenin-
olone-treated animals, were well formed and
showed no sign of precocious extrusion from
the cysts or of abnormal cells despite their
comparatively small size (see PI. I, Figs. 4
and 5) . The picture approximated that of ani-
mals treated with the known androgen preg-
neninolone, in every detail except size. It
seems, then, that estradiol, far from being
an estrogen, acts much like an androgen in
the stimulation of the testes in these fish.
The position of the testes in the abdominal
cavity of the animal and the appearance pre-
sented by the interstitial tissue corresponded
in every way to the pregneninolone-treated
animals.
In order to determine the size relationship
of these stimulated spermatogenic cysts to
those of the normal testis, an adult untreated
male was sectioned and prepared in the same
manner as the experimental animals. As with
the experimentals, 100 cysts of each type,
primary and secondary spermatocytes and
spermatophores, were measured by means
of the ocular micrometer and statistical an-
alyses were made (see Table III and Text-fig.
3). In each case the cysts of the pregnen-
inolone-treated animals were significantly
larger than those of the adult control and
•i
■
o
o
ftj
Q.
mean =73
mean = 49
40
30
20
to
mean = 56
mean = 56
size class midpoints (In micro)
mean = 42
PRIMARY SPERMATOCYTE CYSTS SECONDARY SPERMATOCYTE CYSTS
SPERMATOPHORES
Text-fig. 3. Comparative size ranges of primary and secondary spermatocyte cysts and
spermatophore cysts of pregneninolone- and estradiol-treated animals as compared with
those of a normal adult male.
percentage in each closs
19491
Tavolga: Differential Effects of Sex Hormones on Platyfish
223
with one exception the cysts of the estradiol-
treated animals were smaller. In the case of
the primary spermatocytes the cysts of the
estradiol-treated animals v^ere not signifi-
cantly smaller than those of the control adult.
Control Females.
The typical female ovary of a platyfish of
two to eight weeks of age was located in se-
rial cross sections between two coils of the
intestine, suspended from and approximat-
ing the dorsal peritoneum. The ovary ranged
in size from 0.11 to 1.40 mm. and the mean
value was 0.42 mm. Posteriorly it lay free
of contact with surrounding organs, and far
posteriorly, just anterior to its posterior
margin, it lay almost if not completely free
in the abdominal cavity. At the young stages
studied here, it consisted of a single mass
composed mainly of young circular or ovoid
ovocytes, before, or in the larger animals
during, the process of yolk deposition. The
mean size of these ovocytes was 109 /z plus or
minus 6.5 /x (see Text-fig. 4 and Table IV).
The ovary as a whole was compact, little if
any space being present between adjacent
eggs. Any space which was present was al-
most filled with small amounts of stroma.
Stroma also filled spaces between the outer-
most eggs and the peripheral flattened epi-
thelium (see PI. II, Fig. 6).
In the younger stages the ovocytes were
yolk-free, with slightly reticular cytoplasm
and a lighter-staining nucleus. Each nucleus
contained one or two deeply-staining, prom-
inent nucleoli. The nucleoplasm itself was
reticular in appearance, studded with chro-
matin granules. These larger eggs were lo-
cated mainly at the periphery of the organ
and each was bounded by epithelial cells con-
stituting the follicle. The younger cells,
oogonia, 21 to 45 g in diameter, were situated
50
40
30
20
10
ESTRADIOL BENZOATE
mean * 39
li
CONTROL
mean = 109
ESTRADIOL
freon * 73
hiL
■ ■
PREGNEN/NOLONE
meon * 69
JJlLi ■ ■_
size class midpoints In micro
DIAMETERS of OVA
Text-fig. 4. Comparative size ranges and distributions of eggs of control, estradiol
benzoate-, pregneninolone- and estradiol-treated females.
224
Zooloffica : Nev> York Zoological Society
TABLE IV.
Sizes of Eggs of Treated and Control Females.
[34: 18
Treatment
Number
of
animals
Sample
Mean
diam.
(micra)
± aU
General effect
Control
12
100
109
6.5
Estradiol
benzoate
13
100
39
2.1
Great inhibition
Pregneninolone
12
100
69
3.9
Partial inhibition
Estradiol
16
100
73
4.8
Partial inhibition
Significance Values Calculated from Means in Table IV.
Control
Estradiol benzoate
Estradiol
Pregneninolone
5.2
6.7
0.6*
Estradiol
4.3
6.1
Estradiol benzoate
10.2
* These values are not to be considered significant.
nearer the ovarian cavity. They were some-
times imbedded in the stroma immediately
surrounding it, but usually maintained con-
tact with the epithelium of the cavity. Their
cytoplasm was more deeply stained and pre-
sented a more homogeneous appearance.
In the later stage the situation was much
the same except for the appearance of the
larger eggs. These had now grown consider-
ably in size, measuring 100 to 280 y, and
their cytoplasm presented a far more retic-
ular appearance than was true of the younger
eggs. Near the periphery of some of them
yolk granules were discerned, but this was
true only in the largest of them. In all cases
intermediate stages were present between
the largest and smallest eggs.
Females Treated with Alpha
Estradiol Benzoate.
The ovaries of the animals treated with
alpha estradiol benzoate were more compact
and smaller than those of the controls, meas-
uring 0.11 to 0.61 mm. and averaging 0.30
mm. in width. The eggs appeared tightly
pressed together and were deformed by this
pressure in some cases. The ovaries were
more closely pressed between the coils of the
intestine. Although the arrangement of the
eggs appeared normal and a fairly well
formed duct was present, the eggs themselves
showed an inhibition of development (see PI.
II, Fig. 7) . The larger eggs were peripheral,
gradating to smaller ones in the center of the
organ. In size they ranged from 11 to 96 y
(see Text-fig. 4 and Table IV), and none of
them approached the size of eggs found in the
control fishes of the same size. No evidence
of yolk deposition was present in any of the
eggs in this group. The cytoplasm of these
eggs was more homogeneous than was true
of the ova of the controls, and they compared
in size and structure to a much younger
stage than that which would be expected
from the age and size of the fish. It is evident,
then, that though few acute abnormalities
were present in the structure of the individ-
ual eggs, their size and appearance indicated
that they were greatly inhibited by the treat-
ment.
Females Treated with Pregneninolone.
The ovaries of these animals were again
slightly smaller than those of the controls,
measuring from 0.30 to 0.85 mm. and averag-
ing 0.45 mm. in width. The sizes of the
groups overlap a great deal (see Table I),
but the largest ovaries of the pregnenin-
olone-treated animals did not reach the size
of the largest ovary of the control group. The
main effects, however, were those appearing
in the size and structure of the eggs. The
ovaries of these animals presented an ex-
tremely abnormal appearance. The greater
number of them were shrunken and small,
appearing completely pressed out of shape by
the surrounding coils of the intestine, as in
the estradiol benzoate-treated animals (see
PI. II, Fig. 8). It is doubtful, however,
whether this shrunken appearance was due to
the pressure caused by the intestine, since the
control ovaries were subjected to the same
pressures and did not show the deformities.
Also the pregneninolone ovary, like the con-
trols, lay, at its posterior end, free in the ab-
dominal cavity, and the deformities were
equally great there. Therefore some other
cause must be assigned to this phenomenon,
presumably one due to the hormones in-
volved. The ovaries contained in most cases
little interstitial tissue, and while this was
more deeply stained than the normal, it did
not appear especially abnormal. The eggs
themselves, however, showed definite effects.
They were seldom as large as those of the con-
trol, having a mean size of 69 y plus or minus
1949]
Tavolga: Differential Effects of Sex Hormones on Platyfish
225
3.9 fi (see Text-fig. 4 and Table IV) . In a few
cases large eggs could be found and these
were the most nearly normal-looking ones.
Even they appeared degenerate, however,
showing deeply-staining cytoplasm, slightly
irregular nuclei and a partially deformed ap-
pearance. The remaining eggs were uni-
formly deformed in shape, the main body of
them having irregular depressions in their
sides and usually one concave side, so that
the individual eggs took on the appearance of
pushed-in balls. The nuclei were also mis-
shapen, showing elongations and irregulari-
ties, each one staining deeply. The cyto-
plasm often had a mottled appearance, in
contrast with the even staining of the con-
trol. This resulted, presumably, from some
effect on the cytoplasm, which caused parts
of it to stain deeply and others lightly, with-
out any regularity. Another significant point
was in regard to the size of the eggs. All those
which were not included in the groups of
larger eggs mentioned first were extremely
small as compared to eggs in the same stage
of a control, measuring from 11 to 60 /x. No
evidence of yolk deposition could be seen in
any individual eggs.
In most of these ovaries the duct was poor-
ly formed and the edge of the epithelium was
ragged and abnormal, showing cells and bits
of tissue sloughing off into the duct.
Two variations of these conditions were
found. In two cases the ovary was large but
the eggs were scattered and large spaces
were present between them (see PI. II, Fig.
9) . While the eggs in these specimens were
not usually as deformed as they were in the
cases described above, they were deeply
stained and appeared to be in a state of de-
generation. As above, few eggs could be found
which were as large as those of the control
of the same age and size, but several ap-
peared which had been approaching this size
and condition before treatment with hor-
mones was begun. These eggs showed ao-
nroximately the same irregularities as the
large ones described above. The epithelium
surrounding the gland was ragged and shred-
ded in many places and the cells of the duct
were ragged.
In two other cases a definite bi-partite
ovary was found. In one of these the eggs
were fairly large and normal-looking, meas-
uring between 90 and 130 n (see PI. II, Fig.
10) , and appeared to be comparable to the
eggs of the ma.ioritv of the control ovaries.
In the other case, however, the eggs were
small and degenerate-lookinsr. measuring
about 40 to 60 a. and were stained deeply,
showing deformities. In this ovary there ao-
peared two definite ducts, one for each half
of the gland, which showed fairly regular
epithelium. The first case contained a duct
which was wide and flat horizontally, ap-
parently serving both sides of the eland. Jt
is believed that this remaining evidence of
the b?-lobed nature of the embryonic gonad
mav have been caused bv the inhibitory ef-
fects of the androgenic hormone applied. All
variations of the ovarian conditions were
used together in making the statistical anal-
ysis of the eggs in this group.
Females Treated with Alpha Estradiol.
In general terms, the ovaries of this group
showed the effects expected of an androgen.
The results were very similar to those pro-
duced by pregneninolone. The majority of the
animals possessed ovaries which appeared as
shrunken masses, with deformed eggs such
as those described for the pregneninolone-
treated animals, staining poorly and in a
mottled fashion. The size also, of both the
ovaries and the eggs, was comparable to the
size of those of a pregneninolone-treated fe-
male, since measurements of the ovaries
ranged from 0.20 to 0.88 mm. with a mean
size of 0.45 mm. (see Table I) . The eggs had
a mean size of 73 plus or minus 4.8 y, a size
which is not significantly different from that
of the pregneninolone-treated eggs.
There were several exceptions to this gen-
eral picture. In two cases the ovary showed
the same scattering which appeared in some
of the pregneninolone-treated animals (see
PI. II, Fig. 11). The same larger degenerat-
ing eggs, and the same type of atretic appear-
ance in the small eggs was present. In one
case, there appeared a bi-lobed ovary such as
that described above, which possessed one
duct to serve both parts of the gland. In this
specimen the eggs were small, measuring 40
to 60 fi, and while not as deformed as the typ-
ical eggs of this group, some atypical shapes
were present and the eggs generally stained
more deeply than the controls. In some cases
there appeared a variety of degeneration not
seen in the pregneninolone-treated group. In
these ovaries there were a few eggs which
appeared to be almost normal, both in size
and general appearance. The remainder of
the comparatively large gland was composed
of a substance which at first appeared to be
adipose material. Upon closer inspection,
however, it was concluded that at the places
where this material was seen, there had once
been large eggs in the process of yolk deposi-
tion (see PI. II, Fig. 12). The eggs had ap-
parently degenerated, since the masses con-
tained no recognizable structures, and left
behind them a mass of fatty yolk-filled ma-
terial. Vacuoles were present, which showed
the presence of lipoids. Some yolk granules
were to be seen. At certain points about the
periphery of these masses there appeared
epithelium of a largely degenerate nature
which was broken and sloughed in parts.
Since there had been several eggs of this
nature, it might have been expected that
there would be some type of separation be-
tween them. For the most part, however, this
was not true, and the masses were indistin-
guishable from one another, showing no evi-
dence of where one egg ended and another
began. In some places a portion of the above-
mentioned epithelium remained, to give
some indication of the limits of the egg, but
this was true only in a few cases. The masses
226
Zoologica: New York Zoological Society
[34: 18
of material were of various shapes so that
no indication remained of the original shape
of the ovum. The remaining eggs were of the
small deformed type mentioned above and
were pushed to one side of the organ. The
ovary, because of this peculiar content, was
quite large, although the egg content was ex-
tremely small. The appearance of the organ
as a whole was a degenerate one. The outline
of the organ, even where the eggs were pres-
ent, was ragged and appeared degenerate,
as the epithelium was ragged and uneven in
contrast to the smooth epithelium of the con-
trol.
Effects on the Anal Fin.
Grobstein in 1940 published a complete
description of the developmental stages in
the transformation of the platyfish’s anal fin
into the gonopodium. In 1942, however, he
partially changed the terminology used in
order to conform to prevailing taxonomic
usage. In all the following descriptions, the
terminology used in the 1942b paper will be
employed.
Control Males.
The anal fin of the control male fish at the
ages studied here was one in which no differ-
entiation or growth had taken place in the
change from the undifferentiated fin toward
the typical gonopodium. The fin looked like
the female fin of the same age (see PI. Ill,
Fig. 13) . The fins were well formed, the third
ray slightly thicker than the others. The
fourth and fifth rays, particularly the fourth,
projected slightly beyond the others. No bi-
furcations were present in the younger fins
and the larger ones possessed only primary
bifurcations. Secondary and tertiary bifur-
cations which, according to Grobstein, de-
pend on age, had not yet appeared in any of
the fins studied. No growth of the third ray,
which indicates the beginning of differentia-
tion into the gonopodium, had begun in any
of the fins. The only difference between the
male and female control fins at this age was
the presence of macromelanophoi-es in the
male fin, due to the Sp gene. The females did
not possess these macromelanophores.
Males Treated with Alpha
Estradiol Benzoate.
Fins of the males treated with alpha estra-
diol benzoate presented the same picture as
did the controls. In all the fins, which came
from animals not more than eight weeks old,
no differentiation of any sort tending in the
male direction was seen. The fourth and fifth
rays projected slightly beyond the others,
but no more so than is normal in the female
fin and certainly not enough to give the im-
pression that they are beginning the gono-
podial growth phase (see PI. Ill, Fig. 14).
They appeared as normal fins for this age of
fish, but since no animals were carried to ma-
turity it is not known whether the hormone
would have been enough to prevent gonopo-
dial differentiation entirely.
Males Treated with Pregneninolone.
The anal fins of all animals in this group
were affected by the hormone treatments.. In
the case of the smallest animals, ten days to
two weeks of age and 8 to 9 mm. in length,
which had been treated for one week to ten
days, the transformation had proceeded only
into Phase I, and all these fins possessed third
rays which were segmenting and growing,
producing an elongation of the cephalad por-
tion of the fin. All those treated for three
weeks or longer, however, showed a complete-
ly modified picture. In these groups, every
fish possessed an almost completely differen-
tiated gonopodium (see PI. Ill, Fig. 15).
Most of these were almost perfect, although
a few existed which were lacking in certain
elements present in a normal fin. The usual
element missing in such an incomplete fin
was the spines, which in most cases, if pre-
sent, were flattened and smaller than normal.
Some fins were seen where no spines at all
were present. Since the oldest fish in the
group were not more than eight, or at most,
nine weeks old, it can be assumed that this
precocious development was due to the effects
of the hormones administered. That the mod-
ifications of the fins correlated well with the
growth and differentiation of the gonads is
further evidence for this assumption.
Males Treated with Alpha Estradiol.
Anal fins of males treated with alpha estra-
diol could be placed in two groups. These cor-
responded directly to the division which oc-
curred in the. description given already of
the gonads of this group. In those animals
which were below the size of 18 mm. at the
end of the experiments, the fins, like the go-
nads, did not show the usual effect of the an-
drogen. Each of the anal fins observed in this
group was in Phase I of growth and elonga-
tion of the third ray. The fins appeared as
modifications of the female condition in
which the third, fourth and fifth rays had
grown long enough to project beyond the
others to about one-third of their length. In
the majority of the animals, the third ray
was found to be segmented as in Phase I,
rather than like the control, in which seg-
mentation was far less definite.
The group comprising those animals which
reached a size of 19 mm. or more contained
fins which were modified far more toward
the typical male condition. They were almost
complete but showed more variability than
did the pregneninolone-treated males. Ele-
ments which were present consistently were
the subterminal segments with the terminal
hook, the elongation of the fourth ray, with
its cephalic ramus curved in a cephalad di-
rection, spines and proximal serrae. Ele-
ments which were absent or incomplete in
the imperfect fins were distal serrae, the
spoon and spoon support, and the blade.
These were absent in different combinations.
Although the fins of this group were not
complete, they showed a definite tendency to-
ward the male form (see PI. Ill, Fig. 16) . If,
Tavolga: Differential Effects of Sex Hormones on Platyfish
227
as Grobstein postulates, the gonopodium is
under the control of the testis, the present
evidence supports that view.
Control Females.
The anal fin of the female control animals,
like the males at this age, showed no signs
of differentiation toward the adult form. The
structures possessed by these fins were those
typical of the young female (see PI. IV, Fig.
17). The fins did not differ materially from
the adult female type except that, as in the
male, only primary bifurcations of the
fourth to ninth rays had taken place. The
third, fourth and fifth rays projected slight-
ly beyond the others, as is normal. The
thickening of the third ray present in the
male fin at this age was present to a lesser
degree in the female. No macromelanophores
were present, since the female does not carry
the Sp gene. Except for this last distinguish-
ing characteristic, present only in this strain,
the fins were structurally similar.
Females Treated with Alpha
Estradiol Benzoate.
In animals treated with this estrogen, the
condition of the anal fin was indistinguish-
able from that of the control fin (see PI. IV,
Fig. 18). The third, fourth and fifth rays
showed the same slight extension. The third
ray was again slightly thicker than the
others. Bifurcations of the rays in animals
of the same age were identical.
Females Treated with Pregneninolone.
The fins of the animals in this group pre-
sented a varied picture. None of them posses-
sed a complete and normal gonopodium, but
neither did any possess the typical female
anal fin. All animals possessed fins which
had progressed far beyond the first phase of
gonopodium formation and many had gone
into the third phase (see PI. IV, Fig. 19).
All animals had completed the preliminary
growth phase, in which the 3, 4, 5 ray com-
plex segmented and pushed out in the ceph-
alo-distal portion of the fin to form a pro-
montory there. Elongation of these rays con-
tinued throughout Phase II of gonopodium
formation. At the beginning of this growth
new segments appeared in the third ray, and
at the end of the first phase there were gen-
erally nine segments present (Grobstein,
1940). During the second phase, these seg-
ments increased in number to twenty-two
when the gonopodium had completed its
growth. All animals in the group possessed
at least ten segments and specimens were
found in which the complete number was
present. At the end of Phase II differentia-
tion of the various specialized parts of the
gonopodium began. The great majority of
the animals had arrived at this stage. In
many, however, the differentiation was aber-
rant, showing certain completely differenti-
ated parts, while other parts, which should
have differentiated concurrently, were still
in an undifferentiated or partially differen-
tiated state. Plate I V, Fig. 19, shows a gono-
podium of this group in which the differen-
tiation was almost complete. This fin had pro-
gressed as far as the “blade stage” (Grob-
stein) and shows most of the elements of a
complete gonopodium in a more or less nor-
mal state. The fin possessed proximal and
distal serrae, the blade, the spoon and spoon
support, the terminal hook and subterminal
segments and other elements. Other gonopo-
dia were found which possessed good seg-
mentation of the third ray and a perfect ter-
minal hook, which should not appear until
the time at which the distal serrae differ-
entiate and after the formation of the prox-
imal serrae, but both sets of serrae were
missing. Such varied differentiations were
common but the general rule in this group
was partial or complete differentiation of all
parts, many with slight deformations.
Females Treated with Alpha Estradiol.
Alpha estradiol was found to have almost
the same effects as pregneninolone, although
they were somewhat delayed (see Text-fig.
5). Thus the majority of the fins in this
group had begun differentiation but had pro-
gressed to a lesser degree than those of the
px-evious group. All the fins had entered the
first or preliminary growth phase, since all
of them showed at least the segmentation and
strengthening of the third ray, and the re-
sulting promontory forming on the cephalo-
distal border. Most of them had in addition
entered the second phase, in which growth
had gone on to form a great elongation of the
fin. About one-third of them went on into
the third phase, in which they showed va-
rious stages of differentiation. In this group
no complete gonopodia were found, but many
fins showed the beginnings of the differen-
tiated elements. There were present terminal
hooks together with subterminal segments,
proximal sei'rae, a few sets of distal serrae,
the cephalic tuiming of the fifth ray, and the
cephalic ramus of the foui’th ray. All these,
however, while they appeared together in a
few animals, were usually pi*esent in less
complete combinations. Alpha estradiol,
while it is androgenic to a gi'eat extent, since
it induces the formation of gonopodia, does
not pi'oduce neaiiy as complete gonopodial
structures as does pregneninolone (see PI.
IV, Fig. 20). The presence of an androgenic
effect, however, is easily seen.
Effects on the Liver.
Control Animals.
The liver in the platyfish was divided into
several large lobes. Lobules were not separ-
ated as in the mammal, and the entire gland
appeai-ed as a compactly arranged mass of
cells. Cords of cells were present, though not
as distinctly as in the livers of higher forms.
The cells themselves measui'ed about 8 g in
diameter and wei-e weakly eosinophilic. The
nuclei averaged about 2 p. in diameter. The
stroma was poorly distinguished from the
epithelial cells. The mass of homogeneous-
228
Zoologica: New York Zoological Society
[34: 18
appearing cells was profusely supplied with
blood vessels of all sizes. The nuclei, from
2 to 3 /x in diameter, were nearly circular and
thickly granular. The cells were of three
types, non-vacuolated with weakly eosino-
philic cytoplasm, partially vacuolated with
one or two vacuoles filling about one-third of
the cytoplasm, and a third type in which
larger vacuoles filled most of the cell. In all
the vacuolated cells, the vacuolated ends ag-
gregated together in a direction farthest
from the blood vessels, producing pale
blotches in the structure of the liver from
10 to 20 g in diameter and very irregularly
placed. The predominant type of cell was the
partially vacuolated one. The second cell type
in order of predominance was the one which
was non-vacuolated and weaklv eosinophilic.
See PI. V, Fig. 21.
Animals Treated with
Alpha Estradiol Benzoate.
The liver cells of these animals averaged
8 to 12 g in diameter. Three types of cells
were present, one non-vacuolated and weak-
ly eosinophilic, another containing a vacuole
which filled one-third to two-thirds of the
cytoplasm, and a third which was highly
vacuolated. The predominant type was that
in which one-third to two-thirds of the cyto-
plasm was vacuolated, followed by the non-
vacuolated type. The organ as a whole pre-
sented a compact appearance much like that
of the control. The stroma network resembled
that found in the control. No degenerative
effects could be noted. See PI. V, Fig. 22.
Animals Treated with Pregneninolone.
The cells of the livers of these animals
averaged from 10 to 15 g in diameter, being
generally much larger than those of the con-
trol. The cells were all more or less vacuo-
lated. The type of cell which predominated
was one in which the cytoplasm was occu-
pied almost completely by a large vacuole.
The vacuoles occupied, on the average, about
nine-tenths of the cytoplasm and as a direct
result the nucleus and remaining cytoplasm
were pushed to one side. The general ar-
rangement of the cells about the capillaries
was as in the control and the entire organ
presented a highly vacuolated appearance
(see PI. V, Fig. 23). Vascularization ap-
peared increased, with a capillary for every
RATE OF GROWTH 6 DIFFERENTIATION OF THE GONOPODIUM
CONTROL V
» A < 6 » 4 » ■«
I 2 3 4 5 6 7 8
Weeks of Treatment
Text-fig. 5. Graph showing growth of gonopodium of control, estradiol benzoate-,
pregneninolone- and estradiol-treated animals.
1949]
Tavolga: Differential Effects of Sex Hormones on Platyfish
229
seven to ten cells. The stroma network ap-
peared more distinctly as a result of the ex-
treme vacuolization of the parenchyma cells.
The entire organ showed changes in struc-
ture, fatty in nature as demonstrated by the
application of Sudan IV in frozen sections.
The controls used for these reactions were
normal livers of the same age, which showed
comparatively few sudanophilic globules.
The experimental animals, on the other hand,
showed an abundance of these globules in
their liver cells.
Animals Treated with Alpha Estradiol.
The liver cells in these animals averaged
about the same size as those of the estradiol
benzoate-treated animals, but the great ma-
jority of them showed vacuoles. Occasional
cells showed the extreme vacuolization of the
pregneninolone-treated livers, nine-tenths of
the cell being occupied by a vacuole. For the
most part, however, the cells contained vacu-
oles which occupied about one-half their vol-
ume and there appeared frequently cells with
several small vacuoles instead of one large
one. In general, the organ showed signs of
far greater vacuolization than the control,
and, therefore, partial changes similar to
those shown by the livers of the pregnen-
inolone-treated animals, but in a stage which
was far less advanced. See PI. V, Fig. 24.
Discussion.
Regnier in 1938 made a comprehensive
survey of the history of fishes as experi-
mental animals in sex hormone studies, and
in addition described her own experiments
on Xiphophorus hellerii with anterior pitui-
tary hormones and ovarian and testicular
powders and extracts. Essenberg (1926) de-
scribed sex reversal in Xiphophorus. Blacher
in 1926 found that testicular hormones are
necessary for secondary sex characteristics
in Lebistes. Castration and implants of
gonads in Xiphophorus was carried out by
van Oordt and van der Maas (1926). Berko-
witz, in a series of papers (1937, 1938, 1941a
and b) , described the effects of estrogens and
mammalian gonadotrophins in Lebistes,
while Eversole (1939 and 1941) worked on
the effects of androgens in this animal. In
two papers in 1941 (a and b), Turner tested
the effects of androgens on Gambusia. Scott
(1941 and 1944) worked on the effects of
steroids on the skeleton of Lebistes.
Eversole (1941) mentioned that testes of
Lebistes treated for 42 days with pregnenin-
olone showed all stages of spermatogenesis,
with later stages predominating, that the
spermatids and spermatophores produced
were abnormal, and that in animals treated
for 50 days, later stages in spermatogenesis
were present almost exclusively, spermatids
and spermatophores were atypical and that
the gonad was generally in a degenerate state
because of a rapid maturation of the germ-
inal elements which left few spermatogonia.
He also stated that the epithelium of the
ducts tended to dismember at that time and
that the stroma was hypertrophied. These
observations were not seen in the present
studies. Most of the animals, it is true, were
not treated for more than six weeks, but
some were treated for seven weeks, and even
these did not show the degeneration of the
gonad which was described by Eversole.
Whether this difference was caused by the
difference in experimental animals used, by
the different method of administration of the
hormone, by differences in dosages or ages,
or by different conditions under which the
animals were kept, is not clear. However, the
gonads of the animals in the present work,
while they showed great stimulation, were
seen to contain germinal elements and abund-
ant primary and secondary spermatocytes as
well as spermatophores. This was true
whether the animals were treated for two
weeks or for seven. In the case of the animals
treated for six to seven weeks, the picture
of the gonad was equivalent to a normal ma-
ture testis, both in size and in quality of the
elements contained. Spermatogonia were
present, both primary and secondary sperma-
tocytes were abundant, and the later stages
of spermatogenesis were abundant and ap-
peared normal. No signs of degeneration of
any sort could be distinguished. As was
stated in the results, it is possible that a
longer period of treatment might have pro-
duced the exhaustion effects spoken of by
Eversole.
Winge (1934 ) has shown that sex reversal
may be detected by genetic means in the
guppy. These reversals, however, were al-
ways from female to male. It is evident, how-
ever, that the sex determining mechanism in
the guppy is less stable than that of the platy-
fish, since only two cases of sex reversal in
Platypoecilus have been mentioned in the lit-
erature. Both of these were naturally-occur-
ring phenomena (Breider, 1942; Gordon,
1947a). No sex reversal in the platyfish has
ever been reported as having been induced
by hormonal or any other means. Because of
this relatively unstable sex mechanism in the
guppy, it is easier to understand why Berko-
witz (1937) was able to secure sex reversal
and ovotestes in this form. No such phenom-
ena were found in the present work. Al-
though degeneration of the gonads was com-
mon as a result of hormone introduction, no
sign of any transformation in the gonads was
obtainable, either from male to female, or
from female to male.
Berkowitz (1937, 1938, 1941), in work on
the guppy, mentioned several hormones and
combinations of these which were adminis-
tered, and combined the results into a gen-
eral statement. It is possible, therefore, that
the divergent results of one or more of the
hormones given by him went unnoticed be-
cause of this procedure. Although the results
appear to be consistent, no mention is made
of exactly which results were occasioned by
which hormone and slight differences which
might have led to a suspicion of the present
findings might have been overlooked. A re-
230
Zooloyica: New York Zoological Society
grettable tendency evident in many papers is
to administer “estrogens” without regard to
which estrogen is being administered. The
present work indicates that such a procedure
is not safe.
Essenberg in 1923 stated that the oviduct
in Xiphophorus was derived from a fusion of
the two embryonic components of the ovary
in. such a way as to leave a space between
them which later developed into the oviduct.
Wolf (1931), on the other hand, who worked
out the embryology of the gonads in the
platyfish, stated that the oviduct originated
by what may be considered the classical
method, the degeneration of the medulla of
each embryonic gonad accompanied by the
development of the cortex (Willier, 1939).
Goodrich et al. (1934) found that the oviduct
of the guppy originated in the same way.
Two such opposing views in two forms which
are comparatively closely related seems to
be unusual. Evidence for the double origin
of the oviduct, in which the duct develops in
two parts, one in each embryonic ovary, fusing
to form one duct when the ovary itself fuses
(Wolf’s version), is given by the occurrence
of the degenerate ovary under androgen
treatment found in this work which possessed
two distinct ducts. Such a condition, under
the terms of Essenberg’s hypothesis, would
be unlikely. A further investigation into the
origin of the oviduct in Xiphophorus would
seem to be in order.
Regnier (1938), in her description of the
origin of the oviduct in Xiphophorus, quoted
Essenberg, but since this phase of her work
was a review of the literature, no further evi-
dence was to be found there. Regnier men-
tioned the effects of testis powder as produc-
ing bi-lobed and retarded ovaries in Xipho-
phorus when these animals were treated
when very young. She also mentioned the
comparatively great mortality present when
this treatment was given, but said that with
the addition of anterior pituitary lobe pow-
der to the water in which treated individuals
were placed, the mortality markedly de-
creased. After injections of testosterone for
two months, her animals showed mature
sperm in the testes, but no mention is made
of presence or absence of spermatophores.
