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HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



BREVIORA 



MUSEUM OF COMPARATIVE ZOOLOGY 

AT 

HARVABD COLLEGE, IN CAMBRIDGE 



Numbers 67-120 
1957-1960 



CAMBRIDGE, MASS., U.S.A. 

1960 



Edited 
By 

Nelda E. Wright 



V.^ 



.\ 



CONTENTS 

BREVIORA 

^Museum of Comparative Zoology 

Numbers 67-120 

1957 

No. 67. Notes on Certain Species of Tetragnatha (Araneae, Ar- 
giopidae) in Central America and Mexico. By 
Arthur ]\[. Chickering. 4 pp. January 31. 

No. 68. The Genus Tetragnatha (Araneae, Argiopidae) in Ja- 
maica, B.W.I., and other Neighboring Islands. By 
Arthur M. Chickering. 15 pp. January 31. 

No. 69. A New Zodariid Spider from Panama. By Arthur M. 
Chickering. 7 \)\). January 31. 

No. 70. "Anguimorph" Tooth Replacement in Amphishaena 
alha Linnaeus, 1758, and A. fuliginosa Linnaeus, 
1758 (Reptilia : Ampliisbaenidae). By Carl Gans. 
12 pp. January 31. 

No. 71. Taxonomic Notes on the New World Forms of Troglo- 
dytes. By Raymond A. Paynter. Jr. 15 pp. March 
29. 

No. 72. Is the Ant Genus Tetramorium Native in North Amer- 
ica? By W. L. Brown, Jr. 8 pp. March 29. 

No. 73. Additions to the Mammalian Fauna of Peru and Notes 
on Some Other Peruvian Mammals. By Oliver P. 
Pearson. 7 pp. March 29. 

No. 74. The Discovery of Cerapachyine Ants on New Caledonia, 
with the Description of New Species of Phyracace.s 
and Sphinctomyrmex. By E. 0. Wilson. 9 pp. 
May 1. 



No. 75. Oil a New Oetochaetine Earthworm Supposedly from 
Guatemala. By G. E. Gates. 8 pp. May 1. 

No. 7(). Two New Land and Freshwater Mollusks from New 
Guinea. By William J. Clench. 4 pp. June 18. 

No. 77. Dacetinopa, A New Ant Genus from New Guinea. By 
W. L. Brown, Jr. and E. 0. Wilson. 7 pp. June 21. 

No. 78. The Larva of the Ant Genus Dacetinops Brown and 
Wilson. By George C. Wheeler and Jeanette Wheel- 
er. 4 pp. Jvme 21. 

No. 79. Dasypeltis nicdici lamiiensis, A New Race of Egg-Eating 
Snake (Ophidia, Reptilia) from Coastal East Africa. 
Bj'- Carl Gans. 13 pp. August 9. 

No. 80. A Collection of Drawings of Fishes Ascribed to J. P. 
Kirtland (1793-1877), in the Library of Bowdoin 
College. By James M. Moulton. 4 pp., 2 pis. Sep-^ 
tember 30. 

No. 81. Contributions to a Revision of the Earthworm Family 
Lumbricidae. I. Allolobophora limicola. By G. E. 
Gates. 14 pp. September 30. 

1958 

No. 82. The Trunk ^lusculature of Sanzina and its Beai-ing on 
Certain Aspects of the ]\Iyological Evolution of 
Snakes. By Walter Auffenberg. 12 pp. January 31. 

Xo. 83. Thamnophis hovaUii Dunn Rediscovered (Reptilia, Ser- 
})entes). By Benjamin Shreve and Carl Gans. 8 pp. 
January 29. 

No. 84. Rediscovery of the Australian Chelid Genus Pseudemy- 
dura Siebeurock (Chelidae, Testudines). By Ernest 
E. Williams. 8 pp., 4 pis. January 30. 

No. 85. The Choanal Papillae of the Cheloniidae. By Thomas 
S. Parsons. 5 pp., 2 pis. January 31. 

No. 86. A New Sicistine Rodent from the ^Miocene of Wyoming. 
By Craig C. Black. 7 pp. May 29. 



No. 87. An Enibolonioro J aw from the JMid-Carboniferoiis of 
Nova Scotia. By Alfred Sherwood Romer. 7 pp., 
1 pi. June 20. 

No. 88. A New Species of the Genus iirotheca (Serpentes: Colu- 
bridae) from Venezuela. B}' J. A. Roze. 5 pp. 
June 30. 

No. 89. Remarks on Some Forms of Cinclus (Aves). By James 
C. Greenway. Jr. and Charles Vaurie. 10 pp. 
July 15. 

No. 90. A Fossil X'ampire Bat from Cuba. By Karl F. Koop- 
man. 4 pp., 1 pi. July 30. 

No. 91. Contribution to a Revision of the Earthworm Family 
Lumbricidae. II. Indian Species. By G. E. Gates. 
16 pp. August 13. 

No. 92. A New Genus of Erethizontid Rodents from the Col- 
huehuapian of Patagonia. By Bryan Patterson. 
4 pp. September 17. 

No. 93. A New Barylambdid Pantodont from the Late Paleo- 
cene. Bj^ Bryan Patterson and Elwyn L. Simons. 
8 pp. September 18. 

No. 94. Affinities of the Patagonian Fossil Mammal Necrolestes. 
By Bryan Patterson. 14 pp. September 18. 

No. 95. A New Bolivian Land Snail of the Genus Dryniaeus. 
By Juan Jose Parodiz. 3 pp. September 19. 

No. 9(j. A New Dichobunid Artiodactyl from the Uinta Eocene. 
By C. Lewis Gazin. (i pp. September 19. 

No. 97. Fusion of Cervical Vertebrae in the Erethizontidae and 
Dinomyidae. By Clayton E. Ray. 11 pp., 2 pis. 
October 27. 

No. 98. Two New Species of Bathylagus from the Western 
North Atlantic with Notes on Other Species. By 
Daniel M. Cohen. 9 pp. December 12. 

No. 99. A New Subspecies of Chamadeo jacksoni Boulenger and 
a Key to the Species of Three-llorned Chamaeleons. 
By A. Stanley Rand. 8 pp. December 19. 



No. TOO. On the Pineal Organ of the Tuna, Thijnnus thynnus 
L. By Uno Hohngren. 5 pp., 2 pis. December 23. 

1959 

No. 101. Cervical Ribs in Turtles. By Ernest E. Williams. 12 
pp., 1 pi. March 2. 

No. 102. A New Jamaican Galliwasp (Sauria, Anguidae). By 
Garth Underwood. 13 pp. April 9. 

No. 108. Two New Species of Eleutheroductyius from Puerto 
Rico. By Juan A. Rivero. 6 pp., 1 pi. April 10. 

No. 104. Studies on Fishes of the Family Ophidiidae. III. A 
New Species of Lepophidium from Barbados. By C. 
Richard Robins. 7 pp. April 13. 

No. 105. Bufo (jmidlachi, A New Species of Cuban Toad. By 
Rodolfo Ruibal. 14 pp. April 14. 

No. 106. The Occipito-Vertebral Joint in the Burrowing Snakes 
of the Family Uropeltidae. By Ernest E. Williams. 
10 pp. April 28. 

No. 107. A Revision of the Dacetine Ant Genus Neostruma. By 
William L. Brown, Jr. 13 pp. May 6. 

No. 108. Some New Species of Dacetine Ants. By William L. 
Brown, Jr. 11 pp. May 7. 

No. 109. On the Pineal Area and Adjacent Structures of the 
Brain of the Dipnoan Fish, Protopterus annectens 
(Owen). By Uno Holmgren. 7 pp., 2 pis. May 8. 

No. 110. The Spider Genus Coleosoma (Araneae, Theridiidae). 
By Herbert W. Levi. 8 pp.. 1 pi. June 16. 

No. 111. On the Caudal Neurosecretory System of the Teleost 
Fish, Fundulus heterodiUis L. By Uno Holmgren. 
13 pp., 2 pis. June 17. 

No. 112. A Mounted Skeleton of the Giant Plesiosaur Kronosau- 
rus. By Alfred Sherwood Romer and Arnold D. 
Lewis. 14 i^p.. 1 pi. October 15. 



No. 113. A New Phyllomedusa from Bolivia (Salientia, Hyli- 
dae). By Benjamin Shreve. I^ pp., 1 })1. November 2. 

No. 114. Anomalophis bolccnsis (Massalonj^o), A New Genus 
of Fossil Snake from the Italian Eocene. By Walter 
Auffenberji'. 16 pp. November 23. 

No. 115. The Lemon-Colored Plexaurids from the West Indies 
and Brazil. By Elisabeth Deiehmann and P. M. 
Bayer. 12 pp., 5 pis. Noveml)er 25. 

1960 

No. 116. Insectivores of the Middle Miocene Split Rock Local 
Fanna, Wyoming-. By Katherine Milmine Reed. 
11 pp., 2 pis. January 6. 

Xo. 117. Notes on Hispaniolan Herpetology. 1. Anolis christo- 
phci. New Species, from the Citadel of King Christo- 
phe, Haiti. By Ernest E. Williams. 7 pp. January 
20. 

No. 118. A Survey of the Leptodactylid Frogs, Genus Eupso- 
phus, in Chile. By Jose M. Cei. 13 pp. February 24. 

No. 119. Arctic Archibenthal and Abyssal Mollusks from Drift- 
ing Station Alpha. By Arthur H. Clarke, Jr. 17 pp., 
1 pi. March 8. 

No. 120. Two Species of Tortoises in Northern South America. 
By Ernest E. Williams. 13 pp., 3 pis. March 9. 



INDEX OF AUTHORS 
BREVIORA 

MUSEUM OP COMPARATIVE ZOOLOGY 

Numbers 67-120 

1957-60 

No. 

AUFFENBERG WALTER 82, 114 

Bayer, F. M. and Elisabeth Deichmann 115 

Black, Craig C 86 

Brown, William L., Jk 72, 107, 108 

Brown, William L., Jr. and E. 0. Wilson 77 

Cei, Jose M 118 

Chickering, Arthur M 67, 68, 69 

Clarke, Arthur H., Jr 119 

Clench, William J 76 

Cohen, Daniel M 98 

Deichmann, Elisabeth and F. M. Bayer 115 

Gans, Carl 70, 79 

Gans, Carl and Benjamin Shreve 83 

Gates, G. E 75, 81, 91 

Gazin, C. Lewis 96 

Greenway, James C, Jr. and Charles Vaurie 89 

Holmgren, Uno 100, 109, 111 

KooPMAN, Karl F 90 

Levi, Herbert W 110 



Lewis, Arnold D. and Alfred Sherwood Romp:r 112 

MouLTON, James M. 80 

Parodiz, Juan Jose 95 

Parsons, Thomas S 85 

Patterson, Bryan 92, 94 

Patterson, Bryan and Elwyn L. Simons 93 

Paynter, Raymond A.. Jr 71 

Pearson, Oliver P 73 

Rand, A. Stanley 99 

Ray, Clayton E 97 

Reed, Katherine Milmine 116 

RivERO, Juan A. 103 

Robins, C. Richard 104 

RoMER, Alfred Sherwood 87 

RoMER, Alfred Sherwood and Arnold D. Lewis 112 

Roze, J. a 88 

RuiBAL, Rodolfo 105 

Shreve, Benjamin 113 

Shreve, Benjamin and Carl Cans 83 

Simons, Elwyn L. and Bryan Patterson 93 

Underwood, Garth 102 

Vauree, Charles and James C. Green way, Jr 89 

Wheeler, George C. and Jeanette Wheeler 78 

Wheeler, Jeanette and George C. Wheeler 78 

Williams, Ernest E. 84, 101, 106, 117, 120 

Wilson, E. 74 

Wilson, E. 0. and William L. Brown, Jr 77 



BREVIORA 

Moseiiim of Comparative Zoology 



Cambridge, Mass. Januiiry 31, 1957 Number 67 



NOTES ON CERTAIN SPECIES OP TETRAGNATHA 
(ARANEAE, ARGIOPIDAE) IN CENTRAL AMERICA 

AND MEXICO 

By Arthur M. Chickering 

Albion CollpRp, Albion. Michigan 

In connection with my study of the ^enns Tefrngnafha 
Latreille, 1804 in Panama and the West Indies I have also had 
occasion to examine a number of species from parts of Central 
America north of Panama, and also from Mexico, types of which 
are in the Museum of Comparative Zoology at Harvard College. 
The following notes are offered as a contribution to the further 
clarification of the genus. 

Tetragnatha versicolor Walckenaer, 1841 

T. convexa Banks, 1898 

T. convexa Petrunkevitch, 1911 

T. convexa Eoewer, 1942 

A vial labelled T. convexa Banks and marked "cotypes" now 
contains one male and three females all from San Jose del Cabo, 
Baja California. The chelicerae, palp, and other characters make 
it certain that the male belongs to T. versicolor Walckenaer. The 
females are always more difficult to place with accuracy but I 
feel certain that these belong in the same species with the male. 
There is another male in the collection from Sierra Laguna, Baja 
California, originally identified as a T. convexa Banks l)ut this- 
is also clearly a T. versicolor Walckenaer. 

Tetragnatha guatemalensis 0. P. Cambridge, 1889 

T. fraterna Banks, 1898 

T. mnndihulata Banks, 1898 



BREVIORA 



NO. 67 



T. fraterna Petrunkevitch, 1911 
T. fraterna Eoewer, 1942 

The characteristics of the male palp and the male chelicerae 
definiteh" identify the males collected at San Jose del Cabo, 
Baja California and described as T. fraterna, and I feel confident 
that we may be certain of the correct placement of the females as 
well. Banks identified both sexes collected at Tepic and San Jose 
del Cabo as specimens of T. mnndihiilata Walckenaer, 1841, 
known at present only from Ceylon, India, Australia, and Poly- 
nesia. Re-examination of these shows clearly that they belong 
to T. guatemalensis 0. P. Cambridge. 

Tetragnatha tristani Banks, 1909 
(Figures 1-5) 
There is only a single specimen to represent this species so far 
as I have been able to determine. This is the holotype very 
briefly described by its author. I have carefully searched through 
my collections of Tetragnatha from the regions of Panama closely 
contiguous to Costa Rica, from which country the original was 
collected, without discovering any additional specimens. I have 
also compared the holotype with all other species known to me, 
with the result that I am compelled to regard it as a valid species. 



• ^ 'y / 



e 



D kA 



i * ' 



*■« 






External Anatomy of Tetragnatha tristani Banks 
Figures 1, 2. Cheliceral teeth along the fang groove; promarginal nnd 

retromarginal teeth, respectively. 

Figures 3, 4. Distal ends of cymbiiun, embolus, and eonduetor from two 

different views. 

Figure .t. Paracymbium. 



1957 TETRAGNATHA FROM CENTRAL AMERICA 6 

Since the orig:inal description was so brief I have thoup:ht it 
desirable to furnish a detailed treatment in accord with my usual 
procedure. 

Male holofypc. Total leng-th includinor the chelicerae 4.55 mm. ; 
exclusive of the chelicerae 4.16 mm. Carapace 1.495 mm. long, 
.97 mm. wide opposite second coxae where it is widest ; other 
features as usual in the genus. 

Eyes. Eight in two rows as usual in the genus; viewed from 
above, both rows moderately recurved ; viewed from in front, both 
rows slightly procurved, both measured by centers. Central 
ocular quadrangle wider behind than in front in ratio of about 
14 : 11, slightly wider behind than long. Ratio of eyes AME : 
ALE : PME : PLE = 7 : 5.5 : 8 : 6.5. AME separated from one 
another by 1.5 times their diameter, from ALE by a little more 
than twice their diameter. PME separated from one another by 
1.7 times their diameter, from PLE by slightly more than this 
distance. Laterals separated from one another by the diameter 
of PLE; AME separated from PME by nearly 1.5 times as far. 
Height of clypeus equal to a little more than the diameter of 
AME. 

Chelicerae. Moderately well developed, porrect and divergent; 
basal segment .78 mm. long and, therefore, about half as long as 
the carapace ; the prolateral spur is simple, rather poorly devel- 
oped, and not distally bifid ; the fang is moderately sinuous and 
distinctly bent posteriorly in distal half; the promargin of the 
fang groove has seven teeth but the last three are minute den- 
ticles and would probably be subject to much variation in a large 
population; the retromargin also has seven teeth with the last 
two very small (Pigs. 1-2). 

Maxillae. Nearly parallel ; somewhat concave along outer 
border ; about three times as long as wide at narrowest level. 

Lip. Wider at base than long in ratio of about 15 : 11 ; less 
than one-half as long as maxillae ; sternal suture gently pro- 
curved ; sternal tubercles well developed and robust at ends of 
sternal suture. 

Sternum. As usual in the genus ; with fourth coxae separated 
by about one-half their width. 

Legs. 1243. Width of first patella at "knee" .162 mm., tibial 
index of first leg 4. Width of fourth patella at "knee" .152 mm., 
tibial index of fourth leg 7. 



BREVIORA NTO . 67 



Femora Patellae Tibiae Metatarsi Tarsi Totals 

(All measurements in millimeters) 
2.990 MO 3.250 .3.380 .975 11.245 



2. 


2.080 


.552 


1.755 


1.885 


.700 


6.972 


3. 


1.105 


.390 


.617 


.845 


.325 


3.282 


4. 


2.210 


.390 


1.625 


1.950 


.520 


6.695 


Palp 


.660 


.154 


.176 




.970 


1.960 



All le^s with moderately coarse spines and the normal eoatinp: 
of hair. 

Palp. Both tibia and patella are short with the former only 
slightly longer than the latter; the paracymbium is very trans- 
parent but appears to have the shape shown in Figure 5; the 
cyrabium is long and slender; the conductor and embolus are 
also long, slender, and gently curved (Figs. 3-4). 

Abdomen. Not prolonged posterior to spinnerets; only slightly 
swollen in anterior half; slightly notched dorsally at base; 2.73 
mm. long; .95 mm. wide about one-third of its length from liase 
where it is widest. 

Color in alcohol. Color apparently well preserved. Legs, 
mouth parts and cephalothorax with various shades of yellowish 
to light reddish brown. Abdomen with a well defined folium and 
many closely placed yellowish white silvery spangles on dorsum 
and along lateral sides. Venter plain yellowish. 

There is only one specimen known at present and that was col- 
lected by Prof. J. Fid. Tristan of San Jose, Costa Rica, in his 
home city with no date recorded. 

REFERENCES 

Banks, Nathan 

1898. Arachnida from Baja California and other parts of Mexico. 
Proc. California Acad. Sei., Ser. 3, Zoology. 1, (7): 205-309, 
5 pis. 

1909. Arachnida from Costa Rica. I'roc. Acad. Nat. Sci. Philadelphia, 
April, 1909: 194-234, 2 pis. 



BREVIORA 

Mmseiuiirii of Compsirative Zoology 

Cambridge, Mass. January 31, 1957 Number 68 



THE GENUS TETRAGNATHA (ARANEAE, 

ARGIOPIDAE) IN JAMAICA, B.W.I., AND 

OTHER NEIGHBORING ISLANDS 

Arthur M. Chickering 

Albion College, Albion, Michigan 

For several years before her death in 1953 Miss Elizabeth B. 
Bryant, Museum of Comparative Zoology at Harvard College, 
had been engaged in a comprehensive study of a collection of 
spiders from Jamaica, B. W. I. This collection had come from 
several sources but it had been assembled largely through the 
interest of Mr. C. Bernard Lewis, Director and Curator, Sci- 
ence Museum, Institute of Jamaica, Kingston, Jamaica. After 
Miss Bryant's death this collection was placed in my possession 
for continued study. On my way to Panama in June, 1954, I 
was able to stop in Jamaica for a reconnaissance of the island 
preparatory to what may be a more or less extensive study of the 
spiders of that country. 

As an extension of my study of the genus Tetragnatha 
Latreille, 1804 in Panama, I have been much interested in 
examining the genus in Jamaica and in comparing the species 
found there with the tetragnathids in several of the larger islands 
of that general region. This paper is a result of that study, and 
types of the new species named here are deposited in the Mu- 
seum of Comparative Zoology. 

It is again a pleasure to acknowledge my indebtedness to the 
following persons for their continued encouragement in the pur- 
suit of my studies : Dr. A. S. Romer and Dr. P. J. Darlington, 
Jr., Director, and Curator of Insects, respectively, in the Mu- 
seum of Comparative Zoology at Harvard College, and Miss 
Nelda E. Wright, Editor of Publications in the same institu- 
tion. Without the privileges which have been extended to me 



2 BREVIORA NO. 68 

for many years in this museum the continued progress of my 
studies would have been much more difficult. 

Genus TetraGNATHA Latreille, 1804 

The genus has been well defined by Seeley (1928) and is, in 
general, well understood by araneologists. There are, however, 
certain characteristics of the genus which have in the past made 
it difficult to identify the species correctly and numerous errors 
must eventually be eliminated. It is also my opinion that suffi- 
cient attention has not usually been given to the question of varia- 
tion within species in respect to several of the most important 
structural features used by taxonomists for identification. F. P. 
Cambridge (1897-1905) emphasized the value of the character- 
istics of the male palp such as the form of the paracymbium, 
shape and course of the conductor and embolus as well as the 
features of the eyes, chelicerae, and legs. Petrunkevitch (1930) 
and Wiehle (1939) were the first to appreciate the value of the 
genital area, which lacks an epigynum, in identifying females 
which are often exceedingly difficult to place with certainty. 
Color has been shown to be extremely variable and nearly worth- 
less as a means of identification. Cheliceral teeth are often quite 
variable in number, degree of development, and relative posi- 
tion. Size, when mature, is also subject to great variation in sev- 
eral species. I have tried to take account of all of these salient 
features in making my determinations. 

THE GENTJS IN JAMAICA 

Only the bibliographical references considered essential are 
given in this paper. Extensive bibliographies may be found in 
several sources. 

Tetragnatha antillana Simon, 1897 

T. antillana Petrunkevitch, 1930 

T. antillana Bryant, 1940 

T. antillana Bryant, 1942 

T. festina Bryant, 1945 (male only) 

T. haitiensis Bryant, 1945 



1956 TETRAGNATHA IN JAMAICA 3 

This species appears to be common in Jamaica. It was found 
abundant at Mavis Bank over water by R. P. Bengry. Collection 
records: One male from the Blue Mts., southwest side of Main 
Range, between 3000-4000 ft. elevation, August, 1934 (P. J. Dar- 
lington, Jr.) ; both sexes from Mavis Bank, over water, March, 
1953 (R. P. Bengry) ; one female from Rio Cobre, June, 1954, 

Tetragnatha caudata Emebton, 1884 

Eucta caudata Petrunkeviteh, 1911 
T. caudata Seeley, 1928 
T. caudata Bryant, 1940 

Miss Bryant had a single female from Cuba. The species ap- 
pears but once in the collection from Jamaica placed at my dis- 
posal ; Papine, five miles north of Kingston, April, 1937. 

Tetragnatha exigua sp. nov. 
(Figures 1-5) 

Male holotijpe. Total length including chelicerae 2.795 mm., 
without chelicerae 2.34 mm. Carapace 1.04 mm. long; .67 mm. 
wide opposite second coxae where it is widest ; with the usual gen- 
eral form of the genus ; .209 mm. tall at about the middle ; nearly 
level from PE to posterior declivity; median depression very 
shallow, opposite interval between second and third coxae. 

Eyes. Eight in two rows as usual; lateral ocular tubercles 
rather prominent ; viewed from above, both rows moderately re- 
curved ; viewed from in front, anterior row slightly recurved and 
posterior row slightly procurved, both measured by centers ; cen- 
tral ocular quadrangle wider behind than in front in ratio of 
about 4 : 3, wider behind than long in about the same ratio. 
Ratio of eyes AME : ALE : PME : PLE = 5.5 : 4 : 5 : 4. 
AME separated from one another by about 1.2 times their 
diameter, from ALE by about the same distance. PME sep- 
arated from one another by a little less than twice their diam- 
eter, from PLE by about two thirds as far. Laterals separated 
from one another by about 1.25 times their diameter. AME sep- 
arated from PME by a little more than the diameter of AME, 
hence further from one another than laterals are from one 
another in ratio of about 6 : 5. Height of clypeus equal to nearly 
1.5 times the diameter of AME. 



BREVIORA 



NO. 68 



Chelicerae. Well developed, moderately porrect, quite diver- 
gent in distal two thirds, somewhat swollen in middle ; prolateral 
spur a simple spine not bifid distally; fang- slender, slightly 
sinuate, with a blunt tubercle on inner margin about one-fifth 
of its length from base ; promargin of fang groove with four 
teeth, retromargin with four smaller teeth; with no "large 
tooth" on the promargin (Fig. 1). 

Maxillae. Nearly parallel ; slightly concave in middle of lateral 
border ; somewhat more than twice as long as lip ; three times 
as long as wide in middle. 



r 




n 





External Anatomy of Tetragnatha exigua sp. nov. 

Fig. 1. Chelicerae of male from in front. 

Fig. 2. Paracymbium of male palp. 

Fig. 3. Distal end of cymbium, conductor, and embolus. 

Fig. 4. Cheliceral teeth of female. 

Fig. 5. Genital fold of female. 



Liip. Much widened in basal third where it is wider than long 
in ratio of 22 : 14; sternal suture only slightly procurved; with 
the usual sternal tubercles well developed at ends of sternal 
suture. 

Sternum. Quite convex ; surface finely pitted and granulated ; 
with the usual form ; continued laterally and posteriorly between 
all coxae ; only a little longer than wide ; posterior coxae sepa- 



1956 TETRAGNATHA IN JAMAICA 5 

rated by a little more than their width. 

Legs. 1243. Width of first patella at "knee" .1083 mm., 
tibial index of first leg 4. Width of fourth patella at "knee" 
.0758 ram., tibial index of fourth leg 5. 

Femora Patellae Tibiae Metatarsi Tarsi Totals 

(All measurements in millimeters) 

1. 2.275 .390 2.275 1.755 .718 7.413 

2. 1.625 .325 1.430 1.380 .580 5.340 

3. .910 .198 .445 .550 .308 2.411 

4. 1.430 .260 1.170 1.235 .455 4.550 
Palp .440 .120 .176 .396 1.132 

Spines. True spines appear to be entirely lacking in this 
species (a very unusual feature) ; hair and bristles are sparsely 
present. Trichobotliria are present but have not been accurately 
observed. 

Palp. Both tibia and patella are short with tibia longer than 
patella in ratio of about 3 : 2. The paracymbium is unusually 
broad. The conductor and embolus are shaped and related essen- 
tially as shown in Figures 2 and 3. 

Abdomen. Slender ; broadest near base and gradually tapered 
to a blunt point posteriorly; bluntly truncated at base which 
is not notched; 1.495 mm. long; longer than wide in ratio of 
about 23 : 9 ; not continued posterior to spinnerets. Other 
features as usual in the genus. 

Color in alcohol. First and second femora yellowish ; all other 
segments of legs a dusky yellowish. Palps light yellowish except 
the reddish brown tarsi. Chelicerae : basal segment a deep red- 
dish brown; fang yellowish. Lip a deep reddish brown, lighter 
along distal border. Maxillae yellowish in medial third and 
brown elsewhere. Carapace a deep reddish brown, darker along 
the margins; median region with a narrow dark stripe posteri- 
orly and widening at the median depression and extending to 
PLE ; all eyes except AME surrounded by black pigment. Ster- 
num : a deep reddish brown. Abdomen : nearly white dorsally 
with a few silvery spangles ; in the posterior third there are very 
poorly outlined median gray spots with a series of very narrow 
black transverse lines; a fairly broad gray stripe extends along 



6 BREVIORA NO. 68 

each lateral side ; the venter is generally white with a little gray 
around the genital area and spinnerets. 

Female allotype. Total length including nearly vertical cheli- 
cerae 3.12 mm. Carapace 1.28 mm. long; .715 mm. wide opposite 
second coxae where it is widest ; otherwise essentially as in male. 

Eyes. Central ocular quadrangle wider behind than in front 
in ratio of 5 : 4, wider behind than long in ratio of 5 : 4. Ratio 
of eyes AME : ALE : PME : PLE = 6 : 4.5 : 5.5 : 5. AME 
separated from one another by five-sixths of their diameter, from 
ALE by 1.5 times their diameter. PME separated from one 
another by slightly more than 1.6 times their diameter, from 
PLE by the same distance. Laterals separated from one another 
by the diameter of PLE. AME separated from PME by the 
diameter of PLE, hence as far from one another as the laterals 
are from one another. Height of clypeus equal to about tw^o- 
thirds of the diameter of AME. 

Chelicerae. Moderately well developed ; nearly vertical and 
parallel ; basal segment .454 mm. long and, therefore, about one- 
third as long as cephalothorax ; fang slender and evenly curved ; 
promargin of fang groove with four well-developed teeth fairly 
evenly spaced ; retromargin with four smaller and fairly evenly 
spaced teeth (delicacy of the specimen makes it difficult to ob- 
serve teeth accurately). 

Maxillae, Lip, and Sternum. Essentially as in male. 

Legs. 1243. Width of first patella at "knee" .119 mm., tibial 
index of first leg 5. Width of fourth patella at "knee" .097 mm., 
tibial index of fourth leg 7. 

Femora Patellae Tibiae Metatarsi Tarsi Totals 

(All measurements in millimeters) 



1. 


2.210 


.378 


2.015 


2.015 


.716 


7.334 


2. 


1.625 


.330 


1.170 


1.430 


.585 


5.140 


3. 


.845 


.200 


.520 


.550 


.396 


2.511 


4. 


1.495 


.265 


1.105 


1.170 


.401 


4.436 



Spines, hairs, and trichobothria essentially as in male. 

Abdomen. 1.95 mm. long; broadest near middle where it is 
.910 mm. wide ; slightly notched at base ; genital area essentially 
as shown in Figure 5. Otherwise essentially as in male. 



1956 TETRAGNATHA IN JAMAICA 7 

Color in alcohol. Abdomen : tlorsally the cardiac area is 
nearly colorless; there are numerous silvery spangles and a 
vaguely outlined folium; the venter has a central slightly gray- 
ish stripe with a stripe on each side outlined by silvery spangles. 
Otherwise essentially as in male. 

Type locality. Ilolotype male, allotype female, and three para- 
type males from Hanover, Askenish, Trail to Dolphin Head, 
Jamaica, June 24, 1954. 

Tetragnatha PAiiLESCENS F. P. Cambridge, 1903 

Eugnatha pallcscens Petrunkevitch, 1911 
r. pallescens Petrunkevitch, 1930 
T. pallescons Bryant, 1940 
T. pallescens Bryant, 1945 

Collection records: A male and a female from Ocho Rios, 
January 1929 (W. S. Brooks) ; several of both sexes from St. 
Catherine, Port Henderson, Salina, November, 1949 (Bengry, 
Lewis, Wiles) ; both sexes from St. Thomas, Lysson, June, 1954. 

Tetragnatha tenuissima O. P. Cambridge, 1889 

T. tenuissima Petrunkevitch, 1930 
T. tenuissima Bryant, 1940 
T. tenuissima Bryant, 1945 

Only one specimen, a male, has appeared in the collection 
available to me; St. Elizabeth, Magotty, May, 1953 (G. R. Proc- 
tor). 

Tetragnatha visenda sp. nov. 
(Figures 6-9) 

3Iale holotype. Total length including chelicerae 8.58 mm. ; 
without chelicerae total length 7.475 mm. Carapace 2.60 mm. 
long; 1.495 mm. wide opposite second coxae where it is widest; 
with the usual general form of the genus ; .66 mm. tall opposite 
third coxae just anterior to posterior declivity. 

Eyes. Eight in two rows as usual ; lateral ocular tubercles only 
moderately prominent; viewed from above, posterior row mod- 
erately recurved, anterior row strongly recurved; viewed from 



8 



BREVIORA 



NO. 68 



in front, anterior row moderately recurved, posterior row 
slightly procurved, all measured by centers; central ocular 
quadrangle wider behind than in front in ratio of 6 : 5, wider 
behind than long in ratio of 9 : 8. Katio of eyes AME : ALE : 
PME : PLE = 11 : 5.5 : 8 : 7.5. AME separated from one 
another by slightly more than their diameter, from ALE by 



r 




<^\ 





External Anatomy of Tetragnatha 

Fig. 6. T. visenda sp. nov. ; clieliceral teeth of male from below. 

Fig. 7. Idem; the prolateral spur of male. 

Fig. 8. Idem; the male paracymbium. 

Fig. 9. Idem; distal end of male tarsus. 

Figs. 10-11. T. versicolor Walck.; distal ends of conductors and emboli 

from Cuba and Michigan, respectively. 
Figs. 12-13. T. parva Bryant ; distal end of male tarsus and paracymbium, 

respectively. 

nearly twice their diameter. PME separated from one another 
by 2.5 times their diameter, from PLE by slightly more than 
this. Laterals separated from one another by slightly more than 
the diameter of PLE. AME separated from PME by slightly 



1956 TETKAGNATHA IN JAMAICA 9 

more tlian the diameter of AME, thus are farther from one 
another than laterals are from one another in ratio of about 12 
: 7.5. Height of clypeus equal to a little more than two-thirds 
the diameter of AME. 

Chelicerae. Well developed; moderately porrect; quite diver- 
gent in distal two-thirds of basal segment ; somewhat swollen in 
distal half; prolateral spur well developed and clearly bifid with 
the larger lobe directed inward (Fig. 6); the fang is long, 
slender, only slightly sinuate; the fang groove has the so-called 
"large tooth" with eight others on the promargin and eight on 
the retromargin (Fig. 6). 

Maxillae. Slender; considerably divergent in distal halves; 
a little more than twice as long as lip ; longer than wide in middle 
in ratio of 4 : 1. 

Lip. Only slightly wider at base than long; sternal suture 
clearly procurved; with the usual sternal tubercles well de- 
veloped at ends of sternal suture. 

Sternum. Only slightly convex; somewhat swollen opposite 
second coxae ; with the usual general form ; longer than wide in 
ratio of 12 : 7 ; continued laterally and posteriorly between all 
coxae ; posterior coxae separated by about one-fourth their width. 

Legs. 1243. Width of first patella at "knee" .395 mm., tibial 
index of first leg 5. Width of fourth patella at "knee" .260 mm., 
tibial index of fourth leg 6. 

Femora Patellae Tibiae Metatarsi Tarsi Totals 

(All measurements in millimeters) 



1. 


6.500 


1.105 


6.890 


7.475 


1.560 


23.530 


2. 


4.680 


.910 


4.225 


4.420 


.975 


15.210 


3. 


2.340 


.550 


1.397 


1.755 


.600 


6.642 


4. 


4.940 


.715 


3.835 


4.420 


.845 


14.755 


Palp 


1..560 


.370 


.520 




.850 


3.300 



Spines. All legs with spines of moderate size and length; a 
sparse coating of hair is also present. Trichobothria are present 
on femora and probably other segments of legs but their position 
has not been accurately observed. 

Palp. Tibia and patella both short with tibia longer than 
patella in ratio of about 3 : 2. The paracj^mbium is rather long 



10 BREVIORA NO. 68 

and slender with the chitinoiis knob about one-third of length 
of the structure from base (Fig. 8). The conductor terminates 
in a characteristic manner best shown in Figure 9. 

Abdomen. Slender; only slightly concave at base; widest near 
middle; 5.07 mm. long and about 1.43 mm. wide in broadest 
region ; not continued posterior to spinnerets ; other features as 
usual in the genus. 

Color in alcohol. All legs light yellowish brown, lighter below ; 
first and second somewhat dusky dorsally and dorsolaterally 
with occasional grayish patches ; third and fourth mostly lacking 
the dusky coloring and grayish patches. Chelicerae reddish 
brown, grayish along lateral surfaces. Lip dark brown with 
yellowish distal border. Maxillae yellowish in medial halves, 
darker along lateral halves. Carapace reddish brown with darker 
radiating streaks and an irregular granular border. Sternum 
reddish brown Avith darker irregularly grouped fine dots. Abdo- 
men : dorsum light yellowish because of presence of numerous 
irregular subchitinous yellowish white deposits making this re- 
gion very granular in appearance ; there are also many short 
irregular grayish lines which become more longitudinal in posi- 
tion and prominent along the lateral sides; the venter has a 
median grayish stripe with a broader granular j^ellowish stripe 
on each side together with a white spot just lateral to each an- 
terior spinneret and a smaller white spot just dorsal to the larger 
one. 

Type locality. The male holotype was taken at St. Catherine, 
Port Henderson, June 20, 1954. One male paratype is in the 
collection from a house in St. Andrew, August, 1955 (G. R. 
Proctor). 

THE GENUS IN CUBA 

Tetragnatha antillana Simon, 1897 

There is but one specimen, a male, in the collection in the 
Museum of Comparative Zoology but the species has been re- 
corded from several localities. 

Tetragnatha caudata Emerton, 1884 
Two females are in the collection in the Museum of Compara- 
tive Zoology, both taken at different times in Soledad gardens. 



1956 TETRAGNATHA IN JAMAICA 11 

Tetragnatha elongata Walckenaer, 1805 

This species is well represented by both sexes in the collection 
and appears to be the most common of all of the eight species 
recorded from the island. 

Tetragnatha guatemalensis 0. P. Cambridge, 1889 

T. banhsi McCook, 1893 

T. seneca Seeley, 1928 

T. banlcsi Levi and Field, 1954 

Drs. Gertsch and Levi have apparently agreed that T. seneca 
Seeley is the same as T. hanksl MeCook. I have made careful 
comparisons of the specimens from Cuba identified as T. seneca 
Seeley with my numerous specimens of T. guatemalensis 0. P. 
Cambridge from Panama and other parts of Central America 
with the result that I am convinced that here we have another 
case of synonymy. The characteristics of eyes, several features 
of the male palps such as vermiform distal end of the paracym- 
bium and course and shape of both conductor and embolus to- 
gether with the general characters of the chelicerae all point 
toward this conclusion. Some may object that I am taking too 
much liberty with the cheliceral teeth because the "large tooth" 
is absent in T. seneca Seeley. This "large tooth" is not well 
developed in T. guatemalc7isis 0. P. Cambridge and could very 
well have been reduced to the condition found in T. seneca 
Seeley. The specimens in the Museum of Comparative Zoology 
identified as T. ha^iksi McCook also agree well with T. seneca 
Seeley as concluded by Levi and Field (1954). 

Tetragnatha orizaba Banks, 1898 

There are several specimens of both sexes from several locali- 
ties in Cuba. Also recorded from Hispaniola but from no other 
place in the West Indies so far as know^n to the author of this 
paper. 

Tetragnatha pallescens F. P. Cambridge, 1903 

Both sexes are represented in the collection from Havana and 
Soledad. 

Tetragnatha tenuissima 0. P. Cambridge, 1889 
Both sexes have been recorded from several localities. 



12 BREVIORA NO. 68 

Tetragnatha versicolor Walckenaer, 1841 

T. extensa Emerton, 1884 

T. d&ntigera F. P. Cambridge, 1903 

T. extensa Seeley, 1928 

The specimens from Soledad identified as T. dentigera F. P. 
Cambridge are, I believe, correctly recognized. These are espe- 
cially interesting because of the fact that I have been forced to 
the conclusion that T. dentigera F. P. Cambridge is a synonym 
for T. versicolor Walck. I have examined a large number of 
specimens assigned to the latter species and as many as possible 
of the former. The cheliceral teeth, several features of the male 
palps (paracymbium, conductor, embolus) and the eyes all point 
rather decisively toward the synonymy which I have indicated. 
The tip of the conductor is like nothing else in the genus so far 
as I have seen. I have provided a drawing of the tip of the con- 
ductor from a specimen collected in Cuba and another from a 
specimen of T. versicolor taken in Michigan. There are slight 
differences but the basic plan is the same and the differences are 
well within the normal variation of a species (Figs. 10, 11). 

THE GENUS IN HISPANIOLA 

Tetragnatha antillana Simon,, 1897 

T. festina Bryant, 1945 (male only) 
T. haitiensis Bryant, 1945 (females) 

The males of T. festina Bryant have the bifid paracymbium, 
other features of the male palpi, eyes, and general cheliceral 
characters associated with T. antillana Simon. 2\ haitiensis 
Bryant has the general form, cheliceral characters, and genital 
area characteristic of females of T. antillana. The small differ- 
ences noted by Miss Bryant and myself are all, I believe, within 
the normal variation for a species ranging over a Avide area. 

Tetragnatha confraterna Banks, 1909 

T. eloufjata Bryant, 1945 

One female from Puerta Plata, Dominican Republic, was 
identified as T. elongata Walck., 1805. I have examined this 
specimen very carefully and I am convinced that it has been 



1956 TETRAGNATHA IN" JAMAICA 13 

misidentified. Its slightly extended abdomen, cheliceral char- 
acters, eyes, and f^enital area seem to place it in the species 
7'. confraterna Banks where it is provisionally left. 

Tetragnatha ORIZABA Banks, 1898 

r. Orizaba Bryant, 1945 

The specimens of both sexes from the Dominican Republic 
seem to agree well with our current understanding of this species. 

Tetragnatha pallescens F. P. Cambridge, 1903 
r. pallescens Bryant, 1945 

Numerous specimens of both sexes from Haiti seem to indicate 
that this may be the most common species in Hispaniola. 

Tetragnatha tenuissima 0. P. Cambridge, 1889 

T. tenuissima Bryant, 1945 

T. festina Bryant, 1945 (females only) 

The cheliceral teeth, eyes, general form, lack of leg spines, and 
the genital area all indicate that T. festina females belong with 
T. tenuissima 0. P. Cambridge. 

THE GENUS IN PORTO RICO 

I have had very little opportunity to study the species of 
Tetragnatha from Porto Rico. Petrunkevitch (1930) listed the 
following species from this island : T. antillana Simon ; T. 
elymiquensis Petrunkevitch; T. lahoriosa Hentz; T. pallescens 
F. P. Cambridge ; T. piscatoria Simon ; T. suhextensa Petrunke- 
vitch; T. tenuissima 0. P. Cambridge; T. vicina Simon. There 
appears to be considerable doubt about the occurrence of T. 
vicina in Porto Rico. In 1947 Miss Bryant described T. parva 
from the Luquillo Mountains thus bringing the total number of 
recorded species in this island to nine. Two figures of the tip of 
the conductor and closely related structures have been prepared 
to supplement those provided by the author of the species (Figs. 
12,13). 