Therefore it is not known whether the
treated males in that group were fertile. She
also discussed sex reversal due to hormones
and the prevention of sex reversal by injec-
tions of appropriate hormones, but since it is
known that the sex determining mechanisms
of Xiphophorus are somewhat labile (Essen-
berg, 1926; Witschi, 1939) , these results are
not inconsistent. Mention was made of cer-
tain residual bodies which were derived from
the degenerating follicles of the sex revers-
ing ovaries and which traveled to nearby or-
gans where they established themselves. Al-
though evidence of such bodies was sought in
the surrounding organs of the fish in the
present work, no results were obtained.
Cohen in 1942 andt 1946 treated female
platyfish with pregneninolone and males with
estradiol benzoate. He found at that time
that estradiol benzoate had feminizing ef-
fects on the male platyfish over a twelve-week
period. The other results produced were sim-
ilar to those found in the present experiments
within the time limits used here, except that
Cohen showed evidence that mature ova were
found in normal control ovaries of fishes
only eight weeks old. In the entire group of
control females used in the present work,
only one such ovary was found. This lack of
yolk-filled eggs in the ovaries was not con-
sidered unusual, since, although growth rates
vary with environment, feeding and other
factors, Platypoecilus, even under ideal con-
ditions, does not usually mature until the end
of the fourth month after birth or later, as
will be shown. Under normal conditions, no
mature ova would be expected to occur until
or just before that time. The effects of preg-
neninolone which were repeated in the pres-
ent experiments were in the main more pro-
nounced than those shown in Cohen’s work,
probably because of the larger amount of
hormone actually introduced into the fish as
a result of the different method of adminis-
tration used here. It is believed that this
method has been more effective, since the
main portion of the hormone was introduced
into the fish orally. However, the experiments
run subsequently show that some of the drug
was dissolved into the water, either during
the time when the food lay at the bottom of
the tank or after it was egested or excreted
by the fish in a still potent state. That these
drugs affected the fish within a short time,
through whatever means they became dis-
solved, is also evident. The evidence brought
out by the later experiment showed, however,
that the hormones are not stable under
aquarium conditions for more than about
three weeks, since after that time immature
fish introduced into the tanks with the same
water showed no effects whatsoever. Whether
the hormone was destroyed by the micro-
scopic population of the tank, adsorbed to
the glass, or Otherwise inactivated in some
way is not known, but after that time it was
no longer present in a form which had any
perceivable effect on the fish. Further work
is being done to determine the exact time
when this inactivation takes place, and also,
if possible, what the cause for the inactiva-
tion may be.
As to the effects of estradiol benzoate on
the male, Cohen showed no figures on the de-
velopment of either the control or the treated
testes for eight-week-old fish, and therefore
it is difficult to compare results at that age.
In the present work, however, the testes so
treated were slightly retarded in differentia-
tion though not in size because of the admin-
istration of the hormone. Whether these ef-
fects are similar to those found by Cohen for
an eight-week period is difficult to judge, be-
cause his descriptions did not cover that
period.
Some support for the theory that different
esters of the same hormone may bring about
1949]
Tavolga: Differential Effects of Sex Hormones on Platyfish
231
different effects was given by Grobstein
(1942b) , when he found that different esters
of testosterone may show different effects on
the regenerating anal fin of the platyfish.
Even this paper, though, showed that all the
esters used produced masculinization of the
fin, as might have been expected. That a hor-
mone and its ester should bring about dia-
metrically opposed effects is unique. Grob-
stein also showed that the effects of these
hormones is not to produce a normal gono-
podium, but one that is imperfect. That
evidence is substantiated here. In all cases
there was produced a fin which was not pre-
cisely like the typical male gonopodium as it
is seen in a normal adult animal. Even in
those cases where the differentiated parts
appeared to be almost normal, two differ-
ences in size were noted. The fin as a whole
was smaller than the normal, and within this
smaller fin the proportions existing between
the length and width of the fin were changed.
The 3, 4, 5 ray complex in each smaller fin
was approximately one-third shorter than
would normally have been found in a fin of
the same width. The cause of this difference
is apparently the result of differentiation of
the fin beginning before it had time to grow
to its full length, because of the relatively
rapid action of the hormone. In the normal
fish, the testis develops more slowly and
therefore apparently controls the fin in such
a way as to produce a lower amount of hor-
mone until the fin has reached its maximum
length, at which time the testis releases more
hormone and differentiation takes place. This
theory of hormone levels controlling the
growth and differentiation patterns was pos-
tulated by Turner (1941b) and was adopted
by Grobstein.
It should be established that under normal
developmental conditions the young fish in-
volved in these experiments would not have
matured for about two months after the
termination of the treatment, since they
mature at approximately four to six months
of age. Littermates of the experimental an-
imals matured under conditions equivalent
to those used in the experiments within these
time limits, and averaged five months from
birth, to sexual maturity.
It can be assumed that a testis may be con-
sidered functional when it is producing
spermatophores. Although no correlation
has been found as yet to support this assump-
tion in hormone-treated animals, it is always
found that a normal functional male posses-
ses spermatophores, while a non-functional
male, otherwise normal, or an immature
male, does not. Because of this evidence, it is
assumed that the testes of the animals
treated with pregneninolone, and the larger
animals treated with estradiol were function-
al. Even if free sperm are produced under ex-
perimental conditions, the fish will not be sex-
ually functional because of the necessity for
transferring the sperm in a clump from the
gonopodium of the mqle to the vent of the
female. If this transfer is not carried out
by way of the apermatophore, the sperm
will presumably be lost in the water and fer-
tilization will not result. Therefore, the im-
portant feature of the pregneninolone-
treated testes was the large number of
spermatophores present in both the acini and
the duct. Since the normal testis at this age
shows none of these features, the indication
is that a great stimulation had occurred. An-
other feature to be mentioned is the differ-
ence in reaction of fish of the same age and
size to the two hormones which produced
stimulation of the testes. In the case of preg-
neninolone, the stimulation was a steady one,
producing in every fish some sign of stimula-
tion, the amount of growth and differentia-
tion depending on the size and age of the fish.
It was, however, never completely without
effect. This may be seen from the sizes of
the testes shown in Text-fig. 2 and Table II.
Alpha estradiol, on the other hand, produces
quite a different effect. In all the small fish,
those below and including 18 mm., the effect
was negligible. The testes appeared like nor-
mal control testes of the same age. When,
however, the fish reached the size of 19 mm.,
the effect was different. The testes of the
fishes of this size were immediately and
greatly stimulated (see PI. I, Figs. 4 and 5),
and the testes resulting appeared to be func-
tional, considering the great number of
spermatophores present in the acini and
duct.
As to the difference in size of the sperma-
togenic cysts present in the two types of
treated animals, it is possible that the sud-
den arrival at a threshold level of hormone
in the case of estradiol was responsible for
a rapid differentiation of the gland, causing
the smaller size of the spermatogenic ele-
ments. The pregneninolone-treated animals,
which received a longer and steadier stimu-
lation, were capable of producing cysts
which were larger than those normally seen
(see Text-fig. 3).
In order to suggest an explanation for the
above effects and the others found in the
present work, several assumptions must be
made. First, it is well known that the liver
of mammals inactivates steroid hormones
which pass through the portal circulation
(Biskind and Mark, 1939; Burrill and
Greene, 1942; Cantarow et al., 1943 ; Heller,
1940; Israel et al., 1937; Segaloff, 1943; Tal-
bot, 1939; Teague, 1941; Westerfeld, 1940).
It is assumed that the same action takes place
in the liver of teleosts. Some hormones, how-
ever, are inactivated more than others. Es-
tradiol is inactivated more than estradiol
benzoate because the benzoate ester protects
the molecule from destruction. According to
Heller (1940), the oxidation of the estradiols
takes place at carbon 3 in ring A. Since the
benzoate radical is attached at this position,
its presence protects the molecule from oxi-
dation (Segaloff, 1943). Therefore it can be
assumed that the effective dose of estradiol,
that is, the dose which produces the effects in
the animal, is less than, the effective dose of
232
Zoologica: New York Zoological Society
[34: 18
estradiol benzoate, if identical oral doses are
given.
The toxicity of the hormones must also be
taken into consideration. Plate V, Fig. 23,
shows the appearance of the typical liver of
an animal treated with pregneninolone. The
cells are greatly enlarged and vacuolated and
are presumably in a condition caused by the
relatively great toxicity of the pregnenin-
olone, which may be interpreted as a type of
fatty change. Because of this toxicity, the
liver, which at first probably rapidly inacti-
vated the hormone, was rendered unable to
do so, and the main portion of the hormone
passed through the liver intact, producing a
large effective dose and intense effects. The
estradiol, which is partially inactivated,
causes also a partial vacuolization of the
liver, suggesting a cumulative effect on the
liver, which results in an increase in effective
dose. This eventually has an effect on the
gonads.
If these hypotheses are true, they present
new evidence for the action of abnormal
quantities of metabolic substances on the
liver, since till now the only concusive evi-
dence for the inhibition of inactivation by
the liver has been derived from work on ex-
perimental Vitamin B deficiency (Biskind
and Biskind, 1941, 1942, 1943: Biskind,
1946).
A further assumption concerns the stage
of growth and differentiation in which the
gonads are found during the period of the
experiment. During this period the testes
are, for the purposes of this explanation, in a
relatively undifferentiated state and not yet
under the influence of the pituitary. Gonado-
trophic hormones are known to be present in
fishes (Scheer, 1948). There is evidence to
support the above assumptions. The testes, as
shown by Text-fig. 2, grow very little during
the period of the experiment. They contain
essentially the same elements at the end of
eight weeks as they possessed about one week
after birth. The ovaries, on the other hand,
grow considerably during the same period,
and yolk deposition is begun and progresses
considerably. The ovaries and eggs are much
larger at the end of the period than they
were at one week of age. The growth of the
gonads is known to be under the control of
the pituitary (Matthews, 1939a).
If these assumptions are admitted, at least
as possibilities, a hypothesis may be ad-
vanced as to the method by which the hor-
mones produce their results in these experi-
ments.
In the case of the testis, the first effective
doses of pregneninolone were small because
the substances were largely inactivated by
the liver tissue. These relatively small doses
stimulated the pituitary rather than inhib-
ited it because of the smallness of the dose.
The estradiol had a delayed effect because it
continued to be inactivated for a longer pe-
riod of time, and therefore needed a longer
period of time in which to reach an effective
dose. The dose which was effective in the case
of estradiol was a cumulative one and re-
quired a longer period in which to operate
and a larger animal on which to operate be-
cause of some type of threshold reaction. The
estradioi benzoate went through the liver tis-
tue undestroyed and reached the pituitary
in doses large enough to cause an inhibition
rather than a stimulatory effect. Thus the
testis, which was not yet under pituitary
control, showed little effect from the admin-
istration of this drug.
In the case of the ovaries, which were al-
ready under pituitary control, the effects
were different. The smaller doses of estradiol
and pregneninolone acted as partial inhi-
bitors, shown by the partial inhibition of
the eggs in these specimens, while the estra-
diol benzoate, again passing through the liver
undestroyed, caused an almost complete in-
hibition of growth of the eggs.
To suggest an explanation for the action
on the gonopodium is a more difficult problem.
In both males and females, the effect on the
gonopodium was similar. Pregneninolone
stimulated at least some growth in all gono-
podia, and all older animals treated for a
longer period of time showed almost perfect
transformation of the fin. Estradiol stimu-
lated all gonopodia to a slight growth, and
the largest ones to the same type of differen-
tiation shown by the pregneninolone animals,
though the differentiation was slightly less
advanced. Estradiol benzoate had no effect
on any of the animals. There are a number
of hypotheses which may be advanced.
First, the gonopodium might be under
purely genetic control. It is known that this
is not true because the treated females
showed differentiation to a gonopodium as
readily as did the males.
Second, the ovarian hormone might in-
hibit the gonopodium. If we can assume that
an inhibited ovary is producing little or no
hormone, the above hypothesis cannot be
true because under these conditions a great-
ly inhibited ovary would allow a better dif-
ferentiated gonopodium than a partially in-
hibited one. The estradiol benzoate-treated
ovary was inhibited to the greatest degree,
but there was no gonopodium, while the ani-
mals which possessed partially inhibited
ovaries formed well differentiated gonopodia.
Third, the reactions cannot be due to a
non-specific reaction to steroids because the
different substances produced different ef-
fects.
Fourth, if the reactions are due to the ac-
tion of the fish testis hormone, or to an an-
drogenic effect directly, it must be hypothe-
sized that estradiol has a direct androgenic
effect, while an effect based on dosage differ-
ence would be more plausible, since in mam-
mals the substance has an estrogenic effect.
Fifth, control from the pituitary gland en-
tirely could explain the effects in the males,
where pregneninolone and estradiol stimu-
late the pituitary. In the females, however,
the pituitary, according to the above assump-
tions, and based on its action on the gonads,
1949]
Tavolga: Differential Effects of Sex Hormones on Platyfish
233
inhibits the ovaries, and presumably would
not at the same time stimulate the differen-
tiation of a gonopodium. This of course as-
sumes that the gonadotrophins secreted by
both male and female pituitaries are quali-
tatively identical and stimulate the gonads
of the animals in which they exist. This has
been shown to be true for amphibians (Rugh,
1935) .
Sixth, the theory that pituitary control
plus male gonads or androgenic hormone
cause the effects is the most nearly complete
explanation. In this case, pregneninolone and
estradiol stimulate the pituitary and there-
fore stimulate the gonopodium through the
gonad. Estradiol benzoate inhibits the pitui-
tary. Since the testis is not as yet under pi-
tuitary control, the testis shows no effects.
No androgenic hormones are produced and
the lack of these produces, in turn, lack of
a gonopodium. In the females, however, an
androgenic effect of the substances admin-
istered is necessary to explain the results.
Pregneninolone and estradiol inhibit the
pituitary and through it inhibit the ovary.
The pituitary inhibition plus the androgenic
effects of the hormones cause the differentia-
tion of the gonopodium. Estradiol benzoate
inhibits the pituitary, but, having no andro-
genic effect, does not cause the formation of
the gonopodium.
A detailed cellular examination of the pitu-
itary gland in these fish may reveal signifi-
cant differences between controls and experi-
mental animals, presumably involving the
cells which secrete gonadotrophic hormones.
This may furnish a partial explanation for
the results described above and indicate
whether the action may take place through
the pituitary or is a direct effect of the hor-
mones upon the gonads, as has been shown to
happen in other animals (Nelson, 1937). A
careful examination of the interstitial tissue
of the testes may also aid in determining the
possible effects of the hormones upon this
tissue.
These hypotheses were constructed in an
attempt to correlate the actions of the vari-
ous hormones on the gonads and on the anal
fin. Perhaps the effects on the two are entire-
ly separate, however. The effect of the pitui-
tary may be brought in to account for the ef-
fects on the gonads, but an androgenic ef-
fect of estradiol and pregneninolone would
account alone for the effects on the gonopo-
dium. It cannot be assumed, however, in
view of the evidence brought out by estradiol
benzoate treatment, that the effect is the
paradoxical estrogen effect mentioned above
caused by high dosage with estrogens. The
effective dose of estradiol benzoate is higher
than that of the others because it is protected
in the liver. Therefore, under these condi-
tions, one would be led to expect that it would
produce a more definite effect than either of
the other hormones. Since this is not true,
some other hypothesis must be advanced to
explain this effect. The other possibility
which is most plausible is one in which the
hormone has a directly androgenic effect.
The fact that the hormones which were
used produced uniform results in spite of
more than a tenfold range in dosage is an un-
usual finding. The toxicity which was found
to be present with large dosages has also been
found in mammals, but no sub-maximal re-
sults were found here.
It might have been useful also to treat the
fishes with other benzoates as a control for
the possible action of the benzoate ester ex-
clusive of estradiol. The use of an inactive
free compound with an active benzoate ester
would be helpful in this work.
Turner (1941 a and b) brought to light va-
rious factors affecting the growth of the go-
nopodium. He stated that, first, the growth of
the gonopodium depended on a certain low
concentration of hormone and the differen-
tiation of the fin depended on a higher con-
centration ; second, that there existed certain
dominances in the ray complexes which gov-
erned the differential growth of the various
rays in such a way as to produce what we
know as a complete gonopodium if the fin is
left undisturbed; and third, that castration
at any time during the growth of the gono-
podium would stop its growth, while the ad-
ministration of androgens thereafter would
renew its growth. These findings on Gambu-
sia have important bearings on the present
studies. It was suggested above that the
effect of estradiol was a cumulative one. This
might account, on the basis of Turner’s first
statement, for the anal fin growth shown by
the smaller estradiol-treated animals, where
no differentiation was present. Pregnenino-
lone caused an immediate and sustained
effect on the gonopodium, suggesting that
this hormone reached the threshold level al-
most immediately. Such a hypothesis would
aid in explaining the effects on both testes
and gonopodia.
As to the effect on the females, the hor-
mones, as suggested above, may have had a
direct effect on the fins.
The differential growth of the 3, 4, 5 ray
complex was apparently governed by a low
concentration of hormone. Dominance then
shifted, according to Turner, so that the rays
outside the 3, 4, 5 complex were subordinated
to these three. This might explain the rapid
growth of these rays in the young estradiol
males and others in which a low level of hor-
mone existed.
Castration with the effect of termination
of growth of the fin, followed by androgenic
restimulation, shows that the testis itself is
not necessary for the growth of the fin, but
that a hormone similar to that produced by
the testis is required. This might aid in ex-
plaining the cases in which the females
grew well-formed gonopodia.
The above hypotheses are far from clear
and more work must be done in order to de-
termine the explanation for these seemingly
opposite and confusing effects. Hypophysec-
234
Zoologica: New York
tomy, castration and a combination of the
two performed on animals which were later
treated with hormones would aid in deter-
mining the mechanisms which govern these
effects. Preparations for such work are going
on now.
More work is necessary on the general prob-
lem of the differential effects of these two
estradiol compounds. The exact stage when
the differential effect on the male begins
should be studied in more detail. Smaller
dosages should be used in an attempt to dis-
cover a dose small enough to secure a less
than maximum effect, as such an effect does
not appear in the present work. Finally, an
investigation into the differential effects of
more compounds related to these should be
carried out, since the exact effect of any one
of them is now doubtful, whereas heretofore
they have been used interchangeably, at least
on experimental animals.
Summary.
1. The experimental animal used was a
strain of the platyfish in which males could
be distinguished from females at birth as a
result of a Y chromosome sex linked, spotted
factor, whereas usually the sexes are indis-
tinguishable until maturity, when the anal
fin of the male is transformed into an intro-
mittent gonopodium.
2. The hormones used were alpha estra-
diol, alpha estradiol benzoate and pregneni-
nolone, a synthetic progestogen. These were
administered by mixing the powder or oil
solution with the food.
3. Pregneninolone exhibited a strong
stimulating effect on the males, with preco-
cious maturation of the testes and well-
formed gonopodia. In females, development
of the ovaries was inhibited, and gonopodia
produced.
4. Estradiol benzoate was inhibitory on
the testis and greatly so on the ovary. No
gonopodia were produced.
5. Alpha estradiol had no effect on the
testes of males under 18 mm. in standard
length and produced slight growth of the
anal fin. In males over 19 mm. in length, the
testes were greatly stimulated and large,
well-formed gonopodia were found. All fe-
males so treated showed ovarian degenera-
tion and partial to nearly complete gono-
podia.
6. Studies on the liver showed that preg-
neninolone and estradiol produced great vac-
uolization of the parenchyma cells and re-
sulted in an organ which showed fatty
changes, while the benzoate-treated livers
appeared like those of the controls.
7. It is to be emphasized that although in
the amniotes, estradiol and its ester are used
interchangeably, in this species the two com-
pounds produce diametrically opposed effects
under the conditions of these experiments.
Zoological Society [34: 18
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Tavolga: Differential Effects of Sex Hormones on Platyfish
236
Zoologica : New York Zoological Society
[34: 18
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1949]
Tavolga: Differential Effects of Sex Hormones on Platyfish.
237
EXPLANATION OF THE PLATES.
Plate I.
Fig. 1. Testis of control male eight weeks of
age. X100.
Fig. 2. Testis of estradiol benzoate-treated
male eight weeks of age. X100.
Fig. 3. Testis of pregneninolone-treated male
eight weeks of age. X100.
Fig. 4. Testis of estradiol-treated male eight
weeks of age. X100. Compare with Fig.
3.
Fig. 5. Testis of estradiol-treated male eight
weeks of age. X100. Compare with
Figs. 3 and 4.
Plate II.
Fig. 6. Ovary of control female eight weeks of
age. X100.
Fig. 7. Ovary of estradiol benzoate-treated fe-
male eight weeks of age. X100.
Fig. 8. Ovary of pregneninolone-treated fe-
male eight weeks of age. X100.
Fig. 9. Ovary of pregneninolone-treated fe-
male eight weeks of age. X100. Note
scattering.
Fig. 10. Ovary of pregneninolone-treated fe-
male eight weeks of age. X100. Note
bilobed appearance of organ.
Fig. 11. Ovary of estradiol-treated female eight
weeks of age. X100. Compare with
Fig. 9.
Plate III.
Fig. 12. Ovary of estradiol-treated female eight
weeks of age. X100. Note large degen-
erating eggs and small abnormal eggs.
Fig. 13. Anal fin of control male. X 34.
Fig. 14. Anal fin of estradiol benzoate-treated
male. X34. Compare with Fig. 13.
Fig. 15. Anal fin of pregneninolone-treated
male. X34. Note almost complete dif-
ferentiation of gonopodium. Compare
with Fig. 13.
Fig. 16. Anal fin of estradiol-treated male. X34.
Compare with Figs. 13 and 15.
Plate IV.
Fig. 17. Anal fin of control female. X 34.
Fig. 18. Anal fin of estradiol benzoate-treated
female. X34. Compare with Fig.17.
Fig. 19. Anal fin of pregneninolone-treated fe-
male. X34. Compare with Figs. 15 and
17.
Fig. 20. Anal fin of estradiol-treated female.
X34. Compare with Figs. 16, 17 and 19.
Plate V.
Fig. 21. Liver of control animal. X960.
Fig. 22. Liver of estradiol benzoate-treated ani-
mal. X960. Note similarity to Fig. 21.
Fig. 23. Liver of pregneninolone-treated ani-
mal. X960. Note extensive vacuoliza-
tion. Compare with Fig. 21.
Fig. 24. Liver of estradiol-treated animal.
X960. Note vacuolization — approach-
ing but not equalling that of Fig. 23.
' 'I
VOLGA.
PLATE I.
FIG. 4. FIG. 5.
DIFFERENTIAL EFFECTS OF ESTRADIOL. ESTRADIOL BENZOATE
AND PREGNENINOLONE ON PLATYPOECILUS MACULATUS.
VOLGA.
PLATE II.
FIG. 10.
FIG. 7.
FIG. 8.
FIG. 11.
FIG. 6.
FIG. 9.
DIFFERENTIAL EFFECTS OF ESTRADIOL. ESTRADIOL BENZOATE
AND PREGNENINOLONE ON PLATYPOECILUS MACULATUS.
VOLGA.
PLATE III.
FIG. 12.
FIG. 13.
FIG. 14.
FIG. 15.
FIG. 16.
DIFFERENTIAL EFFECTS OF ESTRADIOL. ESTRADIOL BENZOATE
AND PREGNENINOLONE ON PLATYPOECILUS MACULATUS.
DIFFERENTIAL EFFECTS OF ESTRADIOL, ESTRADIOL BENZOATE
AND PREGNENINOLONE ON PLATYPOECILUS MACULATUS.
/OLGA.
PLATE V.
FIG. 21.
FIG. 23.
FIG. 22.
FIG. 24.
DIFFERENTIAL EFFECTS OF ESTRADIOL. ESTRADIOL BENZOATE
AND PREGNENINOLONE ON PLATYPOECILUS MACULATUS.
Hertlein & Strong : Mollusks of Mexico and Central America
239
19.
Eastern Pacific Expeditions of the New York Zoological Society. XLI.
Mollusks from the West Coast of Mexico and Central America. Part VIII.1
Leo George Hertlein & A. M. Strong.
California Academy of Sciences.
(Plate I).
[This is the forty-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 Templeton Crocker Ex-
pedition (1936) and the Eastern Pacific Zaca
Expedition (1937-1938). For data on localities,
dates, dredges, etc., refer to Zooloqica, Vol.
XXII, No. 2, pp. 33-46, and Vol. XXIII, No. 14,
pp. 287-298.]
Contents.
Page
Introduction 239
Family Semelidae 239
Genus Semele Schumacher 239
Semele corrugata californica Reeve 240
Semele craneana Hertlein & Strong, sp. nov 241
Semele decisa Conrad 242
Semele flavescens Gould 242
Semele guaymasensis Pilsbry & Lowe 243
Semele jaramija Pilsbry & Olsson 244
Semele jovis Reev« 244
Semele laevis Sowerby 245
Semele pacifica Dali 245
Semele pulchra Sowerby 246
Semele quentinensis Dali 246
Semele simplicissima Pilsbry & Lowe 247
Semele sparsilineata Dali 247
Semele tabogensis Pilsbry & Lowe 248
Semele venusta Reeve 248
Semele verrucosa Morch 249
Genus Abra Lamarck 249
A bra palmeri Dali 250
Genus Cumingia Sowerby 250
Cumingia lamellosa Sowerby 250
Family Donacidae 251
Genus Donax Linnaeus 251
Donax asper Hanley 251
Donax assimilis Hanley 252
Donax calif ornicus Conrad 252
Donax carinatus Hanley 253
Donax gracilis Hanley 253
Donax navicula Hanley 254
Donax obesus d’Orbigny 254
Donax punctatostriatus Hanley 255
Donax transversus Sowerby 266
Genws lphigenia Schumacher 257
Iphigenia altior Sowerby 257
Introduction
This is the eighth of a series of papers
dealing with collections of mollusks taken
on the Templeton Crocker Expedition
(1936) and the Eastern Pacific Zaca Expe-
dition (1937-1938). The general plan of
presentation followed in the present con-
1 Contribution No. 859, Department of Tropical Research,
New York Zoological Society.
2 Hertlein, L. G-, and Strong, A. M. Eastern Pacific Ex-
peditions of the New York Zoological Society. XXIII.
Mollusks from the West Coast of Mexico and Central
America. Part II. Zoologica, New York Zool. Soc., Vol. 28,
Pt. 3, December 6, 1943, pp. 149-168, 1 pi. See especially
pp. 149-160.
tribution is that mentioned in Part II of
this series of papers2. Formal headings
and keys are given for the species collected
by the expeditions of 1936 and 1937-1938.
Occasionally additional species are included
in the keys for convenience but in such cases
it is indicated which species do not occur
in the present collection.
Acknowledgment is due Dr. G. D. Hanna,
Curator, Department of Paleontology of
the California Academy of Sciences, Mr. A.
G. Smith, Research Associate of the same
institution, and Dr. A. Myra Keen, Stanford
University, California, for assistance and
suggestions. The photographs used for il-
lustrations on the plate were prepared by
Mr. Frank L. Rogers.
Family Semelidae.
A paper by Dali3 dealing with the west
American Semelidae and one by Lamy4
which contains a revision of the species of
this group in the Natural History Museum
in Paris are useful to anyone studying the
Recent west American forms of this family.
Key to the Genera of the Semelidae.
A. Pallial sinus free from the pallial line
Semele
B. Pallial sinus confluent with the pallial line
a. Length exceeding 10 mm.; moderate-
ly thick Cumingia
aa. Length rarely exceeding 10 mm.;
very thin; usually smooth Abra
Genus Semele Schumacher.
Key to the Species of Semele.
A. Concentrically sculptured with coarsely
corrugated or wrinkle-ribbed ridges
a. Exterior with radial striae or
wrinkled and /granulated sculpture
b. Interior white; dorsal margin
purple; truncated posteriorly
decisa
3 Dali, W. H. Notes on the Semelidae of the West Coast
of America, including some new species. Proc. Acad. Nat.
Sci. Philadelphia, Vol. 67, Issued March 2, 1916, pp. 25-28.
4 Lamy, E. Revision des Scrobiculariidae vivants du
Museum d’Histoire Naturelle de Paris. Journ. de Conchyl.,
Vol. 61, No. 3, March 25, 1914, p. 243-368, pi. 8, also figs,
in text.
240
Zoologica: New York Zoological Society
[34: 19
bb. Interior brown, orange or yellow
c. Interior brown; corrugated
tabogensis
cc. Interior yellow or orange ;
wrinkle-ribbed exteriorly
calif omica
aa. Exterior without radial sculpture
craneana
B. Concentrically sculptured with raised
threads, lamellae, or growth lines only
a. Concentric sculpture of growth lines
only
b. Diagonal striae present
sparsilineata
bb. Diagonal striae absent; smooth,
pure white laevis
aa. Concentric sculpture of raised threads
or lamellae
c. Escutcheon wide, strongly bev-
eled flavescens
cc. Escutcheon very narrow or
lacking
d. Shell without radial sculp-
ture; white
e. Interspaces with fine
concentric striae
pazianau
ee. Interspaces without fine
concentric striae
simplicissima
dd. Shell with radial sculpture
f. Shell with incised radial
grooves on part or all of
valves
g. Incised sculpture
along anterior dor-
sal margin only
h. Concentric ribs
closely spaced,
fine
i. Shell thick,
fairly high;
interior with a
deep purple
blotch pulchra
ii. Shell thinner,
more elon-
gate ; less pur-
ple coloration
quentinensis
hh. Concentric ribs
more widely
spaced, coarse
guaymasensis
gg. Incised sculpture
present along anter-
ior and posterior
margins
j. Concentric rib-
bing fine
far ami ja
jj. Concentric rib-
bing coarse;
entire valve
often reticu-
lately sculp-
tured pacifica
if. Shell without incised ra-
dial grooves; fine radial
striae present, stronger
in interspaces
k. Shell suborbicular
or roundly ovate
l. brownish - red
with white me-
dial streak on
umbos; large
jovis
11. Yellowish or
white
mediamericana:>
kk. Shell elongate
m. Concentric la-
mellae high;
shell large;
white with pur-
ple spots
n.L a m e 1 1 a e
with scal-
loped scale-
like projec-
tions
verrucosa
nn. Lamellae
without
scale-like
projec-
tions; more
rounded
ventrally
formosa 5
mm. Concentric la-
mellae low;
shell small ;
umbos pur-
p 1 i s h or
brownish
o. T h i c k ;
end of
p a 1 1 i a 1
sinus
slightly
atten-
uated
venusta
oo. T h i n ;
end of
pallial
sinus
blunt
incon-
gruan
Semefe corrugate callfornlca Reeve.
Amphidesma calif omica Reeve, Conch.
Icon., Vol. 8, Amphidesma, October, 1853,
species 19, pi. 3, fig. 19. “Gulf of California.'’
s Not represented In the present collection.
1949]
Hertlein & Strong : Mollusks of Mexico and Central America
241
Semele calif ornica A. Adams, Proc. Zool.
Soc. London for 1853 (issued July 25, 1854),
p. 96. “Hab. Gulf of California. Mus. Cum-
ing.”
Type Locality: Gulf of California.
Range : Magdalena Bay, Lower California,
to the Gulf of California.
Collecting Station: Mexico: Cape San
Lucas, Lower California, beach.
Description: Shell ovately oblong, sub-
equilateral, dirty pale brown, radiately stri-
ated; transversely sulcated, ribs elevated,
subcorrugated, ornamented, interspaces
closely longitudinally striated ; anterior side
rounded, posterior subtruncated, very flexu-
ous; interior yellowish, margin yellow. (Free
translation of Adams’ original description.)
The shell of this subspecies is character-
ized by the concentric wrinkle-ridged ribs
and the dense minute radial striae. A small
lunule is present but no appreciable escutch-
eon. Exteriorly the shells are usually yel-
lowish or yellowish-white in color. The um-
bos are often yellow and on some specimens
a few faint brown transverse markings are
present on the dorsal margin both anterior
to and posterior to the beaks. Interiorly the
shells are usually yellow, sometimes a beau-
tiful golden or orange-yellow. The pallial si-
nus ascends gently and is rounded at the end
which extends forward a little over one-half
the length of the shell.
A left valve from Cape San Lucas, in the
present collection, measures: length, 40.2
mm. ; height, 36 mm. ; convexity (one valve),
7.3 mm. A specimen from Magdalena Bay,
Lower California, in the Henry Hemphill Col-
lection of the California Academy of Sciences
measures: length, 37.5 mm.; height, 33.6
mm. ; convexity (both valves together) , 15.2
mm. ; pallial sinus extends anteriorly 22 mm.
from the posterior margin of the shell.
This form apparently is, as stated by Dali,
closely related to Semele corrugata Sowerby6.
The specimens which we have seen from
Magdalena Bay and the Gulf of California
appear to be a little smaller than those of S.
corrugata Sowerby which was described
from Peru. Furthermore these do not have
8 Amphidesma corrugotum Sowerby. Concb. Tllustr..
Catal. issued with Pt. 19. species No. R, issued between
January 18 and March 8. 1833. TNot illustrated]. “Tnuioui.
Peru. Mr. Cuming.” Ref. to “Spec. Conch, f. 18.” TThe
exact dates of issue of the Species Conchyliorum is unknown
to ns. A copy of Volume 1, Part 2, in the librarv of the
California Academv of Sciences is not dated. In Hanley’s
edition of Wood’s Index Testaceolopicus. 1856, p. XIX, the
dates cited for Sowerbv Species Conchyliorum are, "part 1.
1830: part IT. (imperfect) not published until Nov. 1855”!.
—Sowerby. Proc. Zool. Soc. London for 1832. issued March
13, 1833, p. 200. "Hab. in Peruvia et ad Iquiqui.” “Dredged
from coarse gravel in ten fathoms water.”— Reeve, Conch.
Icon., Vol. 8. Amphidesma, October, 1853, species 4, pi. 1,
fig. 4 (as Amphidesma corrugata). [Locality same as in
preceding reference].
Shaw (Proc. Malacol. Soc. London, Vol. 8, No. 6. 1909,
pp. 333-340), in a collation of the Conchological Illustra-
tions, indicated that Parts 17, 18, 19, and the catalogue of
species of Amphidesma issued with Part 19, appeared
between January 18 and March 8, 1833. The species of
Amphidesma named and illustrated in those parts for the
first time take their date of publication from the Concho-
logical Illustrations rather than the Proceedings of the
Zoological Society of London where the descriptions ap-
peared on March 13, 1833.
the purple coloration on the anterior portion
of the hinge as shown on Reeve’s illustration
of Semele corrugata. That species has been
recorded as occurring at Magdalena Bay and
in the Gulf of California, and it is possible
that the present specimen might be referable
to Sowerby’s species. However, for the pres-
ent at least, we are inclined to regard these
northern shells as belonging to a subspecies
of S. corrugata, at least until a comparison
can be made with a series of specimens from
Peru, the type locality of Sowerby’s species.
It appears that in some cases, Semele cor-
rugata calif ornica has been confused with
Semele flavescens Gould, a different shell.
Distribution: A single valve of this sub-
species was taken by the expedition on the
beach at Cape San Lucas. It also occurs in
the Pleistocene of Magdalena Bay, Lower
California. The record “Semele cf. pulchra
Sowerby” in the list of species cited by Jor-
dan, 1936, as occurring in the Pleistocene of
Magdalena Bay, is referable to S. corrugata
calif ornica. Olsson has recorded “Semele cf.
calif ornica Con.” as occurring in the Pleisto-
cene of Panama. Records of the occurrence
of this shell in Asiatic seas are referable to
some other species.
Semele e raneana Hertlein & Strong, sp. nov.
Plate I, Figs. 19, 22.
Shell oval, compressed, thin, with the beaks
a little nearer the posterior end, yellowish,
with faint, pinkish, interrupted radial
stripes; posterior dorsal margin sloping,
slightly convex, forming a distinct angle with
the truncated posterior end, anterior dor-
sal margin more direct, slightly concave,
anterior dorsal margin well rounded; lunule
very small, indistinct; outer surface smooth
near the beaks gradually developing con-
centric ridges which are strongest near
the margins, with deep intersDaces which
about equal the ridges in width; posterior
end with a depression running from near the
beaks to the lower end of the truncation, pos-
terior to which the shell is flattened and
somewhat flexed; radial sculpture entirely
absent; interior white, somewhat iridescent,
showing the concentric sculDture and pinkish
rays quite distinctly; pallial sinus broad,
ascending, rounded at the end and proiecting
about two-thirds the length of the shell; two
cardinal teeth, the Dosterior one the larger,
lateral teeth small. The type measures:
length, 38 mm.: height, 29 5 mm.; convexity
(one valve), 6.5 mm.; pallial sinus projects
forward 24 mm. from the posterior margin
of the shell.
Holotype, a left valve, (Calif. Acad. Sci.
Paleo. Type Coll.), dredged in the Gulf of
California. One left valve was dredged on
Arena Bank, Gulf of California, Station 136-
D-24, Lat. 23° 29' N., Long. 109°23'30" W., in
50 fathoms (91 meters) , mud, Area conglom-
erate; one young specimen and a single right
valve were dredged in the same general local-
ity, Station 136-D-26, Lat. 23°27' N., Long.
242
Zoologica: New York Zoological Society
[34: 19
109° 24' W., in 45 fathoms (82 meters) , sand,
crushed shell; a single valve was dredged 3
miles off Pyramid Rock, Clarion Island, Sta-
tion 163-D-2, Lat. 114° 45' N., Long. 114° 45'
W., in 55 fathoms (100 meters), rock, coral.
This shell is similar in size and shape to
Semele tabogensis Pilsbry & Lowe. The
sculpture is similar but lacks the fine radial
ornamentation of that species and the pos-
terior area is more distinct.
The shell of Semele craneana, although less
arcuate ventrally, is similar to that of S.
martinii Reeve7 which was originally de-
scribed from Brazil.
This species is named for Miss Jocelyn
Crane, Technical Associate, Department of
Tropical Research, New York Zoological
Society, who accompanied the Templeton
Crocker Expedition, 1936, during the course
of which the type specimen of the present
species was collected.
Distribution: This new species is at pres-
ent known only from the southern portion of
the Gulf of California and from off Clarion
Island, in 45-55 fathoms.
Semele deelsa Conrad.
Amphidesma decisa Conrad, Jour. Acad.
Nat. Sci. Philadelphia, Vol. 7, 1837, p. 239, pi.
19, fig. 2. “Inhabits with the preceding”
[which is “Inhabits deep water in the vicin-
ity of Sta. Diego”.] — Reeve, Conch. Icon..
Vol. 8, Amphidesma, 1853, species 24, pi. 4,
fig. 24. San Diego, California.
Semele decisa Conrad, Grant & Gale, Mem.
San Diego Soc. Nat. Hist., Vol. 1, 1931, p. 376,
pi. 14, figs. 13a, 13b. Earlier records cited.
Pleistocene and Recent.
Type Locality: San Diego, California, in
deep water.
Range: San Pedro, California, to Cape San
Lucas, Lower California.
Collecting Station: Mexico: Cape San
Lucas, Lower California.
Description: Shell rounded, thick, sub-
equilateral, the anterior side the longer, the
end rounded; posterior dorsal margin nearly
straight, the posterior end truncated; pos-
teriorly biangulate, the area between some-
what concave; ornamented with numerous,
thick, unequal concentric rugose ribs, the en-
tire surface covered with fine radial grooves
or fine wrinkled and granulated sculpture;
colored exteriorly by whitish-gray with oc-
casional purple in the concentric grooves;
cardinal teeth obsolete, laterals present; pal-
lial sinus wide, rounded at the end, slightly
ascending and extending forward about five-
eighths the length of the shell which is past
a line vertical with the beaks; interior white
with purple around the dorsal margin.
A specimen from Cape San Lucas, Lower
California, measures: length, 45 mm.;
height, 42.5 mm. ; convexity (both valves to-
gether), 19.8 mm.; pallial sinus extends an-
, i
7 Amphidesma martinii Reeve, Conch. Icon., Vol. 8, Am-
phidesma, November, 1863, species 43, pi. 6, fig. 43. “Hab.
Rio.” [A. Adams’ description of this species did not appear
until July 26, 1864].
teriorly 27 mm. from the posterior margin of
the shell. A large specimen of this species
from Magdalena Bay, Lower California, in
the Henry Hemphill collection of the Cali-
fornia Academy of Sciences measures:
length, 94 mm.; height, 86.5 mm.; convexity
(both valves together), 45.5 mm.; pallial si-
nus extends anteriorly 59 mm. from the pos-
terior margin of the shell.
The shell of Semele punctata Sowerby8,
which was described from the Galapagos
Islands, is more elongate in outline and less
truncated posteriorly than that of S. decisa.
Semele nisigotoensis Nomura & Hatai9, de-
scribed from the Miocene of Japan, was com-
pared to S. decisa.
Distribution: A few specimens of this spe-
cies were collected by the expedition at Cape
San Lucas, Lower California. This is an ex-
tension south of the known range of the spe-
cies. It also has been recorded as occurring
in the Pleistocene of Tomales Bay in Central
California, in southern California, and south
to Magdalena Bay, Lower California.
Semele flavescens Gould.
Amphidesma flavescens Gould, Proc. Bos-
ton Soc. Nat. Hist., Vol. 4, November, 1851,
p. 89. “San Diego, Lieut. Green.” — Gould,
Boston Jour. Nat. Hist., Vol. 6, 1853, p. 392.
Original locality cited.
Amphidesma proximum C. B. Adams, Ann.
Lyceum Nat. Hist. New York, Vol. 5, July,
1852, pp. 513, 547 (separate, pp. 289, 323).
“Habitat. — Panama.” — Hanley, Cat. Rec.
Bivalve Shells, p. 341, 1856, pi. 12, fig. 5, 1843
(cited as Amphidesma corrugatum on expl.
to plate). “Mexico.”
Amphidesma proximo Adams, Reeve,
Conch. Icon., Vol. 8, Amphidesma, 1853, spe-
cies 20, pi. 3, fig. 20. “Hab. Panama.” [A re-
production of this figure given by M. Smith,
Panamic Mar. Shells (Tropical Photogr.
Lab., Winter Park, Florida), 1944, fig. 805].
Semele flavescens Gould, Lamy, Journ. de
Conchyl., Vol. 61, No. 3, 1914, p. 358. Gulf of
California; Panama.
Semele proximo C. B. Adams, Olsson, Nau-
tilus, Vol. 37, No. 4, 1924, p. 129. Zorritos,
Lobitos, Negritos, Peru.
Type Locality: San Diego, California.
Range: Catalina Island, California (Dali),
to Negritos, Peru.
Collecting Station: Costa Rica: Golfito
Bay, Gulf of Dulce.
Description: Shell subrotund, compressed,
sculptured by concentric lamellar decussate
striae, orange becoming white in later stages
and covered with a periostracum which is
8 Amphidesma punctatum Sowerby, Conch. Illnatr.,
Amphidesma, Catal. issued with Pt. 19, No. 18, pi. 18, fig. 7,
issued between January 18 and March 8, 1833. * Galapagos
Islands. Mr. Cuming."— Sowerby, Proc. Zool. Soc. London
for 1832 (issued March 13, 1833), p. 200. “Hab ad Insulas
Gallapagos.”— Reeve, Conch. Icon., Vol. 8, Amphidesma.
October, 1863, species 26, pi. 4, fig. 26 (as Amphidesma
punctata). Galapagos Islands.
9 Semele nisigotoensis Nomura & Hatai, Saito Ho-On
Kai Mus. Res. Bull.. No. 10, 1936, p. 131, pi. 16, figs. 8, 9.
Nisigfito. Tanagura Beds, northwest Honsyu, Japan, middle
Miocene.
1949]
Hertlein & Strong : Mollusks of Mexico and Central America
243
brown shaded with gray (fusco) ; beaks
median, acute, not at all elevated; anterior
dorsal area excavated, posterior lanceolate,
concave bounded by a line; interior tinted
yellow, marked with shiny dots; pallial sinus
spatulate, sculptured by close divergent
striae; ligamental pit deep, elongate; ante-
rior lateral teeth approximate to the beaks.
Long. 2% ; alt. 2% ; lat. 1% poll. (Transla-
tion of Gould’s original description).
“Usually found about half the above size;
the concentric lamellae become worn off and
more irregular towards the margin. The in-
terior is faintly tinted yellow when young,
but very richly so when old. It is near A. cor-
rugatum, Sowb.”
The description given by Gould in 1853 is
an enlargement of the original. He stated:
. . . “posterior dorsal edge long lanceolate,
concave, bounded by a distinct angle ; surface
pale orange near the beaks, becoming dingy
white at the older stages, and covered by a
dirty greenish epidermis; marked by con-
centric lamellar striae, crossed by fine radi-
ating striae, especially across the disk.”
A left valve in the present collection meas-
ures: length, 47.5 mm.; height, 43.8 mm.;
convexity (one valve), 11.2 mm. A specimen
from Loreto, Lower California, in the col-
lections of the California Academy of Sci-
ences measures: length, 58.5 mm.; height,
55.4 mm.; convexity (both valves together),
28.3 mm.; pallial sinus extends forward 36
mm. from the posterior margin. A large sin-
gle left valve from Magdalena Bay, Lower
California, in the collection of the same in-
stitution measures : length, 64.4 mm. ; height,
63.4 mm.; convexity (one valve), 15 mm.;
pallial sinus extends anteriorly 40 mm. from
the posterior margin of the shell.
Gould’s type specimen has never been illus-
trated but the foregoing description applies
exactly to specimens of a species in the col-
lection of the California Academy of Sciences
which were collected from Lower California
to Panama. This species is identical with the
one illustrated by Reeve under the name of
Amphidesma proximo, Adams. According to
Dali10 Adams’ species is identical with Sem-
ele flavescens. Amphidesma proximum C. B.
Adams was founded upon a specimen from
Panama 1.8 inches in length. It was said to
be closely related to Semele elliptica Sowerby
and S. lenticulare Sowerby. Carpenter* 11 re-
garded S. proximo as identical with S. ellip-
tica. Whether or not S. proximo is identical
with S. flavescens may be open to doubt but
certainly Reeve’s figure attributed to that
species is referable to S. flavescens.
In some cases Semele flavescens has been
identified under the name of Semele striosa
C. B. Adams12. That species was based upon
10 DaU. W. H., Proe. Acad. Nat. Sci. Philadelphia, Vol.
67, 1915, p. 26.
11 Carpenter, P. P., Proc. Zool. Soc. London, 1863, p. 367.
Reprint in Smithson. Miscell. Coll., No. 252, 1872, p. 203.
12 Amphidesma striosum C. B. Adams, Ann. Lyceum
Nat. Hist. New York, Vol. 6, July, 1862, pp. 515, 647
(separate pp. 291, 323). "Habitat.— Panama.”
a single specimen from Panama .78 inch in
length and it appears uncertain exactly how
it differs from related forms. In the original
description it is mentioned . . . “corselet and
lunule not well defined.” This does not agree
with S. flavescens which has a well developed
escutcheon.
Semele mediamericana Pilsbry & Lowe13
differs from S. flavescens in lacking the
strong escutcheon and in the sculpture in
which . . . “there are narrow, thread-like con-
centric ridges, coarser and more raised than
in S. flavescens, a little less than one mm.
apart on the lower part of the valve, and a
very minute, dense, even radial sculpture
throughout, diverging at both ends, and seen
under the lens to be totally unlike the radial
striation of S. flavescens.”
Distribution: A single left valve of Semele
flavescens was taken by the expedition at
Golfito Bay in the Gulf of Dulce. It occurs
fairly commonly from Magdalena Bay to the
Gulf of California and south to Panama and
apparently to Peru. We have not seen speci-
mens from north of Magdalena Bay, but the
type locality is San Diego, and Dali cited it as
occurring north to Catalina Island. It also
occurs in the Pliocene of the Gulf of Califor-
nia region and in the Pleistocene of Mag-
dalena Bay, Lower California, and it has been
recorded as occurring in the Quaternary of
Ecuador. Olsson, 1932, cited ‘‘Semele cf.
flavescens Gould” as occurring in the Mio-
cene of Peru.
Semele guaymasensis Pilsbry & Lowe.
Semele guaymasensis Pilsbry & Lowe,
Proc. Acad. Nat. Sci. Philadelphia, Vol. 84,
May 21, 1932, p. 92, pi. 12, figs. 8 and 9.
“Guaymas, 20 fathoms.” — E. K. Jordan,
Contrib. Dept. Geol. Stanford Univ., Vol. 1,
No. 4, 1936, p. 145. Magdalena Bay, Lower
California, Pleistocene. Also Gulf of Cali-
fornia, Recent.
Type Locality: Guaymas, Sonora, Mexico,
in 20 fathoms.
Range: Punta Penasco, Sonora, Mexico,
to La Paz, Lower California.
Collecting Station: Mexico: Santa Inez
Bay, Gulf of California (145-D-l, 3), 4-13
fathoms, sand.
Description: The shell is light buff, faintly
mottled or obscurely rayed with dull light
purple, the dorsal borders dark purple. Shape
irregularly oval, nearly equilateral, strongly
compressed; dorsal margin slightly convex
behind, straight in front of the beaks; ends
rounded; ventral margin strongly convex.
Sculpture of strong, concentric ridges gener-
ally a little wider than their intervals, a little
lamellar at the border of the escutcheon, and
on the anterior end cut by about seven radial
grooves. Escutcheon very narrow, flattened,
with weak growth lines only, purple. Lunule
small, sunken, the dorsal area beyond it pur-
13 Semele mediamericana Pilsbry & Liowe, Proc. Acad.
Nat. Sci. Philadelphia, Vol. 84, May 31, 1982, p. 92. pi. 12,
figs. 1, la, 2 (as Semele mcdiamcricanum on e.xpl. to pi.).
"Nicaragua (McNeil).’’
244
Zoologica: New York Zoological Society
[34: 19
pie and smooth except for lines of growth.
The interior is stained with dull purple on
a buff or white ground, with purple markings
on the ventral border. The pallial sinus oc-
cupies about two-thirds of the length. Length,
16 mm.; height, 12.3 mm.; semidiam. (right
valve), 2.6 mm. Length, 22 mm.; height 17
mm.; semidiam. (right valve) 4 mm. (Orig-
inal description.)
This species differs from Semele pulchra
Sowerby and S. quentinensis Dali in the
much coarser and more widely spaced con-
centric sculpture.
Semele anteriocosta Vokes14, described
from the Miocene of Trinidad, is similar to
S. guaymasensis in its general characters
but the strength of the ribbing appears to be
intermediate between that of this species and
S. quentinensis.
Distribution: A few specimens referable
to this species were dredged by the expedi-
tion in Santa Inez Bay, in the Gulf of Cali-
fornia, in 4-13 fathoms, on a sandy bottom.
Seme/e jaramija Pilsbry & Olsson.
Plate I, Fig. 12.
Semele jaramija Pilsbry & Olsson, Proc.
Acad. Nat. Sci. Philadelphia, Vol. 93, Sep-
tember 9, 1941, p. 70, pi. 17, fig. 5. “Canoa
formation, Punta Blanca.” Ecuador, Plio-
cene.
Type Locality: Canoa formation, Punta
Blanca, Ecuador, Pliocene.
Range: Santa Inez Bay, Gulf of Califor-
nia.
Collecting Station: Mexico: Santa Inez
Bay, Gulf of California (145), on shore.
Description: Shell small, suboval in form,
with the beaks placed a little in back of the
middle; but little convex; the sculpture con-
sists of strong, regular, concentric threads,
well developed over the whole shell and at
the anterior-upper end, these concentric
threads are cut by a series of small radial
grooves; at the posterior end the grooves,
about 8 in number, are crossed by the con-
centric threads, the sculpture being beauti-
fully cancellated. Length, 21 mm.; height,
16 mm.; semidiameter, 3.5 mm. (Original
description.)
A left valve of this species in the present
collection measures: length, 16 mm.; height,
12.2 mm.; convexity (one valve), 2.9 mm.;
pallial sinus extends anteriorly 9.5 mm. from
the posterior margin of the shell.
The present specimen agrees exactly with
the illustration of Semele jaramija given by
Pilsbry & Olsson.
As mentioned in the discussion of Semele
pacifica, that species always has radial sculp-
ture on both the anterior and posterior dorsal
areas. The variation in S. pacifica Dali is so
great that it appears quite possible that the
form here cited as S. jaramija may be merely
a subspecies of it.
i * Semele anteriocosta Vokes, Arner. Mas. Novit., No.
988, May 16, 1938, p. 14, fig. 6. Upper Miocene of Spring-
vale, Trinidad, British West Indies.
Semele guaymasensis Pilsbry & Lowe and
S. quentinensis Dali have radial sculpture
usually only on the anterior dorsal area and
when present at all posteriorly it is much less
strongly developed than that on S. jaramija.
Distribution : A single left valve here re-
ferred to Semele jaramija was taken by the
expedition on shore at Santa Inez Bay in
the Gulf of California. This species has also
been recorded as occurring in the Pleistocene
of Panama and in the Pliocene at Punta
Blanca, Ecuador.
Semele jovis Reeve.
Amphidesma jovis Reeve, Conch. Icon.,
Vol. 8, Amphidesma, November, 1853, spe-
cies 34, pi. 5, fig. 34. “Hab.— ?”
Semele jovis A. Adams, Proc. Zool. Soc.
London for 1853 (issued July 25, 1854), p.
94. “Hab. ? Mus. Cuming.”
Tellina barbarae Boone, Bull. Bingham
Oceanogr. Coll. Peabody Mus. Yale Univ.,
Vol. 2, Art. 5, December, 1928, p. 9, pi. 1
(upper figure). “Pearl Islands, depth 12
fathoms.”
Type Locality: Port Parker, Costa Rica
(here designated as type locality). No lo-
cality cited originally.
Range: Kino Bay, Sonora, Mexico, in the
Gulf of California, to the Las Perlas Islands,
Panama.
Collecting Stations: Mexico: Port Gua-
tulco (195-D-2), 3 fathoms, sand; Nicara-
gua: Corinto (200-D-19), 12-13 fathoms,
mangrove leaves; Costa Rica: Port Parker
(203-D-1-3), 12-15 fathoms, sandy mud,
crushed shell, shelly sand, algae, shelly mud.
Description: Shell somewhat roundly
ovate, somewhat ventricose, anterior side
slightly the longer; posterior side with a
flexure, the end truncated ; ornamented with
rather thin, close-set, concentric lamellae;
the interspaces with fine concentric linea-
tion ; extremely fine radial wrinkling present
on fresh specimens but clearly noticeable on
worn specimens; color rose-fawn, beaks red
with a medial white streak; hinge with two
cardinals and laterals in each valve; pallial
sinus broadly elliptically rounded, projecting
forward about four-sevenths the length of
the shell; interior rose and white.
A right valve from Port Guatulco, Mexico,
measures : length, 54.5 mm. ; height, 45 mm. ;
convexity (one valve), 10.3 mm.; pallial
sinus extends anteriorly 32.5 mm. from the
posterior margin of the shell. A specimen
collected by H. N. Lowe at Kino Bay, Sonora,
Mexico, in the Gulf of California, measures
59 mm. in length.
Semele rosea Sowerby15, described from
Peru, is more orbicular in outline than S.
jovis.
is Amphidesma roseum Sowerby, Conch. Illustr., Catal.
issued with Pt. 19, species No. 6, pi. 17, fig. 1, issued
between January 18 and March 8, 1833. “Tumbez, Peru.
Mr. Cuming."— Sowerby, Proc. Zool. Soc. London for 1832
(issued March 13, 1833), p. 199. "Hab. ad littora Peruviae.”
“A single valve was found at Tumbez in Peru."— Reeve,
Conch. Icon., Vol. 8, Amphidesma, October, 1863, species
17, pL 3, fig. 17 (as Amphidesma rosea). Tumbez, Peru.
1949]
Hertlein & Strong : Mollusks of Mexico and Central America
245
According to Verrill the lamellae are more
closely spaced and the plication of Semele
jovis is nearer the outer edge as compared
to that of S. junonia Verrill16 which was de-
scribed from La Paz, Lower California. He
mentioned the presence of radiating striae
in the interspaces of S. junonia, a feature
also present and especially noticeable on
somewhat worn specimens of S. jovis. Ac-
cording to Lamy17 Semele junonia is only a
variety of S. rosea.
Distribution : A few single valves of
Semele jovis were dredged by the expedition
off western Mexico, Nicaragua, and Costa
Rica.
Semele laevlt Sowerby.
Amphidesma laeve Sowerby, Conch. II-
lustr., Catal. issued with Pt. 19, No. 22, pi. 18,
fig. 6, issued between January 18 and March
8, 1833. “Xipixapi. W. Col. Mr. Cuming.” —
Sowerby, Proc. Zool. Soc. London for 1832
(issued March 13, 1833), p. 199. “Hab. ad
Xipixapi, Columbiae Occidentalis.” “A single
specimen of this very delicate species was
dredged from a depth of ten fathoms in sandy
mud.”
Amphidesmalaevis Sowerby, Reeve, Conch.
Icon., Vol. 8, Amphidesma, November, 1853,
species 50, pi. 7, fig. 50. Original locality
cited.
Type Locality : Xipixapi [Jipijapa], Ecua-
dor, in 10 fathoms, sandy mud.
Range: Champerico, Guatemala, to Jipi-
japa, Ecuador.
Collecting Stations: Guatemala: 7 miles
west of Champerico (197-D-1-2) , 14 fathoms,
mud; El Salvador: La Libertad (198-D-1-2),
13-14 fathoms, mud; Costa Rica: Gulf of
Dulce.
Description: Shell elongately ovate, in-
equilateral, the anterior side the longer,
smooth, exterior and interior white ; anterior
dorsal margin nearly straight, sloping, an-
terior end tapering and rounded; ventral
margin curved ; posterior dorsal margin
rounded and highest just back of the beaks,
posterior end rounded; a broad, shallow,
radial groove is present on the posterior area
and where this meets the ventral margin
there is sometimes a vague truncation ; sur-
face smooth except for concentric lines of
growth and an occasional concentric groove
and sometimes with fine submicroscopic
radial striae; hinge of right valve with two
small cardinals, the posterior one bifid, the
anterior one thin, two laterals present, left
valve with two cardinals, the anterior one
bifid, the posterior one thin, also projections
of the margin which fit into corresponding
sockets in the right valve; pallial sinus some-
what elevated above then broadly tapering
to a rounded point which projects forward
about five-eighths the length of the shell.
16 Semele junonia Verrill, Amer. Jour. Sci., Ser. 2, Vol.
69, No. 146, March, 1870, p. 217. "Near La Paz,- Capt.
J. Pedersen.”
17 Lamy, E., Jour n. de Conchyl., Vol. 61, No. 3. 1914,
p. 357.
A large right valve from off La Libertad,
El Salvador, measures approximately:
length, 68 mm.; height, 53 mm.; convexity
(one valve), 13 mm.; pallial sinus extends
anteriorly 48.4 mm. from the posterior mar-
gin of the shell.
Semele laevis var. costaricensis Olsson18
has been described from the Miocene of Costa
Rica and later was cited as also occurring
in the Miocene of Peru.
Semele pallida Sowerby19, described from
Ecuador, bears a resemblance, in general
features, to S. laevis but differs in that it is
less elongate anteriorly, less rounded poste-
riorly, the beaks are more anteriorly situ-
ated and the coloration was described as pale
purple-fulvous.
Distribution: Specimens of this species
were dredged off Guatemala and El Salvador
in 13-14 fathoms and were taken on the
beach of the Gulf of Dulce. These occurrences
extend the known range of the species con-
siderably to the north as heretofore it has
not been reported north of Panama. This
species also has been recorded as occurring
in the Pleistocene of Panama and in the
Pliocene at Puerto Jama, Ecuador.
Semele pacifica Dali.
Plate I, Fig. 11.
Semele pacifica Dali, Proc. Acad. Nat. Sci.
Philadelphia, Vol. 61, March 2, 1915, p. 27.
“Catalina Island, California, to Acapulco,
Mexico, in 9 to 21 fathoms.” — I. S. Oldroyd,
Stanford Univ. Publ. Univ. Ser. Geol. Sci.,
Vol. 1, 1924, p. 180, pi. 3, fig. 5. Original
range cited. — J. Q. Burch, Min. Conch. Club
South. Calif., No. 43, January, 1945, p. 17.
“Dr. A. M. Keen advises ‘Type locality of
S. pacifica is: U. S. B. F. Sta. 2022, off La
Paz, in 21 fms.’ ”
Type Locality: Off La Paz, Lower Cali-
fornia, in 21 fathoms (Keen).
Range: Catalina Island, California, to the
Gulf of California and south to Taboga
Island, Panama.
Collecting Stations: Mexico: Santa Inez
Bay, Gulf of California (145-D-l, 3), 4-13
fathoms, sand, also on shore; Costa Rica:
Port Parker (203-D-3), 12 fathoms, shelly
mud; Golfito, Gulf of Dulce.
Description: The shell of this species is
very similar to that of Semele cancellata
Sowerby20 which occurs in Atlantic waters.
16 Semele laevis Sowerby. var. costaricensis Olsson, Bull.
Amer. Paleo., Vol. 9, Bull. 39, Pt. 2, June 21, 1922, p. 430
(258), pi. 32 (29), fig. 1. "Gatun Stage: Hill No. 3, Banana
River.” Costa Rica, Miocene.
19 Amphidesma pallidum Sowerby, Conch. Illuutr., Catal.
issued with Pt. 19, sp. No. 3, pi. 17, fig. 3, issued between
January 18 and March 8, 1833. “Salango, W. Col. Mr.
Cuming.”— Sowerby, Proc. Zool. Soc. London for 1832
(issued March 13, 1833), p. 199. “Hab. ad Salango,
Columbiae Occidentalis.'* "Dredged in sandy mud at a
depth of seven fathoms.”— Reeve, Conch. Icon., Vol, 8,
Amphidesma, 1853, species 22, pi. 4, fig. 22 (as Amphidesma
pallida). Original locality cited.
20 Amphidesma cancellatum Sowerby, Conch. Illustr.,
Catal. issued with Pt. 19, Bpecies No. 13, issued between
January 18 and March 8. 1833. "Antigua and St. Vincents.”
Ref. to “Spec. Conch, f. 8.”— Reeve, Conch. Icon., Vol. 8,
Aphidesma, 1863, species 44, pi. 7, fig. 44 (as Amphidesma
cancellata). “Hab.— ?.”