14 BREVIORA NO. 68 

THE GENUS IN THE VIRGIN ISLANDS 

Very little opportunity has been afforded me to study the 
spiders of these small islands. I have carefully examined all 
specimens, however, now in the collection of the Museum of Com- 
parative Zoology and am prepared to summarize my findings as 
follows: The vial labelled T. antillana Simon contains specimens 
belonging to this species but it also contains two females which 
I am tentatively assigning to T. confraterna Banks. The same 
vial contains a male palp which I believe was derived from this 
same species. Perhaps we may at least tentatively regard this 
species as being in the West Indies. The single male assigned 
to T. piscatoria Simon is, in my judgment, not this species but 
a specimen belonging to T. visenda sp. nov. 



GENERAL SUMMARY 

At the present time it seems that either fifteen or sixteen spe- 
cies of Tetragnatha may be considered to be known from the 
West Indies under consideration here. T. elijunquensis Petrun- 
kevitch, T. exigua sp. nov., T. parva Bryant, T. suhextensa 
Petrunkevitch, and T. visenda sp. nov. are known only from the 
islands from which they were described. T. antillana Simon has 
been recorded from all islands mentioned here. T. caudata 
Emerton has been recorded from Cuba and Jamaica. T. con- 
fraterna Banks is probably to be regarded as present in both 
Hispaniola and the Virgin Islands. T. elongata Walck. is only 
certainly known from Cuba. T. guatenialensis 0. P. Cambridge 
is now known only from Cuba. T. lahoriosa Hentz is, apparently, 
present only in Porto Rico. T. orizaha Banks appears to be in 
Cuba and Hispaniola. T. pallescens F. P. Cambridge has been 
recorded from all islands considered here except the Virgin 
Islands. T. piscatoria Simon is known only from Porto Rico 
among the islands considered here but it was described from 
St. Vincent. T. tenuissima 0. P. Cambridge is known from all 
of these islands except the Virgin Islands. T. vicina Simon has 
been recorded from Porto Rico but its presence there appears 
doubtful. 



1956 TETRAGNATHA IN JAMAICA 15 

BIBLIOGEAPHY / 

Banks, Nathan 

1898. Aiaclmida from Baja California and other parts of Mexico. 

Proc. California Acad. Sci., Ser. 3, Zoology, 1. (7): 205-309, 

5 pis. 
1909. Arachnida from Costa Rica. Proc. Acad. Nat. Sci. Philadelphia, 

April, 1909: 194-234. 

Bryant, Elizabeth B. 

1940. Cuban spiders in the Museum of Comparative Zoology. Bull. 
Mus. Comp. Zool. at Harvard College, 86 (7): 249-532, 22 pis. 

1942. Notes on the spiders of the Virgin Islands. Bull. Mus. Comp. 
Zool. at Harvard College, 89 (7): 319-363, 3 pis. 

1945. The Argiopidae of Hispajiiola. Bull. Mus. Comp. Zool. at Har- 
vard College, 95 (4) : 359-418, 4 pis. 

Cambridge, O. P. and F. P. Cambridge 

1889-1905. Arachnida-Araneida. Vols. I-II. In: Biologia Centrali- 
Amerieana. Dulau & Co., London. 

Levi, Herbert W. and Hoavard M. Field 

1954. The spiders of Wisconsin. Amer. Midland Natur., 51 (2): 440- 
467, 113 figs. 

Petrunkevitch, Alexander 

1930. The spiders of Porto Eico. Pt. 2. Trans. Connecticut Acad. 
Arts and Sciences, 30: 159-355, 240 figs. 

Seeley, E. M. 

1928. Eevision of the spider genus Tetragnatha. Bull. New York State 
Mus., 278: 99-150. 

WiEHLE, H. 

1939. Die einheimischen Tetragnatha-Arten. Nova Acta Leopoldina 
(Halle), N.F. 6 (41) : 363-386. 



BREVIORA 

Mmseiiinni of Cooipsirative Zoology 



Cambridge, Mass. January 31, 1957 Number 69 

A NEW ZODARIID SPIDER FROM PANAMA 
By Arthur M. Chickering 

Albion College, Albion, Michigan 

Until the discovery of the species described in this paper only 
one zodariid spider was known from Panama, althouoh a few 
others had been reported from Guatemala and Mexico. The 
single species hitherto known from Panama is Store7ia harroana 
(Chamberlin), 1925. 

During the summer of 1954, while sifting decaying hay along 
the roadside near Corozal, Canal Zone, I discovered one im- 
mature and two mature males together with one mature and 
three immature females all belonging to what I regard as a new 
species referred to the genus Leprolochus Simon, 1892. So far 
as I know, only two species in this genus have been previously 
reported. The genotype of this rather remarkable genus, Lepro- 
lochus spinifrons Simon, Avas from Brazil and L. parahyhae 
Mello-Leitao, 1917 was from Venezuela. The latter is known only 
from females. The description of this new species of Leprolochus 
is given in this brief paper in accord with my usual descriptive 
formula, and the types have been deposited in the Museum of 
Comparative Zoology. 

Genus LEPROLOCHUS Simon, 1892 

Leprolochus decoratus sp. nov. 
(Figures 1-6) 

Male holotype. Total length 2.21 mm., including spines pro- 
jecting from head region. Carapace 1.235 mm. long, .770 mm. 
wide opposite second coxae where it is widest; bluntly rounded 
in front and with seven robust and bluntly pointed black spines 




External Anatomy of Leprolochtis decoratua 
Fig. 1. Anterior cephalic region of male from above. 
Pig. 2. Chelicerae, eyes, and special spines of male from in front. 
Figs. 3-4. Tarsus and tibia of male palp; ventral and retrolateral views, 

respectively. 
Fig. 5. Anterior cephalic region of female from above. 
Fig. 6. Ventral view of epigj'num. 



1957 NEW ZODARIID SPIDER FROM PANAMA 3 

projecting forward somewhat below AME (Figs. 1-2) ; the para- 
type male has nine of these spines; with a short and shallow 
median longitudinal thoracic groove opposite the third coxae ; 
finely granulated, especially over pars thoracica; gently arched 
from PME to posterior border with highest part shortly behind 
these eyes. 

Eyes. Eight in two rows; posterior row only slightly longer 
than anterior row; anterior medians dark, all others light. 
Viewed from above, posterior row strongly procurved, anterior 
row moderately so. Viewed from in front, both rows strongly 
procurved. Ratio of eyes AME : ALE : PME : PLE = 5.5 : 
5.25 : 5 : 5. AME separated from one another by about 4/5 of 
their diameter, from ALE by about 8/5 of their diameter. PME 
separated from one another by about 9/5 of their diameter, from 
PLE by nearly twice their diameter. Laterals separated from 
one another by about 7/10 of the diameter of PLE. Central 
ocular quadrangle wider behind than in front in ratio of about 
9 : 7, only slightly longer than wide behind. Height of clypeus 
equal to somewhat more than six times the diameter of AME. 

Chelicerae. Stout, vertical, parallel ; basal segment .352 mm. 
long ; with moderately distinct basal boss and fine striations 
anterior to boss ; the fang is weak, short, and apparently evenly 
curved ; fang groove obscure and with no teeth observed ; the 
pro-margin has a fleshy ridge with a row of stiff bristles. 

Maxillae. Distinctly convergent; tapered distally and nearly 
meeting beyond lip. 

Lip. Nearly triangular; wider at base than long in ratio of 
about 3:2; reaches about 2/5 of the length of the maxillae. Ster- 
nal suture straight. 

Sternum. Cordiform ; widest between first and second coxae 
where it is wider than long in ratio of 25 : 23 ; anterior border 
nearly twice as wide as base of lip ; very convex ; sharply pointed 
at posterior end but not extended between fourth coxae which 
are separated by 4/5 of their greatest width. All coxae moder- 
ately elongated. 

Legs. 4312. Width of first patella at "knee" .108 mm., tibial 
index of first leg 15. Width of fourth patella at "knee" .130 
mm., tibial index of fourth leg 15. 



4 




BREVIORA 






NO. 69 




Femora 


Patellae Tibiae 
(All measurements 


in 


Metatarsi 
millimeters) 


Tarsi 


Totals 


1. 


.660 


.264 .462 




.660 


.528 


2.574 


2. 


.638 


.250 .440 




.660 


.506 


2.494 


3. 


.638 


.275 .450 




.748 


.484 


2.595 


4. 


.811 


.275 .660 




.990 


.690 


3.426 


Palp 


.330 


.110 *.443 






.374 


1.257 



* Including much extended apophysis. 

Spines. All legs with inconspicuous spines difficult in some 
instances to distinguish from coarse bristles. First leg: femur 
dorsal 1-1-1-2, prolateral 1 distal, retrolateral and ventral 1 weak 
proximal ; patella with numerous coarse bristles but probably 
only prolateral 0-1-1-0 spines; tibia dorsal 1-1-1 (all weak), 
prolateral 0-1-0-1, retrolateral 0, ventral only 2 distal; meta- 
tarsus 0. Second leg : essentially as in first except metatarsus 
which has a tuft of ventral bristles at distal end together with 
four spines on ventral and lateral sides. Third leg : femur 
apparently only dorsal 1-0-1-2, patella dorsal 1-1, prolateral 
1-1-0, retrolateral 0-1-0, tibia dorsal 1-1-1, prolateral appar- 
ently 0-1-0-0 and wdth a group of distal bristles forming a comb- 
like structure, retrolateral 1 distal, ventral only distal 2 ; meta- 
tarsus only with distal spines essentially as in second. Fourth 
leg : essentially as in third. 

Trichohothria. Observed as follows : tarsi with a row of two 
or three with the most distal very long ; metatarsi the same ; 
tibiae apparently with two rows of two each in dorsolateral 
positions ; palpal tibia with two dorsal in a row with distal one 
very long. 

Claws. Two claws throughout, each pectinate in a single row ; 
with no claw tufts. 

Palp. Patella very short ; tibia with a short body but with a 
lateral apophysis which extends nearly the full length of the 
tarsus ; also with a pair of retrolateral spines as shown in Figures 
3 and 4. 

Abdomen. Regularly ovoid ; longer than wide in ratio of 17 : 
13; overlaps cephalothorax only slightly; widest slightly behind 



1957 NEW ZODARIID SPIDER FROM PANAMA 5 

the middle ; provided with a moderately well developed scutum ; 
well supplied wdth short appressed and recurved stiff bristles 
both black and transparent ; postabdomen and anal tubercle 
project posterior to scutum ; venter well chitinized ; with a short, 
recurved lip just anterior to genital groove ; tracheal spiracle 
obscure but apparently just anterior to base of spinnerets and 
appears to be marked by a row of relatively long stiff bristles; 
position of probable vestigial colulus obscure. With six pairs of 
spinnerets partly obscured by a group of bristles. 

Color in alcohol. Carapace : cephalic parts a light brown ; re- 
mainder a medium brown with darker streaks. Chelicerae light 
brown with other mouth parts and sternum yellowish brown. 
Legs : all coxae nearl}^ white ; femora brown, lighter beneath ; 
other segments light brown. Abdomen : dorsally light brown 
with three irregularly dark reddish brown cross bars in the 
posterior two thirds and an irregular central stripe reaching 
to base from first cross bar ; the posterior end of the abdomen 
behind the scutum and the postabdomen are white; the venter 
is yellow with varying shades. 

Female allotype. Total length 3.90 mm. Carapace 1.625 mm. 
long; 1.105 mm. wide opposite interval between second and third 
coxae where it is widest ; gently arched from PME to median, 
short, longitudinal thoracic groove from w^liich it descends rather 
gradually to posterior border ; cephalic portion well separated 
from thoracic part ; with eleven robust black spines in the posi- 
tion of the seven similar spines in the male (Fig. 5). 

Eyes. Ratio of eyes AME : ALE : PME : PLE = 6.5 : 5.3 
: 6 : 6. AME separated from one another by about their diam- 
eter, from ALE by twice their diameter. PME separated from 
one another by 13/6 of their diameter, from PLE by 17/6 of 
their diameter. Laterals separated from one another by nearly 
the diameter of ALE. Central ocular quadrangle wider behind 
than in front in ratio of 4 : 3, longer than wide behind in ratio 
of 7 : 6. Height of clypeus equal to about eight times the diam- 
eter of AME. 

Chelicerae, Maxillae, Lip, and Sternum. Essentially as in male. 

Legs. 4321. Width of first patella at "knee" .16245 mm., tibial 
index of first leg 15. Width of fourth patella at "knee" .2166 
mm., tibial index of fourth leg 15. 



BREVIOBA NO, 69 

Femora Patellae Tibiae Metatarsi Tarsi Totals 

(All measurements in millimeters) 



1. 


.975 


.390 


.682 


.715 


.650 


3.412 


2. 


.975 


.390 


.650 


.975 


.650 


3.640 


3. 


1.040 


.455 


.780 


1.235 


.780 


4.290 


4. 


1.170 


.455 


1.000 


1.625 


1.040 


5.290 



Spines. First leg : femur dorsal 1-0-1-2, probably prolateral 
and retrolateral only distal 1 or 2, ventral Ir (weak) near middle ; 
patella only prolateral 0-1-1-0 ; tibia dorsal 1-1-1, prolateral 1-1-1, 
retrolateral 0-1-0-1, ventral O-lr-0-2 ; metatarsus apparently with 
only two distal but with many spiniform bristles. Second leg : 
femur probably as in first; patella dorsal 1-1, prolateral 0-1-1-0, 
retrolateral 0-1-0 ; tibia probably as in first ; metatarsus as in 
first except with a distal ventral brush of bristles and probably 
a group of 3 or 4 weak spines. Third leg: femur, patella, and 
tibia essentially as in second ; metatarsus prolateral 0-1-0, retro- 
lateral 0-1-0-1, ventral with brush of bristles as in second and 
with five clear distal spines on ventral and both lateral surfaces. 
Fourth leg: essentially as in third except brush is absent but 
the spines are retained. Palp : with numerous spines on femur, 
patella, tibia, and tarsus ; tarsal claw finely pectinate with about 
ten slender teeth ; claw is opposed to a chitinous tubercle. Trich- 
obothria essentially as in male so far as observed. 

Abdomen. Well rounded, oviform; 2.405 mm. long, 1.95 mm. 
wide near middle; without scutum such as that possessed by 
male; with six spinnerets, the anterior pair quite robust and 
somewhat the longest ; without definite colulus ; tracheal spiracle 
close to base of the anterior spinnerets. 

Epigynum. Simple but well developed. There is a central 
shallow depression bordered anteriorly by a strongly chitinized 
recurved border. Two internal tubules occur at the posterior 
boundary and on each side there is a relatively large spermatheca 
(Fig. 6). 

Color in alcohol. Carapace essentially as in male except that 
at the base of pars cephalica there is a dorsal, somewhat oval 
brown spot on each side from which a thin irregular line extends 
forward to PLE. The dorsal abdominal region is also very sim- 
ilar to that of male except that the reddish brown markings are 



1957 NEW ZODARIID SPIDER FROM PANAMA 7 

broader and more united. Otherwise essentially as in male. 

Type locality. The male holotype, female allotype, one mature 
male paratype together with four immature specimens from 
Corozal, C. Z., July 10, 1954. 

BIBLIOGRAPHY 

Banks, Nathan 

1929. Spiders from Panama. Bull. Mus. Comp. Zool. at Harvard Col- 
lege, 69:53-96, 4 pis. 

Cambridge, O. P. and F. P. Cambridge 

1889- Arachnida-Araneida. In: Biologia Central- Americana. Dulau 
1905. & Co., London. 

Chamberlin, R. V. 

1925. Diagnoses of New American Arachnida. Bull. Mus. Comp. Zool. 
at Harvard College, 67:211-248. 

Chickering, Arthur M. 

1947. The Male Allotype and Female Hypotype of Storena Barroana 
(Chamberlin). Papers Michigan Acad. Sci., Arts, Letters, 31 
133-140. 

ROEWER, C. Fr. 

1942. Katalog der Araneae. Vol. 1. Bremen. 

Simon, Eugene 

1892- Histoire Naturelle des Araignees. Deuxi^me Edition. 2 Vols. 
1903. Librairie Encyclopedique de Roret, Paris. 



BREVIORA 

Museiiam of Comparative Zoology 



Cambridge, Mass. January 31, 1957 Number 70 



"ANGUIMORPH" TOOTH REPLACEMENT IN 

AMPIIISBAENA ALBA LINNAEUS, 1758, AND 

.1. FULIGINOSA LINNAEUS, 1758 

(REPTILIA: AMPHISBAENIDAE) 

By Carl Gans 

In the course of an investigation into the status of the acro- 
(lont amphisbaenids it proved necessary to prepare the skulls of 
some related forms. When cleaning the mandible of a female 
specimen of Amphishoenn alha Linnaeus, 1758, it was noted that 
six to eight teeth were in the process of being replaced, while 
two or three others had only recently moved into position, and 
were as yet but imperfectly fused to the dentaries. The replace- 
ment teeth appeared to lie interdentally, between, rather than 
below^, their predecessors. 

McDowell and Bogert, in their recent revision of the angui- 
morph lizards (1954, pp. 102, 104, tig. 30), have stated that 
"alternate" tooth replacement was restricted to this group, all 
other lizards possessing "vertical" replacement. The amphis- 
baenids are generally considered to be Scincomorpha (Camp, 
1923, p. 296) and if the distinction between an anguimorph and 
iiou-anguimorph pattern is as clearcut as McDowell and Bogert 
suppose, the occurrence of a pattern descriptively alternate 
would thus be unexpected here. For this reason and because 
certain other aspects of dental replacement in these forms seem 
worthy of special attention, it appears desirable to describe the 
female specimen mentioned above, as well as two specimens of 
A. fuligmosa Linnaeus, 1758, in which a similar replacement 
pattern was noted, in the hope of stimulating further research 
into these matters. 



BREVIOKA 



NO. 70 



The mandibular ilentition of Amphishaena alba 

Figures 1 to 5 show lal)ial and lingual views of the mandil)h> 
of an adult (body length 440 mm.; length of mandibular ramus 
15 mm.i) female specimen (MCZ 54299) from "Brazil." This 
had been cleaned by dissection following controlled applications 
of full strength commercial bleach to selected portions of the 
soft parts. 




Fig. 1. (Upper) Amphisbaena alba. Lingual view of right mandibular 
ramus of MCZ 54299. 



Fig. 2. (Lower) AiiiiiliLsb(n no alba. Labial view of same niamliliular 
ramus as Figure \. 

1 The length of the mamllbulur riinius has been selected as a convenient indi- 
cator of the total length of the specimen and Is hence abbreviated as Imr. 



1957 



AMl'lllSBAENID TOOTH REPLACEMENT 



The first pa it of this description covers the general aspect of 
the tlentitiou and ap])lies equally well to the female cited and to 
larger si)eeinieus listed below. The description of the replacement 
l)attern and of individual or possibly ontogenetic variation fol- 
lows upon this. 

The dentition is weakly pleurodont, the height of the dorsal 
margin above the lingual shelf on which the teeth rest barel}^ 




Fig. 3. AmpMshaena alba. (MGZ 54299) Sketch of lingual view of right 
mandibular ramus showing location of the replacement teeth. 

equaling the width of the tooth base. There are eight teeth on 
each side ; the third is largest, the fourth, fifth, sixth, and seventh 
are slightly smaller, while the second, eighth, and first are pro- 
gressively smaller in that order. All of the teeth are curved, this 
being most noticeable near the tip, and the curvature is directed 
medially and slightly caudad. There is a rotation in the tooth 
alignment so that the planes of curvature of the individual teeth 
do not lie parallel to one another. The teeth are of oval cross- 
section w4th the long axis lying in the plane of curvature. 

The base of the fully formed tooth is hollow, the pulp cavity 
extending two-thirds of the total height of the tooth. About 
midway up the tooth the diameter of the cavity contracts so as to 
continue upwards as a thin cylindrical tube. 

There is no tooth-bearing shelf as in a typical pleurodont den- 
tition; instead, a ridge of bone (Fig. 6) rises to the projecting 
dorsal margin of the dentary between each two adjacent teeth 
(where replacement teeth are present this ridge is hollowed out 
to afford lodgement for the pit of the new tooth). Consequently 



UREVIORA 



NO. 70 



tilt' base of eaeh tootli is almost entirely surrouiulcd by bone, to 
^vhicll it is ankylosed by a ring of cement. The lowest exposed 
])oint lies on the lingual aspect and careful, but thorouph, prepa- 
ration shows here a single round foramen.' The foramen leads 
into the ptdp cavity and ])resumably carries its vascular and 
nerve siipi)ly. A line extending to the level of the tip of the 
interior cavity is visible on the lingual side. Inspection along the 
lingual aspect of even the smallest cap-shaped tooth germs re- 
veals a slight scalloping so that this line may be formed during 




Fig. 4. AinpJtLshacna alhn. Lingual view of the left niaiulilnilar ramus of 
MCZ 54299. 

tooth development by the fusion of the anterior and posterior 
portions. Several of the skulls showed longitudinal cleavage of 
the teeth along this line, which coincides with the long axis of 
the oval tooth cross-section. 

\'arious stages of tooth replacement are shown in MCZ 54299. 
In the subsequent description of it and other specimens, L and 
R will stand for left and right mandibular ramus, and the number 
following this for the particular tooth or alveolus counting from 
front to back. 

At L-7, R-3 and R-4 the replacement tooth is but a small hollow 
conical shell, thicker at the top than at the sides, its lower edge 
somewhat excavated on the lingual side. It lies in a small and 
deep depression in the deutary in line with the posterior edge 

1 These foramina, as shown in Figure 6, are of a larger specimen, which could 
be cleaned completely without danger of dislodging the firmly fused teeth. 



[9i 



)* 



AMPHISBAENID TOOTH KKPI.ACEMENT 



of the tooth it is replaciiij?. As a result, this pit lies on the 
ascendino; ridoe directly between the tooth to be replaced and 
the one posterior to it. Even at this stage there is already some 
erosion into the base of the tooth beinj^- replaced. 

L-4 and L-5 show a slightly more advanced stage with pro- 
gressive erosion into the precursor and an increase in size of 
the enamel cap of the new tooth. 

In R-8 the replacement tooth has reached approximately one- 
third of its final height, though its tip is still below the level of 
the projecting dorsal margin of the dentary. It has, however, 
destroved almost half of the base of the tooth Iving above it. 



8 7 6 5 4 3 




Fig. 5. Amphishacna alba. (MCZ 54299) Sketch of lingual view of left 
iiiniidilmlar ramus showing location of the replacement teeth. 

At R-6 the tooth has just moved into its final position, its 
predecessor having been pushed out at some prior stage. It is 
still only loosely held in place by struts of cement, and appears 
slightly smaller, and thinner-walled than its neighbors. Its dark 
appearance is due to the contents of the large, soft-tissue-filled 
pulp cavity being visible through the translucent walls. This 
may indicate that the buihl up of the internal dentine layers is 
not completed until after the tooth is finally cemented into place. 

The heavier enamel and more opaque, shell-like appearance of 
L-3 may indicate a more advanced stage of development. It is, 
however, still darker than its neighbors, and its base is still far 
from fused to the dentary. Its slightly cocked position seems to 
indicate that it is not fully aligned as yet. 

All of the replacement teeth except the last two lie freely in 
the soft tissues and show no fusion to the dentaries. 



6 



BREVIORA 



NO. 70 



Another interesting point is that the alveolus formation ap- 
pears to start in the interdental ridge and only begins to extend 
into the base of the precursor tooth as growth takes place. No 
pits or alveoli were found at the base of any tooth that Avas not 
undergoing replacement. 

Though the replacement pattern is interdental it differs from 
that described by Camp (1923, p. 329, fig. H) for Gerrhonotus s. 
scincicauda (Skilton), 1849 (and from that seen in a skull of the 
same form in the MCZ). In this specimen there is no trace of 
cavity or alveolus formation and the replacenumt tooth appears 




Fig. 6. Ampliishaena alha. (MCZ 32257) Dorsal view of tip of mandible, 
to show interdental bone ridges, basal foramina, and fusion lines. 

to lie always in the layer of tissue next to the bone. Gerrhonotun 
.s. scincicauda is also definitely pleuro<lont and does not possess 
the interdental ridge that produces what is almost a sub-thecodont 
condition in Amphishaena alba. 

The pattern in aJha is actually closest to that of the snakes 
( Bogert, 1943, p. 327 ff.). Here the replacement occurs in pairs, 
alternate teeth being replaced, while those between them arc 
functional. This results in the characteristic tooth - alveolus - 



1957 AMl'lllSBAENID TOOTH REPLACEMENT 7 

tooth - alveolus succession of the prepared dentate bones of the 
snake skull. Ainphishaena alha differs from this in that at each 
locus only one replacement tooth appears to be present at a time, 
and that this erodes its own pocket into the interdental ridge. 

Four Aiitphisbaena alba skulls and mandibles from museum 
collections were available for comparison (MCZ 4031 — Brazil, 
linr = 23.5 mm; MCZ 32256 — Surinam, Imr = 20 mm; MCZ 
32257 -- Brazil, Imr = 22.5 mm ; AMNH 73233 — no data, Imr 
^ 17 mm). In only one of these was any soft tissue still present 
along- the lingual aspect of the dentary, but in this as in the 
otiiers there was not the slightest indication either of replace- 
ment teeth or of eroded areas in the interdental ridges which 
might have lodged replacement teeth. This lack of evidence of 
tooth replacement may perhaps be related to the fact that all of 
the skidls were larger (see Imr above), and hence probably be- 
longed to older specimens. The tip of one of these lower jaws is 
tigured (Pig. 6) and illustrates not only the basal foramina pre- 
viously referred to, l)ut also the interdental bone ridges and the 
strong cementing of llic tooth bases. The complete absence of 
evidence of replacement activity in these adult specimens, con- 
trasted Avith the large number of teeth undergoing almost simul- 
taneous replacement in the above described smaller specimen, 
may indicate that the ability to replace the teeth is lost in the 
adults. 

It is interesting to observe that there is no evidence for the 
replacement of maxillary oi- pi-emaxillary teeth in any of the 
skulls of .1. aJha examined. 

Tlie mandibular dentition of Ainphishaena fuliginosa 

Ail available dried amphisbaenid skulls were examined for 
evidence of tooth replacement. Only two medium-sized speci- 
mens of Amphishaena fuliginosa (MCZ 2154 — South America, 
lmr-8mm ; MCZ 7799 — Riobamba, Ecuador, lmr-8 mm ; listed 
by Zangerl 1944, p. 426 as specimens A and B) demonstrated a 
discernible tooth replacement pattern.^ 

Figures 7-10 show lingual views of two of the mandibles of 

1 A very clean skull of Rhinriira floridana Baird, (1S5S) (Gainesville. Florida) 
from the' collection of Walter Auffenberg also reveals that some sort of tooth 
replacement occurs in this form. Since there are many reasons such as over- 
cleaning', etc., which might explain the absence of tooth replacement evidence in 
any given specimen, no useful purpose would be served by listing the names or 
numbers of specimens examined with negative resuUs. 



8 



BREVIORA 



NO. 70 



A. fuliginom. The dentition is again pieurodont, with the lateral 
shelf slightly higher than the width of the tooth base. The teeth 
appear to lie in a slight trench formed between the lingual shelf 
and the ascending face of the dentarv. This trench is crossed 
by only faint and occasional ridges. 

There arc seven teeth or lar<>e alveoli in each of the four 




Fig. 7. (Upper) Amphishaena fnliginosa. (MCZ 2154) Lingual view of 
loft iii,niidit)nl;ir ramus. 

Fig. 8. (Lower) Amphisbaena fuliginosa. (MCZ 7799) Lingual view of 
liglit ninnrliliular ramus. 



\'Xu 



AMI'IUSBAENID TOOTH REPLACEMENT 



iiiaiHliliular i-aiui. Ilowcvcr, tlu' broken tooth spacinji' ot" the 
soiiu'wliat luacoratod nuuidil)l(' of one of tlic spccinicns (i\ICZ 
779!)) (Figures 8. 10) leads one to suspect that Ave are dealinj; 
with a row of ei<rht teeth. Tlie third and fourth teeth are a<i'ain 
the hu'ji'est, with numbers five, six and two equal to each other 
and sliji'htly smaller, while seven and one are smallest. In other 
aspects the mandibles elosel}' resemble those of A. aiha though 
the cementing of the teeth is not quite as solid. 

Three replacement teeth are visible in each of the mandibular 
rami of MCZ 2154 and two or three in each of MCZ 77!)!). These 



7 6 5 4 3 2 1 





Fig. 9. (Upper) Amphisbaena fuliginosa. Sketch of mandibular ramus 
shown in Figure 7 to show locations of replacement teeth. 

Fig. 10. (Lower) Amphisbaena fuliginosa. Sketch of mandibular ramus 
shown in Figure 8 to show locations of replacement teeth. 



10 BREVIORA NO. 70 

again lie clearly interclentally and the general pattern of the 
dentition is very similar to that previously described for A. alha. 
The largest replacement tooth, the precursor of which is still in 
jiosition, is but slightly higher than the dorsal margin. 

However, there is one item apparent in these jaws that cannot 
be seen in the specimen of A. alha. where the erosion always takes 
place on the posterior edge of the tooth undergoing replacement. 
In A. fuligmosa there are several instances of teeth eroded on 
the front only or on both front and rear. This is the ease in R-5 
and L-7 of MCZ 2154, as well as in R-3 and R-4, and possibl}' 
L-4 of MCZ 7799. Both in R-5 of MCZ 2154 and in R-3 of MCZ 
7799 the developing anterior replacement tooth is considerably 
larger than the tooth that is eroded. The anterior tooth has thus 
eroded itself an alveolus wider than that of its precursor. The 
general spacing indicates that what is involved here is a re- 
arrangement, respacing or perhaps a change in the tooth number, 
possibly due to the growth of the mandible. 

Both sets of upper jaws show evidence of tooth replacement, 
with alveoli present above both maxillary and in-emaxillary teeth. 
As far as can be seen from the skulls, which are somewhat over- 
cleaned for this purpose, the replacement proceeds alternately 
as described above. The only complications are due to the differ- 
ent alignment of the various teeth which make terms like "alter- 
nate" and "vertical" very difficult to apply here. 



Discussion and Summary 

iu AnipJiisha< 11(1 alba and A. fuliginosa: 

The teeth ai'c plciirodoiit in that they lie against the lingual 
side of the dentary on a low shelf. 

A ridge rises between each two teeth. This character, which 
gives the dentition a sub-thecodont appearance, is most strongly 
expressed in older or larger individuals. 

The teeth when fully grown have a pulp cavity extending 
through more than two-thirds of their total height, and are fused 
to the dentary by a heavy layer of cement around their base. 
Vascular supply reaches the pulp cavity by a foramen located 
on the ventral edge of the tooth 's lingual aspect. 

Tooth replacement is alternate, with the developing tooth lying 
in a deep pit close to the posterior edge of its precursor. During 



U),")7 A.MI'IIISBAKXII) TOOTH REPLACEMENT 11 

jiiowtli it ci'odcs away the posterior aspect of its i)recursor and 
sometinit's the anterior face of the tooth next in line. The latter 
featnre may ])rovide for rearran<iement or ehan<'e in nnml)er of 
teeth. As the mandible lengthens and the individnal teeth in- 
erease in size, it becomes i)ossible for the rei)lacement teeth to 
realign themselves (thus extending the lengtli of the tooth row. 
and maintaining or increasing the interdental gap), rather than 
being restricted to entering the exact alveolus vacated by their 
precursor. Shortly after the new tooth extends higher than the 
dorsal margin it either displaces the tooth above it or the latter 
breaks away due to the dissolving of its base. 

When initially entering the alveolus the new tooth is dark and 
lias but a thin coating of enamel. Internal dentine deposition 
api)ears to continue until the tooth is well cemented into place. 

In AivpJiishacna alho there is evidence that tooth replacement 
t'ither becomes rare or stops altogether beyond a certain size. If 
tiie function of the replacement were related to the provision of 
larger teeth, this phenomenon might result from a flattening of 
the growth curve with age. 

None of the skulls of A. alba examined showed any evidence 
of tooth replacement on maxillaries or premaxillaries. 

Beth taxouDmieally and in a descriptive sense these results 
modify tlie simple j^icture presented by McDowell and Bogert. 
It is hoped that this brief note will stimulate supplementary 
investigation of this problem in every family and genus of the 
Squamata. This seems particularly desirable since even a cur- 
sory inspection of the lizard skeletons in the MCZ shows such 
divergence in the patterns ^ of tooth replacement that clearly 
they cannot be described in terms of just tw'o categories — angui- 
morph or non-anguimorph. 

1 Some of these patterns have previously been referred to in the literature 
(e.g. Camp 1923, p. 329, fig. H). 



12 BREVIORA NO. 70 

1 wish to acknowledge the aid of the following friends, who 
read the manuscript and commented thereon : Walter Auffen- 
berg, Charles ]M. Bogert, Tilly Edinger, Gordon Edmund, Rich- 
ard van Frank, Arthur Loveridge, Samuel B. McDowell Jr., 
M. Graham Netting, Neil D. Richmond, and Ernest E. Williams. 

This paper was completed while working under a National 
Science Foundation Predoctoral Fellowship. 

LITERATUEE CITED 

Bogert, C. M. 

1943. Dentitiouiil phenomena in cobras and other elapids witli notes on 
adaiitive modififotions of fangs. Bull. Anipr. Mus. Nat. Hist., 
vol. 81, ;irt. .S, pp. 285-360. 

Camp, C. L. 

1923. Classification of the lizards. Bull. Anwv. .Mas. Xat. Hist., vol. 
48, art. 11, pp. 289-481. 

McDowKLL, S. B., Jk. and C. M. Bogekt 

19.")4. The systematic position of Laitthanotu.s and the atliaities of the 
anguinomorphan lizai'ds. Bull. Aniei-. Mus. Nat. Hist., vol. 10."), 
art. 1, pp. 1-142. 

Z.\NGERL, R. 

1944. Contributions to the osteology of the skull of the Anipliisbaeni- 
dae. Amer. Midland Nat., vol. 31, no. 2, pp. 417-454. 



BREVIORA 

Miasemei of Comparative Zoology 

Cambridge, Mass. March 29, 1957 Number 71 



TAXONOMIC NOTES ON THE NEW WORLD FORMS 

OF TROGLODYTES 

By Raymond A. Paynter, Jr. 

While preparing a list of the Troglodytidae, for the continua- 
tion of the "Check-list of Birds of the World," it became evi- 
dent that my taxonomic treatment of the family diverged in a 
number of respects from the classification employed by Hellmayr 
(1934), the current standard reference to the New World wrens. 
It seems advisable, therefore, to present, in more detail than 
possible in the Check-list, the reasons which have instigated these 
changes. This paper will be concerned with the New World 
forms of Troglodytes. 

I am greatly indebted to Dr. Dean Amadon of the American 
Museum of Natural History and to Dr. Herbert Friedmann of 
the United States National Museum for loaning me specimens 
under their care, and to Dr. Ira N. Gabrielson for very gen- 
erously allowing me to borrow critical specimens from his private 
collection. 

TROGLODYTES TROGLODYTES 

It has been nearlv 40 vears since the New World forms of 
Troglodytes troglodytes were last revised. At that time, Ober- 
holser (1919) recognized nine races, three of which he described 
in the course of his study. Four races have been designated 
subsequently. After examining nearh^ 450 specimens of the 
species from North America, I find that I cannot recognize five 
of the 13 races proposed. A synopsis of the races is presented 
below. 

One of the puzzling features exposed in this review is that 
the number of male specimens outnumbers that of females by 



2 BREVIORA No. 71 

almost tAvo to one. An unbalanced sex ratio does not seem to 
occur in the wild, at least in the European races (Armstrong, 
1955). I suspect that collectinf? bias is involved in the museum 
material, perhaps caused by the greater ease Avith Avhich the 
vociferous males mav be found. Bias created bv careless sexing 
of specimens is also a strong- possibility and suggested by the fact 
that the non-breeding specimens display a greater disparity in 
the sex ratio than birds taken during the nesting season. Were 
accurately sexed material available, sexual dimorphism in size 
might be more pronounced than now evident and some of tlie 
apparent interracial overlap in size might be diminished. 

Troglodytes troglodytes hiemalis Vieillot 

TrncilDilijIrft hicnuilis Vieillot, LS19, Xouv. Diet, d'llist. Njit., nouv. ed., 
34, p. 514: — Xova Scotia and New York; lestrieted to Nova Scotia by 
Oberholser, 1902, Auk, 19. j.. ITS. 
Troglodytes troglodylc.s aqi(ilonarif! Burleigh and Peters, 1948, Proc. 
Biol. Soc. Washington, 61. p. 116 — Tompkins, Ne\s-foundland. 
T. t. hiemalis is the lightest colored of the continental forms. 
The Newfoundland population was named aquilonaris and was 
described as differing from hiemalis in being darker and less 
rufescent dorsally, and paler and more heavily squamated ven- 
trally — characters which are notoriously variable throughout 
the species. A series of 36 specimens from Newfoundland, in- 
cluding part of the type series, has been examined. Some birds 
possess the characters ascribed to the race, but these individuals 
fall well W'ithin the range of variability of hiemalis and occur 
with the same frequency. T. t. aquilonaris is considered to be 
untenable. 

The race breeds from the southern part of the District of 
Mackenzie and southern Newfoundland south to Pennsylvania, 
Minnesota, and Alberta. It winters in the southeastern half of 
the United States. 

Troglodytes troglodytes pullus (Burleigh) 

Xannu-s hiemalis pullus Burleigh, 1935, Proc. Biol. Soc. Washington, 48, p. 61 
— Mount Mitchell (alt. 6,500 ft.), North Carolina. 
This race, which breeds in the Appalachians, is barely dis- 
tinguishable from hiemalis, but a series of 13 breeding specimens 
from Tennessee, Virginia, and North Carolina shows that 



1957 



XEW WORLD FORMS OF TROGLODYTES 



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4 BREVIORA No. 71 

dorsally it is, almost constantly, slightly darker than birds of the 
northern race collected in the same season. A shorter wing and 
smaller bill were also attributed to pullus. However, 21 adult 
males collected between late May and early August in New- 
foundland, New Brunswick, Quebec, Alberta, Maine, New Hamp- 
shire, New York, and Michigan had a mean wing length of 
49.38±:.28 mm., and 20 of the same series had a mean culmen 
length of 14.25±:.15 mm., dimensions which do not differ sig- 
nificantly from those of breeding males from the Api)alachians 
(Table 1). 

The race breeds in the mountains from Virginia to Georgia. 
It is absent from its breeding grounds during the winter and 
presumably mingles with hiemalis in the southeastern United 
States. Differentiation of the two races in the winter is uncertain. 
The post-nuptial plumage of the species is considerably darker 
than the breeding plumage and although one might assume that 
in this dress pullus remains slightly darker than hiemalis, there 
is no evidence that this is true. 

Troglodytes troglodytes pacipicus Baird 

Troglodytes hiemalis var. pacificiis Baird, 1864, Rev. Anier. Birds, 1, (18(i-t- 
1873), p. 14.5 — Simiahmoo, Puget Sound, Washington. 

The western continental race is much more richly colored 
throughout than hiemalis and pullus and somewhat more fre- 
(piently lacks dorsal barring. Ridgway (1904) stated that the 
bill of pacificus is more slender and less curved, but I cannot 
confirm this. The mean wing length of males of pacificus is 
substantially shorter than that of the males of the eastern races 
(Table 1), although the overlap in measurements is almost 
complete, rendering the character of little taxonomic value. 
Females of pacificus also have shorter wings than their eastern 
counterparts, but the difference is less pronounced than in the 
males. 

The breeding range is from southeastern Alaska and the 
southern Yukon southward through the mountains to Idaho and 
central California. In the winter the race moves to lower 
elevations and occurs casually south to Arizona and New Mexico. 



1957 NEW WORLD FORMS OF TROGLODYTES 5 

Troglodytes troglodytes iielleri (Osgood) 

Anorthura hiemalis hcUeri Osgood, 1!)01, Aiik. 18. \k ]S] — Englisli B;iy, 

Kodiak IsLand, Alaska. 

Kodiak and Afognak Islands are occupied by a nonmigratory 
race which is most similar to pad fie us, but distinguished by its 
less rich coloration, reduced abdominal vermiculations. and 
slightly larger average size. 

Troglodytes troglodytes semidiensis (Brooks) 

Xanniis IiicmaJis semidiensis Brooks, 1915, Bull. Mus. Comp. Zool., 59. p. 
400 — Chowiet Island, Semidi Islands, Alaska. 

It is with reluctance that this form is accepted. It appears to 
be distinguished from kiskcnsis only by the greater average 
length of the bill of the males (19. 66 ±.19 mm. vs. 18. 57 ±.14 mm. ; 
0.05>P>0.02). However, there are only three males in the 
sample from the Semidi Islands, and their bill measurements 
(19.0, 20.0 and 20.0 mm.) fall within the upper range of males 
of kiskensis (17.0 to 20.5 mm.). Under many conditions one 
might ascribe the observed difference to chance and with little 
hesitancy consider the two populations unworthy of separate 
designation. However, it is obvious that the birds are markedly 
different from helleri of the Kodiak region, Avhich is about 100 
miles away, and resemble the Aleutian form, from which they 
are isolated by roughly 400 miles. The extent of isolation makes 
it appear probable that the observed differences are indicative 
of two populations which are morphologically distinct. It is for 
this reason that it .seems best to accept the data at face value and 
maintain semidiensis. 

Specimens have been taken on Chowiet, Aghiyuk, and North 
Semidi Islands. 

Troglodytes troglodytes kiskensis (Oberholser) 

Kannus troglodytes kiskensis Olierholser, 1919, Proc. U.S. Nat. Mus., 55, 
p. 228 — Kiska Harbor, Kiska Island, Aleutian Islands, Alaska. 

Xa7inus troglodytes tanagensis Oberholser, 1919, Proc. U.S. Nat, Mus., 55, 
p. 230 — Tanaga Bay, Tanaga Island, Aleutian Islands, Alaska. 

Kannus troglodytes petrophilus Oberholser, 1919, Proc. U.S. Nat. Mus., 55. 
p. 232 — Unalaska, Unalaska Lsland, Aleutian Islands, Alaska. 



6 BREVIORA No. 71 

Nannus troglodytes stevensoni Obeiholser, 1930, Proc. Biol. Soc. Washington, 

43, p. 151 — Amak Island, Alaska. 
Troglodytes troglodytes seguamensis Gabrielson and Lincoln, 1951, Proc. 
Biol. Soe. Washington, 64, p. 73 — Seguam Island, Aleutian Islands, 
Alaska. 

The Avreiis of the Aleutian Islands present a number of prob- 
lems, the most difficult of which is the condition of the available 
specimens. Almost all collecting has been done during the late 
spring and the summer, when the birds are in their most worn 
plumage, or are immature. Of 110 Aleutian specimens examined, 
only 15 were taken between the months of October and April. The 
condition of wear is frequently variable ; even birds from the 
same island, collected on the same day, maj^ exhibit marked dif- 
ferences. The poor condition of the specimens seems to have 
been the main reason for the description of six races from the 
region, although only two stand up under critical examination. 

On the basis of five adult males and ten young birds, the 
population of Amak and Amagat Islands was named "steven- 
soni." It was defined as differing from "petrophilus," of the 
Fox Islands, in being more gray in both adult and juvenal 
plumages, in having fewer vermiculations below in adult plum- 
age, and in having a slightly shorter bill in the adult. I cannot 
appreciate any of the supposed color characters. The difference 
between the mean lengtji of the bills of three males from Amak 
and Amagat Islands (17.50±.14 mm.) and that of nine males 
from the Fox Islands (18.16±.14 mm.) is without question 
insignificant. (P>.10). 

In the original description, " petrophiliis" was compared with 
alascensis, of the Pribilofs, even though the describer (Ober- 
holser, 1919) admitted that his new race was nearest to "tana- 
<je)isis," a form from the Andrean Islands which he described 
in the same paper. The race " petrophilus" was then casually 
mentioned as differing from " tanacjensis" {op. cit., p. 233) in 
having a shorter bill, shorter wing, more rufescent dorsum, and 
more ochraceous ventral surface. The series available to Ober- 
holser, as well as material collected since that time, has been 
examined but not even a trend toward the characters described 
can be noted. Tlie series is, of course, distinct from the Pribilof 
j)opulation, but a sample from anywhere in the Aleutians would 
have shown the same thing. 



1057 NEW WORLD FORMS OF TROGLODYTES 7 

The most recently' named Aleutian subspecies is "segvam- 
ensis," which was restricted to Seguam, Amutka, and Yunaska 
Islands, with intermediates occun-ino- on the Islands of the 
Four Mountains. It was described, from badly-worn breedinp: 
specimens, as beinp: the palest and grayest of the Aleutian races. 
Reduced barring on the flanks was another supposed character 
and it was indirectly suggested that the race has a longer culmen 
than that of "petrophilus." 

The type series of "scguamensis" is rather lighter, on the 
whole, than the material from the eastern Aleutians. However, 
the series of "seguamensis'' is extremely Avorn and comparable 
coloration maj^ be found among equally-worn specimens from 
anywhere in the Aleutians. The tips of the ventral feathers are 
abraded, resulting in the apparent reduction of barring. As may 
be seen from Table I, no significant difference in the length of 
the bill is evident. 

As has been shoAvn, "tanagensis" is indistinguishable from 
" petrophilus." This leaves only Jtiskensis, to which all of the 
birds of the Aleutians, east of the Near Islands, are referred. 
It is a large race, with only a few of its smallest examples over- 
lapping the largest found on the mainland. It is rather richly 
colored and reminiscent of helleri, although lighter brown dor- 
sally. It most nearly resembles meligerus, Avhich is considered 
below. 

The range may be defined as extending from Amak and 
Amagat Islands, off the western side of the tip of the Alaskan 
Peninsula, west through the Aleutians to Buldir Island. There 
are no records from the Alaskan Peninsula or Unimak Island. 
This may be due to the absence of collectors, since the climate 
(presumably the limiting factor on the mainland) at the end of 
the peninsula and on Unimak is probably not markedly different 
from that slightly farther out on the Aleutians or on Amak and 
Amagat. 



"■fe^ 



Troglodytes troglodytes meligerus (Oberholser) 

Anorthura mcligera OVK-rholser, 1900, Auk, 17. p. 25 — Attn Island, Aleutian 
Islands, Alaska. 
1\ t. meligerus differs from kiskensis in its generally longer 
wing and tail, and in having more and darker vermiculations on 



8 BREVIORA No. 71 

the flanks. It is a fairly well-defined race. 

It has been recorded from the Near Islands (Attn and Agattu 
Islands), 

Troglodytes troglodytes alascensis Baird 

Troglodytes alascensis Baird, 1869, Trans. Chicago Acad. Sci., 1. p. 31o, 
pi. 30, fig. 3 — Saint George Island, Pribilof Islands, Alaska. 

The race differs from kiskensis and meligerus in being darker 
dorsally and in having lighter and fewer vermiculations on the 
flanks. Although the data are scant, it w'ould seem to have a 
somewhat shorter bill and longer wing and tail than either of 
these races. With the limited material at hand, the darker dorsal 
color appears to be the most distinctive character. 

It occurs on the Pribilofs, on Saint George, Saint Paul and 
Otter Islands. 

TROGLODYTES AEDON COMPLEX 

Many taxonomists (e.g., Chapman and Griscom, 1924; Hell- 
mayr, 1934; Sutton, 1951; Bond, 1956) have noted the close 
physical and behavorial resemblances between T. aedon of the 
United States ^nd Canada, T. hrn.nn ei colli s of the mountains of 
Mexico, and T. musculns of Central and South America and the 
Lesser Antilles. The slight differences between the taxa involve 
such characters as the amount of barring on the flanks, minor 
variations in wing-tail proportions, and differences in the general 
shade of the plumage. The groups replace one another geo- 
graphically, witii only minor discontinuities in their ranges. Be- 
cause of their allopatric, orderly distribution, they would seem 
to have arisen as isolates of a common progenitor and later 
expanded their range to form the present pattern. In spite of 
their obvious close affinities, there has been reluctance to con- 
sider them as racial groups of a common species because evidence 
of interbreeding or of bridging forms, has been lacking. Recent 
studies (Marshall, 1956) however, have shown that aedon and 
brunneicollis do interbreed and this evidence, plus previoush' 
known facts, does much toward forming a convincing argument 
for considering musculns conspecific with the other groups. 

The hrunneicollis group, consisting of montane forms which 
range from southernmost Arizona through Oaxaca, may be dis- 



1957 NEW WORLD FORMS OF TROGLODYTES 9 

tinguished from the aedon group, which occurs from southern 
Canada to northern Baja California, by its more heavily barred 
flanks and browner coloration. Ridgway (1904) chose to con- 
sider the more pronounced superciliary stripe an additional 
character, but in reality the stripe is merely accentuated by the 
darker color of the surrounding plumage. These characters 
readily differentiate the birds of southern Mexico from the aedon 
group, but the northern birds are not so distinct, for there is a 
south to north cline toward lighter color and reduced barring, 
or in other words an approach toward aedon, until in Sonora, 
for example, the brown of the ventral surface is restricted to a 
buffy chest band and the barring of the flanks becomes no heavier 
than that which is found in extreme examples within the aedon 
group. The converging of characters is in itself an indication 
that the groups are not specifically distinct, but convincing proof 
has been found by Marshall (1956) in southern Arizona, where 
both types of birds were discovered interbreeding and specimens 
were collected from a population which is intermediate between 
the already only slightly differentiated groups. Marshall also 
noted that the song of the more brownish form does not differ 
appreciably from that of the T. a. parkmanii, the house wren 
of the western United States. Thus, there can be no doubt that 
hrunneicollis should be merged with aedon. 

The problem of the relationship between aedon (now includ- 
ing 'hrunneicollis) and musculus cannot be resolved so simply and 
conclusively. T. musculus is distributed from sea level to high 
altitudes over much of South and Central America and in the 
Lesser Antilles. A moderately dark race (intermedius) with 
barred under-tail coverts and faintly marked flanks extends 
north to Tabasco and to lower elevations in southeastern Oaxaea.^ 
T. a. hrunneicollis, a richly colored, heavily vermiculated form, 
is found at high elevations in Oaxaca. No indication of inter- 
gradation or sympatry between the groups has been noted, but 
the area is ornithologically poorly known and the evidence must 
be considered inconclusive. 



1 Chapmau and Griscom (1924), on the basis of two males from Manuel 
and Rfo Pilon, included Tamaulipas within the ranjip. The wing-tail ratios ot 
these birds, which I have examined, clearly Indicate that they are wintering 
examples of pnrhmnnii. The confusion of the two forms emphasizes their sim- 
ilarity. 



10 BREVIORA No. 71 

Ridgway (1904) distinguished the Middle American forms of 
T. muscidus from T. aedon by the difference in the relative 
lengths of their wings and tails, the tail of muscidus being less 
than five-sixths (83 per cent) the length of the wing and that 
of aedon being five-sixths, or more, the length of the wing. The 
ratios have been recalculated, using more material than available 
to Ridgway, and the distinction between the two groups is still 
maintained (Table 2). But, when the races of musculus from 
South America (which undeniably are representatives of the 

TABLE 2 
Wing-tail Eatios of Males of the Northern Mainland Races of T. aedon 

Ratio 

82.1% 

83.8 

72.7 

69.8 

Middle American forms) are considered, the clear-cut difference 
between the two "species" breaks down, although the majority 
of races are separable. For example, the wing-tail ratios of 
musculus, honaire, and chUensis are roughly 84.5, 87.5 and 89.0 
per cent; the remaining races range between 69.0 (albicans) 
and 82.0 (rex) per cent. When the brumieicollis group is in- 
cluded with aedon, the overlap becomes more nearlj^ complete, 
for these are short-tailed forms with cahooni, hrunneicollis, and 
compositus, for example, having wing-tail ratios of about 75, 78, 
and 78 per cent respectively. It is evident, therefore, that the 
wing-tail ratio cannot serve to distinguish the two "species." 

The absence of ventral barring is useful in distinguishing 
about half of the musculus groups from aedon but within the 
remaining forms there is a range from the faintly barred crissum 
of musculus to the fairly heavily barred flanks and under-tail 
coverts of inquietus. 



Race 


Character 


M 


Om 


Range 


X 


aedon 


wing 


51.74 mm. 


.29 mm. 


49.0—55.0 mm. 


25 


tail 


42.48 


.30 


39.0—46.5 


26 


parTcmanii 


wing 
tail 


52.26 
43.82 


.21 
.28 


49.0—56.0 
38.0—47.0 


53 
48 


. . J. wine 
tail 


50.64 
36.83 


.28 
.25 


43.0 55.0 
31.0—42.0 


55 
53 


inquietua 


wing 


55.00 


.33 


53.0—57.5 


17 


tail 


38.41 


.41 


36.5—40.0 


17 



1957 NEW WORLD FORMS OF TROGLODYTES 11 

Other characters, such as the dorsal and ventral coloration, 
might be cited, but none is more useful in distinguishing the 
groups than those already mentioned. Taken collectively these 
distinguish each taxon from ever}^ other taxon, but no single 
feature, or assemblage of features, can be found which will fully 
separate the aedon group from the muscuUis group. 

Without a mutually exclusive character, or group of characters, 
it is difficult to justify considering musculiis and aedon distinct 
species. When this fact is coupled with what is known of the 
similarity in behavior of the two groups {vide Chapman and 
Griscom, 1924, in particular), with the evidence that the 
groups are allopatric, and with the knowledge that the generally 
more distinct hmnneicollis group has been found conspecific 
with aedon, the case for considering musculus and aedon to be 
conspecific becomes as strong as that which can be made for many 
polytypic species. Although indisputable evidence of conspecifi- 
eity is lacking, it seems far better to accept the evidence as it 
stands than to defer to conservative tradition and the philosophy 
that no change is better than change based on less than inviolable 
proof. 

Twenty-nine subspecies have been recognized. This is three 
less than the number recognized by Ilellmayr (1934). An addi- 
tional form {haldwini) was described after Ilellmayr's synopsis, 
and this is considered to be a synonym. 

T . a. haldwini was described as being darker and grayer than 
the nominate form, with its breeding range in the central north- 
ern United States and adjacent parts of southeastern Canada 
(Oberholser, 1934). Although a large series was examined, in- 
eluding many of the specimens utilized by the describer, not 
even a trend toward the ascril)ed characters could be discerned, 
it appears that the author failed to take into account the post- 
mortem color changes usually associated with wrens, as well as 
the heterogeneity^ in color found even in fresh material from a 
given locality. The race is synonymized with T. a. aedon. 

The race intermedins is now considered to range from southern 
Mexico through Costa Rica, although Nelson (1901) named 
peninsularis from the tip of the Yucatan Peninsula, and Chap- 
man and Griscom (1924) named oreopoliis from the highlands of 
Nicaragua. 



12 BREVIORA No. 71 

Paynter (1955) accepted pcninsularis, with reluctance, and 
stated that it could be distinguished from intermedius only by 
its slightly longer bill. An examination of 54 males from 
throughout the range of intermedius shows them to have a mean 
bill length of 16.16±.08 mm., with a range from 15.0 to 18.0 mm. 
Only one specimen, a bird from Costa Rica which approaches 
inquietus, has a bill as long as 18.00 mm.; the remainder fall 
between 15.0 and 17.0 mm. Six males of 'peninsularis from the 
type locality at Progreso and from nearby Santa Clara, Yucatan, 
have a mean culmen length of 16.83±.02 mm. (not ±.20 mm. as 
published by Paynter, 1955), and range from 16.5 to 17.0 mm. 
While the difference between the two samples is "statistically 
significant," the overlap is much too great to warrant recognition 
of two races. 

Hellmayr (1934) doubted the validity of oreopolus, described 
by Chapman and Griscom (1924) as being a montane race 
slightly darker than intermedius. I cannot recognize the form 
and consider it to be synonymous with intermedius. 

T. a. clarus, of the Guianas, Trinidad, northern Brazil, and 
adjacent parts of Venezuela, Colombia, and Peru, was considered 
by Chapman and Griscom (1924) to be indistinguishable from 
albicans of southwestern Colombia and western Ecuador. Never- 
theless, Hellmayr (1934) chose to recognize the two forms, even 
though he admitted that individual variation frequently bridged 
the slight gap he believed to exist. Approximately 50 specimens 
from the range of each race have been examined and it is con- 
eluded with Chapman and Griscom (1924) that clarus cannot 
be separated from albicans. 

TROGLODYTES SOLSTITIALIS COMPLEX 

At high altitudes from southern Mexico through Central and 
South America, there is a series of isolated populations of short- 
tailed, short-billed wrens of rich, brown coloration, with broad, 
tawny superciliaries, taAvny auriculars, and pronounced, dark 
post-ocular stripes. These have been treated as three species. The 
first is T. rufociliatus, a form with four isolated races ranging 
from Chiapas, Mexico, through El Sah^aclor, which is character- 
ized by heavy barring on the abdomen and under-tail coverts, 
concealed (or obsolete) small, white spots on upper-tail coverts, 



1957 NEW WORLD FORMS OF TROGLODYTES 13 

and a few white spots on the wing coverts. The second species 
is T. solsfitialis (a group of seven or eight subspecies, most of 
which are isolated from one another) which is distributed from 
Costa Rica through the Andes to Venezuela and Argentina. Tt 
may be distinguished from T. rufociliatns l)y its less saturated 
color, lack (or nearly so) of abdominal vermiculation, and 
absence of spots on both the upper tail and wing coverts. The 
third species is T. monticola a monotj''pic form isolated near the 
top of the Sierra Nevada de Santa Marta, Colombia, which is 
much larger than the other two forms. It resembles T. solstitialis 
in that it is not deeply colored and has no spotting on the rump 
or wings, and is similar to T. rufociliatus in that it is heavily 
barred below. 

Hellmayr (1934) saw no specimeiLs of rufociliatus, but thought 
that it was intermediate betAveen hrimneicollis and solstitialis, 
and that eventually it might seem advisable to unite the three 
forms under one species. T. rufociliatus and solstitialis are 
morphologically quite distinct from T. a. irunneicollis, with only 
their rich coloration a point in common. In El Salvador T. a. 
intermedius and rufociliatus occur sympatrically (Dickey and 
van Rossem, 1938) and, as it has been shown, intermedius is 
conspecific with hrunneicollis. Therefore, it is impossible to con- 
sider rufociliatus to be a member of the aedon-hrunneicolUs- 
musculiis complex, even if one ignores their morphological dis- 
tinctiveness. Although hrunneicollis is not the link between 
rufociliatus and solstitialis, the two groups are very similar, with 
only the presence or absence of abdominal barring a conspicuous 
difference. Even this difference is bridged, somewhat, by the 
nominate form of solstitialis, since mature specimens occasionally 
are distinctly, and quite extensively, barred on the flanks. The 
spotting on the wing and upper-tail coverts of rufociliatus is 
not a good distinguishing character either, because within a given 
population there are some individuals lacking these markings. 

The two groups would seem to have arisen from a progenial 
population which at some time became divided into two isolated 
populations and thereupon became somewhat differentiated. 
Further division and isolation within the groups then gave rise 
to the various subspecies now recognized. It may be postulated 
that the primary division occurred during the Cenozoic when 



14 BREVIORA No. 71 

Middle America was divided by various seaways. One population 
(solstitialis) maj^ have been isolated in South America and 
southern Central America while the other population (rufocilia- 
tus) was isolated in northern Central America and southern 
Mexico. Even if the groups did not have their origin precisely 
in this manner, or at this period, the almost certain fact remains 
that they arose from a common ancestral stock which became 
split into two geographically isolated units. 

Since the two groups are so slightly differentiated, there is 
little doubt that their relationship is best expressed by treating 
them as conspecific forms, uniting them under the older name 
of solstitialis. 

T. monticola, the large, heavily barred form of Santa Marta, 
was believed by Hellmayr (1934) to be reminiscent of hrun- 
neicollis. He thought it a specialized offshoot of solstitialis but a 
possible link between solstitialis and hnmneicollis. The resem- 
blance between these two forms is superficial. T. monticola has 
the short tail, short bill, and facial pattern of solstitialis. Its 
large size and barred underparts are the only features which are 
similar to drunneicollis, and even the barred underparts are 
shared by some races of solstitialis. While I do not agree that 
monticola is a link between l>runneicollis and solstitialis, I do 
think that Hellmaja- (oj). cit.) was correct in believing it to be 
an offshoot of solstitialis. The only character which distinguishes 
it from all of the forms of solstitialis is its large size. Being con- 
fined to a small area on the top of a mountain, it is analogous to 
an insular population, in^ which situation a race often is larger 
than its congeners. It would seem best to consider monticola as 
merely another race of solstitialis. 

LITERATURE CITED 

Armstrong, Edward A. 

1955. The Wren. Collins, London, viii + 312 pp. 

Bond, James 

1956. Check-List of Birds of the West Indies. Acad. Nat. Sci. Phila- 
delphia, vi + 214 pp. 

Chapman, Frank M. and Ludlow Griscom 

1924. The House Wrens of the Genus Troglodytes. Bull. Amer. Mus. 
Nat. Hist., 50: 279-304. 



1957 NEW WORLD FORMS OF TROGLODYTES 15 

DiCKFA-, Donald K. and A. J. van Rossem 

1938. The Birds of El Salvador. Field Mus. Nat. Hist., Zool. Ser., 23. 
609 pp. 

Hkllmavk, Charles E. 

1934. Catalogue of Birds of the Americas and the Adjacent Islands. 
Field Mus. Nat. Hist., Zool. Ser., 13. pt. 7, vi + .531 pp. 

Marshall, Joe T. 

19.")6. Summer Birds of the Rincon Mountains, Saguaro National 
Monument, Arizona. Condor, 58: 81-97. 

Nelson, E. W. 

1901. Descriptions of a New Genus and Eleven New Species and 
Subspecies of Birds from Mexico. Proc. Biol. Soc. Washington, 
14: 169-175. 

Oberholser, Harry C. 

1919. Notes on the Wrens of the Genus Nannus Billberg. Proc. U.S. 

Nat. Mus., 55: 223-236. 
193-1. A Revision of North American House Wrens. Ohio Jour. Sci., 

34: 86-96. 

Paynter, Raymond A., Jr. 

1955. The Ornithogeography of the Yucatan Peninsula. Bull. Peabody 
Mus. Nat. Hist., Yale Univer., 9. 347 pp. 

RiDGWAY, Robert 

1904. The Birds of North and Middle America. Bull. U.S. Nat. Mus., 
50. pt. 3, XX 4- 801 pp. 

Sutton, George Miksch 

1951. Mexican Birds. First Impressions. U. of Oklahoma Press, Nor- 
man, XV -|- 282 pp. 



^^Ui) 



BREVIORA 



Meseeei of Cooipsirative Zoology 



Cambridge, Mass. March 29, 1957 Number 72 

IS THE ANT GENUS TETRAMORIUM NATIVE IN 
NORTH AMERICA? 

By W. L. Brown, Jr. 

For some years, a controversy has continued concerning the 
distribution in North America of the ant genus Tetramorium 
Mayr. On opposite sides in this argument ha.ve been Dr. M. R. 
Smith (1943), who believes that all five of the Tetramorium 
species reported from North America have been introduced by 
man from overseas, and Dr. W. S. Creighton (1950), who thinks 
that the evidence points to prehistoric endemicity in the continent 
for at least two of the species mentioned ; T. cacspitnm (Linnaeus) 
and T. riigiventris M. R. Smith. Apparently there is no serious 
disagreement about the origin abroad of 2\ guineense (Fabri- 
cius), T. simillimum (F. Smith) and T. pacificum Mayr. The 
first two of these are very likely African in origin, since they 
occur in wild parts of Africa and have their closest relatives 
among the species of that continent. T. pacificum is apparently 
from the Indo-Australian area, although its exact source has 
never been seriously tracked down. Certainly, it is Old World 
in origin. 

The two species caespitiim and rugiventris are therefore the 
central elements in the discussion, and it seems appropriate at 
this time to review the important facts in connection with these 
two forms and to add whatever significant observations are now 
available. 

Tetramorium caespitum 

This species is widespread in the Palaearctic region and in 
Africa, where it is extremely variable, as attested by an almost 
endless list of infraspecific variants. In terms of modern tax- 



2 BREVIORA NO. 72 

onomic practice, many of these entities would be considered 
good sibling species, as indeed some of them already have been 
treated by various European authors. Many others are mere 
synonyms of caespitum — individual or nest variants that do 
not represent natural, self-maintaining populations. No one 
has challenged the placement of the North American populations 
with the more nearly "typical" caespitum variants. 

Creighton's assertion that caespitum is a native Nearctic ant 
rests on two principal pieces of evidence : first, the species was 
known at a very early date, i.e., 1895, from states as remote as 
Tennessee and Nebraska ; second, there is present in North 
America, at least in the eastern states, a workerless parasite of 
caespitum, the aberrant species Anergates atraiulus (Schenek), 
the transport and establishment of Avhich would seem to present 
special difficulties. 

The difficulty with the first piece of evidence is that the 
exact situation of the collections made in Tennessee, Nebraska 
and elsewhere in the "interior" of North America was never 
specified. All of these collections may have been, and probably 
were, made in or near "culture areas," that is, regions strongly 
disturbed by the presence of man. The experience of several 
practiced myrmecological field workers, including that of Dr. 
Creighton (personal communication) and myself, indicates that 
so far as known, caespitum in North America is known only from 
rather heavily disturbed localities, such as cities, towns, road- 
sides, farmyards, picnic areas and the like. I have been able 
to gather no records at all to show the existence of the species 
in places remote from the works of man in North America. 

This situation contrasts with that holding in the Old World, 
at least so far as my own personal experience goes, and judging 
also from what I have been able to glean from various publica- 
tions dealing with the species, and from personal communications 
with European myrmecologists. To sum up this information, 
it can be stated that T. caespitum in Europe and (although given 
various infraspecific names) in China is often abundant in and 
around human habitations, just as in North America. However, 
it is also to be found, often in abundance, in localities that show 
little or no trace of human disturbance, and that are far from 
the nearest humanly-occupied places. The contrast is very 
marked if one collects, as I have, at intermediate altitudes in 



1957 TETRAMORIUM IN NORTH AMERICA 3 

West China, in pine-oaJc forest, and then compares the collec- 
tions of Tetramorium obtained with the results of a deliberate 
search for Tetramorimn in similar vegetational zones in Pennsvl- 
vania, New Jersey or Massachusetts. In West China, T. caespi- 
ium tends to occur uniformly throughout the pine-oak forest, 
regardless of roads, villages, etc., which are very sparsely dis- 
tributed in the areas I am recalling. In ecologically equivalent 
areas in the eastern United States, I have found the same species 
established only on or near the sites of more or less actively 
maintained human works. Clearly, the density of nests and 
individuals seen in North America is in large part proportional 
to the degree of urbanization of the area occupied, although the 
real extremes of urbanization, where almost all space is covered 
by concrete or asphalt, are certainly not favorable locales for 
colonization by this or any other ant species that lives largely in 
the open. In my opinion, the local, detailed distribution of 
T. caespitum in North America is that expected of an historically 
introduced, not a native ant. 

The second point of evidence, that concerning the presence of 
tlie workerless, and therefore obligatory parasite, Anergates atra- 
iulus, certainly seems on the face of it a real sign of long 
occupancy of North America by both host and parasite. Creigh- 
ton emphasizes the difficulties facing trial colonists of the 
parasite species : first, the species seems to be relatively rare in 
Europe ; second, it is unlikely that the parasite female could 
survive a long trip; third, a female arriving in North America 
would be hard put to find a suitable nest of the host species to 
enter. To take these difficulties one at a time, we should first 
recognize that, while Aner gates is not the commonest of ants in 
collections, it is nevertheless likely to be locally very common 
in restricted localities. Even in Europe, host populations are 
normally concentrated in gardens and waste places within towns 
and cities, including seaports, and these are accordingly \evy 
likely places for Anergates to occur unnoticed by primarily 
country-searching myrmecologists. In the United States, Aner- 
gates is known chiefly from East Coast localities in urban areas 
near the sea — exactly the kind of place from w'hicli colonists 
might be expected to be exported most easily. It should be added 
that Anergates females may be produced in very large numbers 



4 BREVIORA NO. 72 

from a single nest, so that a given locality may be heavily 
saturated with them during the period of nuptial flights. 

The difficulty of transport of live Aner gates propagules is 
real, but far from insurmountable. Females can be carried 
either as individuals carrying the necessary sperm, or as estab- 
lished inquilines in a Tetramorium colony. There is no reason 
why such a voyage might not be successfully made by a parasite 
queen, especially when one considers the evidence of Lindroth 
(ms., personal communication) for transport of faunal frag- 
ments to North America in ballast originating in Europe. Fur- 
thermore, there is no reason to believe that Tetramorium nests, 
with or without Anergates, cannot flourish on shipboard for at 
least the normal span of these species as colonies, a span which 
seems to be sufficient even for a long voyage under sail. 

The third objection is the least difficult one, for there ap- 
parently has been no shortage of suitable host nests at close 
proximity to the waterfront in at least some of the major western 
Atlantic ports, perhaps as far back as colonial times. If a colony 
of the host parasitized by Anergates arrived in ballast or other- 
wise stowed-away, it had only to release its flight of fertile 
female imagines on the new shore to create a high probability 
of successful establishment. 

A similar series of events may have led to the establishment 
of the workerless parasite Xenometra moriilicornis Emery in the 
West Indies, together with or following the establishment of its 
host, Cardiocondyla cmeryi Forel. A Xenometra of the same or 
a very closely allied species lives with C. clegans Emery in Italy ; 
Menozzi (1919) thought this was the male of elegans, but 
specimens from his collection indicate instead its affinity with 
X. monilicontis. Cardiocondyla is a primarily littoral and riparian 
genus from the warmer parts of the Old World ; records of 
several species from the New World seem to indicate rather 
clearly that it has been introduced by man on many separate 
occasions (M. R. Smith, 1944). 

To conclude the discussion of the bearing of parasites on the 
distribution of Tetramorium, 1 think we may safely consider that 
introduction of an obligatory parasite, while less probable than 
the establishment of the host, is nonetheless entirely possible if 
the opportunities exist for a long enough time, and if a dense 
host population is available to the immigrant parasite. 



1957 TETKAMOHir.AI IX NORTH AMERICA 5 

TeTKAMORIUM RUGIYENTRIS 

The type series and onl.y recorded sample oC T. rugiventris 
was obtained from an upland ponderosa jiine stand about ten 
miles south of Prescott, Arizona, and about one mile off the 
liijihAvay. In empliasiziny his difficulty in accepting Smith's 
hypothesis, namely, that the ant was introduced with camel 
food or stores at the time Avhen camels were imported from 
North Africa during the last century, Creighton wrote : ' ' Ento- 
mologists frequently strain at gnats but it is seldom that they 
are asked to swallow a camel." 

I have checked with care a syntype of T. rugiventris kindly 
sent by Dr. Smith. As a result of this examination, I can agree 
with Dr. Creighton that the ant in cpiestion is almost unques- 
tionably endemic to the locality where it Avas found. However, 
I cannot agree with either Smith or Creighton that the species 
rKgivcufris belongs to the genus Tctrcnnorium as it is now con- 
stituted. Instead, the type I have examined seems to me to be a 
cleareut, if somewhat aberrant, member of the genus Myrmica, 
closely allied to M. striolagaster Cole. M. striolagaster is re- 
corded from several localities in Arizona and New Mexico, and 
I have specimens collected by E. 0. Wilson at or near the type 
localit}- of rugiventris, in the vicinity of Prescott. Although the 
two species are separated hy the extent and strength of the 
gastric sculpturing and by other characters as well, it seems 
clear that they are congeneric, and also that they run rather 
close to the M. pnnctiventris group of Myrmica. 

The rugiventris type actually possesses minute barbulation on 
the posterior tibial spurs, as can be seen at magnifications of 
90 X and better, so that in this character, the species would 
key to Myrmica, rather than Tetramorium, in the standard keys 
to tribes and genera of Formicidae. Actually, however, this 
spur barbulation is not worth much as a tribal character within 
the IMyrmicinae, despite the faith that key-makers have placed 
in it. The false distinction between Hylomyrma Forel (tribe 
Myrmicini) and Lundella (tribe Tetramoriini) appears to have 
been based partly on this character (Brown, 1953), and the 
same may hold true of the supposed difference between Crato- 
myrmex Santschi, purportedly a member of tribe Myrmicini, and 
Messor Forel, of tribe Pheidolini. 



6 BREVIOBA NO. 72 

Dr. Smith is fortified in his opinion that rugivcntris belongs 
to Tetramorium by the presence in his species of raised carini- 
form lateral wings of the clj^peus that border the antennal 
fossae in front, as in Tetramorium; but this character is shown 
with varying degrees of clearness in other, undoubted Myrmka 
species, both in North America and in southern Asia, so that it 
cannot be used as a point of separation between the two genera. 
The question comes to mind, of course, as to whether Myrmica 
and Tetramorium. really are separate genera after all, and this 
is precisely the kind of question that most needs asking in ant 
taxonomy these days. In deciding this particular question, fur- 
ther study must l)e made of the males, since tetramoriine males 
(with the exception of a couple of African forms that require 
closer study) have several funicular segments fused in such a 
way as to reduce the numl)er of antennal segments to ten in 
this sex. 

Unfortunately, the males of rugiventris remain unknown at 
present, so that it is not known whether they meet the strong 
criteria of this caste. From the habitus and lesser details of the 
worker, however, I consider the relationship with Myrmica is 
close enough to call for a new combination: Myrmica rugiventris. 
Whether or not this combination finally i)roves to be the valid 
one, it at least helps to establish strong doubts as to the pre- 
Columbian existence of Tetramorium in North America. 

In fact, were it not for Tetramorium lucayanum Forel and 
the Xiphomyrmex spinosus complex, the New World could h( 
considered free of endemic members of tribe Tetramoriini. 
With the synonymy of Lundella under Hylomyrma (Brown, 
1953), the New World lost its one endemic tetramoriine genus. 
T. lucayanum presents no special difficulty, because its distribu- 
tion (Bahamas, Puerto Rico, etc.) is highly suggestive of intro- 
duced status. In its morphological characteristics, lucayannm 
seems closest to an African group of species, but the species 
itself has not l^een identified with any particular continental 
African population. However, our knowledge of African Tet- 
ramorium is in a very imperfect state, and it seems to me likely 
that lucayanum must have come from the Dark Continent, even 
though it may be rare there. A parallel case involving Strumi- 
genys rogeri Emery has turned out to follow exactly this pattern 
(BroAvn, 1954). 



1957 TETRAMOKIUM IN NORTH AMERICA 7 

The Xiphomyrmex sinnosus complex (which may represent a 
single variable species) is the one example that cannot be ex- 
l^lained away, and it is all the more remarkable, considering its 
isolated position in southwestern United States and through much 
of Mexico, far away from the remainder of the generic range, 
which is entirel}' Old World tropical and warm-temperate. Pre- 
limina.ry examination reveals no reason to consider this complex 
as other than bona fide Xiphomyrmex, and its distribution is 
almost certainly that of a long-established endemic group of 
populations, surely pre-Columbian in North America. Xipho- 
inyrmex is separated from Tetramorium by a very minor charac- 
ter, 11 antennal segments in place of the 12 of Tetramorium. 
Future revisers could well fail to be impressed by the soundness 
of the generic split based on this difference, so that we may 
eventually see a systematic technicality bring back Tetramorium 
as a native American genus. 

SUMMARY 

Of the five species of Tetramorium so far reported as occurring 
in North America, only two are under dispute as possibly having 
existed on this continent prior to the advent of European colon- 
ists ; these are the species heretofore known as T. caespitum and 
T. rugiventris. Evidence is presented to show that T. caespitum 
almost certainly was introduced by man from Europe, this evi- 
dence consisting primarily of the demonstration that T. caespi- 
tum in North America, unlike the Eurasian populations, is 
distributed exclusively in the manner of a man-accompanying 
"tramp" species. The species rugiventris, on the other hand, 
is removed from Tetramorium, where it does not fit well, and is 
transferred to Myrmica. The number of Tetramorium species 
occurring in North America is thus reduced to four, all of them 
likel}' introductions from the Old "World w^ithin historical times. 
The only member of tribe Tetramoriini that can safely be con- 
sidered as endemic to the New World at the present time is the 
Xiphomyrmex sj^inosus complex, widespread in southwestern 
U. S. and Mexico. 



8 15IJEVIOKA NO. 72 

KEFEKENCES CITED 

r>iio\v.\, \V. L., Jk. 

19.").'}. C'haraetois aiul aynoiiymics among the genera of ants. Part II. 
Brcvioia, Mus. Comp. Zool., 18: 1-8. 

1!)'>4. The ant genus Strumigcnys Fred. Smith in the Ethiopian 
and Malagasy regions. Bull. Mus. Comp. Zool., 112: 1-34, cf. 
pp. 4-7. 

CUEIGIITON, W. S. 

1950. The ants of North America. Bull. Mus. Comp. Zool., 104: cf, 
pp. 241-245, 286-294. 

MiNCZZi (Mexozzi), C. 

1919 (1918). Primo contribute alhi couoscenza della fauna mirmecologica 

del Modenese. Atti Soc. Nat. Mat. Modena, (5) 4: 81-88, cf. 

p]). 83-84. 

S.\iiTii, ^r. E. 

1943. Ants of the genus Tetramorium in the United States with the 
description of a new si)eeies. Proc. Ent. Soc. Washington, 45: 1-5. 

1944. Ants of the genus Cardiocondyla Emery in the United States. 
Proc. Ent. Soc. Washington, 46: 30-41. 



BREVIORA 

Mmseiuiini of Comparsitive Zoology 

Cambridge, Mass. Mak( h 29, 1957 Number 78 



ADDITIONS TO THE MAMMALIAN FAUNA OF PERU 
AND NOTES ON SOME OTHER PERUVIAN 

MAMMALS 

By Oliver P. Pearson 

Museum of Vortoljiate Zoology, Berkeley, California 

All earlier report discussed the mammals of the highlands, or 
altiplaiio, of southern Peru (Pearson, 1951). Subsequent col- 
lecting has revealed important additions to the fauna of this 
area as well as considerable extensions of the range of certain 
species. 1 am indebted to Dr. Carl P). Koford, who collected 
several of the specimens mentioned, for permission to report 
upon them and to draw upon information in his field notes. All 
specimens are in the Museum of Vertebrate Zoology, Berkeley. 

Hesperomys sorella (Thomas). This species was not listed in 
the previous report on mammals of the altiplano. Eleven speci- 
mens have now been taken in the Department of Puno in bunch- 
grass terrain at 3 mi. NE Arapa, 12,600 ft., 5 mi. S Asillo. 
13,000 ft. and at Hacienda Calacala, 13,000 ft., which is 7 mi. 
SW Putina. They are longer-tailed, longer-eared, and tawnier 
than H. lepidus ducilla, which also lives in this region, and have 
shorter, more slanting zygomatic plates. The range of measure- 
ments (in mm.) is: total length, 128-147; tail, 60-71; hind foot, 
17-19 ; ear from notch, 18-19 ; and greatest length of skull, 21.5- 
23.2. They do not match the type of sorella perfectly but seem 
to be more closely related to that form than to coJlosus, carillus, 
or frida. 

Since publishing the earlier report in which I used the name 
Hesperomys ducilla for the short-tailed species in southern Peru, 
I have examined the types of H. lepidus (Thomas) and H. ducilla 
(Thomas) and find that despite great difference in age of the in- 



2 BREVIORA NO. 73 

(li\i(liials iliey are (luite .siiiiilai- and can easily l)e included in 
the same species, characterized by short tails and tall zygomatic 
plates. Saiiliorn (1950^ also considered (liirtlJa to lie a subspecies 
of Jepidus. 

Klifjmodoiil l(( iJiKriilus piicruhis (Philippi). Four specimens 
of this desert mouse have l)een reported from Peru (Pearson, 
1951), all from Santa Rosa de -luli, Department of Puno. and 
were listed as E. p. hirtipcs, the type locality of ^vhich is Lake 
Poopo in Bolivia. Our recent collections included 11 specimens 
taken in Peru between 13,000 and 15,300 ft. in the Departments 
of Moquegua, Puno, and Tacna. These specimens have now been 
compared Avith topotypes, in the Chicago Natural History 
Museum, of E. p. puendus from northern Chile and are iudis- 
tinguishal)le. E. p. iarnpacensis Mann has also been described 
from northern Chile, but the new material reveals that the shape 
of the zygomatic ])late and the bicolored nature of the tail, 
features on which farapacensis was based primarily, are un- 
reliable. The Peruvian Elifpiindoiil ia therefore should l)e E. p. 
pKcridus. 

FhyUotis [Galoionnjs) garlcppi Thomas. Two specimens of this 
rare mouse were taken in southern Peru, one at Pichupichuni. 
12,600 ft., 5 mi. NW Iluacullani, and another at Pampa de Anco- 
marca, 13,700 ft., 76 mi. S Have, both in the Department of Puno. 
These are the first records of this species from Peru. Since no 
field measurements have been i)ublished and the type specimen 
has a smashed skull, we record measurements (in mm.) of our 
two specimens and photographs of one of the skulls (Fig. 1). 



Total length 

Tail 

Hind foot 

Ear 

Skull greatest length 

Zygomatic breadth 

Width of braincase 

jMaxillary tootlnow 

Compared with the type and another Bolivian specimen in the 
British Museum of Natural History, and one in the Chicago 
Natural History Museum, the Peruvian specimens are smaller, 



MVZ 115903 


MVZ 11590 


129 


132 


32 


30 


23 


24 


19 


19 


26.7 


27.0 


lo.O 


15.0 


12.8 


13.0 


5.3 


5.2 



li);")? 



PERUVIAN MAMMALS 



3 



have iimcli sliorlcf I'lir, color iniicli less l)ri<ilit, and slioiici' cars, 
riit'oi'tuiialcly both I'cniviaii specimens arc ^ouniicr, although 
one w;is oh] enouji'h lo l)e pi-euiiant (April 10) and llie ollici- 
estrons ( F(>l)ruary 1). Tlie Peruvian s])ecinieiis auree with the 




Fis. 1 (;iliove;. I'liylloti.s (Galcnoviys) (jarleppi ; I'.o x. 
Fig. L' (below ). Fr'li'< jdcohlfd ; ().(> x. 

tyj)e in these <lia^nostic i'eatni'es: incisors slender and nia.i-kedly 
])r()odont. antci'ior hoiv'er of zyg'oniatie plate distinctly convex 
and extendin^.^' almost to the top of the rostrum, and dorsal pro- 
file of ihe skull strons'lv convex. 



4 RREVTORA XO. 7.'^ 

One of the specimens Avas caught in front of a l)ui'i'ow of 
Ctenomys pernayius on a hea.vily "razed pampa of dwarf grass 
and prostrate forbs. The other was taken near a stone wall on 
a similar, grazed pampa on which were also growing scattered 
thornbnshes about one foot tall. Cienomys (yphnus was liviiig 
about TOO yards away. Other mammalian associates were (/(ilea 
niusteloidcs, Alxodon jelskii, PhyUoih darwini, Ph. siihlimis, and 
Hesperomys Jepidus dnciUa. The last two of these are sur- 
prisingly similar in appearance to (/arlcppi. Ph. garlcppi is 
larger than dmiUa (total length of garlcppi more than 120 mm., 
foot more than 20 mm.) and smaller than most subliniis with 
tail usually shorter (less than 46 nun.) and not bicolor, soles of 
hind feet more hairy than in .'>i(bli'}iiis. Using Pearson's (1951) 
key to the rodents of the altiplano west of Lake Titicaca, Galeno- 
mys keys out as either Phyllotis osilae, Ph. darivini, or Akodon 
aruocinis. All of these have much longer tails than does garlcppi. 

Punomys lew minus Osgood. The range of this rare mouse has 
been extended considerably by its capture 55 mi. ENE Arequipa, 
15,300 ft., Department of Are(|uipa, and 12 mi. NE Tarata. 
14,600 ft., Department of Tacna. These new specimens agree 
well with those from Caccachara (Pearson, 1951) and with the 
lype. 

In addition, nine specimens were taken 8 mi. 8SW Limbani, 
15,000 ft., Department of Puno. This is 130 miles north of the 
type locality and separated from it by the grassland of the 
Titicaca basin. A circuitous strip of more favorable habitat may 
connect the two regions by way of the mountains to the west 
and northwest, but no specimens have been taken there. The 
specimens from near Limbani, like so many forms from this 
more humid region of the Andes, are distinctly and consistently 
darker than those from farther south or west. Compared wiih 
the type and with the sj^ecimens mentioned in the preceding 
paragraph, the Limbani Punomys are greyer and darker on the 
back, the feet and hands duskier al)ove, ears darker, tail less 
distinctly bicolor, and belly considerably darker grey with a 
distinct bult'y wash. 