246
Zoologica: New York Zoological Society
[34: 19
Dali stated in the original description that
it . . . “differs from that Atlantic species in
its smaller lunule, shorter and weaker right
lateral tooth, and sharper and more delicate
concentric sculpture.”
A left valve in the present collection from
Golfito, Gulf of Dulce, Costa Rica, measures:
length, 20 mm.; height, 16.6 mm.; convexity
(one valve), 4.6 mm.
Semele pacific is a very variable species.
Young specimens have rather flattened shells
and strong cancellate sculpture with the con-
centric lamellae well developed. In the adult
stage the shells become thicker, more ventri-
cose, and with the radial element in the sculp-
ture as strong or stronger than the concen-
tric. The radial sculpture is always present
near the anterior and posterior dorsal mar-
gins and sometimes covers the whole shell.
Semele pacifica is one of a group of related
species which vary in details of sculpture.
Semele venusta Reeve has no radiating sculp-
ture. Semele guaymasensis Pilsbry & Lowe
has strong concentric sculpture but with in-
cised radiating sculpture only on the anterior
dorsal area. Semele quentinensis Dali has
very fine concentric sculpture with radial
sculpture on the anterior dorsal portion and,
rarely, with a few faint striae along the
posterior dorsal margin. Semele pulchra Sow-
erby has concentric and radial sculpture
similar to that of S. quentinensis but the shell
is higher in proportion to the length as com-
pared to Dali’s species. Semele jaramija Pils-
bry & Olsson, described from the Pliocene of
Ecuador, has concentric sculpture interme-
diate in strength between that of S. quenti-
nensis and S. guaymasensis, but in addition
to similar radial sculpture on the anterior
dorsal area it also has strong, incised radial
sculpture on the posterior dorsal area. Semele
pacifica has strong well developed concentric
sculpture with radials on both the anterior
and posterior portions and sometimes all over
the shell.
Distribution : A few specimens of Semele
pacifica were dredged by the expedition in
4-13 fathoms in Santa Inez Bay, in the Gulf
of California, at Port Parker, Costa Rica, in
12 fathoms, and at Golfito in the Gulf of
Dulce.
Semele pulchra Sowerby.
Plate I, Fig. 15.
Amphidesma pulchrum Sowerby, Proc.
Zool. Soc. London, June 5, 1832, p. 57. “Hab.
in Sinu Caraccensi, Americae Meridionalis.”
— Sowerby, Conchyl. Illustr., Catal. issued
with Pt. 19, species No. 2, pi. 17, fig. 2, issued
between January 18 and March 8, 1833, “St.
Elena W. Columbia.” Var. fig. 2*. Panama.
Amphidesma pulchra Sowerby, Reeve,
Conch. Icon., Vol. 8, Amphidesma, October,
1853, species 2, pi. 1, fig. 2. Original locality
cited.
Type Locality: Bay of Caraccas, Ecuador.
Range: Gulf of Fonseca, Nicaragua, to
Ecuador.
Collecting Stations: Nicaragua: Potosi
and 5 miles SSW. of Monypenny Point, Gulf
of Fonseca.
Description: Shell trigonally ovate, fairly
thick, anterior side the longer, sloping and
rounded at the end; posterior side with a
fold, the end roundly truncated; ventral
margin rounded; sculptured with fine close
concentric riblets, on the anterior end these
are decussated by several (10-15) incised
radial lines; color pale yellowish-gray with
purple blotches and A-shaped markings;
pallial sinus higher in front of the adductor
impression then gently sloping to a broadly
rounded end which projects forward about
three-fifths the length of the shell; hinge
normal; interior colored white with the um-
bonal half and the hinge purple or tinged
with purple.
A typical specimen from Nicaragua mea-
sures: length, 31 mm.; height, 25.4 mm.;
convexity (both valves together), 12.5 mm.;
pallial sinus projects anteriorly 19.8 mm.
from the posterior margin of the shell.
The shells here referred to Semele pulchra
agree exactly with the figures of that species
given by Sowerby and by Reeve. Semele quen-
tinensis Dali, a closely related species which
has usually been cited under the name of
S. pulchra, occurs from southern California
to Central America. It is more elongate in
outline, the anterior dorsal margin slopes
more gently from the beaks, the shell is
thinner and the purple coloration is more
weakly developed.
Distribution: Only three specimens of this
species were taken by the expedition in the
Gulf of Fonseca. It ranges south to Ecuador.
Semefe quentinensis Dali.
Plate I, Fig. 10.
Semele quentinensis Dali, West Amer. Sci.,
Vol. 19, No. 3, June 15, 1921, p. 22. “Pliocene
or Early Pleistocene of San Quentin.” — Dali,
Proc. U. S. Nat. Mus., Vol. 66, No. 2554, Art.
17, 1925, p. 26, pi. 8, fig. 4. “Pliocene (?) of
San Quentin Bay, Lower California.”
Type Locality: San Quintin, Lower Cali-
fornia, Pleistocene.
Range : Point Mugu, Ventura County, Cali-
fornia, to Costa Rica.
Collecting Stations: Guatemala: 7 miles
west of Champerico (197-D-2), 14 fathoms,
mud; El Salvador: Meanguera Island, Gulf
of Fonseca (199-D-l), 16 fathoms, sand,
mud, crushed shell; Nicaragua: Corinto
(200-D-19), 12-13 fathoms, mangrove
leaves; Costa Rica: 1 mile south of Golfito.
Description : Shell small, inequilateral, in-
equivalve, rather compressed, anterior end
longer, terminally rounded; posterior end
obscurely subtruncate, base moderately ar-
cuate; beaks inconspicuous; surface finely
concentrically closely sculptured, with fine
radial threads chiefly visible in the sulci ; at
the anterior end are about a dozen stronger
sulci, cutting and more or less beading the
concentric sculpture, but this feature is not
1949]
Hertlein & Strong : Mollusks of Mexico and Central America
247
repeated at the posterior end ; hinge normal,
well developed ; pallial sinus large, subovate,
nearly reaching the anterior adductor scar,
and entirely free from the pallial line; the
left valve slightly flatter than the right valve.
Length, 24; height, 19; diameter, 8 mm.
(Original description).
A large specimen from the Gulf of Fonseca
in the present collection measures: length,
27.3 mm.; height, 20.8 mm.; convexity (both
valves together), 8.9 mm.; pallial sinus ex-
tends anteriorly 18 mm. from the posterior
margin of the shell.
The shell of this species is very similar to
that of Semele pulchra but the length is
greater in proportion to the height and the
anterior dorsal margin slopes more gently
from the beaks. The specimens in the present
collection are thinner and the purple colora-
tion is less pronounced than that of S.
pulchra. These shells possess fine, even, con-
centric sculpture which along the anterior
dorsal margin is crossed by incised radial
lines giving a beaded appearance to that por-
tion of the shell. These specimens agree
exactly with Dali’s description and illustra-
tion of Semele quentinensis which was based
on a fossil specimen from the Pleistocene of
San Quintin, Lower California. These appear
to be identical with the species occurring in
southern California which generally has been
cited in the literature under the name of
Semele pulchra. The only difference seems
to be in size, those from southern California
seldom exceeding 20 mm. in length. A few
specimens of this northern form also possess
a few incised radial lines along the posterior
dorsal margin but they are fewer and much
weaker than those on the anterior dorsal
margins and thus differ from the sculpture
of Semele jaramija Pilsbry & Olssen which
was originally described from the Pliocene
of Ecuador. Semele guaymasensis Pilsbry &
Lowe has more widely spaced and coarser
concentric sculpture.
Distribution : Specimens of Semele quen-
tinensis were dredged by the expedition in
12 to 16 fathoms from Guatemala to Costa
Rica. It also is known to occur in the Pleisto-
cene of southern California and Lower Cali-
fornia.
Semele slmpllclsslma Pilsbry & Lowe.
Semele simplicissima Pilsbry & Lowe,
Proc. Acad. Nat. Set. Philadelphia, Vol. 84’
May 21, 1932, p. 93, pi. 12, figs. 6, 6a. “Aca-
pulco, 20 fathoms.”
Type Locality: Acapulco, Mexico, in 20
fathoms.
Range : Santa Inez Bay, Gulf of California,
to Acapulco, Mexico.
Collecting Stations: Mexico: Arena Bank
(136-D-2, 5), 33-45 fathoms, mud, Area con-
gomerates, sand, weed; Santa Inez Bay
(143-D-2, 3, 4), 25-35 fathoms, mud, crushed
shell; Santa Cruz Bay (195-D-21), 18 fath-
oms, mud; Costa Rica: Port Parker (203-
D-3) , 12 fathoms, shelly mud ; 14 miles S. X
E. of Judas Point (214-D-l, 4), 42-61 fath-
oms, mud, shell, rocks.
Description: Shell ovate, thin, moderately
inflated at the umbos, beaks near the middle ;
anterior dorsal margin sloping, nearly
straight, end rounded, ventral margin broad-
ly rounded, posterior end a little higher than
the anterior, convex dorsally, in large speci-
mens decidedly truncated at the end where
the fold reaches the margin; sculpture of
very fine, fairly regular, low, concentric
ridges which are covered by such fine, con-
centric lines that they disperse light into
spectral colors; the interspaces are flat and
without either concentric or radial striation ;
lunule lanceolate and rather deeply concave ;
pallial sinus ascending, rounded at the end
and projecting a little beyond the middle of
the shell ; color dingy white and on the inte-
rior of fresh specimens a salmon pink or
dark orange flush covers the umbonal half of
the shell.
Some specimens in the present collection
are much larger than the type of this species.
The largest shell, a left valve, measures:
length, 33 mm. ; height, 24.3 mm. ; convexity
(one valve), 6.4 mm.; pallial sinus projects
anteriorly 19 mm. from the dorsal margin
of the shell. One pair of valves from Arena
Bank measures, approximately: length, 29.5
mm; height, 22.8 mm.; convexity (both
valves together), 12.2 mm.; pallial sinus ex-
tends anteriorly 16.5 mm. from the posterior
margin of the shell.
The present specimens have been identified
after a comparison with paratypes of Semele
simplicissima in the H. N. Lowe collection
in the San Diego Society of Natural History.
This species appears to be very similar to
the one described as Semele regularis Dali21
[=Semele paziana, new name], but differs in
that the intervals between the ribs are
smooth and usually not ornamented by con-
centric striations. However, some specimens
here referred to S. simplicissima bear sub-
microscopic striae in the interspaces.
Semele sayi Toula, 1909, described from
the Gatun Miocene of Panama, and especially
S. quirosana H. K. Hodson, 1931, described
from the Upper Oligocene or Miocene of
Venezuela, bear a resemblance to S. sim-
plicissima.
Distribution : Specimens of this species
were dredged by the expedition from Santa
Inez Bay in the Gulf of California, to off
Judas Point, Costa Rica, at depths of 12 to
61 fathoms. These records of occurrence fur-
nish new extensions both north and south
of the known range of the species.
Semele sparslllneata Dali.
Plate I, Fig. 8.
Semele sparsilineata Dali, Proc. Acad. Nat.
I ■
21 Semele regularis Dali, Proc. Acad. Nat. Sci. Phila-
delphia, Vol. 67, issued March 2, 1916, p. 27. "Gulf of
California, off La Paz, in 10 to 30 fathoms."
Not Semele regularis E. A. Smith, Sci. Res. Voy. Chal-
lenger, Zool., Vol. 13, LamelL, 1886, p. 87, pi. 6, figs. 4,
4a, 4b. East of Cape York, North Australia in 166 fathoms.
A new name Semele paziana Is here proposed for the
west American species named Semele regular i# by Dali.
248
Zoological New York Zoological Society
[34: 19
Sci. Philadelphia, Vol. 67, issued March 2,
1915, p. 26. “Panama, 18 fathoms.” Also
recorded from “Chile, Hupe.”
Type Locality. Panama, in 18 fathoms.
Range: Corinto, Nicaragua, to Taboga
Island, Panama. To Chile (Dali).
Collecting Station: Panama: Gulf of
Chiriqui (221-D-l, 5), 35-40 fathoms, sandy
mud.
Description: Shell ovately oblong, the an-
terior portion much the longer, the end
rounded, ventral margin rounded, posterior
end slightly higher, slightly subtruncated, a
flexure present; sculptured by fine concen-
tric lines of growth which, sometimes ante-
riorly and sometimes medially, are crossed
by fine oblique striations; pallial sinus as-
cending, broadly rounded at the end, pro-
jecting forward about three-fifths the length
of the shell; color dingy white with traces of
brownish-purple stains.
The larger specimen in the collection, a
right valve, measures approximately: length,
25.5 mm. ; height, 20.8 mm. ; convexity (one
valve), 4.7 mm.; pallial sinus extends ante-
riorly 15.8 mm. from the posterior margin
of the shell.
The present specimens are somewhat worn
but they show the oblique striae character-
istic of this species. It was upon the basis
of sparser oblique striae that Dali separated
this species from the east American Semele
purpurascens Gmelin22.
Distribution: Only two single valves of
this species were dredged by the expedition
in the Gulf of Chiriqui, Panama, in 35-40
fathoms.
Semele tabogensls Pilsbry & Lowe.
Semele tabogensis Pilsbry & Lowe, Proc.
Acad. Nat. Sci. Philadelphia, Vol. 84, May 21,
1932, p. 91, pi. 12, figs. 5, 5a, 5b. “Taboga
Island, among rocks near the bathing beach.”
Type Locality: Taboga Island, Panama,
among rocks.
Range: Tangola-Tangola Bay, Mexico, to
Taboga Island, Panama.
Collecting Station: Mexico: Tangola-
Tangola Bay (196-D-7), 6 fathoms, sand.
Description: The shell is thin, orange,
shading through pink into light coral red
near the umbones ; very shortly oval, strongly
compressed, slightly inequilateral. The
broadly rounded anterior end is somewhat
lower than the posterior end, which is no-
ticeably truncate. Dorsal margin somewhat
22 Venus purpurascens Gmelin, Syst. Nat., ed. 13, Vol. 1,
Pars. 6, 1791, p. 3288. Habitat unknown. Ref. to: “List.
Conch, t. 303. f. 144.”; “B (List. Conch, t. 304. f. 145.”;
“Klein, ostr. t. 11. f. 57”.)
Tellina obliqua Wood, Gen. Conch., 1815, p. 152, pi. 41,
figs. 4, 5.
This is not Amphidesma purpurascens Sowerby (Conch.
Illustr., Pt. 19, species No. i9, pi. 18, fig. 6, issued between
January 18 and March 8, 1833. "St. Elena. W. Col. Mr.
Cuming.”— Sowerby, Proc. Zool. Soc. London for 1832
(issued March 13, 1833), p. 199. “Hab. ad Sanctam
Elenam.” "A single valve of this elegant 6pecies was
picked up on the sands at St. Elena”.— Reeve, Conch. Icon.,
Vol. 8, Amphidesma, November, 1863, species 37, pi. 6,
fig. 37), which was renamed Semele sowerby i by Lamy
(Bull. Mu8. Nat. Hist. Nat. (Paris), Vol. 18, No. 3, 1912,
p. 166, footnote).
concave in front of the beaks, convex behind
them. In the right valve these margins are
produced towards the other valve, covering
the ligament. Ventral margin is strongly
convex. Sculpture of regular, recurved con-
centric riblets, which become laminar near
the dorsal margin, and are somewhat darker
colored than their intervals, in which fine,
weak, radial striation is seen. Beaks smooth.
Lunule extremely small, confined to the right
valve. The interior varies in color from car-
nelian red to apricot orange, smooth, with
some scattered glossy dots. Teeth are lighter
or whitish in large individuals. Anterior
lateral short, the posterior long and thinner.
The pallial sinus reaches well past the middle.
Length 37.5 mm., height 30.3 mm., semidiam.
(right valve) 6.5mm. (Original description).
The present specimen, a left valve, mea-
sures approximately: length, 22.3 mm.;
height, 17.3 mm.; convexity (one valve), 4.3
mm.
Distribution: A single left valve of this
species was dredged by the expedition in
Tangola-Tangola Bay, Mexico, in 6 fathoms.
This is an extension north of the known
range of this species.
Semele venusta Reeve.
Plate I, Fig. 13.
Amphidesma venusta Reeve, Conch. Icon.,
Vol. 8, Amphidesma, October, 1853, species
3, pi. 1, fig. 3. “Hab. West Columbia.” — A.
Adams, Proc. Zool. Soc. London for 1853
(issued July 25, 1854), p. 96. “Hab. West
Columbia. Mus. Cuming.”
Type Locality : West Colombia.
Range: Acapulco, Mexico, to west Colom-
bia.
Collecting Stations : Mexico : Port Guatul-
co (195-D-9), 7 fathoms, gr. sand, crushed
shell ; Santa Cruz Bay ; Tangola-Tangola Bay
(196-D-8), 9 fathoms, sand.
Description: Shell oblong, transverse,
ventricose, rather shining, dull flesh-color,
obscurely rayed with rose, anterior side much
the longer, posterior slightly truncated,
flexuous at the ventral margin; concen-
trically grooved; purple within, edged with
white (Reeve).
A left valve in the present collection from
Santa Cruz Bay, Mexico, measures: length,
21.9 mm.; height, 15 mm.; convexity (one
valve), 4.2 mm.; pallial sinus extends ante-
riorly 14 mm. from the posterior margin of
the shell.
The pallial sinus of this shell is a distinc-
tive feature. It extends forward about two-
thirds the length of the shell; it is broad and
higher in front of the posterior adductor
impression then tapers elliptically to a
rounded point.
The ribbing of Semele venusta is some-
what irregular toward the anterior and pos-
terior ends similar to that of S. incongrua
Carpenter23 athough coarser ventrally. The
23 Semele incongrua Carpenter, Rept. Brit. Assoc. Adv.
Sci. for 1863 (issued August. 1864), pp. 611, 640. "Catalina
19491
Hertlein & Strong: Mollusks of Mexico and Central America
249
shell in adult forms is thicker than that of
Carpenter’s species. Fine radial sculpture is
present in the bottoms of the interspaces.
A few small specimens in the present col-
lection from off western Mexico are remark-
ably similar to Semele incongrua. Traces of
the pallial sinus appear to be narrowly ellip-
tical at the end similar to that of S. venusta
rather than broady rounded as in S. incon-
grua.
The form described as Semele pulchra var.
montereyi Arnold24, based on a Pleistocene
fossil from San Pedro, California, is, as
mentioned by Dali, a subspecies of S. incon-
grua. The type specimen has not been illus-
trated but the figure given by Arnold repre-
sents a shell which appears to be a little more
rounded, with sharper, more erect concentric
lamellae and with stronger radial ornamen-
tation than that of S. incongrua.
Verrill 25 considered semele venusta to be
but a young form of S. formosa Sowerby.
The pallial sinus, elliptically pointed in S.
venusta, is quite different from the broadly
rounded ascending pallial sinus of S. for-
mosa.
Distribution’. A few valves of Semele ven-
usta were taken by the expedition at Port
Guatulco, Santa Cruz Bay, and Tangola-
Tangola Bay, Mexico, in 7 to 9 fathoms.
Semele verrucosa Morch.
Plate I, Figs. 21, 24.
Semele {Amphidesma) verrucosa Morch,
Malakozool. Blatter, Bd. 7, December, 1860,
p. 190. “Los Bocorones ad prof. 20 org. spec.
2”. Costa Rica.
Type Locality: Los Bocorones Islands,
Costa Rica.
Range: Los Bocorones Islands, Costa Rica,
to Hannibal Bank, Panama.
Collecting Station: Panama: Hannibal
Bank (Sta. 224) , 35-40 fathoms, rocks, coral,
mud, sand, shells, algae.
Description: Shell elongately ovate, in-
equilateral, whitish blotched with purple;
anterior side the longer, the end rounded,
ventral margin broadly rounded, posterior
end broadly rounded and with a gentle fold;
sculpture consists of close concentric ribs,
these especially anteriorly and posteriorly
are wrinkled and give rise to scalloped scale-
like projections, the whole finely radially
wrinkled; hinge (right valve) with two car-
dinals and laterals; pallial sinus broadly
rounded at the end and gently ascending to
about five-eighths the length of the shell.
Is., 40-60 fm. ; common.” Reprint in Smithson. Miscell.
Coll., No. 262, 1872, pp. 97, 126.— Carpenter, Proc. Cali-
fornia Acad. Nat. Sci., Vol. 3, February, 1865, p. 208 (as
Semele incungrua) . “Hab. Santa Barbara, 16 fm. 1 valve;
Catalina Island, 40-60 fm., not uncommon ; Cooper."— I. S.
Oldroyd, Stanford Univ. Publ. Univ. Ser. Geol. Sci., Vol. 1,
1924, p. 181, pi. 11, figs. 12, 13. "Type locality, Santa
Barbara.” Range, Monterey, California, to the Coronado
Islands, Lower California.
24 Semele pulchra Sowerby, var. montereyi Arnold, Mem.
Calif. Acad. Sci., Vol. 3, 1903, p. 166, pi. 15, figs. 3, 3a.
Lower San Pedro series, San Pedro, California. Pleistocene.
Recent in Monterey Bay.
25 Verrill, A. E., Amer. Jour. Sci., Ser. 2, Vol. 69, No.
146, March, 1870, p. 219.
A right valve measures approximately:
length, 43 mm.; height, 32.4 mm.; convexity
(one valve), 7.1 mm.; pallial sinus extends
anteriorly 26.6 mm. from the posterior mar-
gin of the shell.
Morch pointed out that the shell of Semele
verrucosa is more elongate in outline, more
subtruncate posteriorly, and that the ventral
margin is more gently arcuate in outline than
that of S. formosa Sowerby26. The present
specimen possesses those characters as well
as the pronounced scaly verrucose sculpture
characteristic of Morch’s species.
Distribution: A single right valve of this
species was dredged by the expedition on
Hannibal Bank, Panama, in 35-40 fathoms.
This is an extension south of the known
range of the species.
Genus Abra Lamarck.
Abra Lamarck, Hist. Nat. Anim. s. Vert.,
Vol. 5, July, 1818, p. 492. Species cited : “Am-
phidesma tenuis” in the synonymy of which
was included, “Mactra tenuis. Maton, act. soc.
linn. 8. p. 72. no. 8” and “Abra tenuis. Leach”.
“Habite les mers d’Angleterre. Communique
par M. Leach” ; “Amphidesma prismatica” in
the synonymy of which was cited, “Ligula
prismatica. Montagu, test. brit. suppl. 23. t.
26. f. 3. Ex D. Leach.” and “Abra prismatica.
Leach.” “Habite les cotes d’Angleterre. Com-
munique par M. Leach.” — Gray, Proc. Zool.
Soc. London for 1847, p. 187. Type: Mactra
tenuis.— Dali, Trans. Wagner Free Inst. Sci.,
Vol. 3, Pt. 5, 1900, p. 995. Type: A tenuis
Montagu. — Woodring, Carnegie Inst. Wash-
ington, Pub. 366, 1925, p. 179. Type: Mactra
tenuis Montagu.
Type (designated by Gray, 1847) : Mactra
tenuis [Montagu, Test. Brit., Pt 2, 1803, p.
572, Suppl., 1808, pi. 17, fig. 7. “Southamp-
ton, where it is not uncommon on the shore to
the west of the town.” Also from “Wey-
mouth”— Forbes & Hanley, Hist. Brit. Moll.,
Vol. 1, 1853 (issued 1848), p. 323, pi. 17, fig.
7. Various localities in England], [For dates
of publication of this work see Fisher and
Tomlin, Jour. Conch., Vol. 20, No. 5, August,
1935, pp. 150-151].
Shell small, trigonal ; sculpture consisting
of incrementals ; ligament narrow, resilium
seated on a wide, deeply inset chondrophore ;
hinge of right valve consisting of 2 cardinals
(3a, 3b), the posterior one (3b) heavier, and
slender anterior and posterior laterals ; hinge
of left valve consisting of 2 cardinals (2a,
2b), the posterior one (2b) very small; pal-
lia! sinus deep, very wide, confluent with
pallial line (Woodring).
The genus Abra has been recorded as oc-
curing from Eocene to Recent.
26 Amphidesma formosum Sowerby, Conch. Illustr.,
Catal. issued with Pt. 19, No. 4, pi. 19, fig. 8 [two figs.],
issued between January 18 and March 8, 1833. “St. Elena.
Mr. Cuming.”— Sowerby, Proc. Zool. Soc. London for 1832
(issued March 13, 1833), p. 199. "Hab. ad Sanctam
Elenam.” "Only two odd valves were dredged in seven
fathoms water.”— Reeve, Conch. Icon., Vol. 8, Amphidesma,
1863, species 27, pi. 4, fig. 27 (as Amphidesma formosa).
Original locality dted.
250
Zoologica: New York Zoological Society
[34: 19
Lamy27 cited Mactra tenuis Montagu, the
type of Abra, and similar species, under the
genus Syndesmya Recluz, 1843, with the type
Mactra alba Wood, 1801.
Iacra H. & A. Adams, 1856, a subgenus of
Abra, with the type Scrobicularia seychel-
larum A. Adams, 1856, possesses divaricate
radial sculpture.
Abranda Iredale, 1924, is based upon
Abranda rex Iredale, an Australian species.
Key to the Species of Abra.
A. Shell elongate
a. Posterior end pointed pacifica 28
aa. Posterior end blunt tepocana2S
B. Shell short; high, inflated palmeri
Abra palmeri Dali.
Plate I, Figs. 16, 18, 20, 23.
Abra palmeri Dali, Proc. Acad. Nat. Sci.
Philadelphia, Vol. 67, March 2, 1915, p. 28.
“Ballenas Lagoon on the west coast of Lower
California; the Gulf of California (Dr. E.
Palmer) ; and Panama Bay in 26 fathoms (U.
S.N.Mus.). Type locality, Panama Bay. (U.
S.N.Mus., No. 96,301.)”
Type Locality: Panama Bay, in 26
fathoms.
Range : Ballenas Lagoon on the west coast
©f Lower California, and the Gulf of Cali-
fornia to Panama Bay.
Collecting Station: El Salvador: Mean-
guera Island, Gulf of Fonseca (199-D-l), 16
fathoms, sand, mud, crushed shell.
Description: Shell short, high, inflated,
white, with a silky surface, and a very thin,
polished, pale yellow periostracum; anterior
end and base rounded; beaks subcentral, dor-
sal margins descending, posterior end atten-
uated and with the extremity rounded ; right
valve with a deeply bifid (or double) cardinal
tooth, the laterals obsolete; left valve with
a single cardinal and no laterals. Length 10,
height 8, diameter 5.5. mm. The pallial sinus
rounded, 6 mm. deep. (Original description).
This species is nearest to A. lioica Dali, of
the Atlantic Coast of the United States
(Dali).
The largest specimen in the present collec-
tion measures : length, 10.5 mm. ; height, 9.2
mm.; convexity (one valve), 2.8 mm.
The short and high outline of Abra palmeri
separates it from the two other species de-
scribed from west American waters, A. pa-
cifica Dali, 1915, and A. tepocana Dali, 1915,
both of which were said to be elongate in
outline.
Distribution: Several specimens of this
species, mostly single valves, were dredged
off Meanguera Island, El Salvador, in the
Gulf of Fonseca, in 16 fathoms. This is the
first record of the occurrence of the species
since its original description.
2" Lamy, E., Joum. de Conchyl., Vol. 61, No. 3, 1914,
*p. 268-297.
28 Not represented In the present collection.
Genus Cumingia Sowerby.
Cumingfo lamellosa Sowerby.
Cumingia lamellosa Sowerby, Proc. Zool.
Soc. London, May 17, 1833, p. 34. “Hab. prope
littora Oceani Pacifici.” “Found at Payta in
hard clay at low water; and at Panama in
deep water.” — Sowerby, Gen. Rec. and Foss.
Shells, Cumingia, Vol. 2, No. 40, ?1833, pi.
244, fig. 3. — Sowerby, Conch. Icon., Vol. 19,
Cumingia, 1873, species 5, pi. 1, fig. 5. “Hab.
Chili”. — Lamy, Joum. de Conchyl., Vol. 61,
No. 3, 1914, p. 310. Paita, Peru.
Cumingia coarctata Sowerby, Proc. Zool.
Soc. London, May 17, 1833, p. 34. “Hab. ad
Sinum Caraecensem.” “Dredged from a
sandy muddy bottom in seven fathoms
water in the Bay of Caraccas”. [Ecuador].
Cumingia tngonularis Sowerby, Proc.
Zool. Soc. London, May 17, 1833, p. 35. “Hab.
ad Sanctam Elenam.” “Found among stones
in deep water.” — Sowerby, Gen. Rec. and
Foss. Shells, Vol. 2, No. 40, ?1833, Cumingia,
pi. 244, fig. 2. — Sowerby, Conch. Icon., Vol.
19, Cumingia, 1873, species 4, pi. 1, fig. 4.
“Hab. Chili?”
Cumingia adamsii Carpenter, Proc. Zool.
Soc. London, June 23, 1863, p. 367. Reprint in
Smithson. Miscell. Coll., No. 252, 1872, p. 203.
Name proposed for Cumingia, sp. indet. c of
C. B. Adams, Ann. Lyceum Nat. Hist. New
York, Vol. 5, July, 1852, p. 512 (separate p.
288). “Near Panama.”
Cumingia moulinsii De Folin, Les Melea-
grinicoles (Havre) , 1867, p. 16, pi. 2, figs. 12,
13, 14, 15. . . . “l’Ocean pacifique” . . . “pe-
chees aux environs des Negritos” . . . or . . .
“autour des lies aux Perles, dans la baie de
Panama”. — De Folin & Perier, Les Fonds de
la Mer, Vol. 1, 1867, p. 8. Bay of Panama.
[For dates of publication of this work see H.
A. Rehder, Proc. Malacol. Soc. London, Vol.
27, Pt. 2, September 5, 1946, pp. 74-75].
Type Locality: Paita, Peru, at low water,
in hard clay (here designated as type local-
ity) . Panama, in deep water, also cited orig-
inally.
Range: San Martin Island, Lower Califor-
nia, to the Gulf of California and south to
Paita, Peru.
Collecting Stations: Mexico: Port Guatul-
co (195-D-9), 7 fathoms, gr. sand, crushed
shell; Nicaragua: Corinto (200-D-10, 16, 17,
19), 4-13 fathoms, mangrove leaves, sand,
also on shore ; Costa Rica : Port Parker.
Description: Shell oblong, regularly con-
centrically laminated, laminae narrow, stand-
ing out, distant ; anterior side short, rounded ;
posterior side angular, acuminated, subro-
strated; ventral margin contracted near the
end; dorsal margin sloped (Sowerby, Conch.
Icon., Vol. 19).
Some of the larger specimens in the pres-
ent collection are about 12 mm. in length.
Some specimens attain a length of 20 mm. or
more.
Cumingia lamellosa lives in sand, sponges
and in fissures in rocks. Consequently it
1949]
Hertlein & Strong : MoUusks of Mexico and Central America
251
shows great variation in the shape of the
shell and in the development of the lamellae.
This variation has led to the publication of a
number of different names for this species by
various authors.
The more northern Cumingia califomica
Conrad, has a larger and thicker shell. Cw~
mingia similis A. Adams is a synonym of
Conrad’s species.
Cumingia mutica Sowerby29, which occurs
in Peru and Chile, possesses a large shell for
the genus. It is ornamented by finely decus-
sated scupture. Cumingia clerii A. Adams30,
C. grandis Deshayes31, C. striata A. Adams32
and C. ventricosa Sowerby33 were referred to
the synonymy of C. mutica by Dali.
Cumingia lamellosa Sowerby is not to be
confused with Thyella lamellosa H. Adams,
1885, described from the island of Mauritius,
later renamed Cumingia elegans by Sowerby,
1873.
Cumingia tellinoides Conrad, 1831, C. tel-
linoides coarctata Sowerby, 1833, and C. tel-
linoides vanhyningi Rehder, 1939, occur in
east American waters.
Distribution: Specimens of Cumingia la-
mellosa were taken by the expedition off
western Mexico, Nicaragua and Costa Rica.
Specimens questionably identified as this spe-
cies have been recorded as occurring in the
Pleistocene of Magdalena Bay, Lower Cali-
fornia.
Family Donacidae.
Key to the Genera of the Donacidae.
A. Inner margin crenulated Donax
B. Inner margin smooth Iphigenia
Genus Donax Linnaeus.
Key to the Species of Donax.
A. Shell more than twice as long as high
a. Anterior dorsal margin concave
transversus
aa. Anterior dorsal margin straight or
convex
b. Posterior dorsal area flattened or
rounded
c. Shell flattened ; very elongate
gracilis
29 Cumingia mutica Sowerby, Proc. Zool. Soc. London,
May 17, 1833, p. 34. "Hab. prope littora Maris Pacifici.”
Obtained "at Conception in seven fathoms, sand and mud ;
at Iquiqui in nine fathoms, gravel and mud ; at Payta in
hard clay at low water ; and at Muerte.”— Sowerby, Conch.
Icon., Vol. 19, Cumingia, 1873, species 3, pi. 1, fig. 3. "Hab.
Chili, Peru.”
30 Cumingia clerii A. Adams, Proc. Zool. Soc. London,
November 12, 1850, p. 24, pi. 8, fig. 3. "Found at Talcuhano,
Chili, by Capt. Clery, French Marine, attached to fuci in
shallow water. (Mus. Cum.) .’’—Sowerby, Conch. Icon.,
Vol. 19, Cumingia, 1873, species 2, pi. 1, fig. 2. "Hab. Chili."
31 Cumingia grandis Deshayes, Journ. de Conchyl., Vol.
5, 1857, p. 281, pi. 8, figs. 4 and 6. . . . "provient des mers
du Chili”.— Sowerby, Conch. Icon., Vol. 19, Cumingia, 1873,
species 11, pi. 2, fig. 11. “Hab. Chili”.
32 Cumingia striata A. Adams, Proc. Zool. Soc. London,
November 12, 1850, p. 25, pi. 8, fig. 6. "Hab. Conception ;
seven fathoms, sandy mud; H. C. (Mus. Cuming).”
38 Cumingia ■ ventricosa Sowerby, Conch. Icon., Vol, 19,
August, 1873, species 10, pi. 2, fig. 10. "Hab. Probably
Chili.”
cc. Shell moderately inflated;
higher calif omicus
bb. Posterior dorsal area concave ; pos-
terior area smoky-black; shell sub-
rhomboidal navicula
B. Shell less than twice as long as high
a. Shell with a sharply angled umbonal
carina posteriorly
b. Thin ; anterior end acutely rounded ;
highly polished carinatus
bb. Thick; anterior end more broadly
rounded rostratus 34
aa. Shell with a rounded umbonal angula-
tion posteriorly ; strongly sculptured
c. Interspaces punctate (with a
row of fine pits)
d. Shell subtriangular (typi-
cal), high ; posterior margin
sloping rather steeply ; ven-
tral margin sometimes
slightly expanded medially
punctatostriatus
dd. Shell elongate, lower ;flatter;
posterior margin sloping
more gently, posterior end
rostrate; base more broadly
rounded
e. Moderately elongated
contusus3i
ee. Extremely elongated ;
posterior dorsal margin
somewhat rounded
culter 34
cc. Interspaces not punctate
f. Length not exceeding
15 mm.; sculpture
finely cancellate
obesus
ff. Length exceeding 15
mm. ; sculpture
coarsely cancellate ;
thick
g. Beaks subcentral ;
shell high, trigonal
h. Ribs on poste-
rior area of
about equal
size asper
hh. Ribs on poste-
rior area with
1-3 coarser
than the others
dentiferus 34
gg. Beaks decidedly
posterior; shell
more elongated
assimilis
Donax asper Hanley.
Donax asper Hanley, Proc. Zool. Soc. Lon-
don, Pt. 13, April, 1845, p. 14. “Hab. Tumbez,
Peru (Cuming).” — Reeve, Conch. Icon., Vol.
8, Donax, September, 1854, species 12, pi. 2,
34 Not represented in the present collection.
252
Zoologica: New York Zoological Society
r34: 19
fig. 12. Original locality record cited. — Sow-
erby, Thes. Conch., Vol. 3, 1866, p. 307, pi.
280 ( Donax , pi. 1), fig. 24. Tumbez, Peru.
Donax ( Hecuba ) asper Hanley, Romer,
Syst. Conchyl.-Cab. Martini-Chemnitz, Bd.
10, Abt. 3, Donacidae, 1869, p. 14, Tab. 3, figs.
7-10. Tumbez, Peru; Puntarenas, Costa Rica,
in the Gulf of Nicoya.
Donax aspera Hanley, Dali, Proc. U. S.
Nat. Mus., Vol. 37, 1909, pp. 159, 273, pi. 28,
fig. 7. Central America to Tumbez, Peru.
Type Locality: Tumbez, Peru.
Range: Tangola-Tangola Bay, Mexico, to
Tumbez, Peru.
Collecting Stations: Mexico: Tangola-Tan-
gola Bay; Costa Rica: Port Culebra, beach;
Culebra Bay; Cedro Island, Gulf of Nicoya,
beach; Gulf of Dulce, beach; Panama: Bahia
Honda.
Description: Shell triangular, beaks sub-
central, elevated, and rather gibbous; the
anterior end is rather acutely rounded, the
posterior end somewhat concavely truncated ;
ornamented by radiating riblets which are
especially well developed toward the poste-
rior end and on the posterior area where they
are crenated by concentric lines; inner mar-
gin crenulated; the color is ashy-white or
purple.
A large right valve in the present collec-
tion from the Gulf of Dulce measures, ap-
proximately: length, 35 mm.; height, 26
mm. ; convexity (one valve), 8.5 mm.
Compared to Donax assimilis Hanley, the
shell of D. asper is much higher in propor-
tion to the length and the beaks are much
more centrally located. The shell of D. asper
differs from that of D. dentiferus Hanley35 in
that it is thicker, more acutely rounded ante-
riorly and lacks the raised ribs (1-3) which
are coarser than the others on the posterior
area of that species.
Distribution : This species was collected by
the expedition from Mexico to Panama but
nowhere in large numbers. Several single
valves were taken on the beach in the Gulf of
Dulce, Costa Rica, and at Tangola-Tangola
Bay, Mexico.
Donax assimilis Hanley.
Donax assimilis Hanley, Proc. Zool. Soc.
London, Pt. 13, April, 1845, p. 17. “Hab.
Panama. Mus. Cuming, Hanley, &c.” — Reeve,
Conch. Icon., Vol. 8, Donax, September, 1854,
species 10, pi. 2, fig. 10. Panama.
Donax panamensis Philippi, Zeit. f. Mala-
kozool., Jahrg. 5, No. 10, 1848, p. 145. “Pat-
ria: Panama.” [According to Romer, 1869,
this species is a synonym of D. assimilis.]
Donax cayennensis Lamarck, Reeve,
Conch. Icon., Vol. 8, Donax, September, 1854,
species 22, pi. 4, figs. 22a, 22b. “Hab. Panama
and St. Elena, West Columbia; Cuming.”
Not Donax caianensis Lamarck, Anim. S.
Vert., Vol. 5, July, 1818, p. 550. “Habite
35 Donax dentifera Hanley, Proc. Zool. Soc. London , Pt.
11, July, 1843, p. 6. “Hab.— T”— Reeve, Conch. Icon., Vol.
8, Donax, September, 1854, species 2, pi. 1, fists. 2a, 2b.
“Hab. Panama.”
l’Ocean de la Guyane.” — Delessert, Rec. Coq.
decrites par Lamarck et non encore figurees,
1841, pi. 6, figs. 13a, 13b. See also Hanley,
Cat. Rec. Biv. Shells, 1843, p. 82, footnote. —
Lamy, Bull. Mus. Nat. Hist. Nat. (Paris),
Vol. 20, No. 6, 1914, p. 339.
Donax reevei Bertin, Nouv. Arch. Mus.
Hist. Nat. (Paris) , Ser. 2, Vol. 4, 1881, p. 85.
[Name based upon Reeve’s pi. 2, fig. 10.
Panama (Reeve).]
Donax sowerbyi Bertin, Nouv. Arch. Mus.
Hist. Nat. (Paris), Ser. 2, Vol. 4, 1881, p. 85,
pi. 4, figs. 2a, 2b, 2c. Based upon Sowerby’s
(Thes. Conch., Vol. 3, 1866, p. 307), pi. 280
{Donax, pi. 1), fig. 21. “Panama.”
Type Locality : Panama.
Range: Mazatlan, Mexico (Carpenter), to
Santa Elena, Ecuador.
Collecting Stations: Nicaragua: Isla En-
cantada, Corinto; Costa Rica: Culebra Bay;
Cedro Island, Gulf of Nicoya, beach; Gulf of
Dulce, beach; Panama: Isla Parida, Gulf of
Chiriqui.
Description: Shell elongately triangular,
beaks posterior to the middle, anterior end
the narrower, rounded, posterior end trun-
cated ; ornamented with radial riblets which
are stronger toward the posterior end and on
the posterior area which sometimes bears a
faint subangulation ; inner margin crenated ;
color, usually some shade or combination of
gray and purple.
A large specimen of this species in the col-
lection of the California Academy of Sci-
ences, collected by James Zetek at Chame
Island, Panama, measures: length, 41 mm.;
height, 26.8 mm. ; convexity (both valves to-
gether), 17.5 mm.
Compared to Donax asper the shell of D.
assimilis is much more elongate, the beaks are
more posteriorly situated and the posterior
area is more rounded. The more posteriorly
situated beaks and more elongated shell are
characters which serve to separate the pres-
ent species from D. dentiferus.
Distribution: This species was taken by
the expedition on the beach at a few localities
from Nicaragua to Panama. It occurs com-
monly at Panama where it is used for food by
the natives.
Donax eallfornleus Conrad.
Plate I, Figs. 2, 5.
D[onax]. calif ornica Conrad, Jour. Acad.
Nat. Sci. Philadelphia, Vol. 7, 1837, p. 254, pi.
19, fig. 21. “Inhabits the coast of California
in sand, near Sta. Barbara.”
Not Donax californica Conrad, Reeve,
Conch. Icon., Vol. 8, Donax, September, 1854,
species 40, pi. 6, fig. 40. “Hab. Gulf of Califor-
nia.” [Carpenter, 1855, referring to some of
the shells in the Gulf of California region
labeled as D. calif omicus, stated, “The shells
wrongly called D. calif omicus are simply the
white variety of the forms contusus and
culter”].
Donax calif omicus Conrad, Arnold, Mem.
Calif. Acad. Sci., Vol. 3, 1903, p. 170, pi. 13,
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
253
fig. 9. Lower San Pedro Series at Deadman
Island, and San Pedro, California. Lower
Pleistocene. Also upper Pleistocene and Re-
cent.—Weymouth, State of Calif. Fish &
Game Comm., Fish Bull. No. 4, 1920, p. 47,
pi. 16, fig. 1. Localities cited from San Pedro
to False Bay, California.
Type Locality: Near Santa Barbara, Cali-
fornia, in sand.
Range: Santa Barbara, California, to Mag-
dalena Bay, Lower California.
Collecting Station: Mexico: Cedros Island,
Lower California.
Description: Shell elongated, somewhat
pointed at both extremities ; disks with very
minute radiating lines; color yellowish, ob-
scurely rayed; a brown stripe on the anterior
and posterior sub-margin ; within white and
purplish brown; margin beautifully crenu-
lated. (Original description.)
A specimen of this species in the collec-
tions of the California Academy of Sciences,
collected by Henry Hemphill at Cape San
Lazaro, Lower California, measures : length,
24.6 mm.; height, 11 mm.; convexity (both
valves together) , 7.4 mm. ; from beaks to pos-
terior end, 10 mm. Large specimens attain a
length of about 30 mm.
Donax calif ornicus can be referred to the
subgenus Serrula Chemnitz in Morch.
The posterior dorsal area of the shell of
this species is flattened or gently rounded
rather than concave as in D. navicula.
Donax gracilis Hanley is a similar south-
ern species whose shell is more elongated and
whose posterior dorsal margin slopes more
gently ventrally.
Distribution: A few small specimens of
Donax calif ornicus were dredged off Cedros
Island by the expedition. We have not seen
specimens from south of Cape San Lazaro,
Lower California36. It is also known to occur
in the Pleistocene of southern California and
western Lower California.
Donax carinatus Hanley.
Plate I, Fig. 9.
Donax carinata Hanley, Proc. Zool. Soc.
London, Pt. 11, July, 1843, p. 5. “Hab. — ?
Mus Stainforth, Metcalfe.” — Hanley, Cat.
Rec. Bivalve Shells, p. 84, 1843, p. 349, pi. 14,
fig. 28, 1856 (as Donax carinatus on expl. to
plate). [No locality cited.] — Reeve, Conch.
Icon., Vol. 8, Donax, September, 1854, species
11, pi. 2, fig. 11. “Hab. San Bias, California.”
Donax carinatus Hanley, Sowerby, Thes.
Conch., Vol. 3, 1866, p. 305, pi. 280 {Donax,
pi. 1), figs. 4 and 5. “California.” — Romer,
Syst. Conchyl.-Cab. Martini-Chemnitz, Bd.
10, Abt. 3, Donacidae, 1869, p. 10, Taf. 3, figs.
4, 5, 6. “Fundort: Der Stille Ocean bei Cali-
fornien, (St. Bias, Tumaco, Mazatlan).”
Donax culminatus Carpenter, Cat. Mazat-
as According to Miss Viola Bristol, specimens of this
species from Magdalena Bay, Lower California, are in
the collections of the San Diego Society of Natural History
{Min. Conch. Club South. Calif.. No. 47, back page, April,
1945). On the same page of this paper Eyerdam is cited
as having found this species at Corinto, Nicaragua.
Ian Shells, September, 1855, p. 43. “Hab. — 1
young specimen ; L’pool Col.”
Type Locality: San Bias, Mexico (here
designated as type locality) . No locality cited
originally.
Range: Altata, Mexico, to Tumaco, Colom-
bia.
Collecting Station: Nicaragua: Corinto
(200-D-16, 19), 4-13 fathoms, mangrove
leaves.
Description: Transversely elongated, con-
vex, very inequilateral, purplish-brown, with
more or less distinct radiating striae, (usu-
ally with obsolete darker rays and polished) ,
ventral edge little arcuated and forming a
very acute point with the nearly straight
edge of the obliquely truncated and almost
flattened anterior [posterior] slope, which
is sharply carinated and sculptured by close
decussated radiating striae: inside purple,
two lateral teeth in each valve, the ventral
edge crenated, anterior crenulated. 4/5 .. .
1 2/5 [inches] (Hanley, Cat. Rec. Biv. Shells,
1843). 1
The largest specimen in the present col-
lection measures 33.2 mm. in length and 18
mm. in height. A large right valve collected
by the senior author at Corinto, Nicaragua,
measures approximately: length, 39.4 mm.;
height, 22 mm.; convexity (one valve), 7.6
mm.
This species can be easily recognized by
the comparatively thin, polished shell with a
sharply angled umbonal ridge posteriorly.
The only other west American shell that
might be confused with Donax carinatus is
Donax rostratus C. B. Adams37. The shell of
Adams’ species is thicker, less sharply cari-
nated, and is less sharply pointed where the
carina joins the ventral margin, the anterior
end is more broadly rounded, and the color of
the exterior is lighter.
A subspecies, Donax carinatus galveston-
ensis Harris38, has been described from an
artesian well in Galveston, Texas, and was
considered to be of upper Miocene age. Harris
did not consider Reeve’s plate 2, figure 11, as
representing Hanley’s species.
Distribution: This species was dredged by
the expedition off Corinto, Nicaragua, at
depths of 4-13 fathoms. It also has been col-
lected by the senior author at Corinto and at
Mazatlan, Mexico. Romer cited the occur-
rence of the species as far south as Tumaco,
Colombia.
Donax gracilis Hanley.
Plate I, Figs. 4, 6.
Donax gracilis Hanley, Proc. Zool. Soc.
London, Pt. 13, April, 1845, p. 15. “Hab. Bay
of Guayaquil. Var. b. Chiriqui. Var. c. Bay
of Caraccas (Cuming).” — Reeve, Conch.
37 Donax rostratus C. B. Adams, Ann. Lyceum Nat. Hist.
New York, Vol. 5, July, 1852, pp. 602, 646 (separate p p.
278, 321). "Panama.”— Romer, Syst. Conchyl.-Cab. Martini-
Chemnitz, Bd. 10, Abt. 3, Donacidae, 1869, p. 11, Taf. 3,
fig9. 1-3.
38 Donax carinata var. galvestonensis Harris, Bull. Amer.
Paleo., Vol. 1, No. 3, December 2, 1895, p. 92 (10). Well at
Galveston, Texas, depth, 2,652 to 2,920 feet. Upper Miocene.
254
Zoological New York Zoological Society
[34: 19
Icon., Vol. 8, Donax, September, 1854, species
38, pi. 6, fig. 38. “Hab. Gulf of Guayaquil;
Cuming.” — Sowerby, Thes. Conch., Vol. 3,
1866, p. 314, pi. 282 {Donax, pi. 3), figs. 76,
77, 78, 79. Gulf of Guayaquil. — Romer, Syst.
Conchyl.-Cab. Martini-Chemnitz, Bd. 10,
Abt. 3, Donacidae, 1869, p. 80, Taf. 14, figs.
4, 5, 6. [?7, 8]. “Fundort: Der Stille Ocean
bei Mittelamerika und Ecuador, (Guayaquil,
Chiriqui, Panama).”
Type Locality: Bay of Guayaquil, Ecuador
(here selected as type locality). Chiriqui
[Panama] and Bay of Caraccas [Ecuador]
also cited originally for varieties of this
species.
Range: Lat. 24° 18' N., west coast of Lower
California, to the Gulf of California and
south to Negritos, Peru.
Collecting Station: Nicaragua: Corinto
(200-D-10, 11, 16, 17, 19), 7-13 fathoms,
sand, mangrove leaves, also in beach drift.
Description: Shell narrowly elongate, pol-
ished, rather compressed, beaks nearer the
posterior end; anterior end acutely rounded,
posterior end acutely roundly pointed; pos-
terior dorsal margin straight or slightly pro-
duced; inner margin finely crenulated; color
of the exterior is usually brown and that of
the interior brownish-purple.
A large left valve from off Potosi and
Monypenny Point, Nicaragua, measures ap-
proximately: length, 22.5 mm.; height, 9.4
mm.; convexity (one valve), 3 mm.; distance
from beak to posterior end, 9 mm.
The shell of Donax gracilis differs from
that of D. navicula in the much more elon-
gate outline and in that the posterior dorsal
margin is straight or slightly produced
rather than concave. Compared to D. calif or-
nicus the shell of D. gracilis is longer in pro-
portion to the height, more compressed, the
posterior dorsal margin slopes more gently
and the posterior end is more acutely pointed.
It can be referred to the subgenus Serrula.
Donax punaensis Pilsbry & Olsson39, de-
scribed from the Pliocene of Ecuador, is a
very similar species but the beaks appear
to be more centrally situated and the pos-
terior end is more broadly rounded.
Donax petersoni Olsson, described from
the Oligocene of Peru, is a somewhat similar
species.
Donax owenii Gray in Hanley40 of the At-
lantic fauna appears to be somewhat similar
to D. gracilis but the posterior end appears
38 Donax punaensis Pilsbry & Olsson, Proc. Acad. Nat.
Sci. Philadelphia, Vol. 93, September 9, 1941, p. 72, pL 12,
fig. 2. "Pliocene of the north end of Puna Island.’’ Ecuador.
40 Donax owenii Gray in Hanley, Cat. Rec. Bivalve Shells,
1843, p. 81. "Africa ?”— Reeve, Conch. Icon., Vol. 8, Donax,
September, 1864, species 37, pi. 6, fig. 37. "Hab. West Coast
of Africa.”
Dali ( Nautilus , Vol. 6, No. 4, August, 1891, p. 44),
recorded this species from Montevideo and Maldonado.
Maury (Serv. Geol. & Min. Brasil, Mon. No. 4, 1924, p.
466) also cited the species as occurring at Montevideo,
Uruguay. Melvill & Standen recorded a species under the
name of Donax ( Machaerodonax ) owenii Gray, from
Karachi, India (Proc. Zool. Soc. London, November 13,
1906, p. 826). Carcelles recently cited Donax owenii as
occurring along the coast of Argentina (Rev. Mus. de
La Plata (New Ser.), Sec. Zool., VoL 8, 1944, p. 303).
to be shorter and the margin is said to be
smooth not crenulated.
Distribution : Specimens of Donax gracilis
were taken by the expedition in the beach
drift as well as dredged in 7 to 13 fathoms
at Corinto, Nicaragua.
Donax navteula Hanley.
Plate I, Fig. 1.
Donax navicula Hanley, Proc. Zool. Soc.
London, Pt. 13, April, 1845, p. 15. “Hab. Gulf
of Nicoya, Central America (Cuming)”. —
Reeve, Conch. Icon., Vol. 8, Donax, Septem-
ber, 1854, species 18, pi. 4, fig. 18. Original
locality cited. — Sowerby, Thes. Conch., Vol.
3, 1866, p. 314, pi. 282 {Donax, pi. 3), fig.
80. Original locality cited. — Romer, Syst.
Conchyl.-Cab. Martini-Chemnitz, Bd. 10,
Abt. 3, Donacidae, 1869, p. 56, Taf. 10, figs.
1-3. “Fundort: Der Stille Ocean bei Cali-
fornien und Mittelamerika, (Nicoyia, Pana-
ma, Mazatlan, Reallejos).”
Type Locality : Gulf of Nicoya, Costa Rica.
Range: Gulf of California to Panama.
Collecting Stations : Mexico: Santa Inez
Bay, Gulf of California; Cape San Lucas;
Cape San Lucas Bay; Nicaragua: Potosi and
Monypenny Point, Gulf of Fonseca; Corinto
(200-D-10, 16), 4-7 fathoms, mangrove
leaves, also in beach drift.
Description-. Shell elongately rhomboidal,
moderately inflated, fairly thick, obsoletely,
finely radially grooved; anterior end the
longer, acutely rounded, posterior end
acuminately truncated, the extremity round-
ly pointed; posterior area concave; ventral
margin rounded and somewhat expanded just
anterior to the middle, often with a com-
pressed area between this portion of the
valve and the posterior angulation; inner
margin crenulated; color white or brownish
with the posterior and anterior dorsal areas
black or grayish-black; periostracum green-
ish.
One of the largest specimens, a right valve,
from the beach drift at Corinto, Nicaragua,
measures approximately: length, 21 mm.;
height, 10 mm.; convexity (one valve), 3.9
mm. ; distance from beaks to posterior end,
8.5 mm.
The shell of Donax navicula differs from
that of D. gracilis in the more rounded base
and rhomboidal form, more inflated valves
and especially in that the posterior area is
concave. It can be referred to the subgenus
Serrula.
The concave posterior area and higher
more triangular form are features which
serve to separate Donax navicula from D.
calif ornicus.
Distribution -. This species was taken abun-
dantly by the expedition in the beach drift
at Corinto, Nicaragua. A few specimens were
also taken as far north as Santa Inez Bay in
the Gulf of California.
Donax obesus d’Orbigny.
Plate I, Fig. 7.
Donax obesa d’Orbigny, Voy, Am6r.
1949]
Hertlein & Strong: Mollusks of Mexico and Central America
255
Merid., Vol. 5, 1846, p. 541, pi. 81, figs. 28,
30. “Elle a ete pechee a Payta (Perou) par
M. Fontaine.” — Reeve, Conch. Icon., Vol. 8,
Donax, October, 1854, species 49, pi. 7, fig.
49. “Hab. Real Llejos, Central America;
Cuming.”
Donax obesus d’Orbigny, Sowerby, Thes.
Conch., Vol. 3, 1866, p. 310, pi. 281 {Donax,
i pi. 2), figs. 42, 43. “Real Llejos, Central
i1 America.”
Type Locality. Paita, Peru.
Range: Corinto, Nicaragua, to Paita,
Peru.
Collecting Station: Nicaragua: Corinto
(200-D-ll, 19), 8-13 fathoms, mangrove
leaves.
Description: Shell small, subtriangular,
inflated; the anterior end the longer, slop-
ing, acutely rounded at the extremity, pos-
terior end broadly and roundly truncated ;
umbonal ridges rounded; ornamented with
fine radial grooves which are crossed by
J somewhat flexuous concentric grooves form-
ing a fine cancellated pattern of punctate
appearance ; inner margin finely crenulated ;
color white with dark purplish-brown on
' the posterior end and on the anterior dorsal
margin.
The largest specimen in the collection, a
right valve, measures: length, 13.4 mm.;
height, 10.9 mm.; convexity (one valve), 4.2
mm.
The shell of Donax obesulus Reeve41 is
much more abruptly truncated than that of
D. obesus, and the posterior umbonal ridge
is decidedly angulated rather than rounded.
Distribution: This species was dredged by
the expedition at two localities off Corinto,
Nicaragua, at depths of 8-13 fathoms.
Donax punctatostriatus Hanley.
Plate I, Fig. 17.
Donax punctato-striata Hanley, Proc. Zool.
Soc. London, Pt. 11, July, 1843, p. 5. “Hab. —
? Mus. Stainforth, Metcalfe, Hanley, &c.” —
Hanley, Cat. Rec. Biv. Shells, 1843, p. 84,
pi. 14, fig. 24 (as Donax punctato-striatus
on expl. to plate) . [Not the record “China”].
— Reeve, Conch. Icon., Vol. 8, Donax, Sep-
tember, 1854, species 16, pi. 3, figs. 16a, 16b.
“Hab. Mazatlan, Gulf of California.” —
Sowerby, Thes. Conch., Vol. 3, Donax, 1866,
p. 310, pi. 281 {Donax, pi. 2), figs. 49, 50 (as
Donax punctato-striatus). Reeve’s locality
cited on expl. to pi.
Type Locality: Mazatlan, Mexico (here
designated as type locality) . No locality cited
originally.
Range: San Ignacio Lagoon, Lower Cali-
fornia, to the Gulf of California and south
to Negritos, Peru.
Collecting Stations: Mexico: Santa Inez
Bay, Gulf of California; Cape San Lucas;
Chamela Bay; Tenacatita Bay; Sihuatanejo;
•11 Dona x obesula Reeve, Conch. Icon., Vol. 8, Donax,
September, 1854, species 30, pi. 5, fig. 30. “Hal). Peru.”-
Deshayes, Proc. Zool. Soc. London, 1854 (issued May 16,
1855), p. 352. “Hab. Central America.”
Tangola-Tangola Bay; Nicaragua: Gulf of
Fonseca; Potosi and Monypenny Point; Co-
rinto (200-D-19), 12-13 fathoms, mangrove
leaves, also beach.
Description: Subtriangular, very convex,
pale livid brown, with strong radiating punc-
tated striae, becoming very fine and close on
the anterior [posterior] slope, whose edge
is rounded, posterior and anterior edges
much sloping, ventral arcuated in the middle ;
inside stained with violet, the ventral mar-
gins dentated, the anterior [posterior]
crenulated : two cardinal and lateral teeth
in each valve. Long. 4/5 — 1-1/5 [inches]
( Hanley, Cat. Rec. Biv. Shells, 1843) . “When
full grown less inequilateral than most of
this genus.”
A very large specimen of this species in
the Henry Hemphill collection in the Cali-
fornia Academy of Sciences, collected at
Magdalena Bay, Lower California, mea-
sures: length, 44.8 mm.; height) 30 mm.;
convexity (both valves together), 17.4 mm.
The subtrigonal form and the row of fine
pits, which occur in the radial grooved striae,
are characteristic features of this species.
It belongs to the subgenus Chion Scopoli.
Hanley once reported Donax punctato-
striatus from China but it was later recog-
nized as occurring commonly in tropical west
American waters.
The variety described by Carpenter as
Donax punctatostriatus var. caelatusi2 ap-
pears not to have been recognized since its
description. It was described as possessing
short impressed lines rather than pits in the
interspaces.
Sowerby (1866) pointed out that there are
specimens which intergrade between Donax
punctatostriatus and Donax conradi Reeve43.
The latter species is now known to be iden-
tical with D. contusus Reeve44 (see our PI. I,
fig. 14), as pointed out by Tomlin45. Donax
vellicata Reeve (fig. 66) and D. bitincta
Reeve (fig. 68), both described without in-
formation as to the locality from which they
came, are likewise identical with D. contusus
according to Tomlin.
Typical forms of Donax punctatostriatus
are more subtrigonal and higher than those
of typical D. contusus, which are lower and
more elongate in outline. Large specimens
of D. punctatostriatus are sometimes slightly
expanded medially along the ventral margin
and the posterior dorsal margin often slopes
more steeply than that on D. contusus, but as
mentioned by Sowerby, there is intergrada-
tion between the two forms. The most elon-
gate form of this variable group is Donax
42 Donax ? punctatostriatus, var. cuelatus Carpenter, Cat.
Mazatlan Shells, September, 1855, p. 46. "Hab.— Mazatlan :
very rare.”
13 Donax conradi Reeve, Conch. Icon., Vol. 8, Donax.
Sentember, 1854. species 20, pi. 5, fife. 29. “Hab. Gulf of
California.”
Donax contusus Reeve, Conch. Icon., Vol. 8. Donax,
September, 1854, species 24, pi. 4, fig. 24. “Hab. Mazatlan,
Gulf of California.”
45 Tomlin, J. R. leB., Nautilus, Vol. 40, No. 2, October,
1926, p. 52.
256
Zoologica: New York Zoological Society
[34: 19
culter Hanley46, as pointed out by Car-
penter47. Very elongate forms of D. culter
are somewhat reminiscent of Amphichaena
kindermanni Philippi48 as is Donax petal-
linus Reeve49.
Donax aricanus Dali50, recorded as occur-
ring from Paita, Peru, to Arica, Chile, ap-
pears to be very similar to D. punctato-
striatus and possibly some of the records of
the occurrence of the latter species in South
America may be referable to Dali’s species.
Romer considered D. radiatus Valenciennes
[ =aricanus ] to be only a variety of D. punc-
iatostriatus.
Donax striatus Linnaeus, which occurs in
the Caribbean region, is a similar species.
Distribution : This species was collected
by the expedition on the beach and dredged
at depths of 12-13 fathoms, from Santa Inez
Bay in the Gulf of California to Corinto,
Nicaragua. It is a variable shell found com-
monly from the Gulf of California to Peru.