The Limbani specimens, as well as those from near Are(iuipa 
and Tarata, were captured in barren, broken I'ock areas and, as 
at Caccaciuira, were near fleshy-leaved, luuigent Soiccio plants 
or piles of Scnecio cuttings. 



l!);")? PERU VI AX MAMMALS 5 

Covin tscJnuh'i osgoodi Sanborn. Two s])CM'i('s of guinea piprs 
live on the altiplano of Peru and at some localities ])robably 
oeciir together. Cavia (Galea) mustcloicles is diurnal (Pearson, 
1951) and prefers i-ather open habitat with jiood visibility. At 
Ilaeienda Calaeala the mueh darker colored Cavia (Cavia) 
iscliudii lived in thick grass where it made distinct nuiways and 
was crepuscular. 8teel traps set in the runways failed to catch 
any during the day, but numerous individuals were seen and 
collected in the evening after sunset and in the early morning. 

Lagidinui poKanuDt Eleven. In Peru mountain viscachas 
have been considered to live only at high altitude. It was a 
surjirise, therefore, to find a small population living in the fog 
belt, or loma.-;, at only 2200 ft. at Nana, Department of Lima. 
The hilltops at Nana support scattered clumps of fog-nourished 
TiUandsia (Bromeliaceae). On the lower slopes there is no vege- 
tation, but thei-e are irrigated fields on the floor of the valle3^ 
Among rocks on one of the hilltops were many viscacha scats, 
which probably accumidate for years in the absence of rain, a few 
viscacha bones, and at least one living viscacha. This individual 
was seen at a distance of oidy 6 feet and appeared to be similar 
to the mountain viscacha,-; of southern Peru but with more than 
average buffy color. A maxillary toothrow picked up nearby 
agrees well with specimens of Lagidiuni pcruanum. 

This population living more than 6000 ft. lower than any other 
viscacha population known to me in Peru is isolated by several 
thousand feet of brushy and weedy terrain unsuitable for 
viscachas. 

Mustehi fytnaia Lichtenstein. Weasels have long been thought 
to live on the altiplano of Peru but have been inadequately rep- 
i-esented by specimens. A mounted skin is now available, taken 
in the spring of 1951 at Hacienda CaJ.acala, 13,000 ft., 7 mi. ^S^Y 
Putina, Department of Pnno. 

Orison (Gri^onella) cuja (Molina). The presence of this 
nuistelid on the altiplano of Peru has heretofore been inade- 
(juately documented. We now have the skin and skull of an adult 
male from Ilaeienda Pairumani, 13.000 ft., 24 mi. 8 Have, De- 
partment of Puno. JNIeasurements (in mm.) are: total length, 
570; tail, 145; foot, 70; weight, 1700 grams. This specimen was 
shot at 4 p.m. with its stomach crammed with at least three 
mice and a lizard, indicating diurnal feeding. 



6 BRFA'IORA XO. 78 

Felis jacobita Cornalia. The complex taxoiiomie history of 
this wildcat (see Osgood, 1943) has been based on a half-dozen 
skins, many of them without adecinate locality information, and 
on drawings of a single skull (Pliilii)pi, 1873). Xo previous 
specimens are known from Peru. AVe now have a skin and skull 
of a male, trapped March 30, 1952, among rock outcrops at 
15,500 ft., 57 mi. ENE Arequipa, Department of Arequipa. This 
is a barren region of rocks and bare ground with scattered c]um])s 
of buncligrass [Fcsiuca orihophxiUti) and small tola bushes. 
Other steel ti*aps nearby caught a. fox [Diisicifov ciilpdCHs) and 
a mountain caracara (PlialcohooiKs uicgaJopterus) . A mountain 
lion passed close by several times. Probable prey items in addi- 
tion to small rodents and small birds were mountain viscachas 
{Lagidiuni), tinamous (Tinamotis), and seed snipe {Attagis. 
Thinocorus) . Vicunas, the young of which might be killed by 
jncohito, were abundant. 

The skin matches well the vai-ious published descriptions, but 
several features of the skull, some of them previously considered 
to be diagnostic, do not matcii the illustrations in Philippi's 
rei)ort. The audital bullae of our specimen are not bisected by 
a deep sulcus; in fact, scarcely a trace of the sulcus shows. Com- 
pared to four Felis pajeros from southern Peru and northern 
Chile, the nasals of our jacobita are only slightly larger and the 
incisors not more jn-oodont. The skidl of our jacobita is slightly 
flatter and more i-obust than in pajvros, and has lai-ger teeth, 
but skulls of the two s])ecies are not as different in ai)pearance 
as are those shown in Philipi)i's illustrations. A possible diag- 
nostic cliaracter is the orientation of the upper premolars, in 
pajeros these two teeth on each side, when vie\\t'(l from l)elow. 
lie in a straight line. In jacobita the anterior premolar toes in 
abruptly in front. Since no other skulls seem to bi' available and 
since our specimen differs considerably from the one seen by 
Philippi, two photographs are presented in Figure 2. Measui-c- 
ments (in mm.) are: total length, 990; tail, 413; foot. 133; ear, 
(i3 ; Wright, 4.0 kg. ; condylobasal length, 9(i ; greatest length of 
skull, 100; zygomatic lu-eadth, (i!>.5 : greatest length of upi)er 
carnassial, 14.0. 



lO;")? I'KRirVIAX MAMMAliS 7 

'I'lu' ]»t'la;:t' is Hut'tici- and jLii-cyci- than that of /•'. pujcros and 
is without a spinal crest of loiij^- hairs; tlic ears are rounded 
without tufts, and the tail is non-taperino-, much louf>er, an<i 
more eonspicuously ringed than in pajeros. 

In southern i'ei-u Fclis jacohifa seems to be mueh less ahuiul- 
ant than F. pajeros and probably prefers higher elevations. 

BIBLIOGRAPHY 

( )S(H>OI>. \V. il. 

1943. The iiwmminls of Cliilc. Fiold Mus. Nat. Hist., Zool. Ser., 30: 
1-268. 

I'EARSOX, U. P. 

li>.il. ^[;llnln;lls in tin- liigliliiTxls of southern Pern. Bull. Mus. Comp. 
Zool., 106: 117-174. 

rmi.ippi, R. A. 

1S7.S. Ueber Felis Ciuina Molina and iiber die iScliJidelbildung bei 
Felis Pa.ieios und Felis Oolocolo. Areh. Xaturg., 39 (1): 8-15, 
pis. 2-3. 

Sanhokx. 0. C. 

1 !».■)(•. Small rodents from Peru and Bolivia. Pul). Mus. Hist. Nat. 
"Javier Prado," Ser. A. Zool., No. 5: 1-16. 



BREVIORA 

MeseiJim of Comparative Zoology 



Cambridge, Mass. May 1, 1957 Number 74 



THE DISCOVERY OF CERAPACHYINE ANTS ON NEW 

CALEDONIA, WITH THE DESCRIPTION OF NEW 
SPECIES OF PHYRACACES AND 8PHINCT0MYRMEX 

By E. 0. Wilson 

Biological Laboratories, Harvard University 

During December, 1954, and January, 1955, the author was 
fortunate in being able to spend an uninterrupted five-week 
period on New Caledonia, studying the ant fauna of the island.^ 
During this time three species of Phyracaces and Sphinctomyr- 
mex were collected, all undescribed and representing the first 
cerapachyine ants ever found on New Caledonia. Their presence 
is of some zoogeographic importance, in that they appear to ally 
the ant fauna of New Caledonia more closely with that of eastern 
Australia, as opposed to the remainder of Melanesia. Phyracaces 
and Sphinctomynnex are strongly developed both in species and 
in individual numbers in Australia, but are known from only 
three rather rare species (two Phyracaces and one Sphincto- 
myrmex) on New Guinea, and are unknown from the rest of 
northern and central Melanesia. In an obverse relationship, 
Cerapachys {s. sir.), is the predominant cerapachyine genus on 
New Guinea and the Fijis, but has never been collected in either 
Australia or New Caledonia. Finally, the New Caledonian Phyra- 
caces and Sphinctomyrmex are most closely allied to eastern 
Australian species, as indicated in the descriptions to follow. 

Phyracaces cohici Wilson, new species 

Diagnusis. A shining, black, medium-sized species closely re- 
lated to the turneri group of species of eastern Australia. P. 

1 Field research was supported by grants from the Society of Fellows, Harvard 
University, and the Museum of Comparative Zoology. 



2 BREVIORA No. 74 

cohici can be easily distinguished from the latter group, which 
includes turneri Forel, adamus Forel, and larvatus Wheeler, 
by its more obtuse and rounded dorsal propodeal corners. In cohici 
these corners form an angle of more than 110° when viewed 
from the side, while in the turneri group of species they form 
an angle of 90° or less. P. cohici also bears a fair resemblance 
to the Australian P. senescens Wheeler, but can be separated 
from this species by its longer, flatter petiolar node and 
straighter and more horizontally aligned posterior petiolar teeth. 

Holotype worker. Head width ^ 0.99 mm, head length 1.05 mm, 
scape length 0.63 mm, cephalic index 94, scape index 64, exposed 
length of mandibles 0.21 mm, eye length 0.26 mm, pronotal 
width 0.79 mm, petiole width 0.83 mm, petiole length (measured 
from the midpoint of the anterior border of the node to the 
midpoint of the posterior border of the posterior peduncle) 
0.63 mm, postpetiole width 0.83 mm, postpetiole length 0.67 mm, 
width of next gastric segment 1.00 mm. Occipital border very 
feebly convex in full-face view. Alitrunk viewed from above 
moderately constricted medially, marginate along the entire 
dorsolateral border except for an interval of about 0.20 mm in 
the region of the mesothorax. Petiolar node viewed from 
directly above with strongly concave anterior border, and evenly 
convex lateral borders, its widest point being about in the middle 
(see accompanying figure). In the same view the posterolateral 
teeth extend well beyond the posteriormost point of the posterior 
node border. Seen from directly above, the postpetiole is widest 
in the anterior half and is laterally marginate only in the 
anterior half. 

Entire body covered by scattered piligerous punctures spaced 
on an average of 0.03 to 0.06 mm apart, the interspaces com- 
pletely smooth and shining. On the sides of the alitrunk the 
punctures are unusually small, being barely visible at 40X 
magnification. 

1 Head width and other body measurements were made in the standard fashion 
employed by W. L. I'.rown in his published serial studies of the daeetine ants and 
by myself in my recent revision of the ant genus Lasiug (1955, Bull. Mus. Comp. 
Zool. Ilarv., 113: 1-200). In the past, Brown and I have differed slightly in our 
cleflnitiuus of head length, but in the present paper I have decided for the 
purposes of uniformity to follow Brown's original definition, which was given 
as follows : "critical maximum length of the head, measuring from a transverse 
through the posteriormost point or points along the occipital border to a trans- 
verse through the anteriormost point or points on the anterior clypeal border" 
(1953, Amer. Midi. Nat., 50: 11). With this change I believe that our standard- 
ized measurements now all agree in every detail. 



1957 CERAPACllYINE ANTS OF NEW CALEDONIA 3 

Entire body deep blackish browu to jet black, except for the 
mandibles and gastric apex, which are dark reddish brown. Ap- 
})endag'es variably dark reddish brown. 

Worker variation. Maximum head width of tj^pe series (all 
shown by a single colony, ace. no. 190) : 0.88-1.01 mm. The 
worker tj^pe series shows very little variation in other external 
characters. 

MaJe. Head width (across and including compound eyes) 
1.10-1.14 mm. Antenna 13-segmented. Mandible well developed, 
narrowly subtriangular, its masticatory border bearing a large, 
blunt apical tooth followed basally by an indeterminate num- 
ber of serial denticulae ; the masticatory and basal borders join- 
ing through an even, convex curve. Clypeus narrow, its dorsal 
surface gently convex, its anterior border seen from directly 
above moderately and evenly convex. 

Parapsidal furrows well developed, parallel with each other, 
and extending anteriorly for about half the length of the scutum. 
Notaulices absent. Median notal suture present and extending 
posteriorly for slightly more than one-third the length of the 
scutum. Wing venation generalized, essentially similar to that 
of Cerapacliys (see Brown and Nutting, 1950, Trans. Amer. 
Ent. Soc, 75: 132, pi. VIII), differing primarily as follows: 
Rsf2'3 is lacking, and the second radial crossvein and Bsf5 form 
a single unit coming off the stigma. The crossvein " x" indicated 
by Brown and Nutting in the basal cubital-anal region of 
Cerapacliys manni is missing in Fhyracaces cohici. 

Petiole completeh^ lacking the lateral margination that char- 
acterizes the Fhyracaces worker caste. Seen from above, the 
node is widest in its anterior half, and its anterior and posterior 
corners are gently rounded ; seen from the side, the node forms 
a single, even, strong convexity only weakly demarcated from 
the peduncles. Genitalia completely retractile. Subgenital plate 
relatively large (exposed length about 0.35 mm), sclerotized, 
tapering posteriorly to end in a pair of sharp, upward curving 
hooks each about 0.06 mm in straight length. Parameres short, 
broadly rounded apically. 

Pilosity, sculpturing, and color essentially as in the worker. 

Types. Described from a long series of workers and four males 
collected by the author at Ciu, near Canala, at 300 meters, and 
from two kilometers southwest of Ciu, at 500 meters, New Cale- 



4 BREVIORA No. 74 

donia. The following accessions are included : 190, 246, 263 
(holotype nest series), 267, 275, 278, 298. Each represents a 
separate nest series with the exception of numbers 246 and 275, 
which are from the same nest. All of the collections were made 
in the period from December 21, 1954, to January 2, 1955. 

This species is named for Mr. Francois Cohic, of the Institut 
Frangais d 'Oceanic, an able and enthusiastic student of New 
Caledonian entomology. 

Ecological notes. All of the collections were made in the rich 
subtropical evergreen forest clothing the hills that extend from 
the south bank of the Canala River in the vicinity of the Ciu 
Falls. The holotype colony was found under a rock in a densely 
shaded part of the forest, and was occupying a single cavity and 
adjacent short vertical gallery in the soil. The males were very 
active and attempted to fly when the nest was exposed. Another 
colony (ace. no. 246-275) was nesting in open soil at the side of 
one of the forest trails. The nest entrance consisted of a single 
five-millimeter-wide opening surrounded by a low, indistinct 
turret of excavated earth. Lateral excavation revealed three or 
four galleries leading down from the entrance hole and into soil 
packed between several buried rocks. At about ten centimeters 
down two small adjacent chambers had been excavated in the soil 
against the vertical face of one of the rocks. In these were 
massed all of the brood and most of the workers. The following 
rough population estimate was made at the time of collection : 
80-100 workers, 40 pupae (in cocoons), 30-40 half -grown to fully 
grown larvae, and 30 eggs. The reproductive of this colony 
was unfortunately not found. 

Workers belonging to colony no. 246-275 and other colonies 
were encountered on several occasions foraging during the day, 
always in file, over the surface of the ground, and on one occa- 
sion (no. 190) workers were discovered in the upper layers of 
a moist rotting log. Twice, workers were observed in the act of 
raiding colonies of the ant genus Pheidole. The foraging and 
raiding behavior of this species will be described in greater detail 
in a later paper on the general subject of cerapachyine behavior. 



1957 CERAPACIIYINE ANTS OF NEW CALEDONIA 5 

Phyracaces dumbletoni Wilson, new species 

Diagnosis. Distinguished from all other species of the genns 
known to me by the following combination of characters : mod- 
erately large size (head width of type series 1.16-1.25 mm), 
antennal scapes unusually long, sides of alitrunk non-marginate, 
body surface smooth and shining to shagreened and subopaque, 
body color deep blackish brown to jet black. 

The only other Indo- Australian Phyracaces lacking margina- 
tions along the sides of the alitrunk are aherrans Clark and 
pygmaeus Clark of northern Queensland and hewitti "Wheeler of 
Borneo. These species are easily distinguished from dumhletoni 
by their much shorter scapes, which reach only to about the 
posterior margins of the compound eyes when the head is 
viewed in full face — in dumbletoni the scapes extend beyond 
the eyes by about their own maximum width. Of the three 
species, dumbletoni most resembles aberrans Clark, being very 
close in both size and sculpturing. 

Holotype ivorker. Head width 1.17 mm, head length 1.30 mm, 
scape length 1.08 mm, cephalic index 90, scape index 92, exposed 
length of mandibles 0.29 mm, eye length 0.33 mm, pronotal width 
1.03 mm, petiole width 0.95 mm, petiole length 0.76 mm, post- 
petiole width 1.09 mm, width of next gastric segment 1.26 mm. 
Occipital border in full-face view very feebly convex. Alitrunk 
viewed from above only feebly constricted medially, its dorso- 
lateral area evenly rounded and lacking any trace of margina- 
tion. Petiolar node seen from directly above with moderately 
concave anterior border and evenly convex lateral borders, its 
greatest width located in the anterior half. 

Puncturation as described for P. cohici, except that anterior 
to the postpetiole the punctures are more scattered, the majority 
being 0.09 to 0.12 mm apart. On the postpetiole and anterior 
to this segment the interspaces are for the most part "shag- 
reened," the shagreening in this case in reality consisting of 
regular, minute, contiguous foveolae each about 0.01 mm in 
diameter. The foveolae are deepest on the dorsal surface of the 
head and alitrunk, and render most of the cuticular surface there 
subopaque. They are absent posterior to the postpetiole. 

Color as described for P. cohici. 



BREVIORA 



No. 74 



Worker variation. Maximum head width range, Chapeaii 
Gendarme and Mt. Mou series: 1.16-1.25 mm (all shown by ace. 
no. 65, the holotype nest series). Maximum head width range, 
Ciu series: 1.11-1.20 mm (all shown by ace. no. 245). The Ciu 
workers differ significantly from the Chapeau Gendarme and 
Mt. Mou workers in the intensity of the f oveolar ' ' shagreening. ' ' 
The latter approach the condition described in the holotype, with 
little deviation. The Ciu specimens have the same basic form of 
sculpturing, but on the head and alitruncal dorsum the foveolae 
are much shallower, so that the surface is strongly shining 
under ordinary reflected light. 





cohici 



dumbletoni 



Dorsal view of the petioles of the two New Caledonian species of 
Phyracaccs ; outlines drawn to scale from the holotyjoes. 

Ergatoyyne. Head width 1.14 mm, head length 1.26 mm, scape 
length 0.76 mm, cephalic index 90, scape index 67, exposed 
length of mandibles 0.22 mm, eye length 0.30 mm, pronotal 
width 1.00 mm, petiole width 0.95 mm, petiole length 0.62 mm, 
postpetiole width 1.32 mm, width of next gastric segment 1.55 
mm. Ocelli lacking. More similar to the worker caste than to 
the queen caste of other Phyracaces species, differing primarily 
in the following external features. (1) The alitrunk is very 
worker-like, apparently differing only in the somewhat stronger 
pleural suturation. The posterior metapleural suture, which is 
absent in the dumbletoni Avorker, is present although feebly de- 
veloped in the ergatogyne. (2) The petiole is relatively shorter 



1957 CERAPACHYINE ANTS OF NEW CALEDONIA 7 

in the ergatogyne. (3) The postpetiole and gaster are much 
more massive and more poorly demarcated from each other than 
are the same structures in the worker. 

Types. Described from a long series of workers from Chapeau 
Gendarme (Yahoue), Mt. Mou, and Ciu, and a single ergatogyne 
from Ciu. The following accessions are included, each repre- 
senting a separate nest series: Chapeau Gendarme, no. 65 (holo- 
tj'pe nest series); Mt. Mou, single worker; Ciu, no. 245 and 
' ' observation colony. ' ' 

This species is named after Mr. L. J. Dumbleton, formerly 
economic entomologist for the South Pacific Commission, and an 
expert student of the New Caledonian insect fauna. 

Ecological notes. P. dumhletoni was collected at Chapeau 
Gendarme and Mt. Mou in dry, semi-deciduous "valley-pocket" 
forest and at Ciu in moist broadleaf evergreen forest. In all three 
localities it was limited primarily to the least disturbed portions 
of the forest, and was never encountered in the adjacent open 
Melaleuca woodland. 

At Chapeau Gendarme a large colony (no. 65) was found 
nesting in several spacious galleries and chambers in the upper 
layers of a large, moist, fern-covered log. It contained at least 
200 workers, a single ergatogyne (later lost), over 100 cocoons 
and larger larvae, and an undetermined number of eggs. Among 
the brood were found the hollowed-out propodeum of a worker of 
an undescribed species of Lordomyrma and the mangled remains 
of the larva of an undetermined ant genus. These insects ap- 
peared to be the prey of the Phyracaces, but of course this can- 
not be proven. 

When first disturbed, many of the Phyracaces workers sallied 
out with a display of aggressiveness unusual for cerapachyines, 
and one succeeded in stinging me on the forearm. I think it is 
worth mentioning that this is the only time I have ever been 
stung by a cerapachyine ant, despite the fact that I have ex- 
cavated many nests of Phyracaces and other genera without mak- 
ing any effort to protect myself from the workers. The sting 
caused a prominent welt about six millimeters wide surrounded 
by an erythema about twenty-five millimeters wide. There 
was a persistent, dull, throbbing pain of the sort commonly 
resulting from the stings of ponerine ants. 



8 BREVIOBA No. 74 

Sphinctomyrmex caledonicus Wilson, new species 

Diagnosis. Closely resembling S. steinheili Forel of eastern 
Australia, from which it can be distinguished by the following 
two characters. (1) Sculpturing, which is dominated by punc- 
turation, is overall denser in caledonicus. In steinheili, the inter- 
space distances over the genal surface are mainly as great as the 
adjacent puncture diameters or greater, while in caledonicus 
the genal punctures are contiguous and their borders form an 
even rugoreticulum. In steinheili, the lateral surfaces of the 
alitrunk are in large part feebly shining, while in caledonicus 
these surfaces are entirely opaque. (2) In side view, the dorsal 
posterior corners of the propodeum form a distinct angle of 
about 110° in steinheili, but are evenly rounded in caledonicus. 

Holotype worker. Head width 0.56 mm, head length 0.71 mm, 
scape length 0.41 mm, cephalic index 79, scape index 73, exposed 
length of mandibles 0.06 mm, pronotal width 0.42 mm, petiole 
width 0.35 mm, petiole length (including peduncles) 0.38 mm, 
postpetiole width 0.47 mm, w^idth of next gastric segment 
0.60 mm, length of gaster posterior to postpetiole (measured in 
a straight line) 1.35 mm. 

Worker variation. Maximum head width range (internidal) 
0.50-0.57 mm; (intranidal; aec. no. 195) 0.50-0.56 mm. In size 
as well as other external characters the worker type series is 
remarkably uniform. 

Ergatogyne. Head width 0.62 mm, head length 0.76 mm, 
scape length 0.42 mm, cephalic index 82, scape index 68, exposed 
length of mandibles 0.09 mm, pronotal width 0.44 mm, petiole 
width 0.42 mm, petiole length 0.41 mm, postpetiole width 0.60 
mm, length of gaster posterior to postpetiole 1.96 mm. This 
specimen is very worker-like, and can be distinguished only by 
its slightly larger size, proportionately shorter head and scapes, 
broader petiole, and larger postpetiole and gaster. In addition 
the postpetiolar-gastric constriction is somewhat weaker than in 
the Avorker. Compound eyes and ocelli are completely lacking, 
as in the worker, and the structure of the alitrunk is essentiallj- 
the same. 

Types. Described from a long series of workers and a single 
ergatogyne from Ciu, 300 meters, New Caledonia. The following 
two accessions, representing separate nest series, are included : 



1957 CERAPACHYINE ANTS OF NEW CALEDONIA 9 

no. 195, no. 225. The holotype was chosen from no. 225. 

Ecological notes. The two type colonies were found in a 
small, isolated patch of broadleaf evergreen forest on the farm of 
Mr. D. Fere, about half a kilometer northwest of the Ciu Falls. 
This little woodlot did not exceed two acres in extent, and its 
floor had been badly disturbed by cattle. Most of the ants 
present, including the Sphinctomyrmex, were found underneath 
rocks embedded in the soil. It is curious that the Sphinctomyr- 
mex was not found in the undisturbed forest on the south bank 
of the Canala River a short distance away, despite intensive 
collecting there. 

Both colonies were quite large, one containing over 500 work- 
ers and the other over a thousand. That the colonies may have 
been in migration is suggested by the fact that they occupied 
poorly defined galleries in the soil which bore no sign of lengthy 
occupation. The brood of colony no. 195, collected on December 
21, consisted of large numbers of mature larvae ; two days later, 
about three-quarters of a sample of these larvae kept alive in a 
bottle had spun cocoons. The brood of colony no. 229, collected 
on December 31, consisted of large numbers of cocoons, all of 
which contained prepupae of indeterminate caste. These data 
suggest a high degree of synchronization of brood development, a 
condition usually associated in ants with a nomadic way of life. 
An account of the behavior of the w^orkers of this species will 
be given in a later paper. 



BREVIORA 

Museiuoi of Comparsitive Zoology 



Cambridge, Mass. May 1, 1957 Number 75 



ON A NEW OCTOCHAETINE EARTHWORM 
SUPPOSEDLY FROM GUATEMALA 

By G. E. Gates 

INTRODUCTION 

Tweuty-four species of earthworms have beeu reported from 
Guatemala. One of them is widely distributed in North America 
where it presumably originated. Another, "pantropical," may 
have come from the south. Four are obviously exotic, probably 
introduced accidentally by man since 1500 A.D. The eighteen 
supposed endemics are known only from the original material. 

As a result of one of those accidents that occasionally advance 
our knowledge of an infrequently collected group of animals, 
another presumed endemic now can be added to the list of species 
known only from a single specimen. 

The author's thanks are extended to Dr. P. W. Oman for 
providing the present specimen and for assistance rendered in 
connection with problems arising from the study of various 
other species. 

Family MEGASCOLECIDAE 
Subfamily OCTOCHAETINAE 

Genus RamiELLA Stephenson 1921 

RaMIELLA AMERICANA n. Sp. 

Guatemala. In soil with Ceiba pentandra in cargo on plane arriving 
at San Pedro, November 25, 1955, 0-0-1. U. S. Bureau of Plant 
Quarantine. 



2 BREVioRA No. 75 

External characteristics. Length, 133 mm. Diameter, 8 mm. 
(through clitellum). Segments, 217 (+? possibly a recent 
posterior amputee). Body slightly compressed dorsoventrally 
and transversely elliptical in cross section behind clitellum. Un- 
pigmented (formalin preservation). Prostomium, prolobous. 
First two segments with numerous fine longitudinal grooves in 
both ventrum and dorsum. Secondary annulation, one presetal 
furrow each on iv-xii, one postsetal each on vi-xii, first presetal 
furrows deep but from ix less marked and like the postsetal; 
secondary furrows lacking behind the clitellum. Setae small, 
closely paired, only very slightly or not at all protuberant from 
epidermis, unrecognizable (lacking?) anteriorlj^; ventral couples 
first visible on vi, the lateral on xi, CD<^AB, J-A<BC, DD 
ca.=|C, both a and b present on left side of xrsdii, h only on 
right side, ventral couples of xvii and xix penial. Follicles of 
ventral couples larger than the others, ventral setae probably 
also larger. Nephropores unrecognizable. First dorsal pore on 
11/12. 

Clitellum reddish brown, saddle-shaped, reaching ventrally 
about to B, intersegmental furrows obliterated, dorsal pores 
occluded, setae retained, on xiii-xx. Epidermis 3-4 times as thick 
as rest of the body wall, cracking into blocks on slight tension. 

Spermathecal pores transverse slits of about the same width 
as AB, centered at A, on 7/8-8/9. Female pore in a transverse 
groove in A A just in front of eq/xiv. Male pores minute, about 
half way between B and a furrow circumscribing a male field, at 
17/18. Prostatic pores minute, on lateral wall of a common 
lumen containing both a and h penial setae, on xvii and xix. 

Male field longitudinally and broadly elliptical, demarcated by 
a distinct circumferential furrow somewhat lateral to B and 
just behind eq/xvi but less obvious in front of eq/xx. Two small 
but conspicuous conical protuberances on each of xvii and xix, 
about in region of AB, might be either localized temporary eleva- 
tions or eversions of slight parietal invaginations. The large 
aperture at the tip of each is filled by two penial setae. No other 
genital markings. 

Internal anatomy. Septum 4/5 membranous and transparent: 
5/6-9/10 thickly nuiscular, funnel-shaped, posteriorly directed; 
10/11 and 11/12 much thinner though slightly strengthened, ap- 
posed at parietes laterally and ventrally but easily separable till 



1957 NEW OCTOCHAETINE EARTHWORM 3 

just lateral to hearts of xi and there strongly adherent, ap- 
parently united dorsally and without insertion on body wall so 
that gut, dorsal trunk and hearts of xi are invisible at first. 
Septum 12/13 and those following all membranous. 

Progizzard portion of gut ca. 10 mm. long, the thin-walled 
oesophagus sigmoid. Gizzard large, in v. Gut in vi-xiv slender; 
caleiferous glands and lamellae unrecognizable. Inte.stinal origin 
in XV. Caeca vertically placed, 2.5-3 mm. long (high), slightly 
narrower dorsally, protuberant above dorsal face of gut so that 
the large dorsal trunk at first glance appears to be within a 
depression of intestinal roof, nine pairs in xxi-xxix. Typhlo- 
sole begins in xix-xx though a nuu'h lower ridge extends forward 
into xvi, reaching a height of 2 mm., lamelliform, anteriorly for 
a short distance Avith vertically lamellar lateral protuberances, 
ending abruptly in clxii. A fairly high and rather thick longi- 
tudinal ridge on each side in xxi-xxviii, between apertures into 
caeca and the median typhlosole, appears to be a lateral typhlo- 
sole. It probably does not function as a valve to close off lumen 
of gut from the caeca as the latter are filled with soil. (No 
supra-intestinal glands.) 

Dorsal blood vessel single throughout, large, gradually dwin- 
dling anteriorly and disappearing into tissues of pharyngeal 
bulb. Supra-oesophageal vessel recognizable only in xi-xii. Ven- 
tral trunk unrecognizable anterior to viii. Subneural unrecog- 
nizable (lacking?). Extra-oesophageals or other longitudinal 
vessels not found. Hearts of xi-xii large, latero-oesophageal 
though presumed bifurcations to dorsal trunk have no blood. 
Hearts of x somewhat narrower and lateral, those of ix-viii still 
slenderer and also lateral. Paired vessels from dorsal trunk in 
vii-v cannot be traced to the ventral trunk. ("Lymph glands" 
lacking.) 

Nephridia of iv represented by a large cluster of looped 
slender tubules on each side of anterior face of 4/5 (ducts not 
found nor any funnels, possibly closed, enterouephric and open- 
ing into pharynx). Paired, smaller clusters of similar slender 
tubules present on anterior faces of septa ventrally in v-xi, in 
viii-ix also attached to posterior faces of spermathecal ducts 
(ducts and funnels not found). Nephridia, from xii posteriorly 
on the parietes, on each side one transverse row of short loops 
from A nearly to mD. Nephridial tubules are as slender as 



4 BREVioRA No. 75 

anteriorly in xii-xvi but from xvii much thickened and flattened 
without marked increase in length. The thickening appears to be 
due to presence of the same sort of tissue as conceals the slender 
tubules of the micronephridia in certain species of Dichogaster. 
Individual loops, 9-10 on each side in each segment, are con- 
nected by a slender transverse cord (blood vessel? or?) but 
similar cords may be recognizable passing from the loop into the 
parietes or to the gut. Nephrostomes apparently are lacking 
until well posteriorly. From the medianmost loop on each side, 
beginning about at cxxv, a slender cord passes anteromesialh^ to 
and through the anterior septum, the anterior free end median 
to A very slightly widened. This cord and its slight terminal 
thickening presumably are neck and funnel respectively but 
even under highest power of the binocular definite funnel char- 
acters cannot be distinguished. Posteriorly the medianmost 
component of each transverse series becomes somewhat larger 
and of more than a single loop, and the larger widening of the 
free anterior end of the cord (neck) becomes definitely funnel- 
like. Typical, stomate exonephric holonephridia certainly are 
lacking throughout the entire length. Brain and circumpharyn- 
geal connectives well anteriorly in iii. 

Metandrie (no funnels in x). Male funnels large, plicate. 
Male deferent ducts large (with muscular sheath?), with large 
lumen, on parietes in BC, passing into body wall about at level 
of 17/18, slightly narrowed in xvii and still more so within the 
body wall. Seminal vesicles, in xii, medium-sized, cut up into 
very many fine lobes. Prostates tubular, looped and coiled, ca. 
20 mm. long, lumen central and recognizable clear to ental end, 
in xvii-xviii and xix-xx. Ducts 3-|- mm. long, slightly narrower 
than gland, with smooth surface, muscular sheen, slit-shaped 
lumen, muscular layer much thicker than the lining epithelium. 
Penisetal follicles at first separated from prostatic duct in body 
wall by a thin vertical sheet of longitudinal muscle, united more 
deeply. 

Penial setae have been studied by Mrs. Dorothy McKey- 
Fender. Her description is as follows : 

"Penial setae (Fig. A) curved, especially proximallj^ and 
distally, flattened in the plane of curvature and somewhat 
twisted, the apical, more strongly curved portion somewhat 
widened, forming an asymmetrical, spoon-shaped blade (Fig. C) 



1957 



NEW OCTOCHAETINE EARTHWORM 



with attenuate, often spirally twisted, tip. Distal seventh of 
shaft, including tip, sculptured on sides and both faces. Sculp- 
ture (Fig. B), of interrupted rows of fine teeth that become 
slightly longer and fewer per group toward the apex of the 
shaft. Setae a and h of both xvii and xix alike, except that the 
tip of the & setae may be more strongly twisted. An apparently 
mature, unworn seta is 3.2 mm. long (disregarding curvature), 
0.1 mm. wide and (being rectangular in section) 0.05 mm. 
thick." 




Penial setae of Bamiella americana. 

A. Camera lucida drawing of an entire seta. 

Arrow indicates location of detail shown in B. 

B. Detail of edge of blade of same seta, showing character of sculpture 

(More highly magnified.) 

C. Face view of distal portion of a fc seta from segment xix (sculpture 

omitted). 
Drawings by Mrs. Dorothy McKey-Fender. 

Spermathecae adiverticulate, ca. 5 mm. long, a band of con- 
nective tissue passing from each duct to the septum just in 
front. Ampulla narrowing somewhat entally, with several annu- 
lar constrictions externally, ridging of inner wall and large 
lumen. Duct ca. li mm. long, thicker than ampulla but asym- 
metrical, somewhat and gradually narrowed in parietes, lumen 
slit-like, wall thick, muscular. Seminal chambers, seven or eight 
on the bulged side, one or two smaller on the other side, com- 



6 BEEVIORA No. 75 

municate narrowly with duct lumen. Ovaries very small. (No 
ovisacs found.) 

Reproduction. Testis sac with little coagulum and that within 
a forward pocket-like, median bulge of 10/11. Male funnels with 
iridescence recognizable only at margins of the folds. Flecks of 
red iridescence recognizable in each lobule of the seminal vesicles 
(sheen of muscles in connective tissue binding lobules together, 
green-red-orange). Clitellum probably at or near maximal tu- 
mescence. However, seminal chambers in spermathecal duct and 
the ampullae are empty. Iridescence is unrecognizable ter- 
minally^ in male deferent ducts. Obviously copulation had not 
taken place. Accumulation of mature sperm on male funnels 
presumably had begun only fairly recently though the clitellum 
appears to be already at maximal development. Sexual, and bi- 
parental, reproduction is anticipated. 

Remarks. The gut, through xiii-xiv where coelomic cavities 
were filled with coagulum, is macerated but probably not enough 
so that intramural calciferous lamellae would have become un- 
recognizable. Extramural glands, since the intestine begins in 
XV, would have to be in xiii-xiv. In those segments, small 
slenderly-stalked glands, judging from experience with poorly 
preserved material of other species, could have been lost in wash- 
ing out the coagulum, if disintegration had not been previously 
completed. Site of stalk-gut junction, in either case, probably 
would be indistinguishable. As there is no reason for believing 
that extramural glands had been present in the holotype, 
absence in the species must, for the present, be assumed. 

The excretory system obviously is meronephric. 

DISCUSSION 

Absence of calciferous glands in ix-x, location of the male 
pores on xviii at some distance from external apertures of all 
prostatic ducts, and the meronephry require (cf. key in Pick- 
ford, 1937, p. 98) the Guatemalan species to go into the Octo- 
chaetinae. In that subfamily, only two genera now appear to 
require consideration. 

The genus Ramiella, hitherto recorded only from India and 
Burma, is defined by its author (Stephenson, 1930, p. 845) as 
follows: "Setal arrangement lumbricine. All septa present (be- 



1957 NEW OCTOCHAETINE EARTHWORM 7 

hind their commencement). One oesophageal gizzard in one 
simple segment. No calciferous glands. Excretory system purely 
micronephridial, the micronephridia relatively large and few in 
number, from seven to one pair per segment. Sexual apparatus 
purely acanthodriline. ' ' 

The phrase "purely micronephridial" was intended to convey 
the idea that holonephridia are lacking from one end of the body 
to the other. One word, meronephric, is sufficient for that pur- 
pose. Further characterization of the excretory system is not 
warranted at present^ any more than previously, the supposed 
reduction of meronephry to a single pair of nephridia per seg- 
ment being based, in one species at least, on the number of 
opaque white masses that were externally recognizable through 
a semi-transparent body wall. The definition accordingly should 
read, in part, only as follows: Excretory system meronephric. 

Ramiellona Michaelsen 1935, also meronephric, differs from 
Ramiella only in having calciferous glands. The latter are 
characterized (p. 53) as follows: "segmental Erweiterungen des 
Osophagus mit blutreichen Falten der Wandung im 12 Segment 
und einigen vorhergehenden." The small vascular folds on the 
inner wall of the oesophagus in Ramiellona stadelmanni Michael- 
sen 1935 (the only species) are not known to differ from 
similarly located ridges in Ramiella hishamhai'i (Stephenson, 
1914) and R. nainiana Gates 1945. As Ramiellona is not dis- 
tinguishable morphologically, the Honduran stadelmanni, as 
well as the Guatemalan americana, must go into Ramiella. 

The huge distributional discontinuity thus established is not 
exceptional in a system with geographical ranges such as the 
following: South India-Ceylon and Australia-New Zealand 
(Notoscolex and Megascolex), Australia-New Zealand and the 
Pacific coastal strip of the United States {Plutellus and Mega- 
scolides), temperate-zone North America and peninsular India 
(Diplocardia) . 

The two Central American species of Ramiella do nut appear 
to be more closely related to each other, than to any of their 

1 Specimens of Ramiella nainiana Gates 1945, tixed aud preserved in accordance 
with his instructions, were forwarded, along with other material, to K. N. Bahl 
for study of the nephridia. If his failure to publish thereon had been anticipated, 
:jome data would have been included in the original description while the worms 
were still available. Ever since, live individuals of any species of Ramiella have 
been unavailable. 



8 BREVIORA No. 75 

oriental congeners. The anatomy, as known, lacks the uniformity 
one should expect in a good genus. Ramiella thus provides one 
more illustration of the inadequacy of the classical system and 
the futility of patchwork changes in generic definitions . 

EEFEBENCES 

Gates, G. E. 

1945. On some Indian earthworms. Proc. Indian Ac. Sci., 21: 208-258. 

MiCHAELSEN, W. 

1935. Die opisthoporeu Oligochaten Westindiens. Mitt. Mus. Hamburg, 
45: 51-64. 

PiCKFORD, G. E. 

1937. A monograph of the acanthodriline earthworms of South Africa. 
Cambridge, England, 612 pp. 

Stephenson, J. 

1930. The Oligochaeta. Oxford, 978 pp. 



BREVIORA 

Museniim of Coimparative Zoology 

Cambridge, Mass. June IS. U).")? Number 76 



TWO NEW LAXI) AXl) FKESllWATEK MOLLl'SKy 
FROM NEW GUINEA 

By AViLLiA.M -I. Clench 

The tirst species was received from Mr. James Poling of New 
York City, the second from ]Mrs. Marjorie Kleckham of Darn, 
l'ai)na. New Gninea. 

]\Ir. Poling pnrchased all of the remaining stock of the land 
and marine mollusks from Ward's Natural Science Establish- 
ment in Rochester, New York, after the death of Mr. F. H. Ward. 
Contained in this miscellaneous material were a few lots of land 
shells, originally obtained from some collector in New Guinea. 
Subsequent correspondence by ^Ir. Poling with Ward's has failed 
to add to the meager data accompanying the specimens. Just 
wlio the collector was or the exact locality in New Guinea still 
remains unknown. The locality is somewhere in western New 
Guinea, to judgi^ by th(» associated specimens from the same 
('()ll(H'tor. 

VeR1)ICHL01[1T1S new subgenus 

Tliis subgenus differs from all other species in the subfamily 
Chloritinae by being a dull jade green. All other species, now 
inunbering over 200, are uniforml}' brown, red-brown, or banded 
with some shade of broAvn or red-brown. The usual arrangement 
of the periostracal "hairs" is in staggered or offset rows; in this 
subgenus the rows are regular. 

T]ipr aperies. Eusfrnxopsis (VerclicJdo)-)fis) polijufi Clench. 

EUf^TOMOPSIS ( \'e1;I)U IILOKITISJ POLINGI. uew S])ecies 

Figure 1 
1)( scnptioii. Shell small. i-cMching 1") mm. in gi-eatest diameter, 



BREVIOKA 



NO. 76 



plauorboid in shape, thin, fragile and minutely hirsute. Color 
a uniform and dull jade green. Whorls 31/2, convex, and coiled 
in a single plane. Spire depressed below the body Avhorl. Aper- 
ture subcircular. Outer lip thin and very nan-owly reflected. 




Fig. 1. Kiistoniopsifi (I'crdichlorili.s) poivuiii Clciu-li (llolotyi)e, 4X). 
Fig. '1. ll'e.sfralunio <ill)r)-ti.'<i Clcncli (Holot.vpc, 1.5X ). 

Parietal area covered with a vei-y thin glaze. Scul])turc consist- 
ing of numerous rows of very short, gold-colored periostracal 
"hairs" which arc arranged axially and slightly diagonally. 



1957 TWO NEW 1,A.\I) AM) KKKSI 1 WATEIi .MOIJ.ISKS 3 

i'ollowiu<i' the liiu's of <ir()\vtli. These rows nvv I'ejiular aiul not 
staggered or offset as in other species of Eustomopsis. UmbiliciLs 
narrow but deep, the nueleai' whorl being visil)h'. Periostraeiim 
jad(^ green overlaying a nearly glass-like shell. 

Height (it. (liametor Less, diameter 

7.5 mm. 15 mm. 11.5 mm. Ilolotype 

Type. The holotype is in tlie ]\Iuseum of Comparative Zool- 
ogy, No. 212319, from western New Guinea. 