It has been recorded as occurring in the
Pleistocene of southern California, Magda-
lena Bay, Lower California, and at Oaxaca.
Mexico.
Donax transversus Sowerby.
Plate I, Fig. 3.
Donax transversa Sowerby, Cat. Shells
Tankerville, 1825, Ap., p. IV. [No locality
cited]. — Reeve, Conch. Icon., Vol. 8, Donax,
September, 1854, species 36, pi. 6, fig. 36.
“Hab.— ?”
Donax transversus Sowerby, Sowerby,
Thes. Conch., Vol. 3, 1866, p. 306, pi. 280
(Donax, pi. 1"), fig. 11. “Hab. — ?” — Car-
penter, Cat. Mazatlan Shells, September,
1855, p. 44. Mazatlan, Mexico.
Type Locality : Corinto, Nicaragua (here
designated as type locality) . No locality cited
originally.
Range: Mazatlan, Mexico, to San Juan del
Sur^ Nicaragua.
4fi Donax culter Hanley, Proc. Zool. Soc. London, April,
1845, p. 14. ‘‘Hab. Var. a. Matzellan [Mazatlan] Gulf of
California (Cumins). Var. b. Acapulco (Cuming).”—
Reeve, Conch. Icon., Vol. 8, Donax, September, 1854, species
21, pi. 4, fig. 21. "Hab. Gulf of California.”
47 Carpenter, P. P., Cat. Mazatlan Shells, September,
1855, pp. 47-48.
4S See Palmer. R. H., and Hertlein, L. G., Bull. South.
Calif. Acad. Sci., Vol. 35, Pt. 2, May-August (issued Sep-
tember 10), 1936, p. 71, pi. 18, figs. A, B, C. pi. 19, figs. 5,
6. 7, 8, 9, 10. Mazatlan ; Petatlan Bay ; Tenacatita Bay,
Mexico, Recent. Also Oaxaca, Mexico, Pleistocene.
49 Donax petallina Reeve, Conch. Icon., Vol. 8, Donax,
October, 1854, species 51, pi. 8, fig. 51. "Hab.— ?”— Deshayes,
Proc. Zool. Soc. London for 1854 (issued May 16, 1855), p.
350 (as Donax petalina) . “Hab.— ? Coll. Cuming.”
This species was described without information as to the
locality from which it came. Bertin ( Nouv . Arch. Mus.
Hist. Nat. (Paris). Ser. 2, Vol. 4, 1881, p. 84). stated
that he found an indication in Deshayes’ collection that the
species came from Chile. Dali, 1909, and Gigoux, 1934, also
cited it from that country. Pilsbry & Lowe, 1932, and Bales,
1938, cited it from Acapulco, Mexico.
50 Donax radiata Valenciennes, Rec. d’Obser. Zool. Hum-
boldt & Bonpland, Vol. 2, 1832, p. 221, pi. 1, figs. 3a, 3b, 3c,
4. "Habitat in Oceano Pacifico ad Americae calidioris
litora.” Not Donax radiata Gmelin, 1791.
Donax aricana Dali, Proc. U. S. Nat. Mus., Vol. 37,
November 24, 1909, p. 273. New name for Donax radiata
Valenciennes, 1832, not D. radiata Gmelin, 1791. Paita,
Peru, to Arica, Chile.
Collecting Station: Nicaragua: Corinto
(200-D-19) , 12-13 fathoms, mangrove leaves.
Description: “D. testa transversim elon-
gata, laevi; latere postico brevi, biangulato,
carinato, oblique truncato, longitudinaliter
sulcato; extusalbida; fulvo obsolete radiata”.
(Original description).
The shell of this species is very elongated,
very inequilateral, rather thin, moderately
inflated, gaping at each end, polished and
obsoletely radially striated ; anterior dorsal
margin slightly concave; anterior end ellip-
tically rounded obliquely joining the slightly
rounded ventral margin ; posterior end trun-
cated, set off by a carina, the area fairly
broad, concave, with a faint rounded angula-
tion medially, the whole area finely radially
striated and at the end obliquely truncated ;
color yellowish-white with purple rays; ven-
tral margin finely crenated.
Specimens of this species in the present
collection are small. A large left valve col-
lected at Corinto, Nicaragua, by the senior
author measures: length, 36.4 mm.; height.
14.5 mm.: convexity (one valve), 4.5 mm.;
distance from beak to posterior end, 14 mm.
This appears to be the species which
authors have cited from western Mexico and
Central America under the name of Donax
scalpellum Gray. Donax scalpellum Gray51
was originally described without informa-
tion as to locality. Hanley52 later cited it from
“California” and Reeve53 cited it from the
Gulf of California. E. A. Smith54 in 1891
cited it from Aden, in the Gulf of Aden, and
stated : “The above named locality, given by
Reeve (Conch. Icon. sp. 39), has never been
confirmed, and I think there is little doubt
that it is incorrect. The specimens from Aden
agree in form, color, sculpture, and every
other respect with that figured by Reeve.”
Later Melvill & Standen55 cited the species
as occurring at Karachi, India, in the Ara-
bian Sea.
The concavity of the anterior dorsal mar-
gin, obliquely elliptically rounded anterior
end, and much wider and radially striated
posterior area are features separating Donax
transversus from D. scalpellum. Donax
transversus belongs to the subgenus Machae-
rodonax Romer, the type of which is D. scal-
pellum Gray.
Distribution: Three small specimens of
Donax transversus were dredged by the ex-
pedition in 12-13 fathoms off Corinto, Nica-
ragua. It also has been collected by the senior
author on the beach at the same locality.
si Donax scalpellum Gray. Ann. Philos., Vol. 25, Feb-
ruary, 1823. p. 136. TNo locality cited]. -Wood, Index Test.,
Suppl., 1828, p. 4. pi. 2, Donax, fig. 1. [No locality cited].
52 Index Test, by W. Wood. edit, by S. Hanley, 1856,
p. 202, Suppl. pi. 2, Donax fig. 1. "California.”
53 Reeve. L. A.. Conch. Icon., Vol. 8, Donax, September,
1854, species 39, pi. 6, fig. 39. "Hab. Gulf of California.”
54 Smith, E. A., Proc. Zool. Soc. London, 1891, p. 427.
55 Melvill, J. C., and Standen, R., Proc. Zool. Soc.
London, November 13, 1906, p. 826.— Melvill, Proc. Malacol.
Soc. London. Vol. 18, Pt. 3, 1928, p. 115.
19491
Hertlein & Strong: Mollusks of Mexico and Central America
257
Genus Iphigenla Schumacher.
Iphigenia Schumacher, Essai Nouv. Syst.
Test., 1817, pp. 51, 155. Sole species, Iphi-
genia laevigata ( Donax laevigata Chem-
nitz.). Ref. to Chemnitz, Vol. 6, p. 253, pi. 25,
fig. 249. Illustrated by Schumacher on pi. 17,
figs. 4a, b. — Dali, Trans. Wagner Free Inst.
Sci., Vol. 3, Pt. 5, 1900, p. 962. Type : Donax
laevigata Chemnitz. — Pilsbry & Bequaert,
Bull. Amer. Mus. Nat. Hist., Vol. 53, Art. 2,
May 9, 1927, p. 369. Donax laevigata “Chem-
nitz” Gmelin accepted as type.
Type (by monotypy) : Donax laevigata
Chemnitz TNeues Syst. Conchyl.-Cab., Bd. 6,
1782, p. 253, Tab. 25, fig. 249. “Es wohnet
diese Muschel in den ostindischen Meeren.
Bey Tranqueber wird sie nur selten ge-
funden”. Also illustrated by Schumacher,
pi. 17, fig. 4a, b].
Shell large, subtriangular, subequilateral,
without radial sculpture; thick, with entire
ventral margins; two cardinals, the larger
bifid, in each valve and two obsolete laterals
in the right valve. (Dali).
The genus Iphigenia is known to occur in
the Miocene of Venezuela and Peru. At the
present time it often occurs in estuarine or
brackish water conditions and is known to
occur in greatest abundance from the coast
and rivers of West Africa and from the
tropical Atlantic and Pacific coasts of Central
and South America. Iphigenia centralis
Germain, an African species, ranges from
the middle Niger river to strongly saline
waters. One species occurs on the coast of
Florida and one species occurs in tropical
west American waters.
Iphigenia altier Sowerby.
Capsa altior Sowerby, Proc. Zool. Soc. Lon-
don, Pt. 2. for 1832 (issued March 13, 1833,)
p. 196. “Hab. in Peruvia et America Cen-
trali.” “Dredged among coarse gravel, in
twelve fathoms water, in the Gulf of Nocoiyo.
A smaller variety, which is also rather high-
er, was found at Tumbez, at a depth of five
fathoms, in thin mud.” — Hanley. Cat. Rec.
Bivalve Shells, o. 86, 1843. pi. 14, fig. 34,
1844, p. 349, 1856. Peru and Central Amer-
ica.— Romer, Syst. Conchyl.-Cab. Martini-
Chemnitz. Bd. 10, Abt. 3, Donacidae, 1869.
p. 114, Tab. 21, figs. 1-4. Earlier records
cited.
Iphiaenia ambigua Bertin, Nouv. Arch.
Mus. Hist. Nat. (Paris). Ser. 2. Vol. 4. 1881.
p. 120, pi. 4, figs. 4a, 4b, 4c . . . “habite l’ocean
Pacifique, sur les cotes de l’Amerique cen-
trale.”
Iphigenia altior Sowerby, Dali, Proc. U. S.
Nat. Mus.. Vol. 37. 1909. p. 159, pi. 25. fig. 8.
Capon to Tumbez, P^ru. Range, Gulf of Cali-
fornia to Tumbez, Peru.
Type Locality : Gulf of Nicova, Costa Rica,
In 12 fathoms, coarse gravel. Tumbez. Peru,
also cited originally for a small variety of
this species.
Range: Gulf of California to Tumbez,
Peru.
Collecting Stations: Mexico: Chamela
Bay, beach; Nicaragua: Potosi and Mony-
penny Point; Corinto, beach; Costa Rica:
PortCulebra; Culebra Bay; Golfito Bay; one
mile south of Golfito Bay.
Description: Shell subtriangular, the an-
terior side the longer, thick, moderately in-
flated, base rounded, anterior dorsal margin
gently arcuate and sloping, rounded at the
end, posterior dorsal margin more steeply
sloping, more flattened and subtruncated at
the end, a slight depression often present
anterior to the posterior umbonal ridge;
smooth except for lines of growth and submi-
croscopic radiating striae ; two cardinal teeth
in each valve, the right posterior and left
anterior bifid or medially grooved; inner
margin smooth; pallial sinus extends for
about five-eighths the length of the shell,
rounded at the end, and along the base for
about a third of its length confluent with the
pallial line ; color yellowish or purplish-white
under an olive periostracum, the umbos dark,
the interior white and violet.
A very large right valve in the present
collection from one mile south of Golfito Bay,
Costa Rica, measures: length, 79 mm.;
height, 57.3 mm.; convexity (one valve), 18
mm.; pallial sinus extends anteriorly 45 mm.
from the posterior margin of the shell. A
specimen in the collection of the California
Academy of Sciences collected in Panama
Bay by F. M. Anderson, measures: length,
68 mm.; height, 51.5 mm.; convexity (both
valves together), 31 mm.; pallial sinus ex-
tends anteriorly 39 mm. from the posterior
margin of the shell.
The specimen described as Iphigenia
ambigua by Bertin may be slightly longer
in proportion to the height as compared to
some specimens of I. altior. In the absence
of any other differences and in view of the
variation shown in a series of shells, we have
nlaced Bertin’s species in the synonymy of
I. altior. Carpenter56 mentioned an elongate
specimen from Mazatlan with a strong nos-
terior ventral sinus which he referred to
“Iphigenia flaevigata, ?cujus.” According
to Carpenter, Gray considered the snecimen
to be an abnormal variety of I. altior. Tn
general features I. altior is quite similar to
I. laevigata, the type of the genus, from West
Africa.
Compared to Iphigenia brasiliana La-
marck, which occurs in the Caribbean region,
the shell of I. altior is considerably higher
proportionately from beak to base and it has
a much fainter depression anterior to the
posterior truncation.
Distribution: Specimens of Iphigenia
altior were collected by the expedition on
beaches from Chamela Bay, Mexico, to Gol-
fito Bay, Costa Rica. Dali mentioned that this
species was found at a depth of 4 to 6 inches
in sand, on flats and tidal lagoons of Peru.
38 Carpenter, P. P., Cat. Mazatlan Shells, September,
1856, p. 42.
258
Zoologica: New York Zoological Society
[34: 19: 1949]
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
EXPLANATION OF THE PLATE.
Donax navicula Hanley. Hypotype,
left valve, from Corinto, Nicaragua.
Length, 18.8 mm.; height, 9.2 mm. P.
Donax calif ornicus Conrad. Hypotype,
left valve, from San Pedro Bay, Cali-
fornia. Length, 22 mm.; height, 9.8
mm. P
Donax transversus Sowerby. Hypo-
type, left valve, from Corinto, Nicara-
gua. Length, 36.6 mm.; height, 14.5
mm. P
Donax gracilis Hanley. Hypotype, left
valve, from Potosi and 5 miles west of
Monypenny Point, Nicaragua. Length,
22 mm.; height, 9.7 mm. P
Donax calif o miens Conrad. View of
right valve of the specimen shown in
Fig. 2.
Donax gracilis Hanley. Hypotype, left
valve, from Loc. 27588 (C.A.S.), about
13 miles southeast of Cape Tosco,
Santa Margarita Island, west coast of
Lower California. Collected by the
Templeton Crocker Expedition, 1932.
Length, 15.8 mm.; height, 7.3 mm. P.
Donax obesus d’Orbigny. Hypotype,
left valve, from Station 200-D-19, Lat.
12° 28' 03" N., Long. 87° 12' 39" W.,
Corinto, Nicaragua, in 12-13 fathoms
(22-24 meters). Length, 11.1 mm.;
height, 8.9 mm. P
Semele sparsilineata Dali. Hypotype,
left valve, from Station 221-D-l, Lat.
7° 54' 45" N., Long. 82° 04’ 32" W.,
Gulf of Chiriqui, Panama, in 35 fath-
oms (64 meters). Length, 22 mm.;
height, 17.4 mm. P
Donax carinatus Hanley. Hypotype,
left valve, from Corinto, Nicaragua.
L. G. Hertlein, coll. Length, 36.3 mm.;
height, 18.2 mm. P
Semele quentinensis Dali. Hypotype,
right valve, from Station 199-D-l, Lat.
13° 08' N., Long. 87° 43' W., Mean
guera Island, Gulf of Fonseca, El Sal-
vador, in 16 fathoms (29 meters).
Length, 26.4 mm.; height, 21.1 mm. P.
Semele pacifica Dali. Hypotype, right
valve, from Golfito, Gulf of Dulce,
Costa Rica. Length, 19.4 mm.; height,
15.4 mm.
Semele jaramija Pilsbry & Olsson.
Hypotype, left valve, from Santa Inez
Bay, Lower California, in the Gulf of
California, on shore. Length, 16.1 mm.;
height, 12 mm. P
Fig. 13. Semele venusta Reeve. Hypotype,
right valve, from Station 196-D-8, Lat.
15° 45' 37" N., Long. 96° 05' 54" W.,
Tangola-Tangola Bay, Mexico, in 9
fathoms (16.3 meters). Length, 17.8
mm.; height, 13 mm. P
Fig. 14. Donax contusus Reeve. Hypotype, left
valve, from Loc. 27230 (C.A.S.), Pe-
tatlan Bay, Mexico, about 6 miles south
of Sihuatanejo. L. G. Hertlein, coll.
Length, 39.1 mm.; height, 20 mm. (Il-
lustrated for comparison with Donax
punctatostriatus, Fig. 17). P
(in text).
Fig. 15. Semele pulchra Sowerby. Hypotype,
right vaive, from Potosi and 5 miles
SSW. of Monypenny Point, Nicaragua.
Length, 31 mm.; height, 26 mm. P. . . .
Fig. 16. Abra palmeri Dali. Hypotype, right
valve, from Station 199-D-l, Lat. 13°
08' N., Long. 87° 43' W., Meanguera
Island, Gulf of Fonseca, El Salvador,
in 16 fathoms (29 meters). Length, 9.8
mm.; height, 9 mm. P
Fig. 17. Donax punctatostriatus Hanley. Hypo-
type, left valve, from Loc. 4859 (C.A.S.
H. Hemphill coll.), Magdalena Bay,
Lower California. Length, 44.4 mm.;
height, 28 mm. P
Fig. 18. Abra palmeri Dali. View of the inte-
rior of the specimen shown in Fig. 16.
Fig. 19. Semele craneana Hertlein & Strong,
sp. nov. Holotype, left valve, dredged
in the Gulf of California. (Exact sta-
tion unknown but probably in the
southern portion of the Gulf of Cali-
fornia). Length, 38 mm.; height, 29.5
mm. View of the interior. P
Fig. 20. Abra palmeri Dali. Hypotype, left
valve, from Station 199-D-l, Lat. 13° I
08' N., Long. 87° 43' W., Meanguera I
Island, Gulf of Fonseca, El Salvador, I
in 16 fathoms (29 meters). Length, |
10.5 mm.; height, 9.3 mm. View of the I
interior. P
Fig. 21. Semele verrucosa Morch. Hypotype, Ij
right valve, from Station 224, Lat. 7° fl
23' 30" N., Long. 82° 03' W., Hannibal I
Bank, Panama, in 35-40 fathoms (64-
73 meters). Length, 43 mm.; height, fc
32.8 mm. View of the interior. P
Fig. 22. Semele craneana Hertlein & Strong, |
sp. nov. Holotype. View of the exterior I
of the specimen shown in Fig. 19.
Fig. 23. Abra palmeri Dali. View of the exte- I
rior of the specimen shown in Fig. 20. I
Fig. 24. Semele verrucosa Morch. View of the I
exterior of the specimen shown in Fig. 1
21.
All the specimens illustrated on this plate are I
in the type collection of the Department of j
Paleontology of the California Academy of >|
Sciences.
HERTLEIN & STRONG.
PLATE 1.
MOLLUSKS FROM THE WEST COAST OF MEXICO AND CENTRAL AMERICA. PART VIII
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Metcalf : Tettigellidae and Gyponidae of Kartabo
259
20.
Tettigellidae and Gyponidae (Homoptera) of Kartabo,
Bartica District, British Guiana.
Z. P. Metcalf.
College of Agriculture and Engineering, University of North Carolina, Raleigh, North Carolina.
(Text-figures 1-8).
[This contribution is a result of various
expeditions of the Department of Tropical Re-
search of the New York Zoological Society to
British Guiana dui’ing the years 1917, 1919,
1920, 1921, 1922 and 1924, under the direction
of Dr. William Beebe. For maps and ecological
data refer to Zoologica, Vol. VI, 1925, pp. 1-193.]
Contents.
Page
Introduction 259
Family Tettigellidae 259
Subfamily Tettigellinae 260
Erythrogonia Melichar 260
Erythrogonia bicolor n. sp 260
Amblyscarta Stal 260
Amblyscarta aurulenta Fabricius 260
Orectogonia Melichar 262
Orectogonia flavoscutellata Signoret 262
Subfamily Proconiinae 262
Acrocompsa Stal 262
Acrocampsa pallipes Fabricius 262
Acrocampsa rufa Melichar 264
Dichrophleps Stal 264
Dichrophleps despecta Melichar 264
Poeciloscarta Stal 264
Poeciloscarta quadrifasciata Linnaeus 264
Poeciloscarta nigrofasciata nom. nov. 266
Raphirhinus de Laporte 266
Raphirhinus phosphoreus Linnaeus 266
Raphirhinus fasciatus Fabricius 268
Capinota Melichar 268
Capinota virescens n. sp 268
Rhopalogonia Melichar 268
Rhopalogonia purpurata Germar 269
Family Gyponidae 269
Key to Genera of Gyponidae 269
Gypona Germar 272
Gypona fusiformis Walker 272
Gypona thoracica Fabricius 272
Gypona bigemmis Spangburg 272
Gypona flavolimbata n. sp 273
Gypona translucens n. sp 273
Gypona picturata n. sp... 273
Gypona opaca n. sp 275
Gypona castanea n. sp 275
Ponana Ball 275
Ponana fulva n. sp 275
Clinonaria gen. n 277
Clinonaria bicolor n. sp 277
Marganalana gen. n 277
Marganalana testacea n. sp 277
Scarisana gen. n 277
Scarisayia variabilis n. sp 277
Scaris Le Peletier and Serville 279
Introduction.
In 1945 the writer (Metcalf, 1945b) re-
viewed the Fulgoroidea from Kartabo col-
lected by Dr. William Beebe and his asso-
ciates. Included with these were a small num-
ber of leafhoppers belonging to the groups
Tettigellidae (fomerly Cicadellinae) and
Gyponidae. While the number of species in
the present collection is not large, they are
of sufficient interest to be worth reporting. A
number apparently belong to species previ-
ously described and in these cases I have at-
temped to redescribe and illustrate these so
as to bring them into line with modern taxo-
nomic practices. A disproportionate number
of species are apparently new. This number,
however, does not seem to be out of line with
the apparent number of new species from
other parts of the world where, likewise, only
incidental collections of these small and often
inconspicuous insects have been made. No
species has been described as new, neverthe-
less, until a thorough search has been made
of the literature and careful comparisons
made with the original descriptions of these
species. As is well known to the students of
these groups, the older descriptions are often
totally inadequate. The writer is under nq
delusions as to his ability to place these older
species accurately from the descriptions.
However, since so many of these types are
not readily available, the best the modern
taxonomist can do is to try to correlate these
older described species with the material at
hand.
For the present the writer proposes to
treat these two groups, Tettigellidae and
Gyponidae, along with some other groups of
leafhoppers, as families. Whether they are
coordinate with other families of Homoptera
or with families in other orders has not as
yet been determined. But until the groups of
the Homoptera have been studied more thor-
oughly, it is deemed best to treat them as
distinct families of the superfamily Ias-
soidea.
Family TETTIGELLIDAE.
Most of the species of this family are large
or medium sized leafhoppers, very few are
small. The body is cylindrical or more or less
fusiform. The head is usually large and in a
few’ species is produced into a definite ce-
phalic process. The anteclypeus is large. The
lora conspicuous. The postclypeus large,
sometimes inflated, definitely projected onto
the crown, facial portion usually with dis-
tinct transverse ridges. The lateral post-
clypeal sutures usually extending to the
crown. Crown large, usually distinctly tri-
angular in outline. Cephalic process some-
times distinct, short, triangular or elongate
260
Zoologica: Nerv York Zoological Society
[34: 20
terete. Paired ocelli on the crown remote
from the anterior border of crown. Pronotum
usually large, flat, more or less quadrangular
in outline; anterior margin usually broadly
curved; posterior margin usually broadly in-
cised ; lateral margins usually distinct. Meso-
notum small, triangular. Tegmina elongate,
usually narrow, coriaceous; venation usually
distinct, sometimes strongly reticulate api-
cally. Radius with two distinct branches;
media unbranched; cubitus 1 unbranched,
joined to the claval suture (cubitus 2) by a
short but usually distinct cross vein. Usually
two, (sometimes three or more) radial-
medial cross veins and one (sometimes two)
medial-cubital cross vein. Thus there is
formed usually two (sometimes three) basal
cells, two or three ante-apical cells and four
or five apical cells, with the ante-apical and
apical cells sometimes strongly reticulate.
Wings transparent with a distinct submar-
ginal vein which is incomplete on the costal
margin. Radius with two branches, the first
incomplete. Media with two branches. Media
1 plus 2 with a distinct radial cross vein and
media 3 plus 4 with a distinct cubital cross
vein. Cubitus 1 unbranched.
Subfamily Tettigellinae.
Head, including compound eyes, usually
narrower than the pronotum. Antennal ledges
indistinct, not conspicuously separated from
the coronal portion of the postclypeus. Lat-
eral margins of the pronotum divergent
caudad ; or rounding, not distinctly separated
from posterior lateral margins. Anterior
tibiae not sulcate or expanded. Tegmina
broader, usually covering the lateral mar-
gins of the abdomen.
Erythrogonia Melichar, 1926a :373.
Orthotype Erythrogonia laeta Fabr.
The species of this genus are generally
small slender tettigellids with the head in-
cluding the eyes wider than the pronotum.
The tegmina elongate, narrow with nearly
parallel sides.
Ci'own shorter than the greatest width be-
tween the eyes ; the ocelli usually on the pos-
terior half; face somewhat angled to the
crown, strongly impressed. Pronotum usually
about as long as the crown, width usually
greater than the median length. Mesonotum
large. Legs slender. Tegmina elongate,
rounded at the apex with three elongate ante-
apical cells and three shoi'ter apical cells.
Erythrogonia bicolor n. sp.
Text-fig. 1 A-E.
This species closely resembles Tettigonia
rutilans Walker ( Tettigonia carminata Sig-
noret) in general coloration but differs as fol-
lows: the apical margin of the tegmina
translucent, not blackish, and the head, more
elongate. Since neither the male nor female
genitalia of rutilans have been described and
since the head characters of the present
species are decidedly different, together with
the fact that rutilans has been recorded only
from Brazil, it is believed to be better to de-
scribe this as a new species.
Crown about as long as broad, slightly im-
pressed from the posterior lateral margins
around the anterior margin, leaving the ante-
rior margin as a broad slightly elevated bor-
der; ocelli on a line with the anterior margin
of the compound eyes; face strongly pro-
duced, longer than its greatest width. Prono-
tum as long as crown, about one and a half
times as broad as the median length; the
anterior margin broadly curved; the poste-
rior margin nearly straight; mesonotum
large, nearly as long as pronotum.
Female last ventral segment four times as
long as the penultimate, longer than broad,
curvingly acuminate to the obtuse apex with
a distinct median carina. The male genitalia
with the genital plates with a broad base,
slender, triangular acuminate apex about
half as long as the pygofers; the pygofers
when viewed ventrad, elongate, triangular,
gradually acuminate from the base to the
apex, not as long as the anal segment; when
viewed laterad, broad with the apex broadly
rounded; genital styles short, acuminate,
broadly curved outward. Aedeagus shorter
than pygofers with a strongly developed dor-
sal lobe and two elongate acuminate spines at
the apex.
This is a small slender species with the
head including the compound eyes, the pro-
and mesonotum, and the basal angle of the
tegmina blackish fuscous. Beneath, including
the legs and abdomen, blackish fuscous. Teg-
mina bright red with the venation brown.
Wings transparent with scarlet red veins.
Crown black, almost as long as its width be-
tween the compound eyes.
Length to apex of tegmina: 9.75 mm.
Holotype: $ Kartabo, Bartica District;
May 10, 1924. Allotype: 2 Kartabo, Bartica
District; July 4, 1922. Paratypes: 1 $ July 4.
1922; 1 3 May 10, 1924; 3 29 May 10, 1924.
Amblyscarta St&l, 1869a :71.
Logotype Amblyscarta modesta Fabr.
In this genus the head is broader than the
pronotum. The crown is short and broad,
length on the median line more than half the
width between the eyes, broadly rounded to
the face ; face somewhat elongate. Pronotum
about twice as long as the crown. Mesonotum
large, almost as broad as the pronotum. Teg-
mina elongate, apical margin somewhat ob-
tuse ; antei’ior tibiae with a distinctly ciliated
inner margin.
Amblyscarta aurulenta Fabr.
( Cicadella aurulenta Osborn, 1926b :199).
(Text-fig. 1 F-H).
This common and well-known species is
appax-ently widely distributed in South
America, having been previously recorded
fx'om Bx-itish, French and Dutch Guiana, Bo-
livia and Matto Grosso and Rio de Janeiro,
Brazil. Thex*e are numex-ous specimens in the
1949]
Metcalf : Tettigellidae and Gyponidae of Kartabo
261
Text-fig. 1. Erythrogonia bicolor : A, head thorax; B, face; C, male genitalia ventral;
D, male genitalia lateral; E, female genitalia. Amblyscarta aurulenta : F, head thorax;
G, face; H, female genitalia. Orectogonia flavoscutellata : I, head thorax; J, face; K, fe-
male genitalia.
262
Zoologica : Neiv York Zoological Society
[34: 20
present collection from Kartabo, Bartica Dis-
trict, British Guiana.
While the color is quite variable, the head,
pronotum and mesonotum are chiefly ochra-
ceous orange with the legs and venter bright
yellowish, the abdomen usually carmine red.
The tegmina vary considerably in color and
color markings. More typical specimens in
the present collection have the basal third of
the tegmina velvety brown with numerous ir-
regular light bluish spots, the apex testa-
ceous brown, the posterior border of the
crown irregularly irrorate with brownish
and the posterior third of the pronotum usu-
ally marked like the tegmina with a broad
brownish band more or less irregularly
spotted with light bluish. The brown spot at
the apex of the head usually distinctive.
Female with the last genital segment more
than twice as long as the penultimate, pro-
jecting caudad in a broad triangular lobe;
the lateral margins usually projecting as a
pair of small lateral teeth.
Length to apex of tegmina: 9.5-11.0 mm.
Orectogonia Melichar, 1926a :345.
Logotype Orectogonia sparsuta Sign.
This is one of the genera established by
Melichar in his key that has never been fully
described. It may be ehax*acterized as fol-
lows: head including the compound eyes
broader than the pronotum; crown elongate,
conical, flat with the disc distinctly impressed
with a very faint longitudinal furrow on the
median line. Face somewhat inflated. Prono-
tum large, lateral margins parallel; anterior
margin broadly curved; posterior margin
triangularly incised. Mesonotum large, near-
ly as broad as pronotum. Caudal area dis-
tinctly produced. Tegmina translucent; vena-
tion distinct with three ante-apical and four
apical cells, the fourth apical cell elongate.
Legs short and slender, posterior tibiae with
about six short stout spines on the outer
margin; the anterior margin with numerous
slender hairs, those on basal half short, those
on apical half elongate, posterior margin
with numerous short hairs.
Orectogonia flavoscutellata Signoret.
( Tettigonia flavoscutellata Signoret,
1855c: 509).
(Text-fig. 1 I-K.)
This species is appax-ently very close to
Orectogonia sparsuta Sign., but it differs
decidedly in coloration and in size.
Crown longer than the width between the
eyes, the median impressed line rather in-
distinct. Ocelli large, close to compound eyes.
Face strongly inflated ; anteclypeus small.
Pronotum slightly wider than long. Meso-
notum large, distinctly bi-impressed behind
the middle. Tegmina elongate, rather nar-
row; venation distinct, the ante-apical cells
neax'ly three times as long as the apical cells.
Claval veins distinct. Legs short and slendex-.
Female genitalia with the last ventx-al seg-
ment elongate, deeply incised on the poste-
rior border, the incision almost x'eaching the
anterior max-gin of the segment.
Genex-al color of the head, including the
compound eyes and the thorax, blackish with
numerous small x'ound spots, yellowish ox-
pale bluish-gx-een,with a few larger irx-egular
spots on the cx-own and the px-onotum, and a
lax-ge central area on the mesonotum, pale
yellowish. Tegmina tx-anslucent, the veins
blackish. Beneath genex-al color blackish with
a large median yellowish spot on the apex of
the face and a pair of large yellowish spots
at base of antennae, irx-egular yellowish spots
on the sides of the thox-ax and the venter of
the abdomen. Legs chiefly yellow with the
postex-ior tibiae and the base of the posterior
tarsi chiefly blackish.