Neinarks. This new species is (piite remarkable. The dull jade 
green color sets it well apart from all other species in the sub- 
family Chloritinae. So far as I am aware, all other species in 
the Chloritinae are uniform brown, red-brown, chocolate brown, 
or light brown to yellowish and banded witii brown. Many 
species are hirsute, that is, the periostracum forms little "hairs" 
in regular or offset rows, such "hairs" emanating from small pits 
impressed in the calcium of the shell. 

It is quite possible that this species is arboreal and, as in many 
other groups, the species which become arboreal also become 
somewhat brightly colored, as, for example, species in Licjims, 
PoJijmiia, Placostylus, Helicostyla, Amphidromus and many 
other genera. 

A factor of considerable interest is the green color, a rare 
color even in arboreal mollusks. A few species are banded with 
green, such as Liguus and Amphidromus but nearly solid green 
is exceedingly rare. Helicina viridis Lamarck and H. castilloi 
Clench from Hispaniola, Papustyla p^ilcherrima Rensch from 
the Admiralty Islands, two or three species of Helicostyla and 
Ckloraea from the Philipiiines are about all the known species to 
po.ssess this color predominantly. 

AVestralunio albertisi, new species 
Figure 2 



'^p " 



Description. Shell inequilateral, subcircular in outline, 
rounded anteriorly, subtruncate posteriorly and reaching 53 mm. 
in length. Umbos rather small, slightly elevated above the dorsal 
margin and anterior to the center. Shell moderately compressed 
and rather light in structure. Color a dark blackish brown. 
Posterior slope fairly well marked and slightly concave. Po.s- 



4 BREVIORA XO. 76 

tcrioi- i-i(l<z'e not well defined. Lio-ament lon^' and very narrow. 
I'eriostracnm shining- on tlie disc Init somewhat scaly and dull 
on the posterior slope. 

Nacre white, shining and slightly iridescent ]X)steriorly. 
^luscle scars well outlined. Hinge plate long- and narrow, liight 
valve with two pseudocardinal teeth, the innermost being the 
larger. In addition, there is a single and long, narrow, lateral 
tooth. Left valve with a single pseudocardinal tootli and two 
long: and narrow lateral teeth. 



Length 


Height 


Width 




53 mm. 


42 mm. 


20 nnn. 


Ilolotype 


51 


44.5 


22.5 


Paratype 


57 


50.0 


25 


Paratype, Lake ^lurray, 
Flv River 



Types. The holotype is in the ^luseum of Comparative Zool- 
ogy, Xo. 212!)08, inland from Darn, Western Division, Papua. 
New Guinea. A single paratj^pe from the same locality is in the 
Australian Museum, Sydney. An additional paratype is from 
Lake JMurray, Herbert River, Fly River System. All specimens 
were collected by ^Ir. Kleckham of the Department of Agricul- 
ture, Papua. 

This new species is related to Wcstralioiio tl!i( )isis Tapparone 
Canefri from the upper Fly River, Papua. It differs from W. 
ffyensis by being far more circular in outline and in being i)ro- 
portionately somewhat greater in width. In addition, the ventral 
margin is rounded, the posterior portion of the dorsal margin 
is somewhat wing-like and the posterior ridge is well defined. 

This species is named for L. M. d'Albertis. a naturalist-ex- 
plorer who was in New (iuinea in 1872-73 and again in 1875. 
Much of the mollusk work accomplished l)y Tapinu-one Canefi-i 
was hascd upon iiuitei-ifd collected by d'Albei-tis. 

REFEBENCES 

Haas, F. 

1924. Uiiserc hishciigcii Kenntnisso der X;i,);Ml('nf;iini;i Xcu-diiiiicns. 
Xnvn (iiiincii, 15: 65-7(), 1 i)l:ite. 

Mc.Mkuaki., I). F. 

19.16. Note.s on the t'lcsh watei- imissel.s of New (Uiinen. Xautihis. 70: 
38-48, 1 plate. 



n 



BREVIORA 



m of Comparative Zoology 

Cambridge, Mass. June 21, 1957 Number 77 



DACETINOPS, A NEW ANT GENUS FROM 
NEW GUINEA 

By W. L. Brown, Jr. 

and 

E. 0. Wilson 

In the course of an extended field trip to the southwestern 
Pacific area during 1954-1955 (supported by the Museum of 
Comparative Zoology and the Society of Fellows of Harvard 
University), one of us (Wilson) was able to collect in the rich 
tract of rain forest at the Busu River, near Lae, Australian 
Mandated Territory of New Guinea. Among- the samples ob- 
tained here were several of a small myrmicine ant with the 
habitus, both in life and in preservation, of certain short-mandib- 
ulate members of tribe Dacetini. However, further examination 
revealed that this species could not be a member of the Dacetini, 
even though it possessed certain features, especially the spongi- 
form appendages of the petiole, postpetiole and gaster, not 
otherwise known among ants outside the Dacetini. This remark- 
able pseudo-dacetine is described below and its affinities dis- 
cussed. 

DaCETINOPS gen. nov. 

Diagnosis, worker. Small myrmicine ant, apparently closest 
to Lachnomyrmex Wheeler of the American tropics. Body com- 
pact, integument hard and thick. Head subtriangular (with 
mandibles triangular), widest across occipital region, tapering 
toward mandibles. Antennal scrobes present, deep and distinctly 



2 BREVIORA XO. 77 

bounded; compound eyes moderate in size, situated ventrad of 
scrobes near midlenp^th of head. Antennae 11-segmented, with 
a fairly distinct 8-segmented club. Both sets of palpi 2-se<i- 
mented. Clypeus convex, obscurely bicariuate, its anterior 
border forming- a broadly rounded translucent apron. Mandibles 
triangular, broad, with straight, crenulate masticatory margins. 
Alitrunk compact, arched and sutureless dorsally ; propodeal 
dorsum and declivity almost continuous; a pair of small but 
distinct propodeal teeth present. Petiolar node sessile (a small 
condylar extension is visible from above), rounded above, and 
sloping from a median summit ; postpetiole transversely ellipsoi- 
dal, rounded above. Gaster broad and somewhat depressed ; first 
segment taking up nearly all the length, nearly circular in out- 
line as seen from above, bluntly margined along the sides in 
front; remaining segments small, retracted. Sting sclerotized, 
acute, exsertile. The lower halves of the petiole, postpetiole and 
anterior part of the first gastric segment bearing and covered 
i;y bulky festoons of whitish spongiform tissue. Sculpture con- 
sisting predominantly of very coarse longitudinal costation, be- 
coming more reticulate on head, nodes and sides of alitrunk : 
posterior half of gaster smooth and shining, as is also the pro- 
podeal declivity. Mandibles striate. Legs and antennal scrobes 
finely and densely punctulate. Pilosity consisting of sparse, 
long, flexuous tapered hairs, mostly situated at definite bilateral 
positions over the body. Pubescence forming a thin cover mostly 
only on appendages. Color deep brownish red. 

Female. Dealate, but with thoracic flight segments well de- 
veloped, as usual for winged myrmicines of small size ; wing 
stumps present. Size only slightly larger than worker ; form, 
except as mentioned, similar to that of worker, with only the 
usual slight dilTerences of caste. The compound eyes are larger 
than in the worker, but are still modest in size. Scutum and 
scutellum forming together a nearly plane platform, crossed by 
the distinct transseutal suture ; prescutellum reduced to two 
small lobes, one on each side of the suture, so that the scutum and 
scutellum meet at the suture. Scuto-scutellar platform evenly 
costate longitudinally. 

Type of genus : Dacetinops cibdela sp. nov. 



1957 



DACETINOPS, A NEW ANT GENUS 




LAB 





Dacetinops cihdeln, geu. et sp. uov., worker. Fig. 1, holotype, head in 
full-face view; most of sculpture omitted, and only the hairs nearest the 
periphery shown. Fig. 2, holotype, body in lateral view; hairs and all 
sculpture except the prominent dorsal costation omitted. Fig. 3, paratype,. 
detail of maxillary and labial palpi. Drawings by Nancy BufiBer. 



4 BREVIORA NO. 77 

Dacetinops cibdela sp. nov. 

Holotype worker. TL 2.7, HL 0.64, HW 0.59 (CI 91), ML 
0.16 (M2i5), WL 0.76, petiole L 0.22 mm. Index of cephalic 
depression (ICD) is about 62. Measurements and indices are the 
ones used in recent works on Dacetini by Brown (see especially 
Brown, 1953, Amer. Midi. Nat., 50: 7-15). 

Form of head and mandibles as shown in Figures 1 and 2. 
Scrobes sharply defined, each divided partially by a fine anterior 
carina, and able to receive the entire antenna except the apical 
part of the club ; upper scrobe margins narrowly lamellate. 
Pronotum margined in front, submarginate along the sides ; 
liumeral angles obtuse, but distinct, subtuberculate. Propodeal 
region of alitrunk short, tapering rapidly liehind promesonotum ; 
propodeal declivity weakly concave from side to side, bounded 
above by an ogival margin, and marginate laterally. Propodeal 
teeth acute, feebly downcurved toward their apices, their bases 
separated by about 4 times their length. Propodeal spiracles 
not far under the teeth on the lateral margins of the declivity, 
facing posterolaterad. Petiolar node seen from above approxi- 
mately as broad as long, with strongly rounded sides, widest 
near midlength. Postpetiole about half again as broad as long, 
ellipsoidal or subreniform as seen from above, the more strongly 
rounded margin posterior. Gaster with a short, straight anterior 
margin opposite the postpetiole, the sides rounded away on each 
side. 

Spongiform appendages of petiole, postpetiole and gaster finely 
areolate, apparently composed of cuticular material, not ar- 
ranged in distinct lobes like those of the dacetines, but narrowed 
medially below, where masses of opposite sides are joined 
(Fig. 2). 

About 10 coarse, fairly distinct longitudinal costae across the 
head between the ends of the scrobes, these converging and 
anastomosing anteriad between the frontal lobes, and joined by 
short transverse ridges to form a coarse reticulum (not shown 
in the figures). The bottoms of the large pits or foveolae thus 
enclosed are sculptured finely and are subopaque. Disposition 
of costulae on clypeus shown in Figure 1 ; interspaces here finely 
sculptured and subopaque. Underside of head coarsely rugo- 
reticulate. Mandibles coarsely striate at base, becoming smooth 



lf)57 DACETINOPS, A NEW ANT GENUS 5 

and shining apiead, with scattered punctures. Dorsum of ali- 
trunk with smooth, straight, heavy longitudinal costae running 
from anterior pronotal margin to the beginning of the propodeal 
declivity (10 costae across pronotal dorsum), shining and with 
more or less shining interspaces. Sides of alitrunk and dorsal 
surface of petiolar node coarsely rugo-reticulate. Basal half of 
postpetiole and of gaster with coarse longitudinal costae (9-10 
across gastric base), remainder of gaster, postpetiole, declivity 
and concave lateral faces of propodeum smooth and shining, 
often with scattered punctures. Legs, antennae, scrobes and a 
band along each side of the first gastric sternum densely punc- 
tulate, opaque. 

Long, fine, tapered hairs scattered over dorsum of head (those 
nearest the cephalic borders are shown in Figure 1), alitrunk, 
both nodes and gaster as well as a few on the gular surface of 
the head, on the coxae and on the underside of the femora ; 
length ranging about 0.12-0.19 mm. Short, fine, appressed to 
decumbent hairs forming a pubescence on the legs, antennae and 
sternum and apex of gaster; similar hairs are sparsely dis- 
tributed over both surfaces of head, mostly one hair to a foveola, 
and over mandibles. 

Basic body color deep brownish red ; mandibles, legs and 
antennae, especially the club and the first funicular segment, 
more yellowish. 

Holotype worker (deposited in the Museum of Comparative 
Zoology) taken in rain forest at the lower Busu River, near Lae, 
New Guinea (E. 0. Wilson leg., No. 1058), May 17, 1955, either 
as a stray or in berlesate from under bark of a large Zoraptera- 
stage log. Three additional workers (paratypes) were taken in 
this collection and bear the same number. 

Paratypes, workers. 9 mounted dry, 3 in alcohol, all collected 
in the same tract of forest as the holotype (Wilson Nos. 899, 
942, 978, 1052, 1058, 1113). Deposited in Museum of Comparative 
Zoology, U. S. National Museum, Coll. 6. C. Wheeler, and one 
or more each in Australian and European collections as yet un- 
selected. Total maximum variation for all series: TL 2.2-2.6, 
HL 0.54-0.63, HW 0.50-0.57 (CI 91-93), ML 0.15-0.16 (MI 24-27), 
WL 0.65-0.76, petiole L 0.18-0.22 mm. Greatest intranidal varia- 
tion occurs in series No. 1052: TL 2.2-2.5, HL 0.54-0.62, HW 



6 BREVIORA NO. 77 

0.50-0.56 (CI 91-92), ML 0.15 (MI 25-27), WL 0.65-0.75 mm. 
Very little variation among the workers of these lots. Sculpture, 
particularly of cephalic dorsum, varies in minor details ; spongi- 
form appendages a trifle more voluminous in some specimens 
than in others. 

Paratypes, dealate females. 3 specimens, taken with workers 
(Nos. 942, 1052, 1113, the last in alcohol, not measured). De- 
posited in the Museum of Comparative Zoology and elsewhere 
with the workers. Queen variation (Nos. 942 and 1052) : TL 
2.6-2.8, HL 0.58-0.62, HW 0.56-0.60 (CI 96-97), ML 0.17-0.19 
(MI 29-30), WL 0.79-0.85 mm. 

Larva. A single medium larva was preserved in alcohol. This 
specimen is short and thick, with head turned ventrad. Without 
proper preparation, none of the details of this larva can be made 
out, except that the hairs are varied in length, with some of 
the longer dorsal ones anchor-tipped. This larva has been sent 
to Dr. G. C. Wheeler for expert study (See Breviora No. 78). 

Biology. The six collections were all made on the forest floor 
in heavy to medium rain forest. Nos. 978 and 1052 were strays 
taken from soil-leaf litter berlesates. No. 899 was a stray sample 
from the forest floor, beneath a log. No. 942 was a small nest in 
a cavity under the bark of a Zoraptera-stage branch about 6 cm. 
in diameter, buried in the leaf litter ; the queen and two workers 
were taken. No. 1113 is a queen with one worker, originally 
taken with a few larvae and two eggs in a small cavity in the 
middle of a small piece of rotten wood buried in the leaf litter. 
This apparently incipient colony was kept in an artificial nest 
for 10 days, but showed no signs of predatory or nest-founding 
behavior during that time. All of the collections were made 
during the first three weeks in May, 1955. 

From these observations, we may perhaps conclude that D. 
cibdela is normally a dweller in the leaf litter of the rain forest 
floor, where it frequently, perhaps usually, nests in or beneath 
the bark of rotting logs or other large or small masses of rotting 
wood. Apparently the nests are small in volume and in popula- 
tion, like those of other specialized small myrmicines with simi- 
larly slight differences separating the female and worker castes. 
Wilson noted that the workers of this species walk in a slow, 
deliberate fashion reminiscent of dacetines and basicerotines. 



1957 DACETINOPS, A NEW ANT GENUS 7 

Relationships. Although from its general habitus this species 
seems to belong to the Dacetini, a closer look does not bear out 
placement with that tribe. In the first place, its resemblance is 
general, and not particular to any one dacetine genus or sub- 
tribe. The shape of the head and mandibles are fundamentally 
different from those of any dacetine, as is also the arrange- 
ment of the spongiform appendages, which at first sight are so 
dacetiue-like. The antennae are 11-segmented, like those of the 
primitive dacetines, but the distinct club is not a dacetine fea- 
ture; the palpi of Dacetinops are segmented 2, 2, against 5, 3 for 
the primitive dacetines and 1, 1 for the higher dacetines. The 
details of structure of the alitrunk and petiolar node are also not 
like those of any dacetine, and the sculpture recalls that of the 
dacetines only at the gastric base, but even here, the costulae are 
of a basically different type. Ties with the Basicerotini are even 
fewer, the position of the eyes being one important difference ; 
there seems to be no need to compare further with basicerotines. 

As already mentioned, the closest similarity holds with the 
neotropical genus Lachnomyrmex Wheeler (with three described 
species). The best account of Lachnomyrmex is that of M. R. 
Smith (1944, Proc. Ent. Soc. Washington, 46:225), which gives 
excellent figures of the genus. Lachnomyrmex also has an 11- 
segmented antenna, but the club is 2-segmented. This difference 
in club segmentation is really not a very important character, 
since only a slight change in size of the antepenultimate seg- 
ment would be needed to make the Lachnomyrmex club 3-seg- 
mented. Lachyiomyrmex has no posterior expansion of the head, 
as in Dacetinops, and the form of the node is somewhat dif- 
ferent, as well as the proportions of the parts of the alitrunk. 
but the differences here are not as serious as those between 
Dacetinops and the dacetines. Dacetinops appears to belong, 
with Lachnomyrmex, to a group of small myrmicines that in- 
cludes also the New World Kogeria Emery (the so-called Rogeria 
from the Melanesian area are not true members of the genus, 
but are closer to Lordomyrnia), Apsychomyrmex Wheeler, and 
Adelomyrmex Emery, as well as the primitive Agroecomyrmex 
Wheeler of the Baltic Amber. Apparently these are specialized 
relicts of an ancient and widespread myrmicine fauna that still 
retain some marks of their eetatommine ancestry. 



BREVIORA 

Mmseiiim of Cooiparsitive Zoology 



Cambridge, Mass. June 21, 1957 Number 78 



THE LARVA OF THE ANT GENUS DACETINOPS 
BROWN AND WILSON 

By George C. Wheeler aud Jeanette Wheeler 

Department of Biology 
University of North Dakota 

Genus DaCETINOPS Brown and Wilson 

LARVA 

Body hairs sparse, of two types; (1) with short-bifid tip and 
(2) anchor-tipped, with sinuate shaft. Cranium subcordate. 
Antennae minute. Head hairs few (about 26), with short-bifid 
tip. Posterior surface of labrum with numerous spinules. Man- 
dibles long and slender; apex forming a long sharp-pointed 
tooth which is curved medially ; with a narrow blade projecting 
medially from the anterior surface and bearing two long coarse 
medial teeth. Maxillae small, with the apex paraboloidal ; palp 
a stout peg; galea digitiforra. Anterior surface of labium with 
numerous spinules. Hypopharynx spinulose and with numerous 
longitudinal ridges near the pharynx. 

Dacetinops cibdela Brown and Wilson 

(Text figure 1) 

Immature Larva. Length through spiracles about 1.25 mm. 
Short and stout; prothorax directed ventrally; head ventral; 
anterior end formed from the dorsum of the mesothorax; dorsal 
profile C-shaped; abdominal somites II and III produced ven- 



BREVIORA 



NO. 78 



trally into rounded transverse welts. Anus ventral, with a small 
posterior lip. Spiracles small, mesothoracic slightly larger than 
the rest. Integument apparently without spinules. Body hairs 
sparse and uniformly distributed, of two types: (1) 0.036-0.15 
mm. long, slightly curved, with short-bifid tip, some on each 
somite, longest on the thorax and ventral surface; (2) anchor- 
tipped, with sinuate shaft, about 0.16 mm. long, four in a row 
across the dorsum of each abdominal somite II-V. Cranium 
subcordate in anterior view, clypeus bulging. Antennae minute, 




Text figure 1. Dacetinops cibdela Brown and Wilson, larva. A, head in 
anterior view, X93 ; B, left mandible in anterior view (stippled to show 
thickness), X271 ; C, left mandible in posterior view (shaded to show con- 
tours), X271 ; D, immature larva in side view, X54; E, labrum (left half 
of drawing in posterior view, right half in anterior view), X271; F, 
hypopharynx in anterior view, X271 ; G, anchor-tipped body hair, X185; IT 
;ind T, two bifid-tipped body hairs, X185. 



1957 LARVA OF THE ANT GENUS DACETINOPS 3 

each with three sensilla, each of which bears a minute spinule. 
Head hairs few, 0.036-0.09 mm. long, slightly curved, with 
short-bifid tip. Labrum short, wide and bilobed ; each lobe with 
about 13 sensilla on and near the ventral border; posterior sur- 
face of each lobe with a central cluster of 4 or 5 sensilla, lateral 
to which the spinules are large, isolated and sparse, but medially 
and dorsally the spinules are much more numerous, smaller and 
arranged in short transverse rows which are grouped into longer 
rows. Mandibles heavily sclerotized, rather narrow and elongate, 
subtriangular in anterior view; with one large apical tooth 
which is curved medially and posteriorly; with a median blade 
arising from the anterior surface and bearing two large medial 
teeth. Maxillae small, apex with a few short encircling rows 
of minute spinules; palp a stout peg with five apical sensilla 
(three minute and bearing a spinule each, two larger and en- 
capsulated) ; galea digitiform, with two apical sensilla. Anterior 
surface of labium spinulose, the spinules numerous and in short 
arcuate rows, the rows arranged in a longer subtransverse pat- 
tern, the spinules longer ventrally ; palp a low knob with five 
apical sensilla (three minute and bearing a spinule each, two 
larger and encapsulated) ; opening of sericteries in a wide de- 
pression on the anteroventral surface of the labium. Hypo- 
pharynx spinulose, the spinules numerous, minute and arranged 
in subtransverse rows; numerous longitudinal ridges near the 
pharynx. (Material studied : one damaged larva from New 
Guinea, collected by E. 0. Wilson; courtesy of W. L. Brown.) 

A single damaged immature larva is a frail foundation on 
which to base the taxonomic relationships of a genus. Neverthe- 
less we must hazard a few guesses, for that is all the material we 
have. Such guesses are the more difficult because of the nearly 
diagrammatic structure of this larva. If only it had a few (or 
even one) striking peculiarity — something overdeveloped or 
something lacking. If we extrapolate the mature body shape, 
enlarge the antennae to normal size, inflate the bases of the 
mandibles a little and reduce somewhat the size of the medial 
mandibular teeth, the result might be considered the synthetic 
type for the subfamily Myrmicinae. 

AVe are now in the process of synthesizing our studies of the 
larvae of the Myrmicinae. Consequently we do not yet have clear 



4 BREVIORA NO. 78 

ideas as to what characters are phylogenetically basic. However, 
we do feel that a few provisional opinions about the relation- 
ships are not out of order. 

The larva of Dacetmops resembles the larva of Rog&ria so 
closely that the same generic description would apply to either — 
except for the hairs : Rogeria lacks the anchor-tipped hairs which 
we suspect will prove to be of basic phylogenetic importance. 
The larva of Dacetmops also resembles the larvae of the tribe 
Basicerotini but differs in having anchor-tipped hairs and by 
lacking spinules on the mandibles. The worker of Dacetinops is 
convergently similar to the Dacetini, but its larva is not. In 
fact, the larva of Dacetinops shows no close affinity with any 
known genus. 



BREVIORA 

Muiseimm of Coimparative Zoology 



Cambridge, Mass. August 9, 1957 Number 79 



DASYPELTIS MEDICI LAMUENSIS, A NEW RACE OF 

EGG-EATING SNAKE (OPHIDIA, REPTILIA) FROM 

COASTAL EAST AFRICA 

By Carl Gans 

Museum of Couii);iiativc Zoology and Carnegie Museum, Pittsburgh, Pa. 

Some time ago an analysis of the African egg-eating snakes 
of the genus Dosypeltis indicated the presence of an undescribed 
race from coastal East Africa. However, as the entire revision 
of this genus will not be published for several months, I am 
descril)ing the new' race here in order that Mr. Arthur Loveridge 
may include it in his checklist of the reptiles and amphibians 
of East Africa, which is now in press. 

In 1942, Loveridge revived Dasypelfis medici as a subspecies 
of the wide-ranging D. scahra. Examination of additional speci- 
mens from supplementary localities convinced me that two forms 
are involved, and furthermore that D. scahiri and medici, though 
sympatric in many areas, are distinct species which, besides 
their color patterns, show differences in a number of other 
characters. Among these are ventral and caudal counts of both 
male and female species, body proportions, etc. 

In his 1942 paper Loveridge mentioned in passing that north- 
ern specimens of medici were unicolored and did not possess 
the characteristic color pattern upon the basis of which this 
species was initially described by Bianeoni. The ''uniform" 
coloration has been found to be associated with significantly 
lower ventral counts and the northern specimens are here 
recognized as a distinct race. 

The detailed acknowledgments to the many who helped with 
the analysis Avill be given in the main paper. Here I restrict 
myself to thanking Ernest E. Williams and Arthur Loveridge 



BREVIORA 



NO. 79 



of the Museum of Comparative Zoology (MCZ) for checking 
this paper, the John Simon Guggenheim and National Science 
Foundations for supporting the research upon wliicli this paper 




Figure 1. Dasypeltis medici. Dorsal views showing pattern atop head. 
Top. D. m. medici — BM 12-1-30-6 from Matemo, Mozambique. Bottovi. 
I), m. lamuensis — BM 51-1-3-73 from Kilifi, Kenya. Note that the pattern 
is almost completely faded out on the parietal scales of lamuensis, and has 
disappeared in the nuchal region. The pustulosity or pitting on the head 
shields and the dark pigmentation of the pits is clearly apparent. 



I9;i7 DASYPELTIS MKDKI LAMUENSIS 3 

is based, and i\lr. J. C. Battersby of the British Museum (Natu- 
ral History). London (BM), as well as Dr. Heinz Wermuth of 
the Zoolooisehen Museums der Universitat, Berlin (ZMTH, for 

the loan of paratype material. 

DASYPELTIS MEDICI (Bianeoni), 1859 

1. rattern, (>onsisting of narrow lateral liars (of which at least the first 
liirce to eight fuse to form narrow Vs on the nape), extends the entire 
length of snake from the head on to the tail; ventrals more than 229 

in males, more than 236 in females D. m. medici. 

2. Pattern, if present, restricted to top of head (see Fig. 1) ; ventrals 
229 or less in males, less than 233 in females D. m. lamucnsis. 

Dasypeltis MEDICI MEDICI (Bianconl), 1859 

Dip.su.s Medici Bianeoni, 1859, p. 277. No locality designated, luit 

Mozambique b.v inference. Type in Bologna Museum. 
Dasiipeltis scaher var. fasciolata Peters, 1868, p. 451. Type (ZMU 5737) 

from " Angeblich aus Zanzibar". 
Dasypeltis elongata Moequard, 1888, p. 131. Type locality: Zanzibar. Type 
in Museum d 'Histoire Naturelle, Paris. 
Diagnosis. Three to eight narrow Vs commencino- on the nape, 
followed by a series of narrow, lateral, dark red-brown bars. 
These bars wall encircle a pink to silvery-white vertebral dot, if 
they coalesce with their fellows from the opposite flank. In some 
specimens the posterior ])ands are situated betw^een more or less 
clearly expressed oval dorsal saddles, while various intermediate 
patterns (see Fig-. 2) have also been observed. The dorsum is a 
light reddish-brown, which shows a considerable amount of 
mottling under the binocular microscope. The apical scale pits, 
as well as those of the head region, are a dark brow-n, this pig- 
mentation being found on all scales and very sharply set off. 
The ventrum is a clear pink, more or less regularly stippled with 
grey. Beneath the tail of many specimens, particularly^ in 
males, this stippling is arranged in two to four lines. In other 
individuals, hoAvever, the distribution of the stippling appears 
to be haphazard. Three lateral scale rows are definitely reduced 
and inclined, and they, as well as some of the adjacent rows, 
generally have strongly serrated keels. The frontals shoAv pitting 
or pustulosity over their entire surface as do most of the other 
cephalic scales (see Fig. 1). The inter-prefrontal suture is not 



BREVIORA 



NO. 79 



depressed. Veiitrals in males 235-253, in females 237-259 ; eau- 
dals 82-109 and 71-80, respectively. 

Range. Coastal East Africa. Extreme southern Kenya, Tan- 
ganyika, northern and central INIozambiqnc ; inland to Xyasaland. 




Figure 2. Dasypeltis m. medici. Dorsal views at midbody showing color 
pattern variants of this race. Top. BM 12-1-30-6 from Matemo, Mozam- 
bique. Bottom. BM 97-6-9106 from between Nkata Bay and Ruarwe, 
Xyasaland. Note lighter middorsal spots between the saddles, the regular 
pigmentation of the apical scale pits, and the general speckling of the 
ground color. 



1 ;).■)( DASYFELTIS MEDICI LAMUENS18 5 

Disfribidio)! records. (Maj) 1 shows Die relation of these 
localities to eaeh other. Records from the literature are starred.) 
KENYA: Takannon. Between Voi and Ndi, Taita. TANGAN- 
YIKA: Zanzibar Coast. Kibonp:oto' Kilimanjaro ( Intergrade"? 
specimen^. Amani, Usamljara Mts. Usambara. Makindn Kiver. 
Morogoro. Nyange, lUugurn Mts. Northern Ukutu (Kuthu) 
Steppe. Tendaguru. Mikindani. Nchingidi. Liwale. ZANZI- 
BAR. JMAFIA. MOZAMBIQUE: Matemo. Ribaue, Nyassa 
Prov. NYA8ALAND: Between Nkata Bay and Rnarwe. Zomba. 
Cholo Mtn. 

DaSYPELTIS MEDICI LAMUENSIS Subsp. IIOV. 

Uasyiniiis pahnarum I'etors (not of Loach), 1878, p. 20G. 1 ex. Taita, 

Kenya (J. M. Hildebrandt) : ZMU. 
DasypcUis scahra var. F (part), Boulenger, 189-4, p. 356. 1 ex. Mt. Kili- 
manjaro, Tanganyika (F. J. Jackson) : BM. 
Dasyprltis scahrr Uthnioller (part), 1934, p. 113. 1 ex. nr. Goniberi, Kili- 
manjaro, Tanganyika (Uthmoller) : Zool. Staatssanimlung, Miinchen. 
DasypcUis scahcr Loveridge (part), 1936, p. 256. 1 ex. Mt. Mbololo; 2 ex. 

Lamu Island, Kenya (Loveridge) : MCZ. 
DasypcUis scahcr Scortecci (part), 1939, p. 276. 1 ex. Belet Aniin, Ital. 
Somaliland = Somalia (S. Patrizi) : Genoa Civ. Mus. Stated to be 
nnifonn grey dorsally. 
DasypcUis scahcr mcdici Loveridge (part), 1942, p. 283. Mention of uni- 
formly colored northeastern material : MCZ. 
Diagnosis. Uniform reddish-brown dorsally, fading to butf in 
some specimens after preservation (Loveridge). Pink ventrally 
with a fine speckle of a slightly darker pinkish-brown denser on 
the sides (Fig. 4, bottom). Some specimens are a uniform olive 
grey, fading to a plumbeous grey after preservation. These have 
a light grey ventrum, minutely flecked with white. A vague or 
distinct mottling of the ground color may be seen under the 
binocular microscope (see Fig. 4, top). Apical pits of body and 
marginal pits of head scales distinctly pigmented with a darker 
reddish-brown (see Fig. 1). With proper illumination faint to 
clear V-shaped markings can be discerned on the parietal scales 
of some specimens. Three lateral scale rows are distinctly reduced 
and inclined, and they, as well as several of the adjacent rows, 

1 The status of this specimen is discussed helow and its data are not included 
in the above ranges tor tliis subspecies. 



BREVIORA 



NO. 79 




Figure 3 



1957 i\\s\i'i;i/ris medici hAMUENSis 7 

generally have strongly serrated keels. Each frontal scale shows 
pnstulosity or pitting aroniul tlie ])(M'ipluM'v and in the center. 
The suture lietwcen the pi'efroutals is not depressed. V^nitrals 
in males 226-22!), in females 22G-232; caudals 84-94 and 72-84, 
respectively. 

Holoti/pc. jMuseuni of Comparative Zoology No. 40582, an 
adult male from Lanui Island, Ken\a, collected by Arthur Lovei-- 
idge 12 May 1934. 

Allotype. Museum of Comparative Zoology No. 40588, an 
adult female collected with the type. 

Paratypes. British Museum (Natural History) Nos. 51-1-3-72 
and 51-1-3-73 from Kilifi, north of Mombasa, Kenya. Also Brit- 
ish Museum (Natural History) No. 98-1-8-15 from Maungu, 
near Voi, Kenya ; iMuseum of Comparative Zoology No. 40580 
from Mt. Mbololo, Kenya ; Zoologisches Museum der Universitat 
(Berlin) No. 9244 from Taita (region), Kenya; and British 
Museum (Natural History) No. 87-11-3-31 from Mt. Kilimanjaro, 
Tanganyika. 

Description. Both the Lamu holotype and allotype are a uni- 
form brownish-red dorsall}^ and a speckled pink on the venter. 
Under proper illumination a very faint, posteriorly-directed V 
may be discerned on the parietal scales (see Fig. 3, top). Scale 
pits on head and body with sharply defined darker brown pig- 
mentation, though this is only slightly darker than the dorsal 
color (Fig. 3, bottom) and not as clearly visible as on lighter 
specimens (Fig. 4, middle). The frontals are pitted around 
their margins and pustules are also present in their centers, with 
the other head shields showing a similar pattern. The suture 
between the prefrontals is not depressed, the ocular-temporal 
formula is l+2-(-2-|-3, and the upper labials and eye contact are 
7(34). Counts of ventrals are 226 for both type and allotype, 
caudals are 94 and 84, dorsals 23 and 24 at midbody respectively. 
The third, fourth and fifth rows of dorsal scales from each side 
are reduced, inclined and serrated, and the keels of the dorsal 



Figure 3. Dasypellis m. hnniiensis. Views of the holotype (MCZ 40582). 
Top. Dorsal view of head. Middle. Lateral view of head. Bottom. Dorso- 
lateral view of specimen in the midbody region. Note the very faint pattern 
on the parietals, and the fact that the prefrontals, oculars, anterior labials, 
etc. are covered with pigmented pits. 



8 



BREVIORA 



NO. 79 







Figiiie 4 



1957 DASYPELTIS MEDICI LAMUENSIS 9 

scales are serrated around the body in the anal ref^ion of the 
type, hut not of the pai-atype. Body and tail lengths are 498+ 
132 nun. and 570-|-128 nnn. respectively . 

Range. Coastal East Africa from Somalia soutli to the Kenya- 
Tanganyika frontier. 

Disiribufion records. SOMALIA: Belet Amin (Scortecci, 
hm). KENYA: Lamu Island (Loveridge, WSd) ; MCZ 40582, 
40583. Kilifi, north of Mombasa. BM 51-1-3-72, 51-1-3-73. 
Mauno-u near Voi. BM 98-1-8-15. Mt. Mbololo (Loveridge, 
1936); MCZ 40580. Taita (Peters, 1878); ZMU 9244. TAN- 
GANYIKA : Mt. Kilimanjaro (Bouleno-er, 1894) ; BM 87-11-3-31. 
Near Gomberi, Mt. Kilimanjaro (Uthmoller, 1934). 

Discussio)t. The map shows the relative position of collectino' 
localities for both patterned and unicolored forms of D. medici. 
The species appears to be restricted to the coastal regions below 
1000 meters, generally characterized by their reddish laterite 
soils (Loveridge, 1942) . As may be seen from the map, all but two 
of the Kenya specimens are definitely unicolored. Pattern is 
present in the Voi-Ndi juvenile, but the Takaungu individual is 
an almost completely faded adult, which lacks even the pigmenta- 
tion of the apical pits so characteristic of the species. Two of the 
three specimens from the vicinity of Mt. Kilimanjaro (seen, or 
reported on in the literature) were unicolored, while the dorsal 
pattern of the third is faded out posteriorly. All other speci- 
mens and records from Tanganyika have, or are said to have, a 
color pattern. 

Of the other characters examined only ventral counts show 
elinal variation. Figure 5 demonstrates these counts for male and 
female specimens of the two color phases, showing the sharply 



Figure 4. Dasypeltis ?h. lamucnsis. Views of the midbody region of 
different specimens to demonstrate certain characteristics of the pigmenta- 
tion. Top. BM 51-1-3-73 from Kilifi, Kenya. Lateral view of dark grey 
specimen, showing the extensive speckling of the ground color, which almost 
masks the pigmentation of the apical pits. Middle. BM 98-1 8-15 from 
Maungu, Kenya. Lateral view of light colored specimen which clearly 
shows both pigmentation of apical pits and lateral scale arrangement. Note 
that the ground color is still speckled. Bottom. BM 51-1-3-72 from Kilifi, 
Kenya. Ventrolateral view of specimen showing color invasion and speckling 
of light colored ventrum. 



10 



BREVIORA 



xo. 7!) 




LAMU ISLAND 



MT. KILIMANJARO Q^ CY^ <^ 

I ® Okilifi 
VaitaO Y"^ka"ngu 



USAMBARA ^ 
AMANI 

MAKLNDU^ 

MOROGORO 

NYANGE0 
UKUTU* 




ZANZIBAR ISLAND 



MAFIA ISLAND 



KILWA ( 
TENDAGURU( 

LI WALE # 

NCHINGIDI V •MIKINDANI 




Jjasi/pt'ltis medici. Map showing loc-alities for all specimens acliiaUy 
examine.! as well as for the literature record from Belet Amin. Solid dots 
stand for records of m. medici, open circles for records of m. lamuoi.si.s. 
Divided circles refer to the paiis of immediately adjacent records discussed 
in the text. 



195'; 



DASYPELTIS MEDICI LAMUENSIS 



11 



distinct ranfjcs. The only exception is furnished by the faintly 
patterned specimen from Kibonjroto' Kilimanjaro. This has a 
ventral count of 218, i.e. 8 less than the lowest count for a 
unicolored specimen and 19 less than the lowest count for a 
patterned specimen. 

In view of the fact that Ihe two populations replace each other 
jreographically, that they ditfer in two characters which break in 
the same zone, and that there exists a possibly intermediate speci- 
men from a geographically intermediate region, they are here 
considered to be subspecifically distinct. This decision has been 

MALES 



lamuensis 



me 



FEMALES 




lamuensis 



medic 



220 230 240 260 

V E N T R A L S 



Figure 5. Dasypcltis vtcdici. Graph showing ventral counts of all speci- 
mens of the two races actually examined. Note the anomalous count of 
the Kibongoto specimen. 



12 BREVIORA NO. 79 

consciously influenced by a desire to focus attention upon this 
population in the hope that this will permit re-examination of 
the matter on the basis of more nearly adequate collections. 

It has been suf>'0'ested that this situation represents two adja- 
cent species and that the patterned Kilimanjaro specimen is a 
hybrid. While the material is insufficient to decide the point, 
there is some evidence against this idea. Thus both of these 
color patterns feature a sharply defined pigmentation of the 
apical scale pits and are the only forms within the genus that 
have this characteristic. Besides which, 45 out of 46 specimens of 
the combined sample have frontals that are entirely, rather than 
marginally, pustulated or pitted, a frequency not approached 
by any other population, with the exception of Dasypcltis scahra 
from South Africa. In view of these facts and in the absence of 
more detailed data, the two forms are considered to belong to a 
single polytypic species. 

LITERATURE CITED 

BiANCONi, J. Joseph 

1850- Specimma Zoologica Mosambicana. Bononiae, fasc. 1-1(5, ii 
1862. +406(?) pp., (fasc. 14 (1859), pp. 277-278, tab. 14). First 

published in parts in Mem. R. Accad. Sci. Inst. Bologna. (Vol. 

10, pp. 501-2, pi. 26.) 

BOULENGEK, GEORGE ALBERT 

1894. Catalogue of the snakes in the British Museum (Natural His- 
tory). Volume II, containing the conclusion of the Colubridae 
Aglyphae. London, xi -[-382 pp. (353-357). 

LovERiDGE, Arthur 

1936. Scientific results of an expedition to rain forest regions in 

Eastern Africa. V. Reptiles. Bull. Mus. Conip. Zool., vol. 79, 

no. 5, pp. 209-337 (256-57, pi. 4, fig. 1). 
1942. Scientific results of a fourth expedition to forested areas in 

East and Central Africa. IV. Reptiles. P>ull. Mus. Com]). Zool., 

vol. 91, no. 4, pp. 237-373 (282-86). 

MocQu.vRD, Francois 

1888. Sur une collection de reptiles et de batraciens rapportes des 
Pays Comalis et de Zanzibar par M. G. Revoil. Mem. pui)!. Soc. 
Philomat. Occas. Centcnnaire Foundation, i)p. 109-134 (131, 
plate XII, figs. 2, 2a, 2b, 2e). 



1957 DASYPELTIS MEDICI LAMUENSIS 13 

Peters, Wilhelm Carl Hartwig 

1868. liber eine neuc Nagergattung, Chiropoilowi/.s prninllatus, so 

wie iiber ciiiigo neuc odcr weniger bekannto Ainphibien und 

Fisehe. Monatsbcr. kgl. Akad. Wiss. Berlin, pp. 448-461 (451). 
1878. Ubcr die von Hrn. J. M. Hildebrandt wiihrend seiner letzten 

ostafrikaniscluMi Reise gesanimelten Saugetliiere und Aniphibien. 

Mouatsber. konig. Preuss. Akad. Wiss. Berlin 1878, pp. 194-209 

(206). 

ScoKTEoci, Giuseppe 

1939. Spedizione zoologica del Marchese Saverio Patrizi nel Basso 
Giuba e nell 'Oltregiuba. Giugno-Agosto 1934 XII. Eettili. 
Ofidi. Ann. Mus. Civ. Storia Nat. Genova, vol. 58, pp. 263-291 

(276). 

Uthmoller, Wolfgang 

1934. Beitrag zur Kenntniss der Schlangenfauna des Kilimandscharo. 
(Tanganyika Territory, ehenialiges Deutsch-Ostafrika). Teni 
niinckia, Leiden, vol. 2, pp. 97-134 (113-115). 



BREVIORA 

Mmseiuiinti of Comnpsirative Zoology 



Cambridge, Mass. September 30, 1957 Number 80 



A COLLECTION OF DRAWINGS OF FISHES ASCRIBED 

TO J. P. KIRTLAND (1793-1877), 

IN THE LIBRARY OF BOWDOIN COLLEGE 

By James M. Moulton 

Department of Biology 

Bowdoin College, Brunswick, Maine 

During- the latter part of 1956, a pamphlet box removed from 
the shelves of the Bowdoin College Library was found to contain 
61 uncatalogued drawings of fishes, nearly all bearing the sig- 
nature 'M. P. Kirtland, del." Of the drawing^s, all of which are 
in good condition, 57 appear to be originals of various of the 
illustrations accompanying a series of articles by Jared Potter 
Kirtland on the fishes of Ohio, which appeared in the Boston 
Journal of Natural History from 1840 to 1846. Most of the draw- 
ings are in pencil ; a few are finished in watereolors ; all appear 
to have been drawn by the same hand. Of the 69 figures accom- 
panying Kirtland 's articles, corresponding: drawings are lacking 
for but 11 which constituted 4 of the 26 plates and 2 of the 3 
figures of a fifth plate. Various bits of evidence indicate that 
these drawings were executed b}^ Kirtland for his article. How 
might they have come to Bowdoin College? Probably through 
Dr. D. H.' Storer. 

David Humphreys Storer (1804-1891) and Jaretl Potter Kirt- 
land had much in common : New Englanders by birth, educated 
for the medical profession, eminent teachers and practitioners of 
medicine, among the founders of medical schools and active civic 
leaders, prolific authors in diverse fields, both had a lifelong in- 
terest in natural history and especially in ichthyology. Storer 
was a native of ]\Iaine and a graduate in 1822 of Bowdoin College ; 



2 BREVIORA No. 80 

he moved to Boston to begin his medical education at Harvard 
I^niversity where he later taught for many rears. Kirtland moved 
to Ohio in 1823 from New Haven where he had graduated from 
Yale Medical School in 1815. 

Gehr's (1952) biography of Kirtland states that he was a 
member of the first class to matriculate in the medical school of 
Yale University, and that he pursued a long career as a physician 
and teacher of medicine. He was a founder of, and a proponent 
of legislative action which established the medical school of 
AVestern Reserve ITniversity. He was for most of his life a con- 
tributing student of natural history — of mollusks, insects, fishes 
and birds. He was the author of man}' j)ublications and articles 
on subjects as diverse as medicine and horticulture. He was a 
correspondent of Professor Louis Agassiz and collected Ohio 
fishes for him. He Avas a frequent lecturer in many areas of 
knowledge, and included the duties of an editorship among his 
other responsibilities. He was elected a member of the American 
Philos()i)hical Society at the age of 82. Legislator, teacher, physi- 
cian, biologist, noted naturalist, Jared Potter Kirtland led a full 
and most productive life. 

Indicative of their stature as naturalists is the fact that Kirt- 
land and Storer were each appointed in 18.'57 by the government 
(jf their respective states (Storer by iMassachusetts and Kirtland 
by Ohio) to assume an important role in natural history surveys 
of those states. Storer 's report on the fishes and reptiles of Mas- 
sachusetts was published in 1839 by the Commissioners on the 
Zoological and Botanical Survey of the State. Of his report, 
Storer wrote to Nehemiah Cleaveland of Bowdoin College prob- 
abl}' sometime between 1855 and 1867, in a letter preservetl by 
the College, "In 1837, Commissioners were appointed l\v the 
Governor and Council to prosecute the Botanical and Zoological 
Survey of the Commonwealth. The departments of ichthyology 
and herpetology were entrusted to me. I concluded my report in 
1839 ... In 1842, Dr. DeKay published his 'Zoology of the State 
of NeAV York. ' You will excuse me for copying from that report 
the following: 'In 1839, Dr. D. II. Storer, who had previously 
furnished several valuable papers on ichthyology in tbe Boston 
•Journal of Natural History, published a nuisterly report on the 
fishes of Massachusetts. In this rejioi-t, the author has enumerated 
one hundred and nine species, of whicli one liundi-ed and four 



1!I57 KIRTI.AXn's DKAWING!^ OF FISHES 3 

arc accompanied by oi-igiiial aiici carefully drawn up descriptions. 
This report is an invaluable document to the American ichthyolo- 
irist and in every way worthy of its eminent author." 

The Ohio survey was abandoned in 1839, after submission of 
the second annual report to the legislature. This report, which 
contained Kirt land's descriptions of the fishes of Ohio, was not 
l)ublished by the legislature. 

Storer had been an active member of the Boston Society of 
Natural History for nearly a decade when the Journal of that 
Society began in 1840 the publication of Kirtland's (1840, 1841, 
1842a, b, 1843. 184rm, b, 1846) descriptions of Ohio fishes. It was 
perhaps during the period of publication from 1840 to 1846 that 
Kirtland and Storer became acquainted ; their friendship is indi- 
cated by Kirtland's (1845a, p. 82) description of Lenciscus 
Storerianus: "It atfords me pleasure to dedicate it to my friend, 
D. Humphreys Storer, M. D., to whom I am indebted for essen- 
tial aid, and many important suggestions, in arranging and de- 
scribing our western fishes." 

Storer 's connection witii the Boston Society of Natural His- 
tory, and his own interest in fishes, would certainly have made 
him cognizant of Kirtland's work on the fishes of Ohio, and pos- 
sibly it was through an earlier acquaintance with Storer that 
Kirtland decided to publish in the Journal of the Society. Kirt- 
land's figures would presumably have accompanied his manu- 
scripts to Boston, where Storer might well have received them, 
after publication, as a gift from Dr. Kirtland. (ielir (1952, p. 17) 
indicates that Kirtland's figures for the Ohio report were drawn 
by himself, and indeed that he learned to draw for the express 
purpose of illustrating his work on fishes. 

Dr. Storer maintained a lifelong interest in Bowdoin College. 
As late as the year of his death in 1891 he contributed money and 
pamphlets to the College, and at various times submitted infor- 
mation requested for alumni records of the College. After his 
death in 1891, his daughter sent to the College several items in the 
form of books, journals and miscellaneous pamphlets which had 
been in her father's library. The box in which the drawings w^ere 
found contained several pamphlets inscribed to Dr. Storer by 
various authors. It is not improbable that the drawings executed 
by Kirtland came to Bowdoin College with material donated after 
Dr. Storer 's death. 



4 BREVIORA No. 80 

111 view of their historical significance, I liave felt it worth- 
while to inform ichthyologists of the location of drawings which 
I ascribe to Dr. Jared Potter Kirtland, early describer of the 
fishes of Ohio. 



REFEBENCES CITED 

Gehr, a. R. 

1952. Jared Potter Kirtland. The Explorer, 2. No. 7: 1-33. 

Kirtland, J. P. 

1840. Descriptions of the fishes of the Ohio River and its tributaries. 
Boston Jour. Nat. Hist., 3: 338-352. 

1841. [Same title.] Boston Jour. Nat. Hist., 3: 469-482. 
1842a. [Same title.] Boston Jour. Nat. Hist., 4: 16-26. 

1842b. Descriptions of the fishes of Lake Erie, the Ohio River, and their 
tributaries. Boston Jour. Nat. Hist., 4: 231-240. 

1843. Descriptions of the fishes of the Ohio River and its tributaries. 
Boston Jour. Nat. Hist., 4: 303-308. 

1845a. [Same title.] Boston Jour. Nat. Hist., 5: 21-32. 

1845b. Descriptions of the fishes of Lake Erie, the Ohio River, and their 
tributaries. Boston Jour. Nat. Hist., 5: 265-276. 

1846. [Same title.] Boston Jour. Nat. Hist., 5: 330-344. 



FIGURES 




Figure 1. Probably Acipenser fulvescens Eafinesque, named A. nudus by 
Kirtland due to loss of the bony plates with age. The pencilled signature on 
this figure, lacking in the Boston Society article, was retouched by the 
l)hotographer. 






.'^?HHBt?^Tr^fT^Tr^.'^'*.-Arr*<; '.•-■rjr-:-.'i^-f*^:^c^';^-''>Tr^r'?W.V/" 



<»■ v.-;^s'w ■>"!r-u'~.--';'-rar'ici>*">r"> ' 







//».,., ,/ / 



HraosKal 



Figure 2. A second figure of Acipenser fuJvescens Eafinesque, designated 
by Kirtland Accipenser mncrostonius Eafinesque. This figure, also unpub- 
lished in the Boston Society article, was signed in ink. 







w 



■■-''''^"-'^4^1 



\ 



^ 

^*<*«;' 



ipft»i„y iUlm 



'■' //■! r''r/ /■/ 



Figure 3. Kirtland 's drawing of Pomoxis annularis Eafinesque, appar- 
ently intended to be named after Storer, CichJa Sioreria. The signature is 
printed, unlike other signatures in the collection. The drawing was not in- 
cluded in the article of the Boston Society of Natural History. 



BREVIORA 

MuseiLiioi of Comparative Zoology 



Cambridge, Mass. September 30, 1957 Number 81 

CONTKIBUTIONS TO A REVISION OF THE EARTH- 
WORM FAMILY LUMBRICIDAE. I. ALLOLOBOPHORA 

LIMICOLA 

By G. E. Gates 

INTRODUCTION 

External characteristics and the internal anatomy of many 
earthworm species are known only from the original descriptions, 
all too often based on a single specimen. Hence little information 
as to variation, even in common forms, is available. Such records 
as have been provided usually are for the genital system. More 
necessary at present are data for somatic systems that are now 
being found to be phylogenetically more conservative. 

This contribution is the first of a series that will provide, inso- 
far as circumstances permit and from lumbricid material pre- 
served in the field without benefit of special techniques, informa- 
tion as to characters used in the obsolescent taxonomy of the 
past as well as those that may be employed in the more natural 
classification of the future. 

The author's thanks are extended to Harold Davies for pro- 
viding three of his New Jersey specimens. 

Family LUMBRICIDAE 

Genus AlLOLOBOPHORA Eisen 1874 
Allolobophora LIMICOLA Michaclseu, 1890 

Allolobophora limicola Michaelsen, 1890, Jahrb. Hamburg. Wiss. Anst., 7, 
p. 10 (Type locality, Eolfshagen, Hamburg, Germany. Types probably 
in the Hamburg Museum.) 



2 BREVIORA XO. 81 

Allolobophora {Allolobophora) limicola, Bosa, 1893, Mem. Ac. Sci. Torino, 

43, pp. 424, 450. 
Allolohoplwra limicola, Beddard, 1895, Monograph, p. 716. 
Allolobophora limicola, Bretscher, 1896, Rev. Suisse Zool., 3. p. 52."^. (Swit- 
zerland) 
Helodrilus (Allolobophora) limicola, Michaelsen, 1900, Das Tierreich, 10, p. 

484. 
Ilelodrilus (Allolobophora) limicola, Piguet & Bretscher, 1913, Cat. Inv. 

Suisse, 7, p. 177. 
Allolobophora limicola, Ude, 1929, Tierwelt Deutschlands, 15 (1), p. 120. 
Allolobophora limicola, Michaelsen, 1931, Bull. Mus. Hist. Nat. Belg., 7 (1), 

p. 6. (Belgium?) 
Allolobophora limicola, Cernosvitov, 1936, Bull. Mus. Hist. Nat. Belg., 12 

(21),pp. 5, 12, figs. 8-9. (Belgium) 
Allolobophora limicola, Tetry, 1940, Bull. Mus. Hist. Nat. Belg., 16 (31), 

p. 21. (Belgium) 
Allolobophora limicola, Wilcke, 1949, Senckenbergiana, 30, p. 178. 
Allolobophora limicola. Gates, 1953, Bull. Mus. Comp. Zool. Harvard, 107, 

p. 518 ; Breviora, 15, p. 2. (Massachusetts) 
Allolobophora limicola, Graff, 1953, Zool. Anz., 151, p. 26. Regenwiirmer 

Deutschlands, pp. 26, 60, fig. p. 61. (Germany) 
Allolobophora limicola, Davies, 1954, Breviora, 26, p. 3. (New Jersey) 
Allolobophora limicola, Satchell, 1955, Ann. Mag. Nat. Hist., (12) 8, p. 224. 

(England) 

Material examined 
Massachusetts 
Boston. Arnold Arboretum, peat bog. May 18, 1956, 1-10-62-1. 

New Jersey 
(Three specimens supplied by Harold Davies. Tide, Davies, 1954, p. 3.) 

External characteristics. Segments (c/. Table 1), of worms 
that are not obviously recent amputees, 83-136. 

Setae begin on ii where all usually are present. \'entral 
setae larger, slightly- more protuberant than lateral ones, more 
closely paired on xv. Apertures of follicles are very slightly 
farther apart than elsewhere in the genital tumescences. Inter- 
setal intervals varj^ in absolute width from one region of the body 
to another as well as on the same segment in individuals of dif- 
ferent sizes. Relative widths of the intervals are as previously 
indicated (Gates, 1953). 



1957 ALLOLOBOPHORA LIMICOLA 

Table 1 





Segment 


number 


in A 


. limicola 






Number 


af specimens 


from 




Segment 


Mass. 












Number 


1956 1952 


(b) 




N. J. (a) 


Totali 


83 


1 










1 


95 


1 










1 


98 




1 








1 


104 










2 


2 


105 


1 










1 


107 




1 








1 


108 


3» 










3 


110 


1 










1 


111 










1 


1 


112 


1 








1 


2 


113 


2 


1 






1 


4 


114 










3 


3 


115 


2* 










2 


117 












2 


118 





2 








4 


119 


1* 










1 


120 


3 


2 








5 


121 


4 








o 


6 


122 


4 


1 






1 


6 


123 


8 


5 






o 


15 


124 


8 


3 






1 


12 


12.- 


6 


1 








i 


126 


2 


1 








3 


127 


1 


1 








2 


128 


4 










4 


129 




1 






1 


2 


136 


1 










1 



(a) Cited from Da vies (1954) 

(b) Cited from Gates (1953) 

Nephropores usually closed and not certainly recognizable 
even after softening, treatment with picric acid, or use of other 
techniques, but probably so situated as to permit the following 
characterization. Nephropores alternating irregularly, with fre- 
quent asymmetry, between two levels, one well above D, the 
other somewhat above B, pores of xiv usually in DD, of xv-xvi 
usually at the lower level. 

* One of the worms, in each case, may be a posterior amputee. 



4 BREVIORA NO. 81 

Clefts in male tumescences, though obvious, go down only 
slightly, if at all, below general level of outer surface of the body. 

The clitellum always covers xxx-xxxv, occasionally extending 
into xxix (sometimes to the equator but never in front of it), 
and/or into xxxvi or even to 36/37. The clitellum reaches 
slightly farther posteriorly, in variant specimens, on that side 
which has the longer tuberculum. Tubercula pubertatis have a 
distinct margin, after appearance of clitellum, only on the me- 
dian side and occasionally are unrecognizable even under optimal 
illumination. The median margin is concave or incised at 33/34 
(also at 34/35, 35/36, in variant specimens). Tubercula usually 
are in xxxiii-xxxiv (57 specimens) but on some of the variant 
worms are also in xxxv as follows .- on the left side (5), and right 
side (2), on both sides (8). Locations in the remaining specimens 
are : on left sides of xxxiii-xxxv and right sides of xxxiii-xxxvi 
(1), on left sides of xxxiv-xxxvi and right sides of xxxiii-xxxv 
(1), in xxxiv-xxxvi on both sides (2). Incidence of worms with 
variant tubercula in previous lots (of 129 aclitellate and clitel- 
late specimens), 14.7 per cent; in present lots (of 73 specimens), 
26 per cent. 

Genital tumescences (cf. Table 2) may be present on any of 
.segments ix-xiii, xvi-xvii, xxvi-xxxii, xxxv-xxxviii. Locations, be- 
hind XV, on the 19 variant specimens (as regat"ds tubercula), are 
as follows: on xxix-xxxii, xxxv-xxxvi (1), on xxix, xxxi-xxxii, 
xxxv-xxxvii (1), xxx-xxxii, xxxv (3), xxx-xxxii, xxxv-xxxvi (2), 
xxx-xxxii, xxxv-xxxvii (2), xxx-xxxii, xxxvi (2), xxx, xxxii. 
xxxv (2), xxxi-xxxii, xxxv (2), xxxi-xxxii, xxxv-xxxvi (2). 
xxxii, xxxv (2). 

Internal anatomy. Septa 6/7-9/10 muscular, 5/6 and 10/11- 
11/12 or 12/13 slightly muscular. Pigment unrecognizable in 
longitudinal muscle strip at niD and in sections of body wall, 
also in circular muscles after stripping off longitudinal layer. 
Epidermis thickened two to three times in male tumescences, 
less so in genital tumescences. 

Oesophagus widest in x, narrowing gradually through xiv, 
constricted fairly deeply at in.sertions of 10/11-11/12 so that 
portions in xi and xii appear subspheroidal. Additional but 
slighter constrictions, at 12/13-13/14, occasionally are recogniz- 
able. Lumen verticallv slit-like in xi-xii or widened in middle of 



1957 



ALIiOLOBOPIIORA LIMICOLA 



xii (internal constriction of gut wall). An internal constriction 
in xii usually is confined to one side but in xiii usually is on both 
sides. Oesophageal wall much thinner in xiii-xiv than in xi-xii, 
calciferous lamellae recognizable only in xii-xi and the posterior 
half of x. In x the dorsal blood vessel at first appears to be within 
a groove in dorsal face of gut but this appearance results from 
presence of two rounded protuberances (77 specimens) which, 
with a similar pair on the ventral face, provide the only evi- 
dence, in these strongly contracted worms, for existence of calci- 




Fig. 1. Oesophagus in segments x-xiv, dorsal view. 

Fig. 2. Oesophagus in segments x-xiv, horizontal section. 

ferous sacs. Intestinal origin in xv (77). Gizzard in xvii (77), 
the layer with brilliant muscular sheen in horizontal section 
abruptly narroAving and ending shortly behind insertion of 
17/18. Posteriorly the gut wall is much thinner, weaker, breaking 
most easily in both wet and dry material just behind 17/18. 
Gut narrowing slightly through xix, rather valvular in region 
of insertion of 19/20. The typhlosole, shortly after its beginning. 



BREVIORA 



NO. 81 



is nearly circular in transverse section and deeply grooved on 
each side clear to level of the small median ridge on the ventral 
face. The grooves appear to be exactly opposite each other (not 
alternate) and produce, on each side, a series of laterally di- 
rected, haU'moon-shaped leaflets. Superficially, as previously 
suggested, the appearance suggests a stack of coins with a cord 
running along it. Behind the clitellum the lateral grooving 
gradually disappears and the typhlosole becomes more nearly 
thickly laraelliform. The grooving is typical, characterizing 
each of the 77 worms of the 1956 series. The typhlosole ends in 
front of the 119th segment as shown in Table 3. 

Dorsal and ventral blood vessels have not been traceable an- 
terior to 5/6, probably because they are empty. Extra-oesopha- 
geals, median to the hearts and in contact with the gut only in 





mftmmim^fimmifammm^mwm 




Fig. 3. Typhlosole. Anterior end from ventral side. 

Fig. 4. Typhlosole. Horizontal section slightly posterior to level shown 
in Figure 3. 

regions of some septal insertions, pass up to the dorsal trunk in 
xii (77). Hearts present in vi-xi (35), usually slenderer in vi 
and there often empty or nearly so. Inability to see hearts in 
vi of the other 42 worms proljably not due to absence but to 
difficulty of distinguishing the vessels, when empty, from 
nephridial and other tissues with which they are associated. 
Last hearts in xi (77). Nephridia, behind xiv and except in cli- 
tellar region, large, back from xix reaching up to dorsal blood 
vessel. A bladder is present but its condition (not improved by 
various attempts to render nephropores visible) obviates accurate 
characterization, possibly XT-shaped. The supposed duet, slender 
and very fragile, always seems to pass into parietes close to B 



1957 ALLOLOBOPHORA LIMICOLA 7 

Brain, left by a transverse section exactly alonj? 3/4, in iii. 
Usually all of each circumpharyngeal commissure also is left in 
iii, at most only very short stubs remain in iv. 

Holandric (77). Seminal vesicles, four pairs, in ix-xii (77). 
Male deferent ducts, just behind funnel septum, variously looped, 
the loops (epididymis) or all but one of them bound together in 
a disc (77) often of about the same size as a testis. Ducts of a 
side come into contact posteriorly in xii but may not unite until 
into xiv. They disappear into longitudinal musculature about 
at eq/xv. 

Ovaries, each with a single distal egg-string containing one to 
three ova, erect discs with shape approximating to vertically 
oblong. Oviducts pass under a muscle inserted on h follicle of 
xiv and then into parietes just lateral to B. Ovisacs always 
present (77) and in the usual positions on posterior face of 
13/14. Spermathecae, two pairs, always in ix-x (77) and open- 
ing to exterior posteriorly. 

Atrial, TP and supraparietal GS glands are lacking in the 
coelom (77) and no special glands are recognizable (in unstained 
free-hand sections) Mdthin the muscular layers. Follicles of 
ventral setae, from xv into postclitellar region at least, with two 
exceptions, much larger and more protuberant into the coelom 
than those of the lateral pair in the same segment. Exceptions 
are provided by segments xxxiii-xxxiv where the ventral follicles 
usually appear to be little if any larger than the lateral ones. 
Ventral follicles of xv usually appear to be little if any larger 
than in xvi. Follicles opening through genital tumescences al- 
ways (77 specimens) are especially enlarged and unusually pro- 
tuberant into coelomic cavities. Setae in all examined follicles 
have been genital (GS) and may be 1+ mm. long. 

Reproduction. Spermatophores were not found on any of the 
1956 series. Iridescence of male funnels more or less brilliant 
in each worm except the juvenile and one of the aclitellates, 
localized in a few small spots in several aclitellates, more general 
in others and most l)rilliant in those worms with clitellar tumes- 
cence nearer the maximum (which probably had not been 
reached by any of the present lot) . Iridescence in the contents of 
some or all of the spermathecae (both spermathecae of a side 
empty in several worms) of 26 clitellate specimens shows that 
sperm are present there too and hence that copulation had taken 



8 BREVIORA NO. 81 

place. As sperm had been, or were being, matured so generally, 
and as they are exchanged during copulation, reproduction can 
be assumed to be biparental. 

Regeneration. A tail regenerate at 78/79 has ca. seven seg- 
ments but the metamerism is quite abnormal. 

Ahnormality. No. 1. Extra male tumescences, with equatorial 
cleft, present on left side of xvi. No pore is recognizable in that 
cleft and both male gonoducts of the left side apparently pass 
down through the muscular layers in xv. 

No. 2. Left anterior spermatheca bifid down to parietes. Both 
ampullae preseptal, filled with sperm. 

No. 3. Eight segments in front of anal region are involved in 
a complicated spiral metamerism. 

Nos. 4-6. One helicometamere, in the postclitellar region of 
each of two worms, was observed while counting segments. Two 
spirals were noticed in the same region of another worm. 

Table 2 
Genital tumescences in Massachusetts 
collections of A. limicola 
Number of specimens in 



In 

segment 

9 


previous 
lots 
100 


present 
lot 
74 


Totals 
174 


10 




5 


5 


11 


97 


68 


165 


12 


112 


74 


186 


13 




3 


3 


16 




2 


2 


17 




1 


1 


26 




1 


1 


27 




2 


2 


28 




6 


6 


29 


31 


23 


54 


30 


88 


58 


141 


31 


81 


51 


132 


32 


118 


72 


190 


35 


98 


71 


169 


36 


39 


25 


64 


37 


18 


6 


24 


38 




1 


1 


I'sent lot is from the very 


same spot at which a majoritj- of the 



previous specimens were obtained. 



1957 ALLOLOBOPHORA LIMICOLA 

Table 3 

Typhlosole termination and segment number in 

American collections of A. limicola. 



1- 
<v 

.2 = 

u a 
v 

m 


Typhlosole 
ends in 
segment 


Pj IS 

< 


Number of 
segments 




1 


76 


7 


83 


S 


2 


85 


10 


95 


S 


3 


86 


19 


105 


S 


4 


94 


14 


108 


S 


5 


95 


18 


113 


V 


6 


96 


12 


108 


V 


7 


96 


14 


110 




8 


99 


16 


115 




9 


100 


8 


108 


V 


10 


102 


19 


121 


V 


11 


103 


18 


121 


V 


12 


103 


19 


122 


V 


13 


103 


21 


124 


V 


14 


104 


14 


118 




15 


104 


16 


120 




16-17 


104 


18 


122 


V(l) 


18 


104 


19 


123 


s 


19 


104 


21 


125 




20 


105 


13 


118 


s 


21 


105 


17 


122 




22-24 


105 


18 


123 


S(l) 


25 


105 


19 


124 


s 


26 


105 


20 


125 




27 


106 


13 


119 


V 


28 


106 


17 


123 




29 


106 


18 


124 


V 


30 


106 


19 


125 




31-32 


107 


17 


124 


V(l) 


33-34 


108 


17 


125 


V(l) 


35 


108 


20 


128 




36 


110 


18 


128 




37 


118 


18 


136 





Old posterior amputee? 



New Jersey 



Old posterior amputee! 



Other from New Jersey. 



S Small specimen, not obviously a recent posterior amputee. Nos. 1, 2, 9, may 

be old posterior amputees, also No. 26. 
V Variant specimens, i.e.. with tubercula pubertatis extending beyond xxxiii- 

xxxiv. 
(1) One specimen only. 

Except as otherwise indicated all specimens are from the same spot in the 
Arnold Arboretum. 



10 BREVIORA NO. 81 

Table 4 

Individual patterns of genital-tumescence 

location in 1956 series of 

A. limicola. 

Other Number of 

Segments segments specimeng 

9 10 11 12 29 30 31 32 35 36 37 38 

»««•»« «** 1 

 *■)♦****** Q 

»»•*«** 1 

«««#»•♦* T 

»  » * » « 26 1 

»*»**««»* jg 2 

* ******* 26-28 1 
» •>»«*«»* ^7-28 1 

» « « * s » « 90 1 

******** A 

********* A 

****** 1 

»»«»»*» 1 

»*«-»*«* 9 

****** 1 

»♦»«*««* 2 

«»»»»♦**« 2 

» •» * » 2 

»»«**■» J 

»«♦»«»* 27 

««***•»■»» 5 

»*«*»*««• ^ 

*«*»»» 2 

*»*»•♦>■ 2 

«»»•■»* 2 

»»*«»»* 2 

* « « > « g 

* * « ^ « * » * 13, 17, 27, 28 
******* 23 28 

» »•» **** TO 

» » » » * oo 



 Genital tumescences present In the segment thus indicated, but disregarding 
asymmetry, i.e., absence on one side. 



1957 ALLOLOBOPHORA LIMICOLA 11 

liemarks. A dark coloration with a bluish metallic lustre in the 
body wall of the New Jersey worms is an artifact that results, 
in appropriate conditions, when formalin around the worms has 
remained for some time in contact with cork. 

Precis. Clitellum, saddle-shaped, on 5i^-8i/2 segments, xxviii/2, 
xxix, xxix/n, xxx, xxx/2-xxxv, xxxvi/n, xxxvi. Tubercula puber- 
tatis with median margin concave or incised at 33/34, in xxxiii- 
xxxiv (xxxv, xxxvi), rarely lacking on xxxiii. Genital tumes- 
cences around apertures of A'entral follicles only, usually in ix, 
xi, xii, xxx-xxxii, xxxv, occasionally in one or more of x, xiii, 
xvi-xvii, xxvi-xxix, xxxvi-xxxviii. Male pores equatorial on xv. 
in BC, within transverse clefts in tumescences that obliterate 
14/15-15/16. Female pores just lateral to B, equatorial on xiv. 
Quadrithecal, pores at C, on 9/10-10/11. Setae, closely paired, 
more so laterally, ventral setae larger ( ?), CD < AB, BC <! AA, 
DD ea. = y^C Nephropores inconspicuous, irregularly alter- 
nating, with asymmetry, betw^een a level somewhat above B and 
another well above D (usually at upper level on xiv but at lower 
level on xv-xvi?). First dorsal pore at 4/5 or 5/6. Prostomium 
epilobous, tongue usually open. Unpigmented. Segments, 80-136. 
Size, 60-100 x 3-5 mm. 

Calciferous sacs of x lateral and vertical, opening widely into 
oesophagus but reaching slightly below and above it. Calciferous 
gland more or less constricted at insertions of 10/11-12/13 so 
that gut in xi-xii or xiii may be rather moniliform. Intestinal 
origin in xv. Gizzard mostly in xvii. Typhlosole deeply incised 
anteriorly, by opposite lateral grooves, into half-moon shaped 
leaflets, ending in the region between 95th and 118th segments 
so as to leave 14-21 segments atyphlosolate. Extra-oesophageal 
trunks, median to hearts, pass up into dorsal trunk in xii. 
Hearts in vi-xi. (Nephridial bladders U-shaped, ducts passing 
into parietes near Bl) Holandric. Male deferent ducts with 
discoidal epididymis. Seminal vesicles four pairs, in ix-xii. 
(Atrial, TP and coelomic GS glands lacking.) Genital setae in 
enlarged follicles opening through genital tumescences. 

Reproduction. Presumably sexual and biparental. 

Disfrihution. Germany (Hamburg, Steinweiler, Worth/Rhein. 
Bienwald). Switzerland (Zurich). Belgium (Bouge, Esneux, 
Bastogne, Comblains-la Tour). England ( Grange-over-Sands, N. 



12 BREVIORA NO. 81 

Lancashire). Massachusett.s (Boston). New Jersey (Morris- 
town, Dover, Beatystown). 

Presence in America certainly is due to transportation, pre- 
sumably by man. The same agent may well be responsible for 
presence in England. 

Hahitats. Marshes, marshy places, meadows, peat bogs, very 
damp and loamy soil, mud. In soils with pH of 4.5-6.6. 

The specimen found in an English pasture on glacial drift, 
away from water, presumably was a stray. 

The Boston peat bog supports a population that certainly 
can be called dense. 

Associated earthworm species. Seven species, in previous 
samplings of the Boston peat bog, were found along with the 
dominant limicola. Five of those seven again were obtained, in 
1956, from the few shovels-full of earth that provided the above- 
described series of limicola. The species and numbers obtained 
are as follows : 

Allolohophora limicola 1-10-62-1. 

tuherculata 0-0-4. 

turgida 0-0-1. 

Litmhricus ruhellus 6-0-1. 

terrestris 0-0-1. 

Octolasium lacteum 1-0-3. 

Individuals of species of Lumhricus are of about average size. 
Individuals of tuherculata, turgida and lacteum are all smaller 
than average and, indeed, smaller than many of the individuals 
of limicola. 

DISCUSSION 

Only one specimen of .1. limicola has been reported from Eng- 
land, and one from Switzerland. How many worms from Bel- 
gium and Germany have been identified is not know^n but if lack 
of data as to differences from the type is any clue, the number 
is small. Paucity of records may be due, at least in part, to failure 
to collect in the proper habitats where dense populations can be 
expected. 

A clitellum, of SVo segments, shorter than on the types (where 
it was of 71/2-8V2 segments) was noted by Cernosvitov for his 
Belgian specimen. Otherwise, nothing of importance is known as 



1957 ALLOLOBOPIIORA LIMICOLA 13 

to the variability of European material. In contrast, records 
now have been published of the variation in 270 individuals 
from four American localities. Those records make possible for 
the first time in the Lumbricidae, an examination of the taxo- 
nomic value of somatic versus sexual characters. 

In the genital system of a species presumably with obligatory 
biparental reproduction, no variation whatever was found as to 
number and location of : gonads, gonoducts, seminal vesicles, 
spermathecae, ovisacs, male and female as well as spermathecal 
pores. An epididymis always was present. Each ovary had but 
one egg-string. Certain structures that have been neglected in 
the past, atrial, TP and supraparietal GS glands, always were 
absent. Exceptions to that invariability can of course be expected, 
more or less rarely, when hypo- or hypermeric ceplialic regenera- 
tion has taken place or when embryonic development has been 
prevented from proceeding normally. Development of an extra 
area of male tumescence on a single individual is a mutational 
aberration that may have more importance in other circumstances 
(to be considered subsequently) than in limicola. 

Variability worthy of note was found only in three portions 
of the reproductive system, the genital tumescences, the tuber- 
cula pubertatis, and the clitellum. Those are the very structures 
that have furnished the characters most used for distinguishing 
and defining lumbricid species. So much importance has been 
attached to one of the organs that ''variant," in a previous 
communication, was unwittingly restricted to mean divergent 
with respect to tubercula. In limicola, the tubercula usually are 
confined to xxxiii-xxxiv but there is enough individual deviation 
therefrom to require mention in a specific definition, if for no 
other reason than to obviate needless erection of species. A 
species norm as to number of segments covered by the clitellum 
and as to its location remains to be determined. Accordingly, 
all that can now be provided in a precis (p. 11) is a list of 
variations as hitherto recorded. 

Genital tumescences appear to provide a still more complicated 
problem in limicola. The number of individual patterns of 
tumescence location (Table 4) in the Massachusetts material 
seems large in view of a possibility that the colony may be highly 
inbred (descendant from a single importation of a few individ- 



14 BREVIORA NO. 81 

iials?). Only seven of the patterns were found in more than 
two individuals and just one in more than five individuals. Al- 
though a species pattern has not been recognized for limicola 
the data of Tables 3 and 4 show that tumescences usually can 
be expected in certain segments as listed in the precis. Those 
metameres, with the exception of xi, are the ones also mentioned 
in the original description of the species. 

In the somatic systems no variability whatever was found (in 
dissections of lOO-f- specimens) as to location of calciferous 
gland, gizzard, intestinal origin, last pair of hearts, and junction 
of extra-oesophageal trunks with the dorsal vessel, as to presence 
of calciferous sacs in x, and presence of an intestinal typhlosole. 
Especially noteworthy is the invariability of the sculpturing 
(into leaflets) at the anterior end of the typhlosole (Fig. 3) and 
the invariability in structure of the calciferous sacs (Figs. 1-2). 
Deviations from conditions described above for any of those or- 
gans can be expected only rarely and then as a result of some 
aberration of embryonic development. 

SUMMARY 

Variability of somatic as well as of genital structure has been 
i-ecorded from an American colony of a lumbricid species, pre- 
sumably with obligatory biparental reproduction. Variation in 
the genital system was found only with respect to those charac- 
ters most used hitherto in identification and definition of species. 



BREVIORA 

Miaseeim of Comparative Zoology 



Cambridge, Mass. January 31, 1958 Number 82 



THE TRUNK MUSCULATUKE OF 8ANZINA AND ITS 

BEARING ON CERTAIN ASPECTS OF THE 

MYOLOGICAL EVOLUTION 

OF SNAKES 

By Walter Auffenberg 

Dept. of Biology and Florida State Museum, 
University of Florida 

Since Mosauer's paper in 1935,^ very little has been published 
concerning the myology of the trunk region in snakes. In that 
paper he outlined three myological types on the basis of an ex- 
amination of a number of species" from several distinctly related 
groups. These types were described as characteristic of (1) the 
Boidae, (2) the Colubridae and (3) the Viperidae, and were 
distinguished on the basis of several correlated peculiarities in 
muscle arrangement. These arrangements were, according to 
Mosauer "surprisingly constant" within each group. 

As a contribution to a much needed wider survey of snake 
myology, the body musculature of Sanzina (a boid genus rarely 
available for this type of study) is described in this paper. It is 
shown that, at least in this genus, there is considerable departure 
from the basic boid pattern described by Mosauer. More impor- 
tant, this departure embodies some aspects of myological arrange- 
ment previously considered to be characteristic of the Viperidae 
or Colubridae. 



1 Walter Mosauer (1935) Univ. Calif., Los Angeles. Publ. Biol. Sci., Vol. 1. 
No. 6, pp. 81-120. 

2 In addition to Constrictor constrictor, Mosauer examined the following boid 
genera and species : Python moltirus, Calabaria rcinhardti, Epieratea striatus. 
Lichanura roseofusca and Charina bottae. 



2 BREVIORA No. 82 

I wish to thank Dr. Ernest E. Williams, Museum of Compara- 
tive Zoology, who made the specimen of Sanzina available for 
study. Dr. William Riemer, Florida State Museum, made it 
possible to study the musculature of a large specimen of Ophio- 
phagus hannah in the University of Florida collections, for which 
I am very grateful. 

The description below is based on a single adult specimen of 
Sanzina madagascarensis, approximately four and a half feet 
long (M.C.Z. 8002) . In addition, the musculature of one specimen 
each of Crotalus durissus terrificiis, Ophiophagus hannah and 
Coliiber constrictor was studied. No differences were found from 
the descriptions of these or similar forms given by Mosauer. The 
myological description of Sanzina is meant to supplement that 
given by Mosauer, and follows, for the most part, his outline of 
presentation to facilitate comparison. 

M. Semispinalis et spinalis 
(Figs. 1,4, ^.SP./SfP.) 

As in Constrictor constrictor, this muscle arises from a tendi- 
nous arch, stretching between the neural spine and postzygapo- 
physis of a single vertebra, with the concavity directed caudally. 
From this arch the muscle extends as a flat ribbon, directed for- 
ward, medially and dorsally. It is inserted by a tendon into the 
caudodorsal edge of the neural spine of a more anterior vertebra. 
From origin to insertion the muscle and its tendons cover ten 
segments. 

As in the typical bold arrangement, the medial portion of the 
muscle seems to represent the spinalis and the lateral part the 
semispinalis fused and inserted by means of a common terminal 
tendon. 

M. LONGISSIMUS DORSI 

(Figs. 1,2, 3, 4, ML) 

This muscle is also similar to that in Constrictor, with the 
origin at the craniolateral portion of a prezygapophysis. At 
about the level of the thirteenth vertebra cranial to its origin the 
muscle is replaced by a flat tendon, which divides in two. The 
medial division inserts on the posterolateral portion of the four- 



1958 



TRUNK MUSCULATURE OF SANZINA 




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BREVIORA 



No. 82 



teenth neural spine anterior to the origin of the muscle. The 
lower division of the tendon extends cranially for an additional 
vertebra, where it sends a small thin tendinous slip to the rib of 
the fifteenth vertebra, closely associated with the origin of the 

Anterior 




ML 



-CVO 



Fig. 2. Diagrammatic ventral view of the musculature of the middle 
thoracic region in Sanzina. Abbreviations: CIS, costalis internus supe- 
rior; CVD, dorsal head of the costovertebrocostalis; CVV, ventral head of 
the costovertebrocostalis; lAI, interarticularis inferior; IAS, interarticularis 
superior; IC, intercostalis proprius ; IQ, intercostalis quadrangularis; LC, 
levator costae; ML, longissimus; TC, tuberculocostalis ; TED, transversus 
dorsalis. 

supracostalis dorsalis and the insertion of the medial belly of the 
retractor costae biceps. Anterior to the fifteenth vertebra, the 
aponeurosis of the longissimus gives rise to the slender, band-like 



1958 TRUNK MUSCULATURE OP SANZINA 5 

medial belly of the retractor costae, which in turn gives rise to 
the more anterior lateral belly, inserting on the twentieth vertebra 
cranial to the primary origin of the entire complex. 

The main differences between this series of muscles and tendons 
in 8anzina and Constrictor lie in the lengths of the myological 
segments. In the former the medial tendinous insertion of the 
longissimus is on the fourteenth neural spine. In the latter it is 
on the ninth. In Sanzina the tendinous slip from the aponeurosis 
of the longissimus inserts on the fifteenth rib ; in Constrictor it 
inserts on the fifth. The entire complex, including the lateral 
head of the retractor costae includes twenty vertebrae in Sanzina, 
and eighteen in Constrictor. Thus, the longissimus is five seg- 
ments longer in Sanzina, and the retractor costae biceps complex 
seven segments shorter. 

M. MULTIFIDUS 

(Figs. 1,4, MM) 

In Constrictor this muscle originates from the ventral surface 
of the tendinous arch forming the origin of the semispinalis et 
spinalis. It extends four to five segments, inserting into the 
caudal border of the laminae of the neural arch. In Sanzina, as 
in most Colubridae and Viperidae studied by Mosauer, the 
muscle arises from the craniodorsal edge of the neural spine, and 
as in Crotahis extends anteriorly for only three to four vertebrae. 

M. Interarticularis superior 
(Pigs. 1,2, 3, 4, 7A^) 

In Constrictor this muscle arises from the lateral tendon of the 
tendinous arch of the semispinalis et spinalis. It then extends 
anteriorly for four vertebrae, where it inserts on the posterior 
lateral border of the postzygapophysis of several successive verte- 
brae. In Sanzina the muscle originates on the craniodorsal 
border of the postzygapophysis and extends cranially for one 
vertebra, inserting on the same region as in Constrictor. The 
muscle is thus considerably shorter in Sanzina, being similar to 
that in the Viperidae. In addition, its origin on the postzygapo- 
physis, rather than on the tendinous arch suggests the condition 
Mosauer found in most colubrids and viperids. On the other 



6 BREVIORA No, 82 

hand, the muscle is joined by a slip from the lateral portion of 
the fused semispinalis et spinalis, suggesting the origin for the 
interarticularis superior found in most boids (Fig. 4, A, B). 

Mosauer takes the medial head of the digastricus dorsalis of 
Colubridae to be homologous with the interarticularis superior 
of the Boidae. In all of the colubrid genera which he studied the 
medial head of the digastricus arises from the superior surface 
of the postzygapophysis and inserts by a thin tendon into the 
caudal border of a postzygapophysis several vertebrae cranially. 
It is joined by the lateral head of the digastricus, which arises 
from the accessory process of the next vertebra posterior to the 
one which forms the origin of the medial head. It is also joined 
by a small raphe of the longissimus (Fig. 4, D). 

In Sanzina the main body of the interarticularis superior in- 
serts on the cranial border of the postzygapophysis of the pre- 
ceding vertebrae as in other boids, but a small tendinous slip 
passes cranially to a slightly more lateral insertion on the postzy- 
gapophysis of the second preceding vertebra. This tendon is so 
similar to the anterior tendon of the digastricus in most colubrids 
that there can be little doubt as to homology. 

In the Anilidae, according to Mosauer, the medial head of the 
digastricus arises on the tendinous arch of the semispinalis et 
spinalis. It joins the lateral head, which covers one vertebra, 
and is apparently homologous to the interarticularis superior of 
Sanzina. 

Mosauer states that in the colubrid snakes the lateral head of 
the digastricus has apparently migrated posteriorly to the ac- 
cessory process of the succeeding vertebra, and has also become 
associated by means of a small raphe with the longissimus of the 
succeeding vertebra. That such migration is possible in the 
Colubridae seems reasonable in view of the fact that in at least 
the single specimen of Coluber which I have examined the lateral 
head of the digastricus is provided with a very thin tendon con- 
necting that muscle with the superior surface of the postzygapo- 
physis; probably this tendon represents the caudal end of the 
interarticularis superior of the Boidae. The caudal tendon of 
the medial head of the digastricus thus seems to represent, at least 
in part, the original semispinalis et spinalis of the tendinous arch. 

It seems reasonable to assume tliat the semispinalis has mi- 



1958 



TRUNK MUSCULATURE OF SANZINA 



grated posteriorly along the medial tendon of the longissimus. 
The Anilidae apparently represent an intermediate condition 
(Fig. 4). 