Length to the apex of the tegmina: 11 mm.
Single female specimen from Kartabo,
Bax-tica District, British Guiana, Max-ch 18,
1922.
Subfamily Proconiinae.
Head, including compound eyes, usually
bx-oader than the pronotum. Antennal ledges
distinct, pi-ojecting, conspicuously separated
fx-om the coronal portion of the postclypeus.
Lateral margins of the pronotum usually
parallel. Anterior tibiae sulcate or expanded
toward the apex. Tegmina narrow, not cov-
ering the latex-al max-gins of the abdomen.
Acrocampsa Stal.
(Melichar, 1925a :337).
Haplotype Fulgora pallipes Fabr.
Head including the compound eyes wider
than the pronotum; crown suddenly con-
stricted in fx'ont of the compound eyes, then
broadly triangularly produced to the apex
which is slightly uptux-ned. Apex with a deep
longitudinal furx-ow both above and below.
Face strongly px-oduced ; anteclypeus viewed
laterad somewhat angular. Pronotum narrow
with the sides parallel; anterior max'-
gin broadly curved; sepax-ated into two dis-
tinct pax-ts — the anterior part depx-essed,
about the same level as the crown ; posterior
pax*t sti'ongly elevated, trilobed. Mesonotum
broad, not vex-y long; apex px-oduced into a
rather slender process. Anterior tibiae broad-
ened at the apex; postex-ior tibiae elongate,
nearly twice as long as the femox*a, with a
few rather stout spines on the anterior and
posterior ventral margins; basitarsi not
longer than the two apical segments. Teg-
mina x-ather narrow, coriaceous; venation
not very distinct; the whole surface finely
rugulose. Costal and commisux-al margins
nearly pax-allel; apical margin obtuse.
Acrocampsa pallipes Fabr.
( Fulgora pallipes Fabx*icius, 1787a: 261).
(Text-fig. 2 A-F).
This species may be x-eadily distinguished
from Catorthorrhinus resimus by the follow-
ing combination of characteristics and dis-
tinctive colox*ation :
1949]
Metcalf : Tcttigellidae and Gyponidae of Kartabo
263
Text-fig. 2. Acrocampsa pallipes: A, head thorax; B, face; C, lateral head thorax;
D, male genitalia ventral; E, male genitalia lateral; F, female genitalia. Acrocampsa
rufa : G, male genitalia ventral.
264
Zoologica: New York Zoological Society
[34: 20
Crown somewhat longer than greatest
width between the eyes, equalling the pro-
notum in length; cephalic process short,
broadly rounded, deeply impressed, semi-
erect; ocelli large, projecting; compound eyes
large ; face elongate, nearly twice as long as
greatest width; postclypeus nearly ellip-
tical in outline; median impression on face
deep.
Female last ventral segment broad, not
quite twice as broad as median length ; caudal
area nearly quadrangular; posterior margin
shallowly incised with a median third broadly
produced with a median incision at the apex.
Male genitalia last ventral segment short
and broad, nearly four times as broad as its
median length ; genital plates nearly as long
as pygofer; aedeagus elongate with pair of
elongate, slender, acute apical processes.
Length to apex of tegmina: 13.5 mm.
Acrocampsa rufa Melichar.
(Melichar, 1925a :339).
(Text-fig. 2 G).
This species may be recognized by the fol-
lowing points: crown, pronotum and meso-
notum chiefly yellowish, more or less clouded
with light brown. Tegmina chiefly bright
reddish-brown, base usually greenish-fus-
cous ; apex transparent bordered with black-
ish basad. Beneath, including the legs and
venter of the abdomen, chiefly pale ochra-
ceous.
Male genitalia last ventral segment about
three times as broad as its median length;
genital plates slightly shorter than pygofers,
obtuse; aedeagus less than half as long as
genital plates.
Length to apex of tegmina: 13.5 mm.
Dichrophleps Stal.
(Melichar, 1925a :327).
Haplotype Cicada aurea Fabr.
This genus has recently been reviewed by
Melichar. He describes this genus and gives
a key to the species including aurea Fabr.
The species of this genus are quite variable
in color markings and it is by no means cer-
tain that there is more than one species in
Central and South America.
Head, including compound eyes, broader
than pronotum; crown flat. Anterior margin
acutely parabolic; face fairly flat. Pronotum
rather cylindric, lateral margins impressed ;
mesonotum large, triangular ; tegmina trans-
parent; venation distinct; three ante-apical
and four apical cells, inner apical cell rather
large. Legs rather slender, hind tibiae elon-
gate, twice as long as the femora.
Dichrophleps despecta Mel.
(Melichar, 1925a: 330).
(Text-fig. 3 A-E).
I place the series of specimens in the
present collection under this name because
they resemble more closely the color mark-
ings described by Melichar for this species
than they do the other species from British
Guiana. Melichar describes the head as
shorter than the pronotum, whereas in the
present series the crown is longer than the
pronotum.
The genital plates of the male when viewed
ventrad are short, triangular, somewhat
asymmetrical at the apex; the pygofers elon-
gate, narrow, nearly as long as the anal
segment, somewhat truncate at the apex, the
whole surface set with short stout spines;
the aedeagus elongate, stout, somewhat mem-
braneous with a pair of ventral spines which
are asymmetrical, the right spine being al-
most as long as the pygofer, the left spine
shorter than the aedeagus; the apical area
of the aedeagus is thin, irregularly broken
into lobes. When viewed laterad the pygofer
is about half again as long as its greatest
width, thickly set with short, fairly stout
spines.
Length to apex of tegmina: 16.3 mm.
Redescribed from 5 males from Kartabo,
Bartica District, British Guiana. Dec. 5,
1920; April 4, 1924; April 6, 1924.
Poeciloscarta Stal, 1869a :73.
Cardioscarta Melichar, 1932a :285).
Logotype Tettigoniella ( Poeciloscarta ) car-
dinalis Fabr.
Stal established this genus as a subgenus
of Tettigonia as indicated above, for thirteen
species from South America. Melichar estab-
lished the new genus Cardioscarta for species
from America and assigned Stal’s genus to
species from Madagascar and Africa. On
what basis this was made has never been
revealed as Melichar did not finish his work
on this group.
The genus may be defined briefly as fol-
lows: head including compound eyes broader
than the pronotum; crown somewhat elon-
gate, usually distinctly broader than long;
anterior margin broadly parabolic; face
elongate, not impressed. Pronotum broader
than long, longer than the crown; anterior
margin broadly curved; posterior margin
nearly straight. Mesonotum large, shorter
than the pronotum. Tegmina with the basal
area of the corium and the clavus coriaceous
with the venation indistinct ; apex beyond the
clavus translucent.
Poeciloscarta quadrifasciata Linnaeus.
(Cardioscarta fasciata Melichar, 1932a:300) .
(Text-fig. 3 F-K).
There has been a great deal of confusion
in the nomenclature of this species. It was
described originally as Cicada quadrifasciata
Linnaeus, 1767a : 706. In this he was followed
by the earlier subsequent writers who merely
repeated his short but distinctive description
and added the reference to the original de-
scription. Goeze, 1778a: 130; Fabricius,
1781a :329, 1787a :274, 1794a :51 and 1803a:
72; de Tigny, 1802a: 152; Donovan, 1820a:
T6] ; and Stoll, 1788a: 82; pi. XXI, fig. 114A
19491
Metcalf: Tettigellidae and Gyponidae of Kartabo
265
Text-fig. 3. Dichrophleps despecta: A, head thorax; B, face; C, tegmen; D, male
genitalia ventral; E, male genitalia lateral. Poeciloscarta quadrifasciata: F, head thorax;
G, face; H, male genitalia ventral; I, aedeagus ventral view; J, male genitalia lateral;
K, female genitalia.
266
Zoologicu : New York Zoological Society
[34: .20
and 1792a : 64 ; pi. XXI, fig. 114A described
and illustrated this species as fasciata Lin-
naeus. In this they were followed by Blan-
chard, 1840a: 190, and again in 1850a: 190.
This species varies somewhat in color from
light yellow to brilliant orange. Marked
dorsad with four broad blackish fasciae, the
first across the compound eyes, the posterior
margin of the crown and the anterior margin
of the pronotum ; the second across the basal
part of the tegmina beyond the sutural angle ;
the third across the apex of the clavus ; and
the fourth at the apex of the tegmina. The
dark bands on the tegmina are quite variable
in width and in color. The band at the apex
of the tegmina is especially variable and is
in some specimens reduced to a small fuscous
spot with the apex of the tegmina translu-
cent fuscous. The wings ax-e translucent
smoky with two blackish fuscous bands simi-
lar to the middle and the apical bands of the
tegmina. The dorsal part of the abdomen is
more or less marked with black; the venter
and legs usually pale yellowish.
Crown broader than long, distinctly im-
pressed in front of the compound eyes, broad-
ly rounded to the face; the anterior margin
parabolic; the posterior margin broadly
sinuate. Pronotum quadrate; the anterior
margin broadly rounded, the postei*ior mar-
gin nearly straight; face narrow, the ante-
clypeus large and protuberant.
Female genitalia with the last ventral seg-
ment elongate, nearly three times as long as
the penultimate, the posterior border deeply
and roundly emarginate; the male genitalia
with the anal segment longer than the pygo-
fers; the pygofers elongate, when viewed
ventrad, broadened from the base to the mid-
dle and then constricted to the narrow apices ;
genital plates nearly as broad at the base as
the last ventral segment, suddenly con-
stricted at the middle and continued as a
bi ramose process for half the length of the
pygofers; the aedeagus elongate, stout with
a biramose process, ventrally at the apex of
the basal third, and a pair of elongate acute
spines at the base of the apical third.
Length to apex of tegmina: 9.5 mm.
Described from numerous specimens from
Kartabo, Bartica District, British Guiana.
Melichar distinguishes another species
under the name of Cardioscarta quadrifas-
ciata Fabr. under the assumption that Cer-
copis quadrifasciata Fabr. was a distinct spe-
cies from Linnaeus’s Cicada quadrifasciata.
This cannot be true, however, as Fabricius,
1781a : 329, merely copies Linnaeus’s desci*ip-
tion and cites Cicada quadrifasciata as a syn-
onym. Melichar distinguishes the two spe-
cies in his key on the basis that fasciata
Linnaeus (quadrifasciata Linnaeus) has two
narrow bands on the tegmina, quadrifas-
ciata Fabr. has three broad bands. This can-
not be correct, however, as all of the de-
scriptions and illustrations of the true quad-
rifasciata Linnaeus show three broad bands
on the tegmina. As shown above quadrifas-
ciata Fabr. is the same as quadrifasciata
Linnaeus and 1 would synonymize quadrifas-
ciata Melichar with quadrifasciata Linnaeus
but for the fact that Melichar describes the
male genitalia of the latter species as having
the genital plates short; whereas in fasciata,
that is, the true quadrifasciata Linnaeus, the
genital plates are described as long. Until we
can know more about these two species, it
would seem best to give a new name to Mel-
ichar’s quadrifasciata Fabr. and I propose
Poeciloscarta nigrofasciata nom. nov. for
Cardioscarta quadrifasciata Melichar nec
Cercopis quadrifasciata Fabr. This species
should not be confused with Tettigonia
fabricii which Signoret proposed for Ful-
gora fasciata Fabr. under the assumption
that fasciata Fabr. was a secondary hom-
onym of Cicada fasciata [sic] Linnaeus. But
since fasciata Linnaeus is a typographical
error for quadrifasciata Linnaeus and be-
longs to the genus Poeciloscarta, and fasciata
Fabr. belongs to the genus Raphirhinus, the
correct synonymy for Fabricius’ species is
as follows:
Raphirhinus fasciatus Fabr.
Fulgora flammea [nec Linnaeus] Stoll,
1781a :28; pi. VI, fig. 29.
Fulgora fasciata Fabricius, 1787a: 261.
Raphirhinus obliquatus de Laporte, 1832d :
415.
Tettigonia fabricii Signoret, 1855c :521;
pi. 21, fig. 12.
Raphirhinus de Laporte, 1832d :413.
Orthotype Fulgora abscendens Fabr.
i.e., Fulgora phosphorea Linnaeus.
This is one of the most distinct genera of
the subfamily Proconinae. It may be charac-
terized briefly as follows :
Head including compound eyes as wide as
the pronotum suddenly constricted in front
of compound eyes, triangularly produced to
the obtuse apex which is continued into an
elongate, terete, erect, somewhat recurved
slender process. Pronotum usually slightly
broader than long; lateral margin slightly
divergent; anterior margin broadly curved,
the posterior margin triangularly incised, sin-
uate. Mesonotum nearly as long as the prono-
tum, triangular. Legs slender; the anterior
tibiae broadly expanded apically. Tegmina
elongate, slender, coriaceous; venation fairly
distinct.
Raphirhinus phosphoreus Linnaeus.
(Melichar, 1925a :354).
(Text-fig. 4 A-F).
This species is very variable in color. The
females are chiefly ferruginous, heavily
speckled with light yellow dots dorsad. Ven-
trad chiefly ochraceous. Males chiefly black-
ish fuscous, dorsad with the median area of
cephalic process, lateral margins of crown,
and pronotum yellowish testaceous. Beneath
chiefly light testaceous yellow with postcly-
peus pro- and meso- plura twice banded
1949]
Metcalf : Tettigellidae and Gyponidae of Kartabo
267
Text-fig. 4. Raphirhinus phosphoreus : A, head thorax; B, face; C, lateral head thorax;
P, male gerntalia ventral ; E, male genitalia lateral; F, female genitalia. Capinota
mresceps ; G, head thorax; H, face; I, male genitalia ventral; J, male genitalia lateral;
K, female genitalia. ■ • o
268
Zoologica : New York Zoological Society
[34: 20
with blackish fuscous; the fore tibiae and
middle tarsi and hind tarsi apically blackish
fuscous.
Female last ventral segment slightly
longer than preceding segment; the apical
margin on the median third shallowly bisin-
uate with a broad obtuse tooth. Male geni-
talia with pygofers, genital plates and anal
segments all approximately the same length.
The genital plates elongate, somewhat tri-
angular, obtuse at the apex. The aedeagus
capitate with the lateral margins strongly
produced.
Length to apex of tegmina: 13.7-18.5 mm.
British Guiana, French Guiana and Suri-
nam. Kartabo, Bartica District.
Raphirhinus faseiatus Fabr.
( Fulgora faseiatus Fabricius, 1787a: 261).
This common and well-known species has
been recorded previously from Surinam,
French Guiana, Peru, Bolivia and in Brazil
from Amazonas, Para and Bahia. It may
be recognized by its large size and its char-
acteristic color and markings. The dorsal
surface is chiefly testaceous brown with a
central vitta on the cephalic process, lateral
margins of the crown, most of the prono-
tum, the basal area of the tegmina, a nar-
row fascia beyond the apex of the mesono-
tum, a broader fascia covering the apex of
the clavus and an irregular fascia just be-
fore the apical cells, grayish or greenish-
yellow. Beneath, the color is pale yellow
with two narrow brownish vittae extending
across the face, the lateral margins of the
head, the pro- and meso-pleura to the base
of the tegmina. The dorsal vitta extends
just below the compound eyes and the lateral
margins of the pronotum; the ventral vitta
just above the clypeal suture and then curved
upward to the base of the tegmina. The tips
of all the tarsi and tibiae blackish fuscous.
The female last ventral segment is longer
than the penultimate with the lateral mar-
gins broadly sinuate; the median area
broadly and shallowly sinuate, with a broad
obtuse tooth on the median line.
Capinota Melichar, 1926a :319.
Orthotype Capinota fowleri Mel.
This genus was described originally for
a single species from Mexico. The present
collection contains a single species which I
assigned to this genus with some hesitation,
but apparently the present species should be
included here.
Head including compound eyes broader
than pronotum. Crown elongate, produced,
suddenly but not strongly constricted in
front of the compound eyes. Apex broadly
upturned, impressed on the median line; lat-
eral margins of the impression slightly car-
inate; a fine median carina from the base to
the indented apex. Ocelli behind the anter-
ior margin of the compound eves. Face
strongly inflated. Anteclypeus rather large,
strongly inflated. Pronotum slightly broader
than long; anterior margin broadly rounded;
lateral margins nearly parallel; posterior
margin slightly indented. Mesonotum large,
nearly as long as the pronotum. Legs slender ;
anterior tibiae not grooved; posterior tibiae
elongate, about twice as long as the femora;
basitarsus elongate, longer than the other
two segments combined. Tegmina narrow;
costal and commisural margins nearly par-
allel, coriaceous; venation regular but not
very distinct: claval veins parallel.
Capinota virescens n. sp.
(Text-fig. 4 G-K).
This species may be recognized by its gen-
eral fuscous color above with a decided green-
ish cast; beneath chiefly ochraceous yellow
with the tibiae of the legs chiefly ochraceous
orange; ventral portion of the postclypeus
clouded with blackish; anteclypeus chiefly
blackish fuscous on the median line. Crown
nearly one and one-half times as long as the
width betwen the compound eyes. Ocelli near-
ly three times as far from each other as from
the compound eyes. Pronotum slightly broad-
er than long; sides nearly parallel; the whole
surface finely but irregularly rugulose.
Male genitalia with the genital plate broad-
ly triangular at the base, deeply incised on
the median line. Inner caudal angles pro-
duced into two elongate, slender processes.
Genital styles elongate, slender, curving in-
ward, hooked at the apex. Pygofers large
with a distinct ridge on the inner ventral
margin. Aedeagal strut elongate, slender,
acuminate and recurved at the apex. Aedea-
gus elongate, needle-like, sharply elbowed on
basal third; basal area broadly Y-shaped
when viewed ventrad. Anal segments elon-
gate, terete, with a pair of ventral hooks at
the base. Anal style elongate, slender, some-
what sagittate with a pair of distinct leaf-
like processes arising at the base of the anal
segment and covering most of the anal seg-
ment and anal style laterad and dorsad.
General color of the male above fuscous
with a greenish cast; lateral margins of the
crown, anterior border of the pronotum. and
costal margin of the male greenish-yellow.
Beneath ochraceous yellow except the con-
spicuous blackish cloud on the anteclypeus
and postclypeus and the ochraceous orange
tibiae and tarsi. Dorsum of the abdomen
blackish fuscous; lateral margins ochraceous
yellow: tips of tegmina translucent.
Length to apex of tegmina: 10.52 mm
Holotype $: British Guiana without def-
inite date or locality.
Allotype 9: Kartabo, Bartica District.
British Guiana, Sentember 20. 1922
Paratvpes: 1 S : Kartabo. Bartica District.
British Guiana. August 13. 1920; 1 o: Brit-
ish Guiana without definite date or locality
Rhopalogonia Mel.
(Melichar, 1926a :341).
Logotype Rhopalogonia scita Walk.
Head broad, including compound eyes
1949]
Metcalf : Tettigellidae and Gyponidae of Kartabo
269
broader than pronotum ; crown much broader
than long, broadly rounded to face. Ocelli
placed near the posterior border of the crown
and near the compound eyes. Pronotum
broad, nearly twice as broad as the median
length. Mesonotum broad, not as long as the
pronotum. Tegmina coriaceous, elongate,
narrow; apical margin broadly rounded ; ven-
ation indistinct, regular with four apical and
three ante-apical cells, claval veins nearly
parallel, complete. Legs elongate, slender; ail
tibiae more or less quadrangular; fore tibiae
slightly longer than femora, middle tibiae
more elongate and hind tibiae nearly twice as
long as the femora.
Rhopalogonia purpurata Germ.
( Tettigonia purpurata Germar, 1821a :63).
( Tettigonia purpurata Signoret,
1853b :325, PI. 8, Fig. 2).
(Tetigonia purpurata Osborn, 1926b :200).
(Text-fig. 5 A-E).
There is a small series in the present col-
lection which I believe represents this spe-
cies as described by Germar and described
and illustrated by Signoret, despite the dif-
ferences in coloration.
Head short and broad, crown nearly three
times as broad as the median length, dis-
tinctly bi-impressed. Impressions including
the large ocelli connected by a broad trans-
verse groove; posterior border broadly cari-
nate; anterior border broadly curved; post-
erior margin nearly straight. Face inflated,
the postclypeus distinctly angulate near the
middle. Pronotum about half again as broad
as the median length, anterior margin broad-
ly rounded, posterior margin nearly
straight; mesonotum large, broader than
long.
Female last ventral segment longer than
broad, more than three times as long as the
penultimate with a broad triangular median
tooth and a pair of small lateral triangular
teeth at the apex. Male genitalia with the
genital plates, anal segment and pygofers
all about the same length. Genital plates elon-
gate, slender, obtuse at the apex; pygofers
elongate, slender, very obtuse at the apex;
aedeagus about half as long as the genital
plates, broadened into two distinct lobes
apically, with the outer posterior angle
broadly produced ; genital styles elongate,
slender, curving outward ; apices reflexed.
General color chestnut brown, usuallv dis-
tinctly lighter on the head, with a distinct
round spot at the apex of the crown, blackish
fuscous. Pronotum chiefly chestnut brown
with a broad fascia across the middle, grav-
ish-white. Tegmina crossed bv two bluish-
white fascia, one immediately posterior to
the apex of mesonotum and the other near the
anex of the clavus. Venter including legs
chiefly chestnut brown.
Length to anex of tegmina: 11.0-11.5 mm.
Redescribed from a series of four speci-
mens, two from Georgetown. British Guiana,
and two from Bartica District, British Gui-
ana. This species has pi'eviously been re-
corded from Surinam, French Guiana, and
various localities in Brazil.
Family Gyponidae.
In this family the body is elongate, de-
pressed. The head is usually broad but nar-
rower than the pronotum at the posterior
lateral angles. The crown is usually longer
than broad with the ocelli on the disc. The
anterior margin of the crown is thin and
foliaceous, or broadly thickened or rounded
to the face. The face is usually broad, usu-
ally suddenly widened at the level of the an-
tennae and forming a distinct antennal ledge.
The postclypeus is narrow and elongate. The
lora conspicuous. The genae broad and flat.
The pronotum is large with the anterior
lateral margins rounding into the anterior
margin with the anterior lateral angles in-
conspicuous. Posterior lateral angles usually
conspicuous. The posterior lateral margins
distinct. The posterior margin usually broad-
ly incised. Mesonotum large, nearly as broad
as long. Anterior and intermediate femora
and tibiae with distinct spines. Posterior
tibiae with the internal and external dorsal
margins and the external ventral margin
with several fairly close set heavy spines;
the internal ventral margins fringed with
numerous more or less hair-like spines. Teg-
mina coriaceous or translucent; venation
distinct, sometimes reticulate over the whole
surface or the apical third only; radius 2
branched; media unbranched; cubitus 1 with
two branches; cubitus 2 unbranched; first
and second anal veins distinct. There are
typically a single cross vein between the
radial sectors; two radio-medial cross veins;
and two medio-cubital cross veins thus form-
ing three ante-apical and four apical cells.
Key to Genera of Gyponidae.
A. Head with a cephalic process.
Ohausia Schmidt, 1911b :299
AA. Head without a cephalic process.
B. Tegmina uniformly deeply pitted,
body dorsoventrally wedge-shaped
anteriorly, transversely wedge-
shaped posteriorly, tegmina verti-
cal, laterally compressed at apex.
Dragonana Ball and Reeves,
1927a: 489
BB. Tegmina not pitted except along
veins, tegmina not appressed at
apex.
C. Tegmina with numerous reti-
culate veins, at least on apical
portion.
1. Tegmina rugose or rough-
ened, often with white mot-
tling, crown and pronotum
usually rugose, two round
black spots on pronotum.
Rugosana DeLong, 1942a: 64
1. Tegmina with venation often
prominent but not rugose or
270
Zoologica: New York Zoological Society
[34: 20
Text-fig. 5. Rhopalogonia purpurata : A, head thorax; B, face; C, male genitalia ventral;
D, male genitalia lateral; E, female genitalia. Gypona thoracica : F, head thorax; G, face;
H, male genitalia ventral; I, male genitalia lateral. Gypona bigemmis : J, female genitalia.
19491
Metcalf: Tettigellidae and Gyponidae of Kartabo
271
roughened, crown and pro-
notum never rugose.
Gyponana Ball, 1920a :85
CC. Tegmina without reticulate
veins, typically with four api-
cal and three subapical cells.
1. Face narrow, deeply exca-
vate; pronotum and tegmina
coarsely rugose.
Rhogosana Osborn, 1938a: 14
1. Face not deeply excavated,
flat or inflated 2
2. Crown acutely angled with
front, margin thin, sharp, or
foliaceous 3
2. Crown with thick margin or
broadly rounding to front 6
3. Crown flat with longitudinal
furrows, ocelli on furrow
each side of median line.
Tegmina usually marked
with brownish dots or lines
in areoles.
Prairiana Ball, 1920a :90
3. Crown without longitudinal
furrows, tegmina usually
without brownish markings
in areoles, or if present with-
out small brown punctate
spots on vertex 4
4. Body usually dorsoventrally
thickened, pronotum and
crown depressed anteriorly,
strongly sloping to anterior
foliaceous margin 5
4. Body usually dorsoventrally
flattened, pronotum and
crown almost flat or gently
sloping to foliaceous margin.
Acusana DeLong, 1942d :57
5. Color black, green, or orange-
yellow; crown with narrow
longitudinal stripes, or two
round black spots on prono-
tum, or both.
Male styles short, blunt, or
truncate at apex.
Gypona Germar, 1821a : 73
5. Color green or brown, with-
out stripes on crown or
round black spots on prono-
tum. Male styles long,
slender, apex with curved,
pointed tips.
Hamana DeLong, 1942d:85
6. Crown with distinct thick
margin 7
6. Crown without definite mar-
gin, broadly and evenly
rounded to front as viewed
laterad 11
7. Face strongly inflated or
bulbous .....8
7. Face not inflated, almost
straight in profile, color
some shade of yellow or
brown usually with distinct
dark markings 9
8. Crown elongate, ocelli near
anterior margin. Mesonotum
not longer than pronotum.
Bnlbana DeLong , 1942d : 107
8. Crown short and broad.
Ocelli about equi-distant
from anterior and posterior
margins. Mesonotum longer
than the pi’onotum.
Scaris LeP. and Serv.
9. Pronotum conspicuously
wider than crown, ocelli
large, located almost half
the length of crown from
anterior border.
Ponana Ball, 1920a :93
9. Pronotum scarcely wider
than crown, ocelli anterior
to middle in the depression
above margin 10
10. Anterior margin of crown
quadricarinate.
Marganalana Metcalf
10. Anterior margin of crown
not quadricarinate.
Marganana DeLong,
1948b: 101
[ Margana DeLong,
1942d:1091
11. Ocelli on anterior border of
crown.
Polana DeLong, 1942d:110
11. Ocelli on disc 12
12. Head including compound
eyes nearly as broad as pro-
notum.
Scaroidana Osborn,
1938a :49
12. Head including compound
eyes decidedly narrower
than pronotum 13
13. Crown short and broad, usu-
ally more than three times
as broad as its median
length 14
13. Crown and pronotum longer,
usually not more than twice
as broad as their median
lengths.
Sc-arisana Metcalf
14. Pronotum short and broad,
more than three times as
broad as median length ;
humeri prominent; the an-
terior lateral and posterior
lateral margins about the
same length.
Clinonana Osborn, 1938a: 13
14. Pronotum not more than
twice as broad as long;
humeri not prominent; an-
terior lateral margins long-
er than the posterior lateral
margins.
Clinonaria Metcalf
272
Zoologica : New York Zoological Society
[34: 20
Gypona Germar, 1821a :73.
Haplotype Cercopis glauca Fabr.
This is a large genus of about 153 species,
widely distributed from Canada, the United
States, to Mexico, Central and South Amer-
ica as far south as Argentina and Chile. The
genus formerly contained many species which
are now distributed in other genera of the
family Gyponidae. Many species now in-
cluded in the genus Gypona undoubtedly be-
long to other genera which have been pro-
posed recently. The correct disposition of
many of the species described by Spangberg
and earlier students has not been determined.
There are in the present small collection
no less than 10 species of which 8 seem to be
new. The genus may be described briefly as
follows: body usually dorsoventrally thick-
ened; pronotum and vertex depressed ante-
riorly, strongly sloping from the posterior
margin of the pronotum to the anterior
foliaceous margin of the crown which may
be slightly upturned. Head narrower than
the pronotum ; crown usually elongate, some-
times almost as long as the width between
the eyes. Ocelli variously placed on the
crown; the anterior margin of the crown
thin and foliaceous; face flat. Pronotum
usually large with the anterior margin about
as wide as the greatest width of the eyes;
lateral margins strongly divergent, usually
distinctly carinate; anterior and posterior
margins strongly curved ; mesonotum large ;
venation of the tegmina simple, the veins
not punctate, with three ante-apical and four
apical cells. Anterior and intermediate legs
slender, short; posterior tibiae elongate with
stout, close-set spines on the lateral and
ventral borders. Male styles usually short
and blunt at the apex. In color the species
are usually decidedly variable with con-
siderable sexual dimorphism. The females
are usually light green or orange yellow,
frequently unmarked, sometimes with nar-
row vittae on the crown or two black spots
on the anterior border of the pronotum or
both. Males are frequently black above, with
or without pale vittae.
Gypona fusiformis Walker, 1858b :257.
There is a single female specimen in the
present collection which agrees with Walk-
er’s short description. It may be described
briefly as follows: head distinctly narrower
than the pronotum; crown elongate, more
than half as long as the pronotum; the an-
terior margins nearly parallel for a short
distance in front of the compound eyes, then
broadly and obtusely angulate. Ocelli behind
the middle about as far from each other as
from the compound eyes. Pronotum large,
the lateral margins strongly divergent, an-
terior margin broadly rounded. Mesonotum
large.
Tegmina translucent with the median
apical cell clouded with blackish fuscous.
Head, pronotum and mesonotum ochraceous
tawny, the anterior margin of the crown
narrowly blackish, veins of the tegmina, ex-
cept the concolorous subcostal vein, blackish
fuscous; wings translucent, the narrow
apical margin blackish fuscous; beneath,
ochraceous yellow, the claws of the anterior
and middle legs blackish; tips of posterior
tibiae and the tarsi blackish.
Female last ventral segment slightly long-
er on the median line than the penultimate.
Lateral posterior angles broadly rounded;
the posterior margin broadly, triangularly
incised for nearly half the length of the
segment.
Length to apex of tegmina: 14.5 mm.
Gypona thoracica Fabr.
(Osborn, 1938a: 21).
(Text-fig. 5 F-I) .
This species was described from South
America without definite locality and has
been recorded from various states in Brazil.
There is a good series in the present collec-
tion from Bartica District, British Guiana,
collected from March through May but none
later in the year.
All specimens in the present collection are
males. Most of them are black in color, dor-
sad, with a narrow longitudinal median vitta
fi’om the apex of the crown to the apex of
the mesonontum. One specimen has the pos-
terior half of the pronotum bright orange
yellow; another specimen has the entire pro-
notum and mesonotum bright orange yellow,
with the crown with a narrow median vitta
and two large pale comma-shaped marks
curving between the ocelli and the compound
eyes. Beneath, all the specimens are pale
yellow with the posterior tibiae and tarsi
and the anterior margins of the segments
black.