In the Viperidae the interarticularis superior is said by 
Mosauer to be very similar to that in the Boidae, except that a 
few fibers originate on the postzygapophysis. I have checked this 
in Crotalus durissvs terrificus. The mnscle extends for only one 

Anterior 




Fig. 3. Diagrammatic ventral view of tlie musculature of the anterior 
thoracic region in Sanzina. Abbreviations as in Figure 2. 

vertebra, as the main portion does in Sanzina. However, in C.d. 
terrificus and in the other viperids examined by Mosauer there is 
no anteriorly directed tendon, and the muscle originates mainly 
on the semispinalis et spinalis. 



8 BREVIORA No. 82 

M. COSTOVERTEBROCOSTALIS 

(Figs. 1,2, 3,072), cyy) 

As in Constrictor, this muscle is composed of two heads. The 
medioventral head arises from the lateral surface of the haemal 
keel and from the caudoventral parapophysial portion of the 
paradiapophysis. It extends caudally and laterally, inserting 
on the costal tubercle and neck of the succeeding rib. The dorsal 
head arises from the cranioventral portion of the diapophysial 
portion of the paradiapophysis and unites with the ventral head 
caudally. 

M. Intercostalis quadrangularis 
(Figs. 1,2, 3,/^) 

This small muscle is not mentioned by Mosauer in his de- 
scription of the typical musculature of the Boidae, but only in 
the description of the arrangement found in the Colubridae. It 
is, however, well developed in Sanzina. As in the Colubridae and 
Viperidae, it arises from the caudal surface of the capitulum 
costae, just lateral to the articular head of the rib, running 
caudally and laterally to be inserted on the inferior cranial 
border of the following rib. The identity of this muscle in San- 
ziTia is so obvious that homology to the similar muscle mass in the 
Colubridae is hardly to be questioned. 

M. TUBERCULOCOSTALIS 

(Figs. 1,2, 3, TO 

Mosauer describes this muscle as being found only in the 
Colubridae. However, it is present in Sanzina, originating, as 
it does in colubrid snakes, from the laterocaudal circumference 
of the dorsocaudal costal tubercle, running obliquely to insert on 
the cranial border of the neck of the following rib. 



'^t?' 



OTHER MUSCLES 

In addition to the muscles described above, the remaining units 
described by Mosauer as being found in the Boidae have all been 
located in Sanzina. Some of these differ in a very minor fashion 
from those found in Constrictor. The retractor costae biceps is 



1958 



TRUNK MUSCULATURE OF SANZINA 



somewhat shorter, as pointed out above. The eostalis intern i 
superior covers eight vertebrae in Sanziyia and nine in Con- 
strictor. The following- muscles were found to be the same in both 
Constrictor and Sanzina: Intervertebralis, levator costae, trans- 
versus dorsalLs, interarticularis inferior, supracostalis dorsalis, 



Anterior 



TTSSPSP 




B 



D 



Fig. 4. Diagrammatic representations of the dorsal musculature of 
(A) Constrictor, (B) Sanzina, (C) Anilius and (D) Coluher illustrating 
the apparent evolution of the somispinalis et spinalis and the interarticularis 
superior to form the digastricus and the highly modified semispinalis in the 
Colubridae. Abbreviations: DDL, digastricus, lateral head; DDM, digastri- 
cus, medial head; IAS, interarticularis superior; ML, longissimus; MM, 
multifidus; MT. ML., median tendon of the longissimus; ECM, retractor 
costae biceps, medial belly ; SP, spinalis ; SSP, semispinalis ; SSP.SP., 
semispinalis et spinalis; TA, tendinous arch; TT. SSP.SP, terminal tendon 
of the semispinalis et spinalis. 



supracostalis lateralis superior, supracostalis lateralis inferior, 
intercostalis proprius, intercartilaginosus, eostalis inferior, trans- 
versus abdominus, obliquus abdominis internus, costocutanus 
superior and costocutanus inferior. 



10 BREVIORA No. 82 

COLUMNAR VARIATION 

Little work has been done regarding the change in position 
and shape of muscles from the middle to the anterior or posterior 
portion of the vertebral column in snakes. Considerable struc- 
tural changes take place along the length of the column. The 
most important of these is the presence of well-developed hy- 
papophyses anteriorly. Such changes would be expected to in- 
fluence arrangement of at least some myological elements in the 
anterior part of the column. Mosauer made only a few comments 
concerning this variation. 

For the most part, myological changes correlated with inter- 
columnar morphological variability involve shortened or length- 
ened muscle segments, so that fewer or additional vertebrae are 
covered in the anterior part of the column than in the middle or 
posterior portions. However, some of the changes involve more 
than mere variability in muscle length, and these are of consider- 
able interest (c/. Figs. 2 and 3). 

As mentioned previously, and also by Mosauer, the transversus 
dorsalis is apparently absent in the anterior part of the column 
of most snakes. The costalis internis superior thus forms the 
inner muscular lining of the body cavity in this region. The 
latter muscle becomes progressively shorter and thicker cranially, 
so that each muscle covers only three segments instead of eight 
as in the mid-dorsal region. The insertion and origin, as well as 
relationship to other muscles, remain the same. 

Of the deeper muscles the costovertebrocostalis becomes longer 
anteriorly, covering from one to four vertebrae, instead of only 
one to two. Occasional fibers may even extend to the fifth vertebra 
anteriorl3^ As in the middle thoracic region, the muscle is pro- 
vided with two heads. The longer, thinner member inserts on the 
cranioventral portion of the hypapophysis of the fourth vertebra 
cranially. The shorter, thicker head inserts on the basal part of 
the hypapophysis and the ventrolateral portion of the body of the 
centrum, including the cranial border of the parapophysis. Of 
particular importance is the fact that in this area this muscle 
also originates in two heads, the larger on the neck of the rili, 
and the smaller on the cranial surface of the parapophysis. The 
fibers of both become interlaced before they divide into the tAvo 
heads at their insertions described above (Fig. 3). 



1958 TRUNK MUSCULATURE OF SANZINA 11 

Mosauer described a muscle, the transversohypapophyseus, 
supposedly peculiar to the Colubridae, Elapidae and Viperidae, 
which originates on the cranial border of the parapophysis and 
extends to the caudolateroventral border of the hypapophysis. 
These fibers are also joined by others from deeper, shorter mus- 
cle segments. The muscle is quite distinct in the single specimens 
of C. constrictor, Ophiophagus hannah and C. durissus terrificus 
I have examined. It seems highly probable that the same muscle 
is represented in the anterior portion of the vertebral column 
of 8an2i7ia as a second head of the costovertebrocostalis, not 
fully separated from a shorter head, which lies ventral to it 
(Figs. 2, 3), and which is found as a distinct unit in the anterior 
vertebrae of the Colubridae and all vertebrae of the Viperidae 
and Elapidae. Mosauer also reports the transversohypapophyseus 
as extending to the anus in colubrids in which hypapophyses 
are present throughout the column. Thus it seems apparent that 
the muscle cannot be considered characteristic of any particular 
family or families, but simply a development of the costoverte- 
brocostalis found only in those regions of the column in which 
hypapophyses are well developed. 

DISCUSSION 

The exact phyletic position of Samina is not at all clear and 
the present myological study sheds little light on this problem. 
The genus is generally placed in the Boinae. Its zoogeographic 
isolation on Madagascar suggests that it may represent a very old 
stock. 

Myologically Samina is not as much like Constrictor as one 
would suppose. Deviations in muscular arrangement are sug- 
gestive of intermediacy between tj'pical boines, such as Con- 
strictor, and the Anilidae, as represented by Anilius. The latter 
seems, in turn, intermediate between Sonzina and the Colubridae. 

It is entirely possible that various members of the Boidae have 
independently evolved colubrid-like modifications several times 
in the past. These radiations may even be represented in rather 
extreme fashion by one or more of the Recent morphologically 
intermediate families (Anilidae, Xenopeltidae and Fropeltidae). 
Sanzina, as a slightly modified general boid type, may represent 
a single minor radiation paralleling the myo-evolutionary se- 



12 BREVIOBA No. 82 

quence leading to the Colubridae. There is, at least at present, 
little reason to suppose that Sanzina is on, or even close to, the 
line leading to the colubrids. 

The musculature of relatively few snakes is known in detail. 
Before the full value of this tool in systematics can be realized, 
it is necessary to study the myology of many important Recent 
genera. Mosauer had intended that his observations would stimu- 
late other workers to examine more closely the trunk musculature 
of snakes. However, the preliminary nature of his study has been 
largely overlooked by later writers. It is hoped that the present 
paper will re-awaken interest in comparative trunk musculature 
of reptiles if only by pointing out the inadequacy of Mosauer 's 
system, which was intentionally over-simplified because of its 
admitted preliminary nature. 



BREVIORA 



Museiuiiii of Contiparative Zoology 

Cambridge, Mass. January 29, lOoS Xumber 88 



THAMXOPHIS BOVALLII DUNN REDISCOVERED 
(REPTILIA, SERPENTES) 

By Benjamin Shreve and Carl Gans 

In the course of a collecting' trip l)y the junior author in ]\Iexico 
and Central America, a garter snake (]\Iuseum of Comparative 
Zoology 54974) was secured at San Jorge, Lake Nicaragua, 
Nicaragua, on July 1, 1956. 

This specimen was found to agree closely with E. R. Dunn's 
original description (1940, p. 190) of TJianinophis hovaUii (type 
locality : Granada on Lake Nicaragua, somewhat farther north 
than San Jorge). The evidence of this additional specimen and 
certain other considerations led to the conclusion that Smith 
(1942, pp. 97 and 110) was wrong in synonymizing hovallii with 
sumichrasti Cope. Later examination of a specimen cited by 
Dunn {Joe. cit. p. 192) as possibly hovallii — American Museum 
of Natural History 12439 from Colorado bar, Costa Rica — con- 
firmed this conclusion and further extended the range of the form 
(map). 

The table gives the scale counts for Dunn's two juvenile females 
(specimen A was expressly designated the type by him), and for 
the two specimens, both males, that we have added. 

The color pattern of the IMCZ specimen is well shown in Figures 
1-3. There are a few differences from Dunn's specimens, as des- 
cribed. The American Museum specimen also differs slightly. 
Thus anteriorly, and probably posteriorly as well, the lateral bars 
are shorter (true also of the American IMuseum specimen) and 
two rows of alternating and sometimes coalescing spots occur 
throughout the vertebral region. In Dunn's two specimens the 
spots were separate only posteriorly. In the AMNH specimen 
the two rows of spots tend to coalesce into one row but in such a 



BREVIORA 



No. 83 



way that the identity of the two rows is not entirely lost over the 
whole length of the animal. The MCZ specimen lacks the light, 
black-edged spots on the parietals found by Dunn in his juveniles 




Figure 1. Ihaninophis bovallii. Dorsal and ventral views of MCZ 54974 
from San Jorge, Nicaragua. 



and present in the American Museum specimen. Both the MCZ 
and the AMNH specimens have seemingly a less clear delimi- 
tation of dorsal coloration on the side of the head, and neither 



1957 



THAMNOPHIS BOVALLII 



shows the reddish in the dorsal coloration mentioned for the type 
series by Dunn. As the s])eeimens l)el\)re us are males and ap- 
parently adult, while both of Dunn's specimens were females and 
young enough to still show the "navel," it is suspected that some 
of the color differences are due to age or sex. 




Figure 2. TJiamnopJii.'i horaUii. Detail of middorsal eolor pattern of MCZ 
54974 slightly posterior to midliody. Note that there e.Kicts a tendency for 
middorsal separation of the two rows of blott-hes which are partly fu?ed 
in this region. 

H. M. Smith [he. cif.) was the first to regard bovallii as in- 
valid (without, however, stating his reasons), placing it in the 
synonymy of the typical race of T. su))iichra>iti. The two cotypes 
of the latter were statetl to have come from Orizaba, Vera Cruz, 
Mexico, but Smith thinks it more likely that they were collected 
on the Atlantic slope of the Isthmus of Tehuantepee. 



SpociniPii 



Goteborg A 
Goteborg B 
MCZ 54974 
AMMI 12439 



Sf.x Vcntrals 
-f 
Cauda Is 



140 + 70 
1-14-f 71 
148 + 79 

14S + .\- 



Dorsals 



21-19-17 
21-19-17 
21-19-17 
21-19-17 



TABLE 

Dfirsals Pro- Post- rpiirr Liiwcr liod.v 

ilrcip to 19 ocular m-nlni- Labials I,abial^^ + Tail 

at ventral Length 

iniml)er in mm. 



11 

ir.-ll 

7 

S-9 



1 



■•4 


8(4,5) 


10 


1 so + RO 


% 


8(4,5) 


10 


180 + 58 


3 


8(4,5) 


10 


485 + 1(57 


3 


8(4,5) 


10 


543 + x 



As Thamnophis bovallii Dunn is known only from southern 
Nicaragua and northern Costa Rica, it would be surprising if it 



BREVIORA 



No. 83 




Figure 3. Thamnophis bovallii. Dorsal, lati'inl and ventral views of the 
head of ^I("Z 54974. Xoto the very irrcsular Ijlaek fleeks in tlie eentor of the 
parietals where the other specimens have the light, black-edged spot. The 
color difference between left and right sides is due to the shedding of the 
scales of the left side. 



1957 THAMNOPHIS BOVALLll 5 

were quite identical with a form from eastern Mexico, the more so 
since two races (later three, Smith, Nixon, and Smith, 1950, 
p. 579) of the same species were recognized by Smith from inter- 
vening Guatemala, and since there appear to be no records at 
all for this group of garter snakes between Guatemala and 
southern Nicaragua. 

W. W. Milstead (1953) has treated this group of Mexican 
and Central American Thamnophis very differently than did 
Smith but has followed him in synonymizing hovallii, again with- 
out discussion. Milstead has made Smith's species, sumkhrasti, 
a race of cyrtopsis {equcs of authors), and has contributed to 
the understanding of the variational range of scale counts and 
color of the populations he treats. However, he is so far from 
giving serious attention to hovallii that after citing it in the 
synonymy of T. cyrtopsis sumichrasti he forgets to mention 
Nicaragua in the range of the subspecies in the section on ' ' Sub- 
specific Variation" {loc. cit., p. 359), although he does include it 
later under the formal presentation of the range of sumichrasti 
{loc. cit., p. 361). 

Several characters persuade us not only to regard hovallii as 
valid, but even provisionally to retain it as a full species until 
the taxonomic uncertainties surrounding this section of Thamno- 
phis are further clarified. In scale characters the most significant 
point is the presence of 21 rather than 19 scale rows at the nape, 
while in coloration the most striking feature is the dark anterior 
border (narrowed medially) of all the ventrals. Both these fea- 
tures serve to separate T. bo vail ii from all T. cyrtopsis as con- 
ceived by Milstead. The shortness of the lateral bars (in our two 
examples), and the presence always of two rows of spots in the 
vertebral region (even if partly coalesced) are additional char- 
acters. 

ECOLOGY 

The San Jorge specimen was collected by two boys under 
lakeshore debris, early on the morning of July 1, 1956. Just above 
the water line there is a windrow of vegetable debris, wood, coco- 
nut husks, etc. This and the small puddles of this zone were 
inhabited at dusk by large numbers of small toads and frogs. 
Series of Leptodactylus melavonotus (Hallowell), and Bufo mari- 



bREVIORA 



No. 83 



nus (Linne), were collected during the visit. During the daytime 
the frogs appeared to hide away from the beach, but large num- 
bers of lizards were found along the bluff beyond the beach 
{Anolis achilles Taylor?, Sceloporus rariabilis oUoporus Smith. 
and CnemidopJiorous d. deppii Wiegmann were collected). In 
addition to the specimen of horallii the beach also yielded a speci- 
men of Thamnophis saurita chalceus (Cope) and one of Lepto- 
deira rhombifera Giinther. 



NICARAGUA 



GRANADA 



SAN JORGE 




COSTA RICA 



Map. Thamnophis bovallii. Sketch map showing the relation of the thioe 
localities from which this species has been recorded. 



Granada, the type locality of bovallii, also lies on Lake Nicara- 
gua (elev. 105 ft.) and the two specimens listed l)y Dunn (loc. 
cit.) probably came from a locality similar to that described. 
"Colorado Bar," on the other hand, refers to the mouth of the 
Colorado river (Boca do Rio Colorado on Costa Kican maps). 
While this lies on the Caribbean coast, it is part of the delta of 
the San Juan River which drains Lake Nicaragua and which has 



1957 THAMNOPHIS BOVALLII 7 

such a low gradient that it was used for navigation in historic 
times. 

Thus all three localities at which T. bovallii has been taken 
are lowland. This is in strong contrast with Milstead's descrip- 
tion of the situation in T. cyrtopsis which he regards as a species 
that does not descend below 1000 feet in altitude. He admits one 
exception, the Edwards Plateau of Texas (minimum altitude 700 
feet), but ascribes this to the temperate, subhumid climate of the 
region. He doubts the records of cyrtopsis from British Hon- 
duras, Yucatan (Mexico), and Escuintla, Guatemala, which 
would bring the form into the tropical zone. Here again is evi- 
dence of his lack of attention to hovallii since he fails to mention 
it or Nicaragua at all in this discussion. 

ACKNOWLEDGMENTS 

The jvuiior author welcomes this opportunity to thank those 
friends, whose unselfish and valuable assistance made this trip 
and the visit to Rivas and San Jorge both possible and successful. 
I should like to express my particular appreciation to Messrs. 
Julio Cesar Moya, Inspector Porestal, and Gustavo Hernandez, 
Agronomo, both of Rivas as well as to the municipal officials of 
this city, and in addition, to my companions Gerardo Budowski 
of the Instituto Interamericano de Ciencias Agricolas of Tur- 
rialba, and Phil Noyce. 

We also wish to thank Arthur Loveridge for comments, and 
Ernest E. Williams for additions to and revision of the manu- 
script, and the staff of the Department of Amphibians and Rep- 
tiles of The American Museum of Natural History for loan of the 
Colorado-bar specimen. 

LITEEATURE CITED 

Dunn, Emmett Reid 

1940. Notes on some American lizards and snakes in the Museum at 
Goteborg. Herpetologica, vol. 1, pp. 189-94. 

MiLSTEAD, William W. 

1953. Geographic variation in the garter snake, Thamnophis cyrtopsis. 
Texas Jour. Sci., vol. o, no. 3, pp. 348-379. 



8 BREVIORA No. 83 

Smith, Hobart M. 

1942. The synonymy of the garter snakes (Thavmophis) , with notes on 
Mexican and Central American species. Zoologica (New York), 
vol. 27, pts. 3-4, pp. 97-123. 

Smith, Hobart M., C. William Nixon, and Phillip W. Smith 

1950. Mexican and Central American garter snakes (Thamnophis) in 
the British Museum (Natural History). Jour. Linn. Soc, vol. 
41, no. 282, pp. 571-584. 



BREVIORA 

MiuseuiirM of Comparative Zoology 

Cambridge, Mass. January 30, 1958 Number 84 



REDISCOVERY OF THE AUSTRALIAN CHELID GENUS 
PSEUDEMYDURA SIEBENROCK (CHELIDAE. 

TESTUDINES) 

By Ernest E. Williams 

In 1954, L. Glauert described a new species — Emydura iv- 
spectata — from western Australia, comparing his new form only 
with species of Emydura. A g:lance at his photograph revealed 
that the new form was congeneric with, and perhaps conspecific 
with, Pseudemydura umhrina described by Siebenrock from an 
unique type specimen in the Vienna Museum with no more pre- 
cise locality than "Australia." and which had come into the 
possession of that museum in 1839. 

Pseudemydura umhrina was named by Siebenrock in a pre- 
liminary note in 1901, but it was not figured nor fully described 
until 1907. The long interval between these two papers may, I 
tliink, be easily explained : The specimen was unique and without 
adequate locality, yet it was made the type of a genus. Only Sie- 
benrock 's wide knowledge of the order as a whole justified such 
a procedure, and even for him it was natural and desirable to 
wait for some years in the hope of procuring additional specimens 
the better to document his case. 

Siebenrock never obtained any further specimens, but in default 
of them his 1907 paper compared Pseudemydura carefully and in 
a key with every other genus in the family. 

Pseudemydura belongs to the group of short-necked chelids, 
and is peculiar among these in having a large intergular plastral 
scute that not only separates gulars and humerals but penetrates 
some distance between pectorals. A somewhat similar condition 
occurs in the long-necked chelid genus Chelodina, in which, how- 
ever, the gulars usually meet in front of the intergular, separat- 



2 BREVIORA NO. 84 

ing the latter from the anterior ])lastral margin. Only in Chclo- 
dina interc/idaris (Fry, 1917) among Reeent turtles is there any 
elose parallel to the condition found in Pscudcrnydura. Among 
known fossil forms, however, a very similar intergular pattern 
occurs in some members of the related family Pelomedusidae — 
KlochcJ]!^ perfect a Nopcsa of the Cretaceous of Europe, and prob- 
ably in some Taphrosphys (Cretaceous and Tertiary of Xoith and 
South America). 

The relationship of the intergular to the other anterior plastral 
scutes in Pseudemydura nmhrina (clearly seen in Glauert's speci- 
mens) is thus striking enough among Recent members of the 
Chelidae and in particular tlu^ short-necked chelids to be l)y itself 
a feature permitting recognition at the generic level. However, 
the judgment that the specimens from Warbrook, 24 miles north 
of Perth, West Australia, described by Glauert as Emydura 
Inspcciata belong not to Emydnra but to Pseudemydura does not 
rest only on this feature but also uj^on a similarity that in crea- 
tures so variable as turtles is surprisingly complete. 

Neither the Vu una tj'pe nor any of Glauert 's material has been 
available to im\ but the similarity of the two described forms can 
be placed beyond all question merely by comparing in parallel 
columns a translation of Siebenrock's 1907 description and Glau- 
('i-t"s 19;14 account. I have given Siebenrock's description in full 
and rearranged Glauert's shorter description to make the paral- 
lels and ditferences more obvious : 

Pseudemydura umbrina Emydura inspectata 

Carapace length: 106 mm Carapace length (two siiecimens) : 

133 mm, 98 mm 

('.■II apace breadth: 81 mm Carapace breadth (two specimens): 

103, 83 mm 

Carapace height: 33 mm 

Carapace very strongly flattened with Carapace slightly depressed with a 
a definite vertebral furrow, most distinct sulcus. 

di.stinct on tlie middle three verte- 
brals. 



U)5b 



KEDISCOVERY OF PSEUDEMYDURA 



I'fsciidriii iidina uinhi'ina 

'1'Ir' postiM-ior carapace rim only 
sliglitly expanded so that the 
greatest breadth (between the 
eighth marginals) only slightly ex- 
ceeds the breadth at the middle of 
the shell. 

Anterior carapace rim truncate and 
only insignificantly incurved. 

Posterior carapace rim projecting 
medially into a keel produced by 
the arching of the supracaudals 
and adjoining eleventh marginals. 
The supracaudals also forming 
ventrally an acute-angled notch, 
which, with the posterior end of 
the plastron, also angularly 
notched, forms a rhomboidal open- 
ing for the tail. 

First vertebral longest, broader an- 
teriorly than posteriorly, narrower 
than the second and third, as 
broad as the fourth and fifth. 



First costal as broad as, the remain 
ing three costals narrower than, 
the corresponding vertebrals. Of 
these the second costal broadest, 
almost twice as broad as the 
fourth. All costals broader at the 
lateral margins than medially, and 
broader than the lateral margins 
of the corresponding vertebrals. 

Carapacial shields leathery, finely 
wrinkled. 

Nuchal rather large, trapezoidal, 
broader in front than behind. 



Kmydura inspcctata 

Slightly narrower in front (96 mm 
as compared with 103 mm) ; mar- 
ginals 5-8 with a distinct flange. 



First vertebral largest, as wide as 
long, obtusely pointed in front, the 
second and third much wider than 
long, fourth and fifth slightly 
wider than long, longer than the 
second and third. 



Second and third vertebrals shorter 
than the adjacent costals. 



Nuchal very small (9 x .5 mm) en- 
tirely free below. 



BREVIORA 



NO. 84 



Pseudemydura umbrina 

Lateral marginals very narrow; the 
sixth narrowest, only about one- 
third as wide as the ninth. The 
form of the lateral marginals is 
correlated with the great size, es- 
pecially breadth, of the second 
costal. 

Plastron flat, about as large as the 
shell opening, with a distinct angle 
at the bridge. 



Anterior lobe broader than posterior 
lobe, truncate. 

Posterior lol)e with an angular notch. 

The two corners of the anal notch 
are bent upward to make contact 
with the posterior rim of the shell, 
thus forming the rhomboidal open- 
ing l)efore mentioneil. 

Breadth of bridge one-third the 
length of plastron and one-half 
the width of the anterior lobe. 

Intergular very large, heart-shaped, 
not much longer than broad, its 
broad anterior margin straight, 
weakly dentated, the pointed pos- 
terior end inserted between the 
pectorals, separating these in their 
anterior third. 

(iulars very small, forming equilat- 
eral triangles, their medial sides 
scarcely one-third the adjacent lat- 
eral rim of the intergular. 

numerals small, widely separated 
from one another, in form and size 
showing great similarity with those 
of Chelodina. 



Emydura inspectata 

Fifth, sixth, seventh and eighth 
marginals with distinct flange, 
fourth to seventh narrowest. 



Plastron (124 x 95^2 nim, measured 
to the edge of the bridge) flat, 
large, almost as wide as the cara- 
pace, bridge rounded. 

Plastron narrower behind, semi-cir- 
cular in front, posteriorly with a 
broad, straight-sided, anal notch. 



Bridge one-third the length of the 
plastron measured along the mid- 
line. 

Intergular very large (29.5 x 25.5 
mm). 



Widely in contact with the pectorals. 
Gulars very small, widely separated. 



Humerals triangular slightly nar- 
rower than the intergular. 



I 



1958 



KEDlt^COVERY OP PSEUDEMYDURA 



Fseudernydu ra ii in hrtit a 

Anal middle suture longer than the 
pectoral and siK"iticaiitly longer 
than the femoral. 



Intergular longer than the pectoral 
middle suture. 

Head broad and tiat, the upper sur- 
face finely wrinkled. 

Parietals strongly expanded, cover- 
ing the whole width of the head, 
their posterior margin not ])ointed 
but somewhat incurved. 

Snout short. 

Interorbital space broad and con- 
cave, its breadth exceeding the 
transverse diameter of tlie orbits. 
Both jaws narrow; the l)readth of 
the lower jaw at the symphysis not 
quite two-thirds the transverse di- 
ameter of the orbit. 

Two small chin barbels, widely sep- 
arated. 

Back of neck with numerous large 
erect conical tubercles. 



Limbs with rather large flat scales. 
Transverse lamellae such as occur 
on the anterior surface of the 
loiver arm in Emydvra entirely 
lacMng. 

Webbing well developed, extending 
to the claws. Fifth toe of hind 
foot clawless. 

Tail short, reaching scarcely to the 
posterior rim of the shell. 



(■'.nijidura tiinptrtalu 

Pectorals slightly shorter than the 
femorals which are shorter than 
Ihe anals, the l;itter forming a 
broad shallow notch behind. 



Head broad and flat, rugulose above. 



Snout projecting. 

Interorbital space twice the width 
of the symphysis, the latter equal 
to the vertical diameter of the 
orbit. 



Two small barbels. 



Neck above and laterally with pro- 
nounced conical tubercles ; tem- 
poral region, chin and throat 
reticulate. 

Forelimbs with three series of trans- 
verse lamellae and a flap of three 
or four enlarged scales. 



Tail very short, hardly projecting. 



6 BREVIORA NO. 84 

Pseudemydura umbrina Emydura inspectata 

Carapace and upper surfaces of head Color of carapace bone brown, 
umbra brown. Young: brighter brown. Head 

above dark olive, sides of face, 
lower lip, chin and throat mar- 
guerite yellow, a few dark mark- 
ings on the throat. 

Plastron dirty yellowish green. All Plastron buffy olive, over the sutures 
sutures of the shell brown. buff. Young: plastral plates edged 

with brown. 

Limbs and back of neck dark brown, Forelimbs olive black, hind limbs 
the jaws horn-colored. darker. 

Siebenrock "s type was intermediate in size between Glauert's 
type and the latter 's young specimen, but closer to the young 
specimen. Careful comparison of the two descriptions discovers 
few differences that are not merely verbal, while there is agree- 
ment in all essentials even in color — the latter remarkable when 
it is realized that CTlauert's fresh specimens are being compared 
with a specimen that had been in the Vienna Museum 62 years 
before it was first described. 

Only one of the few described differences would seem possibly 
significant. Siebenrock makes a point of the absence in umhrina 
of transverse lamellae such as exist in the genus Emydura. Glau- 
ert specifies in inspectata "three series of transverse lamellae and 
a flap of three or four enlarged scales." 

Glauert's photographs accompanying the type description of 
inspectata, however, while excellent for other aspects of the ani- 
mal do not show this area. I have therefore obtained from Dr. 
Glauert two photographs of the forelimb in his type. These are 
reproduced in Plates 3 and 4 and sliow the flap or spur very well 
and in the ventral view exhibit also the three series of scales de- 
scribed as "transverse lamellae" Ijy Glauert. 

Siebenrock 's 1907 figure (Plates 1 and 2) definitely does not 
show any flap or sjiur on this forelimb on the type of Pseudemys 
)())ibri)ia but does show in the ventral view several series of scales 
apparently less regular than those of Glauert's type but easily 
comparal)le to these. These scales are individually rather squar- 
ish, though arranged in transverse rows. In this respect the dif- 



15)58 REDISCOVERY OP PSEUDEMYDURA 7 

ference between Glauert's and Siebenroek's descriptions would 
seem to be primarily verl)Hl. However, the absence of the flap 
would still remain as a serious difference between Pseudemyduro 
}(*iibriua and IJini/diira iitspectata. 

To ehet'k this final point 1 reqvu^sted Dr. Eiselt of the Vienna 
Museum to re-examine Siebenroek's specimen. He reports: "The 
type of Pseudeniydura nmhrina is a stuffed and lacquered speci- 
men. In correspondence with Siebenroek's figures it showed no 
spurs whatsoever. Reing somewhat of a sceptic, I had the lac 
removed from the front leg's and feet, and then soaked them in 
water. Tlie effect was surju'ising : both lower front legs now show 
on their outer (hind) edges a series of three triangular scales 
which had, up to now, been glued firmly to the lower surface of 
the leg. Also the scales on the outer edge of the fifth toe (finger) 
show a tendency to form a flat and serrated lamella. As a whole 
the scalation of the legs shown in Siebenroek's figures is rather 
inaccurate and needs redescription or refiguring. " 

Dr. Eiselt has also provided a sketch of the flap, which appears 
to be not significantly different from that described and photo- 
graphed by Glauert. 

With this evidence the synonymizing of rseudemiidura uni- 
brina and Emydura iiispectata must be considered established. 

Glauert's species name must thus disappear from the list, but 
his discovery is still important. A genus known from a single 
specimen without adequate provenance is an unliappy problem. 
The Glauert specimens provide an exact locality, confirmation 
of the characters described by Siebenrock, and knowledge of the 
peculiar spur-flap, a feature missed by Siebenrock and which, 
though approached in other chelids by a flap of scales along the 
whole external range of the forelimb, is in its peculiar spur-like 
<levelopment apparently peculiar to this genus. 

Dr. Glauert has called to my attention that Western Australia 
was opened in 1S2!> and that the German collector Dr. L. Preiss 
collected there in the late 1830 's and early 1840 's. This accords 
very well with the date of arrival of the Vienna specimen at that 
museum, but Dr. Eiselt could supply me with no information 
about the collector. 

Acknowledgonoifs. 'My indebtedness to Dr. Eiselt and Dr. 
Glauert has been conspicuous throughout this paper. I want to 



8 BREVIORA NO. 84 

make, however, a very personal expression of thanks i'or their 
kindness and courtesy in answering my questions. The photo- 
graphs of the forefoot of E my dura inspcctata were furnished by 
Dr. Glauert ; the reproduction of Siebenrock 's figure is the work 
of Mr. Frank White. 

EEFERENCES CITED 

Fry, D. B. 

1917. Herpetological notes III. On a new Chelodinu from Australia 
witli a key to the genus. Proc. Eoy. Soe. Queensland, vol. 27, 
pp. 88-90. 

Glauert, L. 

1954. Herpetological Miscellanea. IV. A new swamp tortoise from the 
Swan River District. The Western Australian Naturalist, vol. 4, 
no. 6, pp. 125-127. 

Siebenrock, F. 

1901. Beschreibung einer neuen Schildkrotengattung aus der Familie 
Chelydidae von Australien. Sitzber. Akad. Wiss. Wien, math, 
nat. Kl., Jahrg. 1901, pp. 248-250. 

1907. Beschreibung und Abbildung von Psevdemydina umbriiKi 
Siebenr. und ilue systematische Stellung in der Familie Chelyd- 
idae. Sitzber. Akad. Wiss. Wien, math. nat. Kl., vol. 116, pp. 
1205-1211. 







Plate 1. Type of Fseudemydura umbrina, dorsal view. From Siebenrock 
(1907). 




Plate 2. Type of Pseudemydura umbrina, ventral view. From Siebeurock 
(1907). 




Plate 3. Dorsal view of forefoot of type of Emydura inspectata Glauert. 



f?J--^^ 




Plate 4. Ventral view of forefoot of type Emydura inspectata Glaueit. 



BREVIORA 



Museiiimi of Comparsitive Zoology 

Cajibridge, Mass. January 31, 1958 Number 85 

THE CHOANAL PAPILLAE OF THE CHELONIIDAE 

By Thomas S. Parsons 

Harvard Biological Laboratories 
Cambridge, Mass. 

INTRODUCTION 

There appear to be no published descriptions of the papillae 
whirh are found along the lateral margin of the choanae or in- 
ternal nares in the members of the Cheloniidae. They are men- 
tioned in two papers, Derauiyagala (1939) and Carr (1942), but 
in each case only the numl)er present is noted and given as a 
generic character. 

The following note is based primarily on the study of two 
heads of Chelonia my das which were obtained from the Carolina 
Biological Supply Company. Supplementary observations were 
made on specimens in the collections of the Museum of Com- 
|)arative Zoology. Histological investigations were confined to 
Chelonia mydas. The specimens had been preserved in formalin 
and were stained with hematoxylin and eosin and by the Biel- 
schowsky silver method. A third method, Mallory's triple con- 
nective tissue stain, was tried, l)ut did not prove successful. 

1 am indebted to Dr. E. E. AVilliams for allowing me to examine 
the specimens in the Museum of Comparative Zoology, and for 
his advice throughout the course of this study. Thanks are also 
due to Dr. A. B. Dawson and Mrs. T. S. Parsons for advice and 
assistance in making the histological preparations. 

CHELONIA MYDAS 

■■■'■.1. , 
in Clielo7iia my das the papillae form a row along the entire 

lateral margin of the choana. They are long and narrow conical 



2 BREVIORA No. 85 

pi-ojectioiis of irregular length, so that tlie row of papillae ap- 
pears quite uneven. In the anterior two-thirds of the choana, they 
lend to be longer and more prominent, Avith ai)proximatply half 
of the width of the ehoana occupied by the papillae; farther pos- 
teriorly they do not extend out into the choana any great dis- 
tance. Deraniyagala (1939) reports not only a lateral roAV. but 
also a single anteromedial pa{)illary vestige. However no such 
vestige was seen in any of the specimens studied in the present 
investigation. 

The number of papillae in the lateral row is not constant. 
Carr (1942) states only that they are numerous, and Deraniya- 
gala (1989) that there are eight. Unfortunately the latter authoi- 
does not mention the number of specimens which he examiucd. 
In the two adults dissected for the present study there were 
eleven to fourteen pai)illae on either side (one had thirteen on 
each side, the other eleven on the left and fourteen on the right i. 
The only other adult specimen which was examined (MCZ 7150) 
possessed ten papillae on the right side and eleven on the left. 
Thus there are probably ten or more pai)illae along the latei-al 
margin of each choana in most fully mature Chdonia nijidos: 
however this is not true of juveniles. In three specimens under 
six inches in total length (MCZ 1413, 1414, and 29594) there 
was only a single, rather weakly developed papilla near the 
anterior end of the lateral margin of each choana. A larger 
juvenile (MCZ 3454) possessed two papillae on the right side and 
three on the left. 

In section, each papilla can be seen to consist of two major 
histologic elements. First there is a thick, cornified, stratified 
squamous epithelial covering, and secondly a mass of dense con- 
nective tis.sue forming a central core. Plate II shows transverse 
sections of papillae. The upper section, stained with hematoxylin 
and eosin, was tjiken near the center of the i)apilla. while the 
lower one, stained by the Bielschowsky silver method, was taken 
near the base. 

The epithelium closply resembles normal body epidermis, lu 
some regions all four strata — germinativum, granulosum, hu-i- 
dum, and corneum — are Avell developed. The first and last are 
l)resent throughout, but the granidosum and lucidum are not 
always recognizable. There is a gradual increase in the thickness 



1958 CHOANAL PAPILLAE OP CHELONIIDAE 3 

of the stratum eorneum from a minimum at the l)ase of the pap- 
illa to a maxinuim at its tip. The external surface of the epi- 
thelium is relatively even at the distal end of the papilla, hut 
hasally it becomes in-e^ular with numerous small projections 
and. more rarely, indentations. As in typical epidermis, the 
boundary betAveen the epithelium and the underlying connective 
tissue is generally a rather wavy line, although near the tip of 
the papilla it becomes far more even. No basement membrane 
eould be seen ; in fact the tibers of the connective tissue appear 
to continue into the basal epitlielial cells. However in human 
epidermis, which a])pears sinular when viewed through a light 
microscope, a thin l)asement membrane can be demonstrated by 
electron microscopy (Maximow and Bloom, 1957). 

The core of the papilla is a mass of typical dense connective 
tissue. Irregularly arranged bundles of collagenous fibers occupy 
most of the space, but there are numerous included cells and 
small relatively clear areas. The cellular constituents of the con- 
nective tissue were not identified in the present study. Within 
the connective tissue tlicre are (piite numerous very small blood 
ves.sels; however only a single arteriole and venule could be 
found at the base of the papilla, and the tissue is not very heavily 
vascularized. No nerve fibers or endings could be recognized. 
However nerve fibers would resemble most closely the fibers 
of the connective tissue, and thus be very difficult to demonstrate. 
Presumably there are some nerves present; certainly the lining 
of the oral cavity is well innervated in most forms. 

OTHER CHELONIIDS 

Plate I, lower figure shows a specimen of Cantta carcita. In 
this specimen (MCZ 53183) there is a single choanal papilla 
located near tlie anterior end of the lateral margin of the choana. 
This is in agreement with the findings of both Deraniyagala 
(1939) and Carr (1942). The form of the papilla is essentially 
similar to that found in Chelonia mydas, but it appears to be 
slightlv shorter and broader in Carefta. 

The only specimen of Chelonia depressa which was examined 
(MCZ 54712) is a juvenile, and possesses a single choanal papilla 
exactly like that of Chelonia my das juveniles. 



4 BREVIORA No. 85 

In Eretmochelys imhricata, Deraniyagala (1939) found a 
single choanal papilla, and Carr (1942) reports one or two 
papillae. All seven specimens examined for the present study 
(MCZ 1138, 1141, 1416, 1418, 4049, 49406, and 49407) possess 
a single papilla exactly like that of Caretta. 

No specimens of LcpidocJuiys were seen by me. Both Derani- 
yagala (1939) and Carr (1942) report a single choanal papilla 
like that found in Caretta. 

FUNCTION OF THE PAPILLAE 

In adult specimens of Chelonia mydas the form of the papillae 
suggests the possibility that their function is to prevent pieces 
of food from entering the nasal cavities. However the single 
choanal papilla of the other cheloniid genera and of juveniles 
of Chelonia mydas could not be very effective in this regard. 

According to Deraniyagala (1939), all the cheloniids are omni- 
vorous, preferring an animal diet but often subsisting on purely 
plant food. Chelonia in particular is primarily hei'bivorous. Its 
eating habits also differ slightly from the other members of the 
family in that its "stomach contents are never in a finely divided 
condition but appear to be 'bolted' entire" (Deraniyagala, 1939. 
p. 234) ; the other cheloniids tend to break up their food more. 
However it does not appear to me that an unchewed piece of alga 
would cause any more injury than a broken piece of mollusc 
shell if lodged in the choana or nasal cavity. Thus food habits 
do not seem to ex])lain the differences between the genera. 

Another possible function of the papillae is that they might 
bear some special sensory receptors. However the liistological 
observations provide no evidence for such a theory, although they 
are not absolutely conclusive. As was noted above, no nerves 
or sensory endings were observed in the papillae of Chelonia. 
Since the material was not ideally preserved for histologic 
study and since nerve fibers would not be clearly differentiated 
from connective tissue fibers by the stains employed, their ab- 
sence cannot be considered as proven ; in fact, it appears highly 
probable that there is some sensory innervation of the papillae. 
However the absence of any conspicuous nerve trunk entering 
the base of the papilla and of any distinguishable receptors 
renders it most unlikely that the papillae function primarily in 



1958 (CHOANAL PAPILLAE OF CUELONIIDAl': 5 

a sensory capacity. Furthermore, the thickness and high degree 
of cornification of the epithelium would be surprising, though 
not impossible, in a sensory structure. 

Thus I am unable to suggest any function for the single 
papilla in Caretta, Eretmochelys, or Lepidocluhjs. In Chdonia 
(adults) they must serve to prevent the contents of the mouth 
from entering the nasal cavities ; whether or not this is their 
primary function cannot be stated. 

BIBLIOGRAPHY 

Cakr, a. F., .Ik. 

1942. Notes on sea turtles. Proc. New England Zool. Chili, vol. 21. 
pp. 1-16. 

Deraniyagala, p. E. P. 

1939. The tetrapod reptiles of Ceylon. Volume 1. Testiidinates and 
erocodilians. Columbo. 412 pp. 

.Ma.xi.mow, a. a., and W. Bloom 

1957. A textbook of histology, 7th ed. Philadelphia. 628 pp. 





I'LATE I 

Upper, Chelonia mi/da.s. \'iew of part of the palate showini; ilic cho'inal 
papillae. 

Lower, Caretla r(ir(thi. \\v\s of tlii' loft side of llu- palate slidwiiiy 
the single choanal papilla. 




^' '^ 




IM.A'I'K II 

i'pper, Cluloiiia iiii/da.-^. Transverse section of a choanal papilla taken 
near its center and stained with hematoxylin and eosin. 250x. 

Lower, Chelonia mydas. Transverse section of a choanal papilla taken 
near its base and stained by the Bielschowskv silver method. 2o0x. 