Crown elongate, more than half as long
as the pronotum, about two-thirds as long
as the greatest width between the eyes. Ocelli
behind the middle about as far from each
other as from the compound eyes. The an-
terior margin of crown broadly parabolic,
slightly concave. Pronotum nearly twice as
broad as median length, the anterior margin
broadly curved, the posterior margin broadly
sinuate.
Male genitalia with the pygofei's and
genital plates about the same length, shorter
than the anal segment. The genital plates
with parallel sides, obtuse at the apex. Pygo-
fers when viewed laterad about as long as
broad with a distinct ventral apical lobe.
Genital styles elongate, acuminate and di-
verging cauded. Aedeagus elongate, slender
with two pairs of spines at the apex, the
ventral pair short, strongly divergent, the
apical pair elongate, slightly divergent at
the apex, about half as long as the aedeagus.
Length to apex of tegmina: 13.8-14.5 mm.
Gypona blgemmis Spangb.
(Osborn, 1938a :25)
(Text-fig. 5 J).
This species has been reported previously
19491
Metcalf : Tettigellidae and Gyponidae of Kartabo
273
from Rio de Janeiro, Colombia and Guate-
mala. It may be recognized by its small size,
uniform pale greenish color, and distinctive
female genitalia. Head broad and short;
crown broadly rounded before, slightly more
than twice as broad as the median length ;
ocelli slightly before the middle as remote
from each other as from the compound eyes.
Pronotum more than twice as long as the
crown, more than twice as broad as its
median length. Female last ventral segment
slightly longer than the penultimate, deeply
excavated with a broad, round median lobe.
Length to apex of tegmina: 6. 5-7.5 mm.
There is a single female in the present
collection from Kartabo, Bartica District,
British Guiana, March 6, 1924.
Gypona Aavolimbata n. sp.
(Text-fig. 6 A-E).
This is a medium large species closely
resembling Gypona glauca Fabr. in general
coloration but differing decidedly in female
genitalia.
Head broad, only slightly narrower than
the pronotum; crown twice as broad as its
median length; ocelli just behind the middle
as far from each other as from the compound
eyes. Pronotum twice as broad as its median
length; anterior margin broadly rounded;
anterior lateral margins nearly straight,
slightly divergent; posterior margin shallow-
ly rounded almost parallel to the anterior
margin. Mesonotum large.
Female with the last ventral segment only
slightly longer on the median line than the
penultimate ; the posterior margin shallowlv
incised with a broad median tooth with a
small triangular notch at the apex; the pos-
terior lateral angles only slightly produced.
Male genitalia when viewed ventrad with the
pygofer narrow, distinctly appendiculate.
Genital plates shorter than the pygofer with
the apex obtuse; genital styles elongate,
acute at the apex, as long as the pygofer;
aedeagus as long as the pygofer, the apex
with two elongate slender spines. The pygo-
fer when viewed laterad about as broad as
long with the posterior dorsal angle dis-
tinctly produced.
Length $ : 9.75 mm.-lO mm. Length 2 :
10.75-11 mm.
Holotype S : British Guiana without defi-
nite date or locality.
Allotype 2: Kartabo, Bartica District
British Guiana, March 25, 1922.
Paratypes 1 $ : British Guiana without
definite date or locality; 1 2: Kartabo, Bar-
tica District, British Guiana.
Gypona translucent n. sp.
(Text-fig. 7 A-C).
This species resembles Gypona fusiformis
Walk, very closely. Head narrow, broadly
triangularly produced, the crown more than
half as long as the pronotum. The ocelli be-
hind the middle, about as far from each other
is from the compound eyes. Pronotum large,
the lateral margins strongly divergent, the
whole surface coarsely rugulose; tegmina
semitransparent.
Female last ventral segment about as long
on the median line as the penultimate seg-
ment; lateral posterior angles strongly pro-
duced, the posterior margin broadly, para-
bolically incised with the incision slightly
produced either side of the median line in
short, blunt lobes.
General color tawny; the tegmina trans-
lucent; the anterior border of the crown
narrowly blackish; the veins of the tegmina
blackish fuscous except subcosta, which is
chiefly yellowish-brown; beneath, including
the legs, pale ochraceous yellow; the claws
and some of the spines blackish fuscous.
Length to apex of tegmina: 17.0 mm.
Holotype 2: Kartabo, Bartica District,
British Guiana, August 18, 1920.
Paratype 2: Kartabo, Bartica District,
British Guiana, August 9, 1920.
Gypona plcturata n. sp.
(Text-fig. 6 I-M).
This is one of the most conspicuously col-
ored Gyponas known to me. In general struc-
ture it resembles somewhat Gypona peru-
viana Osb. but differs in essential details of
the female genitalia and decidedly in colora-
tion. Head narrow, crown short and median
length about one-half as long as the greatest
width between the eyes. Pronotum short and
broad, about twice as long as the crown, its
median length about one-half its greatest
width; the anterior margin broadly curved,
anterior lateral borders carinate, about twice
as long as the posterior lateral margins;
posterior margin nearly parallel to the an-
terior margin. Tegmina short and broad;
venation regular.
Female last ventral segment nearly twice
as long as the penultimate; the lateral pos-
terior angle strongly produced, broadly
rounded. Median area strongly produced
with a deep notch on the median line. Male
genitalia with genital plates short and broad,
broadly rounded at apex; pygofers shorter
than anal segment distinctly appendiculate;
genital styles slender, elongate, strongly di-
verging caudad; aedeagus elongate, some-
what bulbous with a distinct process apically
directed dorsad.
General color above, pale greenish-yellow
with the crown with a bright red anterior
border and a pair of pale reddish vittae
through the ocelli ; the pronotum with three
pairs of reddish vittae; the tegmina chiefly
translucent greenish-yellow, more or less
clouded with brownish and blackish; the
clavus is chiefly brownish with a broad saddle
of white just before the apex; the claval
border and the sutural border and the veins
chiefly blackish; some of the veins on the
corium are blackish or brownish and the
apex is narrowly bordered with blackish fus-
cous with a cloud of blackish or brownish
fuscous across the apical cross-veins. Be-
274
Zoologica: New York Zoological Society
[34 i 20
'■A
f\ • : \ y
l^~T^ . \
/ \ ; \ . \
( \\ I'l ! \
%/
x ! .y a
V : 1
W
J ... f
\ ^
1
/* Y
r%
1
X J
J
H
Text-fig. 6. Gypona flavolirnbata, : A, head thorax; B, face; C, male genitalia ventral,
D, male genitalia lateral; E, female genitalia. Gypona castanea: : P, head thorax; v, lace;
H, female genitalia. Gypona picturata.: I', head thorax; J, face; K, male genitalia ventral ,
L, male genitalia lateral; M, female ■ genitalia. < --
1949]
Metcalf : Tettigellidae and Gyponidae of Kartabo
275
neath, pale ochraceous yellow with all the
tibiae and tarsi brownish fuscous.
Length 9 to apex of tegmina: 13.5 mm.;
$ : 11.5 mm.
Holotype 9: Kartabo, Bartica District,
British Guiana, March 17, 1922.
Allotype $: Georgetown, November 15,
1933.
Paratype 9: Kartabo, Bartica District,
British Guiana, March 17, 1922.
Gypona opaca n. sp.
This is another large species somewhat
similar in coloration to Gypona fusiformis
Walk, and Gypona translucens Mete, in that
the head and pronotum are generally lighter
than the tegmina. It differs, however, in that
the crown is very much shorter, the tegmina
are opaque and variegated, and the anterior
margin of the crown is without the narrow
black border.
Head narrow; crown broad and short,
about half as long as the short pronotum;
the anterior margin broadly parabolic; the
ocelli slightly behind the middle, about as
far from each other as from the compound
eyes; the anterior margin thin and folia-
ceous. Pronotum short and broad, nearly
twice as broad as the median length; the
lateral margins short, strongly carinate,
slightly divergent; the posterior lateral mar-
gins about half as long as the anterior lateral
margins; anterior margin broadly curved
with posterior border nearly parallel. Teg-
mina opaque, somewhat rugulose. Face very
flat; the dorsal margin distinctly impressed
with a nearly quadrate impressed point.
Female last ventral segment somewhat
longer than the penultimate on the median
line. The lateral postex-ior angles broadly
produced, somewhat obtuse. The posterior
margin broadly, roundly incised either side
of a strongly produced, rounded tooth.
General color of the head and thorax above,
yellowish-tawny; the ocelli and compound
eyes black; tegmina generally reddish-
brown, variegated with pale yellowish over
most of the surface; the costal margin with
the reddish-brown and yellowish spots rather
regularly distributed. The whole area of the
tegmina somewhat rugulose but the venation
regular. General color beneath, yellowish-
tawny with the dorsal margin of the face
infuscated; legs chiefly reddish-brown with
the spines of the hind tibiae blackish fuscous.
Length to apex of tegmina : 15 mm.
Holotype 9: Kartabo, Bartica District,
British Guiana, April 4, 1922.
Gypona c astanea n. sp.
(Text-fig. 6 F-H).
This is an almost uniformly castaneous
brown species of fairly large size with the
venter of the abdomen a little paler and two
conspicuous black spots on the anterior bor-
der of the mesonotum.
Head narrower than the pronotum; the
crown about twice as broad as its median
length ; the ocelli almost equidistant from
each other and the eyes, placed near the
middle of crown ; the anterior margin strong-
ly curved. Pronotum twice as long as the
crown, more than twice as broad as its median
length; anterior margin bi’oadly curved; an-
terior lateral margins nearly straight, not
quite twice as long as the posterior lateral
margins; posterior margin deeply incised;
surface of pronotum strongly rugulose.
Last ventral segment of the female longer
than the penultimate, deeply, almost quad-
rately incised with a broad median tooth
which is notched at the apex; posterior
lateral angles strongly produced, distinctly
rounded at the apex.
Color almost uniformly castaneous above
and beneath, except the basal segments of
the abdomen which are slightly paler. Meso-
notum with two conspicuous black spots on
the anterior border. Tegmina with a row of
inconspicuous brownish-vellow spots in the
costal cell and a few irregular scattered spots
on the corium and the clavus; apex of the
tegmina infuscate.
Length to apex of tegmina: 11.7 mm.
Holotype 9: British Guiana without defi-
nite date.
Ponana Ball.
Gypona ( Ponana ) Ball, 1920a :93.
Orthotype Gypona scarlatina Fitch.
This genus may be recognized by its nar-
row head, narrower than the posterior angles
of the pronotum, crown short with a distinct
thick margin separating it from the face.
Face fiat, distinctly impressed beneath the
broad border. Mesonotum large. Tegmina
coriaceous, venation regular. Pronotum
broad; anterior margin broadly curved, al-
most continuous with anterior latex-al max--
gin ; posterior margin neax-ly stx-aight. Whole
surface finely rugulose.
Ponana fulva n. sp.
(Text-fig. 7 D-F).
This is an almost unifox-mly tawny-colored
species with the thick antex-ior margin
bi-own, the ocelli and compound eyes black.
Dox-sal max-gin of abdomen chiefly bright
carmine red. Crown bx-oad and shox't, nearly
two and a half times as bx*oad as its median
length; antex'ior max-gin bx*oadly thickened.
Ocelli about as far fx-om base as from apex,
almost equidistant fx-om each other and the
compound eyes. Face distinctly impressed
beneath a broad dorsal margin. Px-onotum
not quite twice as bx-oad as its median length;
anterior latex-al max-gins not distinctly
separated from anterior max-gin; posterior
lateral max-gins short; posterior margin
broadly incised. Mesonotum nearly as broad
as the head, bx-oader than long.
Female last ventral segment about as long
as penultimate; posterior max-gin broadly
V-shaped; posterior latex-al angles not pro-
duced.
Length to apex of tegmina: 9.50 mm.
276
Zoologica: New York Zoological Society
[34: 20
Text-fig. 7. Gypona translucens: A, head thorax; B, face; C, female gmritalia. Ponana
fulva: D, head thorax; E, face; F, female genitalia. Clinonana bicolor : G, head thorax;
H, face; I, male genitalia ventral; J, male genitalia lateral.
1949]
277
Metcalf: Tettigellidae and Gyponidae of Kartabo
Holotype 2: British Guiana without defi-
nite date or locality.
Clinonaria gen. n.
Orthotype Clinonaria bicolor n. sp.
This genus may be recognized by its nar-
row head with short broad crown. The pro-
notum is not especially broad and the humeri
are not very prominent. When viewed laterad
the pronotum is strongly declivous and the
crown is broadly rounded to the face. Teg-
mina somewhat rugulose but the venation is
distinct.
Clinonaria bicolor n. sp.
(Text-fig. 7 G-J).
This species resembles Clinonana declivata
Osb. but differs in detail and decidedly in
color. Head narrow; crown very short and
broad with the ocelli near the anterior mar-
gin. Anterior margin broadly l-ounded to the
face. Pronotum short, not very broad ; meso-
notum large.
Male genitalia with the pygofers rather
broad, obtuse at the apex. Genital plates
shorter than pygofers, narrow at the base,
broadly curved on the lateral margins ; apices
obtuse; genital styles elongate, as long as
the pygofers with an obtuse basal expansion ;
aedeagus large on the basal half, gradually
narrow to the apical third with two short
spines at the apex.
General color of the crown and pronotum
ochraceous orange ; compound eyes and ocelli
black; narrow posterior border of the pro-
notum brown, margined anteriorly with
pale green ; mesonotum and tegmina auburn
brown; tegmina with irregular blackish fus-
cous spots along the costal margin and on
the basal and apical thirds; face chiefly
ochraceous orange. General color beneath
yellowish ochraceous with the apex of the
fore, middle and hind femora with a large
blackish spot near the apex; spines on the
middle and hind tibiae with large black spots
at their bases ; apex of the hind tibiae black.
Length to apex of tegmina: $ 11.75 mm.;
2 13.0 mm.
Allotype 2: British Guiana without defi-
nite locality or date.
Holotype $: Kartabo, Bartica District,
British Guiana, July 20, 1922.
Marganalana n. gen.
Orthotype Marganalana testacea n. sp.
This genus is close to Marganana DeLong
( Margana DeLong) but differs in several
respects. Head narrower than the pronotum ;
crown twice as broad as long ; anterior mar-
gin parabolic; ocelli nearly equidistant from
the anterior and posterior margin. Anterior
margin of the head conspicuously thickened,
quinquecarinate, not distinctly impressed
above or below. Face flat, narrow; pronotum
slightly more than twice as long as the crown,
not quite twice as broad as long; anterior
lateral margins elongate; posterior lateral
margins short; anterior margin broadly
curved in an almost continuous line from the
humeri; posterior margin nearly straight.
Mesonontum large. Tegmina with the vena-
tion regular. The main veins of the corium
and clavus with a few indistinct punctures.
Marganalana testacea n. sp.
(Tex-fig. 8 A-D).
Crown distinctly broader than long; ante-
rior margin broadly curved; median length
greater than length next to compound eyes.
Ocelli conspicuous, about as far from each
other as from the compound eyes. Pronotum
not twice as broad as long, scarcely longer
than mesonotum. Female last ventral seg-
ment nearly twice as long as the penultimate
with a broad V-shaped shallow notch on the
posterior border; posterior lateral angles
not produced.
General color above and below tawny, with
anterior margin of crown, compound eyes
and ocelli black. The apex of the tegmina
infuscate. Dorsal segments of the abdomen
chiefly scarlet red.
Length to apex of tegmina : 8.5 mm.
Holotype 2: British Guiana without defi-
nite date or locality.
Scarisana n. gen.
Orthotype Scarisana variabilis n. sp.
This genus may be recognized by the long
broad head which is nearly as broad as the
pronotum. Crown long and broad; anterior
margin broadly parabolic; pronotum short
and broad ; anterior lateral and anterior
borders making a continuous broad circle
from the shoulders; posterior lateral borders
short; posterior border straight. Mesonotum
large. Face strongly inflated. Tegmina coria-
ceous, venation fairly regular, rather indis-
tinct with three ante-apical and four apical
cells; claval veins nearly straight and paral-
lel. Legs rather short and stout; anterior
tibiae with numerous rather long stout
spines on the posterior border. Hind tibiae
half again as long as the posterior femora.
Scarisana variabilis n. sp.
(Text-fig. 8 E-I).
This is a variable species as far as color
is concerned. Some specimens are light och-
raceous buff, others are deep tawny.
Crown rather long and broad, nearly twice
as wide as the median length; anterior mar-
gin broadly parabolic. Ocelli behind the mid-
dle farther from each other than from the
compound eyes. Pronotum about one and
one-half times as broad as its median length ;
anterior lateral and anterior borders broadly
curved; posterior border nearly straight.
Mesonotum large, broader than long. Apex
produced into a distinct spine-like process.
Female last ventral segment about four
times as long as the penultimate with a dis-
tinct V-shaped notch on the posterior bor-
der. Male genitalia with genital plates broad
at the base, gradually restricted to near the
278
Zoolopica : New York Zoological Society
[34: 20
Text-fig. 8. Marganalana testacea: A, head thorax; B, face; C, head thorax lateral;
D, female genitalia. Scarisana variabilis : E, head thorax; F, face; G, male genitalia
ventral; H, male genitalia lateral; I, female genitalia.
1949]
Metcalf : Tettigellidac and Gyponidae of Kartabo
279
middle then nearly parallel to the apex. Py-
gofers with a distinct apical process.
Length to apex of tegmina: 11.25 mm.
Holotype 3 : British Guiana.
Allotype 2: Kartabo, Bartica District,
British Guiana.
Paratypes 3 33, Kartabo, Bartica District,
British Guiana, various dates. 4 2$ Kartabo,
Bartica District, British Guiana, various
dates.
Scans Le Peletier and Serville, 1825a :609.
Haplotype Iassits ferrugineus Fabr.
This genus is unknown to me. I repeat be-
low the more essential portions of the origin-
al description, as there is no modern descrip-
tion. Germar, 1833a: 179, separated it from
his new genus Gypona as having the ocelli
remote from each other, whereas in Gypona
the ocelli are approximate. Subsequent au-
thors added but little. Evans, 1947a :215; fig.
30 G, has an excellent illustration of the head
and thorax of the species Scaris ferruginea
Fabr.
Body somewhat triangular. Crown short,
transverse, much narrower than the prono-
tum but of the same width as the anterior
margin of the pronotum, with anterior bor-
der of the head thick, rounded. Ocelli remote
from each other. Pronotum not dilated later-
ad, transverse, rather long, narrowed an-
teriorly and also a little posteriorly, trun-
cate on posterior margin. Mesonotum trian-
gular, prolonged caudad into an elongate
sharp process.
Literature Cited.
All references to the literature in this
paper are cited by author and date accord-
ing to the references in the author’s Biblio-
graphy of the Homoptera (1942) except the
articles which have been published since
that time.
DeLong, D. M.
1948b. A proposed new genus name Marga-
nana and the allotype description of
Prairiana hirsuta DeL.-Gyponinae.
Ohio Jour. Sci., 48:101.
Metcalf, Z. P.
1945b. Fulgoroidea (Homoptera) of Kartabo,
Bartica District, British Guiana.
Zoologica, 30:125-143.
[1949]
Zoologica: Index to Volume SU
281
INDEX
Names in bold face indicale new
genera, species or varieties; numbers
in bold face indicate illustrations;
numbers in parentheses are the serial
numbers of papers containing the
plates listed immediately following.
A
Abra, 249
palmeri, 250, (19) PI. I
Acrocampsa, 262
pallipes, 262, 263
rufa, 263, 264
Aeronautes montivagus montivagus,
61
Amblyscarta, 260
aurulenta, 260, 261
Ammotragus lervia, 9
Apolymelis, 93
cognata, 93
dombei, 94
Ashtabula furcillafa, 39, 40, 164, 173
B
Ballus depressus, 170
Bunistygnellus beebei, 23, 24
C
Capinota, 268
virescens, 267, 268
Chaetura brachyura brachyura, 57
cinereiventris lawrencei, 58
Chaeturella rutila brunneitorques,
58, (8) PI. I
Chlorura chlorura, 5
Clinonaria, 277
bicolor, 276, 277
Corythalia chalcea, 168, 173, 175
fulgipedia, 168, 173
xanthopa, 168, 173, 177, 190
Creatophora cinerea, 103
Cummingia lamellosa, 250
Cymafoica, 89
Cynorla bromeliaca, 22, 23
estebana, 22
Cypseloides cherriei, 59
cryptus, 60
D
Dichrophleps, 264
despecta, 264, 265
Donax asper, 251
assimilis, 252
califomicus, 252, (19) PI. I
carinatus, 253, (19) PI. I
conlusus, 255 (19) PI. I
gracilis, 253, (19) PI. I
navicula, 254, (19) PI. I
obesus, 254, (19) PI. I
punctatoslriatus, 255, (19) PI. I
transversus, 256, (19) PI. I
Dysschema heliconides, 20
E
Elliptotellina, 87
Ergasilus, 127
caeruleus, 128
centrarchidarum, 130
chatauquaensis, 130
cotti, 130
elegans, 130
elongatus, 130
funduli, 130
labracis, 130
lanceolatus, 130
lizae, 130
luciopercarum, 130
manicatus, 130
megaceros, 131
mugilis, 131
nigratus, 131
osburni, 131
versicolor, 131
Erythrogonia, 260
bicolor, 260, 261
Eucyane bicolora, 19
temperata, 19
Eunica monima, 108
Eurytellina, 73
Eustiromastix, 168
Evarcha falcata, 166
G
Gerlschia noxiosa, 162, 173
Gorilla gorilla berengei, 111
gorilla gorilla, 111, (13) Pis. I & II
Gypona, 272
bigemmis, 270, 272
caslanea, 274, 275
flavolimbata, 273, 274
fusiformis, 272
opaca, 275
piclurata, 273, 274
thoracica, 270, 272
lranslucens, 273, 276
H
Hentzia mitrata, 164
Hyctia nivoyi, 162
pikei,. 162
Hyalurga fenestra, 20
modesta, 20
mysis, 20
partita, 20
sixola, 20
I
Icius elegans, 164
Iphigenia, 257
alfior, 257
K
Kalina tuberculala, 21
L
Lyssomanes bradyspilus, 31, 32, 33,
164, 193
M
Macaliopsis, 81
Macoma, 88
(Cymatoica) undulata, 89
(Macoma) nasuta, 88
(Macoploma) medioamericana, 93
(Psammacoma) elongata, 89
lamproleuca, 90
panamensis, 91
speclri, 91, (9) PI. I
(Psammolreta) aurora, 92
pads, 92
Macoploma, 93
Maevia vittata, 170
Mago dentichelis, 49, 50, 170, 173,
175, 193
Marganalana, 277
teslacea, 277, 278
Marpesia chiron chiron, 108
Marpissa rumpfi, 162
undata, 162
Menemerus bivittatus, 162, 175, 193
Merisca, 82
Metaphidippus galathea, 164
prolervus, 164
Moerella, 67
O
Orectogonia, 262
flavoscuiellata, 261, 262
P
Pandion haliaetus carolinensis, 1
Panyptila cayennensis, 61
Papilio agesilaus agesilaus, 121, (14)
PI. I
anchises osyris, 121, (14) PI. I
anchisiades anchisiades, 121, (14)
PI. I
areas areas, 122, (14) PI. I
belus varus, 122, (14) PI. I
cleotas coroebus, 122, (14) PI. I
crassus, 122, (14) PI. I
erithalion zeuxis, 123, (14) PI. I
lycophron hippomedon, 123, (14)
PI. I
paeon thrason, 123, (14) PI. I
phaon, 123, (14) PI. I
polydamus polydamus, 123, (14)
PI. I
polyxenes americus, 123, (14) PI. I
protesilaus archesilaus, 124, (14)
PI. I
sesostris tarquinius, 124, (14) PI. I
thoas neacles, 124, (14) PI. I
torquaius orchamus, 124, (14) PI. I
Paradilepis simoni, 1, 2
Paraphidippus marginafus, 164
Paruterina candelabraria, 6
chlorurae, 5, 6
morgani, 6, 7
similis, 6
Peckhamia picata, 162
Pericopis catilina angustilineata, 19
calilina catilina, 19
tricolora tricolora, 20
Phiale flammea, 47, 166, 175, 193, 196
Phidippus audax, 166
clarus, 166
purpuralus, 166
whitmanii, 166
Philaeus chrysops, 166
Phoebis eubule marcellina, 108
Phyllodella, 87
Phyllodina, 86
Platypoecilus maculatus, 215, (18)
Pis. I-V
Plexippus paykullii, 168, (17) PI. I
Poecilocranaus gratiosus, 23
282
Poeciloscarla quadrifasciala, 244, 265
Ponana, 275
fulva, 275, 276
Psammacoma, 89
Psammolreta, 92
Pseudothelphusa chacei, 27-29
garmani, 24, 27
R
Raphirhinus, 266
fasciatus, 268
phosphoreus, 266, 267
Rhopalogonia, 268
purpurata, 269, 270
S
Saitis barbipes, 168
Salpinctes obsoletus obsoletus, 5
Salticus cingulatus, 162
scenicus, 162
Santinezia albilineata, 23
Sassacus flavicinctus, 41, 42, 164, 175
ocellatus, 44, 45, 164
Scaris, 279
Searisana, 277
variabilis, 277, 278
Scissula, 84
Scrobiculina, 66
Semele corrugala caliiornica, 240
craneana, 241, (19) PI. I
decisa, 242
flavescens, 242
guaymasensis, 243
jaramija, 244, (19) PI. I
jo vis, 244
laevis, 245
Zoologica: Index to Volume 3b
pacifica, 245, (19) PI. I
pulchra, 246, (19) PI. I
quenlinensis, 246, (19) PI. I
simplicissima, 247
sparsilineata, 247, (19) PI. I
labogensis, 248
venusla, 248, (19) PI. I
verrucosa, 249, (19) PI. I
Semorina brachychelyne, 35, 36, 162
megachelyne, 38, 175
Streploprocne zonaris albicincla, 54,
56, 57, (8) PI. I
Strigilla, 95
cicercula, 95, (9) PI. I
coslulifera, 95, (9) PI. I
disjuncla, 96, (9) PI. I
lenticula, 96, (9) PI. I
Synageles Venator, 162
T
Teliidora, 88
burneti, 88
Tellina, 64
(Elliptotellina) pacifica, 87
(Eurylellina) eburnea, 73
inaequislriata, 74, (9) PI. I
laceridens, 75
mantaensis, 75
panamanensis, 76
planulala, 76, (9) PI. I
prora, 77
regia, 78
rubescens, 78
simulans, 79
(Macaliopsis) lyra, 81
lyrica, 81
[1949]
(Merisca) crystallina, 82
proclivis, 83, (9) PI. I
reclusa, 84
(Moerella) amianta, 67
arenica, 68, (9) PI. I
erythronotus, 69
felix, 70, (9) PI. I
macneilii, 70
paziana, 71
recurvata, 71, (9) PI. I
suffusa, 72
labogensis, 72
(Phyllodella) insculpla, 87
(Phyllodina) pristiphora, 86
(Scissula) cognata, 84
nicoyana, 85, (9) PI. I
virgo, 86
(Scrobiculina) ochracea, 66
viridolincta, 66
(Tellinella) cumingii, 65
zacae, 65, (9) PI. I
(Tellinidella) purpureus, 80
Tellinella, 64
Tellinidella, 79
Tilapia heudeloli, 158
macrocephala, 133, (16) Pis. I-UI
V
Vima plana, 21, 22
X
Xesurus laticlavius, 101
Z
Zygobunus rufus, 21
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS
of the
NEW YORK ZOOLOGICAL SOCIETY
VOLUME 34
Part 1
Numbers 1-6
*
Published by the Society
The Zoological Park, New York
May 16, 1949
;ii’ • •i' '* /\
^ ?IU l till yf,,
CONTENTS
PAGE
1. Paradilepis simoni n. sp., a Cestode Parasitic in the Osprey. (Ces-
toda: Dilepididae) . By Robert Rausch. Text-figure 1 i
2. A Contribution to the Study of North American Cestodes of the
Genus Paruterina Fuhrmann, 1906. By Robert Rausch and
Everett L. Schiller. Text-figures 1-12 5
3. Behavioral Interactions in a Herd of Barbary Sheep ( Ammo-
tragus lervia) . By Irwin Katz 9
4. The Pericopidae (Moths) of Kartabo, British Guiana, and Cari-
pito, Venezuela. By Henry Fleming 19
5. Report on a Collection of Phalangids from Rancho Grande, Vene-
zuela. By Clarence and Marie Goodnight. Text-figures 1-4 21
6. Fresh-water Crabs of the Genus Pseudothelphusa from Rancho
Grande, Venezuela. By Jocelyn Crane. Text-figures 1-3 25
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS
of the
NEW YORK ZOOLOGICAL SOCIETY
VOLUME 34
Part 2
Numbers 7-12
Published by the Society
The Zoological Park, New York
August 10, 1949
CONTENTS
PAGE
7. Comparative Biology of Salticid Spiders at Rancho Grande, Vene-
zuela. Part III. Systematics and Behavior in Representative New
Species. By Jocelyn Crane. Text-figures 1-8 31
8. The Swifts of Rancho Grande, North-central Venezuela, with
Special Reference to Migration. By William Beebe. Plate I;
Text-figures 1-3 53
9. Eastern Pacific Expeditions of the New York Zoological Society.
XL. Mollusks from the West Coast of Mexico and Central Amer-
ica. Part VII. By Leo George Hertlein & A. M. Strong. Plate I.. 63
10. Fishes That Rank Themselves Like Soldiers on Parade. By E. W.
Gudger. Plate I ; Text-figures 1 & 2 99
11. Notes on Seasonal Changes in Creatophora cinerea, the Wattled
Starling. By Lee S. Crandall. Plate 1 103
12. Insect Migration at Rancho Grande in North-central Venezuela.
General Account. By William Beebe. Plates I & II ; Text-figure 1 107
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS
of the
NEW YORK ZOOLOGICAL SOCIETY
VOLUME 34
Part 3
Numbers 13-16
Published by the Society
The Zoological Park, New York
November 30, 1949
CONTENTS
PAGE
13. The Behavior of Two Captive Specimens of the Lowland Gorilla,
Gorilla gorilla gorilla (Savage & Wyman). By B. F. Riess,
Sherman Ross, S. B. Lyerly & H. G. Birch. Plates I & II;
Text-figures 1 & 2 Ill
14. Migration of Papilionidae at Rancho Grande, North-central
Venezuela. By William Beebe. Plate I; Text-figure 1 119
15. Notes on Ergasilus Parasites from the New Brunswick, New
Jersey, Area, with a Check List of All Species and Hosts East
of the Mississippi River. By Roland F. Smith 127
16. An Analysis of Reproductive Behavior in the Mouth-breeding
Cichlid Fish, Tilapia macrocephala (Bleeker). By Lester R.
Aronson. Plates I-III; Text-figures 1-10 133
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS
of the
NEW YORK ZOOLOGICAL SOCIETY
VOLUME 34
Part 4
Numbers 17-20
Published by the Society
The Zoological Park, New York
December 30, 1949
t2jl, . i*
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