BREVIORA 

MiuiseiLim of Comparative Zoology 

Cambridge, Mass. May 29, 1958 Number 86 



A NEW SICISTINE RODENT FROM THE 
MIOCENE OF WYOMING 

By Craig C. Black ^ 

During the course of a study of the Split Rock local fauna of 
Middle Miocene age, several isolated teeth, two partial maxil- 
laries, and one fairly complete lower jaw of a new sicistine rodent 
were discovered. Because of the rather limited occurrence of this 
group in North America and the paucity of previous records, 
it is felt that this form should be described before the study of 
the fauna as a whole can be completed. 

For the loan of material described below I am indebted to 
Drs. P. 0. McGrew, Loris Russell and A. E. Wood and to the 
Chicago Natural History Museum. I would like to acknowledge 
the helpful assistance and suggestions of Professor A. E. Wood 
and Professor Bryan Patterson. 

The following abbreviations are used throughout : 

A.C. — Amherst College 

C.N.H.M. — Chicago Natural History Museum 

E.O.M. — Royal Ontario Museum of Zoology and Paleontology 

U.W. — University of Wyoming 

Family ZAPODIDAE 
Subfamily SICISTINAE 

SCHAUBEUMYS SABRAE,^ U. Sp. 

Holotype. U. W. No. 1405, partial left mandible with the 
incisor. Mi, M2, and the anterior roots of M3. 
Referred Specimens. Partial maxillaries with P'^-M^ A.C. 

1 This study was made while the author was Rufus B. Kellogg Fellow from 
Amherst College. 

2 The specific name is given in honor of my wife. 



2 BREVIORA No. 86 

No. 10490, R.O.M. No. 1950; Mi A.C. Nos. 10489, 10491, 10493, 
10494, 11293, C.N.H.M. Nos. PM 2101, 2102, U.W. No. 1406; Ji 
C.N.H.M. Nos. PM 2105, 2106; Mi A.C. Nos. 10492, 11294; Mo 
C.N.H.M. No. PM 2103 ; M3 C.N.H.M. No. PM 2104. 

Diagnosis. Intermediate in size between Plesiosminthus? 
clivosus (Galbreath, 1953, p. 99) and Shauheumys grangeri 
(Wood, 193ob, p. 1) ; no anteroeonid on Mi ; mesostylid closely 
applied to metaconid on M2 ; mesolophid connected to metaconid 
on M2 ; hypoconulid well-developed on M1-M2. 

Horizon and Locality. Middle Miocene. Vicinity of Brachy- 
crus quarry "Exposure No. 2" (Schultz and Falkenbach, 1940, 
p. 251), seven miles by road northwest of Muddy Gap filling 
station, in a draw about 14 i^ils south of U.S. Highway 280, 
Fremont County, Wyoming. 

Description. The jaw (Fig. 1 H) is broken through the middle 
of the alveolus of M3 and down the outer half of the body so 
that the incisor is revealed beneath Mo. The ridge marking the 
lower border of the masseteric scar is thick and prominent and 
extends forward to below the anterior root of Mi. The mental 
foramen lies almost immediately below the center of the diastema. 

The upper teeth are associated in several cases with fragments 
of palate, the most complete of which is shown in Figure 1 A. 
The posterior boundary of the anterior palatine foramen is on 
a line with the anterior end of P*. A small part of the zygomatic 
arch is preserved, arising somewhat anterior to P"^, with the 
ventral border of the infraorbital foramen preserved. However, 
only that portion of the foramen which transmits the infra- 
orbital nerve is present so that no estimate of the size of the 
opening for the masseter can be made. 

There are a number of isolated upper incisors (Fig. 1 F, G) 
which agree so closely with the description given by Schaub 
(1930, p. 621, Fig. 4) for those of the European sicistine, 
Plesiosminthus schauhi Viret, that they are here referred to the 
North American form, Schauheumys sahrae. They are prom- 
inently grooved with the enamel extending 0.2 to 0.3 mm. over 
the medial side of the tooth. The pulp cavity is Y-shaped, the 
median arm of the Y being the longer. 

The upper fourth premolars (Fig. 1 A, E) are bud-like in ap- 
pearance with two small buccal cusps, a deep central pit, and a 
slight lingual swelling. The upper first molars (Fig. 1 A-E) 



1958 



NEW MIOCENE SICISTINE RODENT 



are almost square. The paracone and metacone are higher than 
the other cusps. The auterocone and anterior cingulum are well 
developed with the latter sloping steeply buccally to join the 







Figure 1. Teeth of Schaubewmys sairae, all X 10 except for F and G 
X 5. A, R.O.M. No. 1D50, part of palate with RP^-Mi; B, A.C. No. 10491, 
LMi; C, A.C. No. 10489, LMi ; D, A.C. No. 10494, LM^; E, A.C. No. 
10490, RP4-M1; F, C.N.H.M. No. PM 2105, Ii, lateral side to right; G, the 
same, lateral view; H, U. W. No. 1405, partial left mandible with Ii, Mi, 
and M2 (type) ; /, U.W. No. 1405, LM1-M2; J, U.W. No. 1405, Ii, lateral 
side to right; E, A.C. No. 11294, LMi ; L, C.N.H.M. No. PM 2103, LMo ; 
M, C.N.H.M. No. PM 2104, RM3. Anterior end to the left except for A, E, G, 
and M where it is to the right, and F and J where it is at the top. 

paracone at its base. The paracone and protocone are usually 
connected through what is probably the protolophule II, either 
directly, or via the mesocone. In one specimen (Fig. 1 B), 



4 BREVIORA No. 86 

however, there is no trace of the protoloph. The mesoloph 
extends to the buccal margin of the teeth and is occasionally 
connected to the paracone, metacone, or both, by low lateral 
ridges. The posterior cingulum lies below the level of the meta- 
loph. 

The lower incisors (Fig. 1 J) are triangular in outline with 
a flat anterior face and a small elliptical pulp cavity on the wear 
surface. The enamel extends slightly over the medial side of 
the tooth. 

There is no anteroeonid on the first lower molars (Fig. 1 I, K), 
but on A.C. No. 11293 (Fig. 1 K) there is a small distinct 
cuspule in the deep basin between the protoconid and the meta- 
conid. This cuspule is not on the anterior cingulum, however, 
and there is no trace of such a cuspule on either of the other 
two first low'er molars. In all the specimens there is a strong 
ridge or arm rising steeply from the anterior cingulum to the 
protoconid while lingually the cingulum abuts against the base 
of the metaconid Avith no indication of such a cingular arm. 
In two instances the mesolophid is w^ell-developed in contrast to 
the distinct mesostylid and mesoconid of U.W. No. 1405 (Fig. 
II). The hypoconulid is distinct in all cases and closely con- 
nected to the hypoconid. The posterior cingulum descends steeply 
from the hypoconulid and passes to the lingual edge of the tooth 
terminating well below the entoconid. 

The second lower molars (Fig. 1 I, L) are in general similar 
to the first molars with the one exception that in the former 
the mesolophid has shifted anteriorly abutting against the meta- 
conid. The mesostylid is thus distinct, lying half way down the 
steep metaconid slope. The protoconid and metaconid are con- 
nected, probably through the metalophulid II. The mesolophid 
and metalophulid II enclose a small pit between the protoconid 
and metaconid. 

The third lower molar (Fig. 1 M) is proportionately smaller 
than any of the first or second molars so far encountered, but 
since so few specimens are available no dependable ranges for 
tooth size can be determined. The anteroeonid is not distinct, 
but blends into the two cingulum arms, one of which passes to 
the lingual margin while the other slopes downwards to the base 
of the protoconid. The mesolophid passes anteriorly to unite 



1958 NEW MIOCENE SICISTINE RODENT 5 

with the metaconid at its base, forming a deep pit between the 
metalophid and mesolophid. The mesoeonid and entoconid are 
not distinguishable within the crests. The hypoconid is large 
and is connected to the entoconid region through the posterior 
cinguluni and the anterior hypoconid arm, the two isolating a 
posterior pit. There is also a central pit between the mesolophid 
and hypolophid. 

Discussion. Schau^eumys sdhrae is the third mid-Tertiary 
species of sicistiue to be reported from North America. The 
species previously described are Schaubeumys grangeri (Wood, 
193ob) from the Lower Rosebud, and Plesiosminthusf clivosus 
(Galbreath, 1953) from Quarry A in NE Colorado, of Middle 
Miocene age. I have assigned the present population to the 
North American genus, Schaubeumys, rather than to the Euro- 
pean genus, Plesiosnmithus, because I do not feel that at present 
there is enough evidence available to warrant considering the 
North American and European forms congeneric. Galbreath 
(1953, p. 99) was of the same opinion when, in his discussion of 
P.? clivosus, he stated, "It seems unlikely that this North 
American zapodid from the middle Miocene is congeneric with 
the lower Miocene zapodid of western Europe." I also feel that 
the three North American species are congeneric and that PJ 
clivosus should be referred to Schaubeumys. 

The North American forms can be distinguished on the basis 
of size: S. clivosus is the smallest; S. sabrae is intermediate; and 
8. grangeri is the largest. S. sabrae is strikingly different from 
either of the others in the absence of the anteroconid on M^ and 
in the fact that the mesolophid abuts against the metaconid and 
fails to reach the mesostylid on M2. 8. clivosus appears to have 
no distinct hypoconulid on any of the lower teeth whereas this 
cusp is relatively large in the other two species. 

It is of interest to note that all three species occur in a rela- 
tively limited time span, no sicistines yet being known in North 
America from any deposits earlier than Miocene (I am excluding 
from the Zapodidae the peculiar Simimys from the Sespe 
Eocene), or later than Middle Miocene. This brings up the 
problem of where the North American sicistines came from. 
No definite answer is possible but some guesses may be hazarded. 



6 BREVIORA No. 86 

All the North American species show a great many similarities 
to contemporaneous forms in Europe, and to forms known from 
the Oligocene of Europe and Mongolia. In fact, if it were not 
for the presence of grooved upper incisors, *S'. sabrae might be 
referred to the Mongolian genus Parasminthus described by 
Bohlin in 1946. Parasminthus in turn is equally close to the 
European Plesiosminthus (Bohlin, 1946, pp. 16-17), from which 
it is separated only on the basis of the non-grooved incisors. 

Wood (1935a, pp. 225-226) has pointed out that it is quite pos- 
sible that the presence or absence of a groove on rodent incisors 
may be controlled by a single gene. He goes on to state that if 
this is the case such grooving could be lost and regained within 
one lineage practically at will. In the case of the sicistines this 
would mean that the North American forms could be derived 
from the Mongolian complex and that the European forms could 
also be so derived, or vice versa. Bohlin (1946, p. 43) points out 
that the structural differences observed between species of Para- 
sminthus are of the same kind as those found between the species 
of Plesiosminthus. This is also true when the North American 
complex is compared with either Parasminthus or Plesiosmiyithus. 
These complexes show the presence of the anteroconid in some 
species and its absence in others. The degree of development of 
the mesolophid and the position of the mesostylid vary in a 
similar manner in each group. The way in which the lophs 
are formed also shows the same variation, and, finally, the same 
size ranges occur in each group. This great morphological 
similarity would seem to indicate that these three complexes are 
basically similar genetically. (No attempt has been made here 
to determine generic limits within the Miocene sicistines.) 

TABLE OF MEASUREMENTS ^ 



11 


Number of 
Specimens 


Me.in 


a 


c.v 


anteroposterior 


8 


1.20 


.10 


8.3 


transverse 


8 


.74 


.05 


6.8 


P4 










anteroposterior 


2 


.55 






transverse 


2 


.65 







1 All measurements in nun. o = Standard Deviation, C.V. = Coefficient of 
Variation. 



1958 NEW MIOCENE SICISTINE RODENT 



Ml 










anteroposterior 


10 


1.32 


,08 


6.0 


transverse 

Ii 

anteroposterior 


10 


1.29 


.10 


7.8 


1 


.85 






transverse 


1 


.60 






^h 










anteroposterior 


3 


1.32 






transverse 


3 


1.02 






Mo 










anteroposterior 


2 


1.28 






transverse 


2 


1.11 






Ms 










anteroposterior 


1 


1.03 






transverse 


1 


.91 







EEFEEENCES 

BOHLIN, B. 

1946. The Fossil Mammals from the Tertiary Deposit of Taben=buluk, 
Western Kansu. Part II: Simplicidentata, Carnivora, Artio- 
dactyla, Perissodactyla, and Primates. Palaeont. Sinica, N. S. 
C, 8b:l-259. 

Galbreath, E. G. 

1953. A Contribution to the Tertiary Geology and Paleontology of 
Northeastern Colorado. Univ. Kansas Publ., Contr. Paleont., 
Yertebrata, 4:1-119. 

SCHAUB, S. 

1930. Fossile .Sieistinae. Ecologae Geologicae Helvetiae, 33:616-637. 

SCHULTZ, C. B. AND C. H. FaLKENBACH 

1940. Merycochoerinae, a New Subfamily of Oreodonts. Bull. Amer. 
Mus. Nat. Hist., 77:213-306. 

Wood, A. E. 

1935a. Evolution and Eelationships of the Heteromyid Rodents with 

New Forms from the Tertiary of Western North America. Ann. 

Carnegie Mus., 24:73-262. 
193ob. Two New Genera of Cricetid Rodents from the Miocene of 

Western United States. Amer. Mus. Novit., 789:1-3. 



BREVIORA 

MuiseiLinti of Comparsitive Zoology 



Cambridge, Mass. June 20, 1958 Number 87 



AX EMBOLOMERE JAW FROM THE 
MID-CARBONIFEROUS OF NOVA SCOTIA 

By Alfred Sherwood Romer 

It is not always realized that our knowledge of Carboniferous 
tetrapods is almost entirely confined to fornLs of relatively late 
age. Apart from a meager fauna from Scotland (ef. Watson 
1929) and a single specimen from England, all described Car- 
boniferous amphibians are of Westphalian or later date — i.e., 
from Middle to Upper Pennsylvanian strata (Romer 1947, p. 
322). The jaw here figured is the first specimen to he described 
from any American pre-Westphalian formation.^ 

in 1956 the Museum of Comparative Zoology undertook a sum- 
mer's exploration of the Carboniferous deposits of Nova Scotia 
in search of vertebrate fossils. Some excellent material of West- 
phalian amphibians was obtained, and will be described later. 
The present specimen is the only pre-Westphalian amphibian 
identified in our collections. 

It was derived from the Point Edwards formation (Hyde 
1913, p. 252), considered by Bell (1938; 1944, pp. 11-12. 16) 
to belong to the Canso group. It is definitely earlier than the 
Riversdale series, with a flora of Westphalian A type, and 
younger than the Windsor series of the Mississippian. It thus 
appears to l)e e([uivalent to some horizon in the Namurian of 
European nomenclature and is currently considered as late 
Mississippian (Weller et al. 1948, p. 173, chart 6). 

Exposures of the Point Edwards formation occur only in 
limited areas adjacent to Sydney Harbor in Cape Breton. They 
ai-e well shown in the series of maps illustrating the geology of 

1 I hoiii' to describe at a later time remains from the Mississippian of West 
Virginia (ef. Romer 1941). 



2 BREVIORA Xo. 87 

the Sydney Coalfield which were published by the Canadian 
Bureau of Geology and Topography in 1938. The best exposures 
are along- the shores of Point Edward. A Gyracmithus spine was 
collected there many years ago. Our group in 1956 discovered a 
beach exposure containing dipnoan and other fish bones about 
one-half mile southwest of the point. The present specimen was 
found on the shore about 100 yards southeast of the tip of the 
])oint, on a detached slab of rock. An incomplete Gyracanthu.s 
spine and fragmentary fish remains were found nearby in the 
ledge from which the jaw slab was derived. On another loose 
slab was a crushed bone, about 50 mm. long and 29 mm. broad at 
one end, probably a Sagenodus quadrate. Intensive search of 
the area failed to joroduce further amphibian material. 

The specimen consists of a slab of calciferous siltstone which, 
with its missing counter])art, had contained the greater part of 
the right lower jaw of a large laljyrinthodont amphiljian. Wave 
action had removed all the bone, leaving, however, an almost j^er- 
fect mold of the outer surface, from which excellent casts have 
been made. The portion preserved extends from the symphysis 
to a diagonal line running down and forward from a point a 
short distance l)ehind the termination of the tooth row ; the great- 
est length as preserved, measured in a direct line, from the 
symphysis dorsally is 235 nun ; the length of the tooth row is 
191 mm. The shape of the jaw, as preserved, is that common to 
many early labyrinthodonts, particularly embolomeres ; super- 
position of our fragment on an outline of such embolomeres as 
Archeria and Ptcroplax suggests a total length of jaw — and 
hence, approximately, of skull — of about 365 mm. This indi- 
cates an animal of considerable size, this measurement being 
within 10-15 ])er cent of that of typical skulls of the familiar 
Permian labyrinthodont Eryops. If outlines of the jaws of such 
familiar embolomeres as Fteroplax and Archeria be superposed 
on that of our specimen by using as a common measurement the 
length of the tooth row, it will be found that the jaw ramus is 
intermediate in proportions between the long- and slender 
Archeria jaw and the relatively deep and short ramus of Ptero- 
plax. There are few non-embolomere jaws of Carboniferous age 
with which comparison can be made, and little reliance can be 
l)laced on propoi-tions of the jaw ramus in attemi)ts to determine 



1!)5S EMBULOMEKK JAW FKOAl NOVA SCOTIA 3 

relationships. If, howevor, a scries of represontative jaws of 
labyrinthodoiits bo exainiiicd (ef., for example, Komer 1J)47. 
figs. 9, 10), it will be seen that in general there is little tendency 
for the dcveloinnent of an "ascending rannis" in tiie snrangnlar 
region in teninospondyls, but some trend in this direction in 
seymonriamorphs, and in embolomeres a strong development of 
depth posteriorly, with a notable nji-swing of the dorsal margin 
of the jaw posteriorly. This last trend is obvions in the specimen 
as preserved, and the edge of the jaw back of the end of the 
tooth row shows a thin broken margin, indicating that in life 
the jaw curved upwaril posteriorly still more strongly. This 
posterior depth strongly suggests that the jaw is that of an 
anthracosaurian (as I have used that term) and ])ro) ably an 
embolomere. It is regrettable that the inner surface, Avliere posi- 
tive embolomere features are to be found is not available. 

When the cast is viewed dorsally, it is seen that although the 
mold is slightly incomplete in the symphysial region, the jaw 
was not widely bowed outward as in such a broad-snouted type 
as Eryops, but turns sharply backward to run a straight course 
for almost the entire length of the portion preserved. 

The surface has an ornamentation of pits anteriorly and ex- 
tending back along dorsal and ventral margins; tlorsally, the pits 
give way posteriorly to deeply incised striations. Between dorsal 
and ventral margins the surface shows a lighter series of antero- 
posterior striations. At the posterior end of the ventral margin 
there is part of a deep groove carrying the mandibular lateral- 
line canal ; farther forward the mold does not extend quite to 
the ventromedial line of the ramus and hence the anterior pro- 
longation of the canal is not visible on the cast. 

The jaw was crushed before fossilization. As is frequently the 
ease in labyrinthodont specimens (and well shown in an 
Archeria jaw before me at the moment), the outer surface was 
crushed downward (i.e. medially) in the area between the thick- 
ened dorsal part of the jaw carrying the tooth sockets and the 
stout ventral margin. Sutures are, for the most part, not ap- 
parent and since we are dealing with a mold only, further devel- 
opment is impossible. It seems certain that the uncrushed upper 
margin is entirely, or almost entirely, included in the dentary, 
and it mav be that the longitudinal line of breakage below this 



4 BREVIORA No. 87 

reofion is on or close to the suture between the dentary and the 
infra-dentary series. An area below this line of breakage, in 
which parallel striae extend forward and slightly downward, 
appears to be part of the surangular, which thus extends re- 
markably far forward (as in Kotlassia) . Below this region, 
posteriorly, is one in which the striae extend diagonally an- 
terodorsally ; this seems surely part of the angular. The rugose 
ventral margin was surely occupied by postsplenial and splenial. 
The pertinence of the wedge-shaped anterior part of the finely 
striated area is uncertain. 

A sympliysial tooth pair is present, with both members de- 
veloped and striated in labyrinthodont fashion. Much of the 
lateral tooth row is preserved. The most anterior teeth seen on 
the cast, lying close to the symphysis, appear to be somewhat 
crowded and irregularly placed. These are conical teeth of 
modest size, striated basally and slightly recurved at the tips. 
One complete tooth extends 8 mm. above the jaw margins; an 
adjacent incomplete tooth appears to have been several milli- 
meters higher. For 5 cm. back of this region the teeth are 
l)roken oft', but the alveolar surface of the dentary bone is pre- 
served. Here there is seen a typical labyrinthodont arrangement 
of alternate teeth (broken) and empty alveoli; the interval be- 
tAveen successive elements of the row is about 4 mm. The tooth 
bases suggest that the teeth are comparable in size to those seen 
at the anterior end, without the development of any large 
"fangs." For the next 10 cm. the mold does not extend inward 
beyond the outer rim of the dentary, so that the alveolar region 
is not visible and there are impressions of only 5 teeth in all, 
some of them obviously loose from their sockets. The most 
anterior of these teetli appears to be comparable to those seen 
near the symphysis; the others give the impression of greater 
slenderness, although this may be due to incompleteness of the 
mold. For the last 3 cm. of the series the alveolar surface is 
again visible, and 5 of the 6 most posterior teeth are present, 
tilthough none is complete. Naturally these are smaller than the 
more anterior teeth, and the interval between successive elements 
is reduced to about 8 mm. There is little indication that the teetli 
were markedly recurved or had the chisel-like tijis seen in 
Archcria. Calculation suggests that the total marginal row con- 
sisted of about ')() teetli, which were rather closely si)aced and 
of modest size. 



1958 EMBOLOMERE JAW PROM NOVA SCOTIA 5 

AVhat arc tlic relationships of tliis specimen? Thei-e are no 
described American i)re-Westphalian amphibians witli which 
cominii-ison can be made. It was at one time 1 elieved that the 
older teti-apod faunas of Europe and North America were as 
radically distinct as are tlie reptile and amphil)ian faunas today, 
and tliat trans-Atlantic comparisons were invalid. However, 
such recent work as my study of the Kounova fauna (1945) and 
Baird's current (unpublished) studies of the Linton fauna indi- 
cate that eastern North America and western Europe were ex- 
tremely similar in faunas in the Carboniferous, and that the 
supposed contrasts in genei'a present in the two areas are due in 
"Teat measure to false assumptions as to faunal contrasts and 
lack of comparison of American and European materials. 

But even so. the number of pre-Westphalian specimens from 
Europe, with which this jaw can be lefjitimately compared, are 
few (Watson lf>29). There are several loxommids from the late 
Mississippian and early Pennsylvanian of Scotland, but this 
jaw is (^uite different from that of known loxommids. in which 
the jaw is shallow posteriorly, and in which the dentition con- 
sists of a smaller number of much more rol)ust teeth. The straigrht 
ramus of the jaw indicates an animal with a more or less pointed 
snout (altliouji'h the skull may have broadened posteriorly, with 
a triangular sha]^e). This appears to eliminate from considera- 
tion Otocrat'd and Palaeogyrinus and perhaps Crassigyrinns. 
The massive upper teeth of Antliracosaurus (in the proper 
sense) make it improliable that our jaw is pertinent. Two Scot- 
tish forms remain — Pholidogastcr from the Mississippian (Gil- 
merton Ironstone), and a skull attributed by Watson to 
Pholidrrpeton from the Lanarkian of Airdrie. Both have a 
pointeil snout and, appropriately, a marginal dentition includ- 
ing a long series of small teeth. Of Pholidogastcr, the type skele- 
ton (Huxley 1862; Watson 1929, pp. 230-233, tigs. 7-9) is that 
of an animal much smaller than the present one ; a skull from 
tlie same beds as the type, and which may l)e a specimen of the 
same form, is of somewhat larger size, with a length, measured 
to the (piadrate of about 156 nun. The lower jaws are present of 
the type specimen ; they do not appear to show the increasing 
depth posteriorly which is characteristic of this jaw, as of em- 
holonieres (which PJwIidogaster is not). Of Pholiderpeion, the 



6 BREVIORA Xo. 87 

type (Huxley 186!)) is of Westplialiaii age, but Watson (1929, 
223-224, pi. I, fig'. 2) assigned to this form a skull of earlier date. 
The latter skull appears to have had a length of approximately 
405 mm., a length comparable to that estimated for our speci- 
men. I have not seen the Manchester Mviseum slab containing 
part of this specimen but from Watson's description and the 
Museum of Practical Geology specimen it -would appear that 
the jaw (definitely of embolomere type) is comparable to that 
described here. Plence, although the matter lies at two removes 
from certainty, our specimen may be provisionally assigned to 
PhoJidcrpcton. It may be reasonably questioned whether a frag- 
ment of this sort should receive a specific name. iSince, however, 
it is of considerable interest from stratigraphic and topographic 
points of view, 1 will, nevertheless, describe it as Pholiderpe- 
louC^.) hrrtonensis, sp. nov. A specific diagnosis is, of course, 
valueless, but since this is required by the ''regies," I will men- 
tion as a supposed sjiecific character the observable gently 
striated nature of the extei-nal jaw scul]iture between the coarser 
sculpture bands of dorsal and ventral borders. Tlie holotvpe is 
M.C.Z. Xo. 2772. 

Althorgli it now seems probable that many of the Carbon- 
iferous labyrinthodonts once thought to be embolomeres are 
members of other groups — rhachitomes, ichthyostegalians, or 
pre-seymouriamorph anthracosaurians — there nevertheless re- 
mains a "hard core" of typical embolomeres which carry the 
story of the group downward through much of the Carboniferous. 
Archcria is the last survivor; Pteropla.v {'' Eoyyrinus") is a 
typical Westphalian representative; the skull assigned to PhoJi- 
derpeton by Watson and the present jaw carry the story down 
to or across the Pennsylvanian-Mississippian boundary, and 
undescribed materials from West Virginia show the presence of 
a true emlolomere in Upper ^lississippian times. The trend 
within the group appears to 1 e the development of persistently 
a(|uatic fish-eating ampliibians with an elongated body and 
tail and short limbs, and, as in piscivorous vertebrates of many 
groups, a trend toward development of a long slendei- snout and 
a long row of relatively small marginal teeth. 

In this first of several projected papers on our Nova Scotia 
materials, 1 wish to express our thanks to the X'^ational Science 
Foundatiou, Avhose grant for fiekl expenses made the summer's 



1908 EMHOLO.MKIiK .lAW I'I{()M NOVA SCOTIA 7 

work {)(>ssil)le, and. further, our gratitude to others who aided 
and facilitated our work. Dr. II. D. Smith, of tlie Nova Scotia 
Research Foundation, was most helpful, as was Mr. John Ilaslam 
of New Glasji'ow. in charge of mining operations at Stellarton 
and Springhill. Dr. W. A. Bell gave us the benefit of his expert 
knowledge of the Nova Scotia Carboniferous. To the Nova Scotia 
Bureau of Mines and the Massachusetts Institute of Technology 
we owe thanks for generous hospitality at the summer geological 
camp at Crystal Clitfs, conducted by Dr. Walter 0. Whitehead. 

LITERATURE CITED 

Bell, W. A. 

1938. Fossil rtoia of Sydney Coalfield, Nova Scotia. Canada, Dept. 

-Mines, Mem. Geol. Surv., 215: 1-115. 
1944;. Carboniferous rocks and fossil floras of northern Nova Scotia. 

Canada, Dept. Mines, Mem. Geol. Surv., 238: 1-119. 

IlUXLEY, T. II. 

1862. On new labyrinthodonts from the Edinburgh Coal-field. Quart. 

•Tour. Geol. See. London, 18: 291-296, pi. 11. 
1869. On a new labyrinthodont, from Bradford. Quart. Jour. Geol. 

Soc. London, 25: 309-310, pi. 11. 

Hyde, J. E. 

1913. The Carbonifei'ous sections on Sydney Harbour ; Xlllth Internat. 
Geol. Cong., Excursion in Eastern Quebec and the Maritime 
Provinces, Guide Book No. 1, pt. 2, pp. 251-262. 

ROME», A. S. 

1941. Earliest land vertebrates of this continent. Science, 94: 279. 
1945. The late Carboniferous vertebrate fauna of Kounova (Bohemia") 

compared with that of the Texas redbeds. Amer. Jour. Sci., 243: 

417-442. 

1947. Review of the Labyrinthodontia. Bull. Mus. Conip. Zool., 99: 
1-368. 

Watson, D. M. S. 

1029. The Carboniferous Amphibia of Scotland. Palaeont. Hungarica, 
1:221-252. 

Wkller, J. M. (Chairman) and others 

1948. Correlations of the Mississippian fornuitions of North America. 
Bull. Geol. Soc. Amer., 59: 91-196. 




I 



h- 



BREVIORA 

Miiseimi of Comnparative Zoology 



Cambridge, Mass. June 30, 1958 Number 88 



A NEW SPECIES OF THE GENUS UROTHECA 
(SERPENTES: COLUBRIDAE) PROM VENEZUELA 

By J. A. RozE 

Escuela de Biologia 
Universidad Central de Venezuela 

While touring several U. S. museums in 1957 I discovered 
among the Venezuelan herpetological material in the Museum 
of Comparative Zoology, Harvard University, a very peculiar 
undescribed species of the genus Urotheca. Later I found that 
the U. S. National Museum (USNM), the Carnegie Museum, 
Pittsburgh (CM), and Museo de Biologia, Universidad Central 
de Venezuela (MBUCV) also have one specimen each of this 
form. I take the opportunity to describe this new species. 

I express my appreciation to Dr. E. E. Williams, Dr. D. M. 
Cochran and Mr. N. D. Richmond, not only for permitting me 
to examine the collections at their museums, but also for other 
kindnesses I received during my stay in the United States. 

I would also express due thanks to the Fundacion Creole, 
Caracas, Venezuela, and to the Council Research Fund, American 
Museum of Natural History, whose help enabled me to take this 
study tour. 

Urotheca williamsi n. sp. 

Leimadophis bimaculatus opisthotaenia, Marcuzzi, 1950, Nov. Cient. La Salle, 
Ser. Zool., No. 3, p. 8. 
Holotype. 9 MCZ No. 51329, collected by G. Marcuzzi in 
October, 1949. 



2 BREVIORA No. 88 

Type locality. El Junquito, Distrito Federal, Venezuela, 1900 
meters. 

Paratypes. $ USNIVI No. 121206 collected in Colonia Tovar, 
estado Aragua, Venezuela, 1800 meters, by E. G. Holt on March 
30, 1929; $ CM No. 7393, collected in Colonia Tovar, estado 
Aragua, Venezuela, 1800 meters, by E. G. Holt on April 30, 1929 ; 
9 MBUCV No. 3044, collected in Rancho Grande, estado Aragua, 
Venezuela, 1200 meters, by J. Racenis on August 23, 1949. 

Distribution. The Central Cordillera de la Costa in northern 
Venezuela, between 1000 and 2000 meters. This zone is covered 
by tj'-pical cloud forest (high rain forest). 

Diagnosis. Urotheca with one scale pit on the dorsals ; 17-18+2 
maxillary teeth ; 146 to 158 ventrals ; 53 to 61 subcaudals ; 17-17 
dorsals, without any reduction ; 1 or 2 preoculars, and a Leima- 
dophis-like coloration : dark longitudinal lateral lines beginning 
on the anterior part of the body and extending to the tip of the 
tail. 

Description of the holotype. Maxillary teeth 17+2, the last 
two somewhat enlarged and separated from the rest by a gap. 
Mandibular teeth subequal. Head squamation that of Urotheca, 
rostral scarcely visible from above, nasals divided, the nostrils 
situated on the dividing suture. A pair of internasals and pre- 
frontals, the latter a little broader than long and longer than 
the internasals. The frontal shield twice as long as the prefrontals 
and approximately as long as the parietals. A loreal shield, 
deeply penetrating the preocular, and reaching almost to the 
orbit. One preocular, trapezoidal, twice as wide above as below. 
Two postoeulars, the upper much larger than the lower. Tem- 
porals 1+2 on both sides. Seven upper labials, the third and 
fourth entering the orbit. Nine infralabials, the first four in 
contact with the anterior pair of chin shields, of which the first 
pair is longer than the second. Ventrals 150, anal divided, and 
53 pairs of subcaudals. The dorsal formula 17-17, without any 
reduction. 

Coloration. Head and upper part of the body grey (brown in 
life?). Upper lip black, crossed by a white line, which begins 
on the rostral shield and disappears on the neck. Infralabials 
and other ventral shields white, heavily mottled with black. This 
mottling is present also on the ventrals, where it concentrates 



1958 NEW SPECIES OF UROTHECA 3 

mainly along the borders of the ventral scales, but fades gradu- 
ally posteriorly on body. On mid-line of the subcaudals the 
black coloration appears again as an irregular ill-defined dark 
line. 

The black supralabial coloration continues on the neck as a 
series of black blotches, which on the anterior third of the body 
form a lateral black line, at first occupying the whole of the 
scales on the fourth dorsal row, but more posteriorly shifting 
to the lower part of the scales of the fourth and upper part of 
the scales of the third dorsal row. 

Anteriorly on the mid-dorsum there are irregular black spots, 
which gradually unite to form two black lines on the eighth 
scale row on each side. These lines are well defined on the pos- 
terior part of the body, the zone between them being darker 
than the general coloration. On the tail the two middorsal lines 
fuse, so that there are only three black lines (one middorsal and 
two lateral) on the tail. 

Total length : 420 mm., tail 85 mm. 

Notes on the paratypes. The general characteristics are those 
of the holotype. The total variation of the scale counts in all 
specimens, including the holotype, is as follows : 146 to 158 
ventrals {$ $ 155-158; 9 5 : 146-150), 53 to 61 subcaudals 
{ S $ : 58-61; 9 9 : 53-59), anal always divided, dorsal formula 
17-17. Supralabials 7 (3, 4), infralabials are 9 (4) or 10 (5). 
There is one preocular on both sides in one specimen, two pre- 
oculars on both sides in two specimens, and one specimen has 
one on the left and two on the right side. There are 1+2 or 
l+l-|-2 temporals. 

The coloration of the paratypes is similar to that of the holo- 
type, with exception of the MBUCV No. 3044, which seems to 
be a somewhat more melanistic and aberrant specimen. It has 
the middorsal black line broader, covering the superior part of 
the seventh and the whole of the eighth and ninth rows, and the 
zone between the middorsal and lateral black lines is lighter 
(pale brown). The MBUCV specimen is darker in general: the 
mottling on the underside of the head being almost black and 
the dark streak on the middle line of the subcaudals being 
darker. 



4 BREVIORA No. 88 

The CM specimen has the following hemipeneal characters : 
The sulcus spermaticus is bifurcated, the bifurcation taking place 
rather close to the base. At the base are several large spines or 
hooks. About 12 longitudinal rows of larger spines run from 
the base to the place where the bifurcation of the hemipenis 
occurs, each row containing 5 to 7 larger spines ; among them are 
placed several smaller spines, which eventually may form addi- 
tional longitudinal rows, between the big rows. At the apex, 
where the sulcus ends, there is a calyculated zone, formed by 
several larger and smaller calyces. 

Generic position. This is one of the most complicated cases in 
the already sufficiently complicated Leimadophis-Liophis-TJ roth- 
eca, etc. group of genera. This species has the following charac- 
teristics of the genus Urotheca: 

1. Absence of any reduction in the dorsal squamation. 

2. Calyculated and simple bifurcated sulcus of the hemi- 
penis, as well as the presence of large spines or hooks at the 
base. 

3. Dark lip coloration, with a white line crossing it. 

It has, on the other hand, these characteristics of the genus 
Leimadophis: 

1. The presence of one scale pit on dorsals. 

2. The general coloration, i.e., a longitudinal dark lateral and 
dorsal line beginning to form at midbody, occupying all the 
posterior part of the body and the tail. 

The maxillary dentition is in general intermediate between 
the two, although with a slight predominance of the Urotheca-like 
characteristics, i.e., there is a gap, but it is rather small, and 
also the two last teeth following the gap are somewhat smaller 
than usually is the case in Leimadophis. 

The hemipenis and the dorsal scale formula were the charac- 
teristics which caused me to put this new species in the genus 
Urotheca, although the presence of one dorsal pit has been con- 
sidered as a very decided Leimado phis-like character. However, 
on several occasions I have observed the absence of pits in speci- 
mens of several species of Leimadophis (L. mclanotus, L. regiyiae), 
and I have found scale pits occasionally present in specimens 
of Lygophis and — more important — in Liophis. It seems that the 
presence or absence of scale pits in this group of genera and 
occasionally in others (cf. Amaral's genus Barhourina!) can 



]f)58 NEW SPECIES OF UHOTIIECA 5 

be used only as an auxiliary generic character. Moreover, scale 
pits are present in practically all reptiles, although usually so 
small that they are not seen except under high magnification. 
The problem thus arises : when are they sufficiently visible to 
be useful as a character? 

As to the question of Vrotheca vs. Rhadinaea, there can be 
little doubt, as pointed out by Dunn, 1957, that the earlier name 
should be used. Vrotheca dumerilii, the genotype, apparently 
is not a Cuban form, as originally presumed, but most probably 
a species from Central America (the description suggests this 
very strongly), described later as a species of Rhadinaea {R. 
pachyura fnlviceps). On this view, the genus Rhadinaea Cope, 
1863, is a synonym of Vrotheca Bibron, 1843. To use Vrotheca 
will certainly simplify the taxonomy greatly, since in the past 
various authors have distributed species to one or the other 
genus, without any clear concept of the differences between 
them. 

The holotype and the MBUCV specimen have been reported 
by Marcuzzi, 1950, as Leimadophis himaculatus opisthotaenia, 
but this is an error, since the latter species is a form from the 
Andes of Merida, Venezuela, and is a true Leimadophis (I have 
seen several specimens of that form), and very distinct indeed 
from V. ivilliamsi. 

I take pleasure in naming this species for my friend and col- 
league. Dr. Ernest E. AVilliams, from whom I received much 
kind help and cooperation, while staying at the Museum of 
Comparative Zoology. 

BIBLIOGEAPHY 

BOULENGER, G. A. 

1908. Descriptions of new South American reptiles, Ann. Mag. Nat. 
Hist., (8) 1: 111-115. 

Dunn, E. R. 

1957. Neotropical frog genera: Prostherapis versus Hyloxalu.s with 
remarks on Phyllobates. Copeia, 1957 (2) : 11-18. 

.Marcuzzi, G. 

1950. Ofidios existentes en las colecciones de los museos de Caracas 
(Venezuela). Nov. Cient. La Salle, Ser. Zool., 3: 1-20. 



I 



BREVIORA 

Mimseiuiinii of Comparative Zoology 

Cambridge, Mass. July 15, 1958 Number 89 



REMARKS ON SOME FORMS OF CINCLUS (AVES) 

By James C. Greenway, Jr. 
and Charles Vaurie 

The genus Cinclus was studied by us independently. Green- 
way studied the collections in the Museum of Comparative Zool- 
ogy and in the American Museum of Natural History in prep- 
aration of a part of the Check-list of the Birds of the World. 
Vaurie studied the collection of the American Museum in 
preparation of a cheek-list of the Palearctic avifauna (1958, in 
press), and published some notes on this genus on two occa- 
sions (1951 and 1955). Greenway deferred publication. Later 
he sent the draft of his paper to Vaurie, who in 1957 examined 
the material in several European museums. We have decided 
to present our conclusions jointly. 

Cinclus cinclus 

Two main types are represented on the European continent: 
one which has a blackish breast, and the other in which the 
breast is several shades of brown. Some populations are con- 
stant or relatively constant, as in Scandinavia and northern 
Europe where the birds (nominate cinclus) are blackish but 
other populations vary individually. The latter may consist of 
individuals that are blackish and very similar, if not identical, 
with nominate cinclus or of individuals in which the color of 
the breast varies from dark and dull brown to rather bright 
rufous. In some regions, as in the Alps and to some extent in 
Spain, the birds inhabiting a watershed may be blackish, whereas 
in a contiguous watershed they are brown and these blackish, or 



2 BREVIORA No. 89 

brown, populations may resemble other populations from which 
they are widely separated. For instance, blackish birds inhabit 
Scandinavia and northwestern Spain and brown birds inhabit 
Germany and southern Spain. On the other hand, the popula- 
tions may grade more or less smoothly into one another, as from 
southeastern Germany southeastward through central Europe. 

These complicating factors render a division for nomencla- 
tural purposes somewhat arbitrary and Vaurie was, at first, in- 
clined not to recognize any subspecies. However, all the birds of 
Scandinavia and northern Europe are blackish and identifiable 
at a glance from the brown-breasted ones from farther south, and 
the geographical variation, though not constant in central and 
southern Europe, shows several trends. In the end, Vaurie 
(1955) recognized five subspecies. These five subspecies are also 
widely recognized by conservative authors and are as follows : 
nominate cinclus Linnaeus, 1758, type locality, Sweden ; aquati- 
cus Bechstein, 1803, type locality, Germany; pyrenaicus Dresser, 
1892, type locality, Pyrenees ; meridionalis Brehm, 1856, type 
locality, Carinthia, Austria; and orientalis Stresemann, 1919, 
type locality, Macedonia. 

There is, however, little agreement about the limits of their 
ranges. Hartert (1910) stated that the range of nominate cinclus 
was Scandinavia and western Russia south to East Prussia ; 
Witherby (1922, p. 341) added the Cantabrian Mountains of 
northwestern Spain, and in 1928 (pp. 618-620) the mountains 
of central Spain; while Vaurie (1955) referred all the birds 
of the Iberian Peninsula to pyrenaicus which Hartert had re- 
stricted only to the Pyrenees. Hartert (1910) stated that the 
birds of France were aquaticus (except in the Pyrenees and in 
the southwest where they are replaced by meridionalis) but 
Mayaud (1953, p. 49) includes Brittany and the Massif Central 
of France in the distribution of pyrenaicus, stating that in the 
southeast the populations show a tendency toward meridionalis. 
Hartert (1922) recognized orientalis for southeastern Europe, a 
region which he had formerly included in the range of meridion- 
alis, while Rokitansky (1939) extended the range of orien- 
talis westward to Moravia in central Europe. Finally, von Burg 
(1924) and Troller (1935) described six new forms from the 



1958 REMARKS ON CINCLUS 3 

Alps, and Floericke (1926, Mitt. Vogelw., p. 78. Non vid.) one 
from northern Portugal which he named atroventer. The latter, 
the type locality of which is Serra do Gerez, is a very black form. 

It is evident from the foregoing that students of this group 
have often disagreed (especially regarding the classification and 
distribution of the populations of France, the Alps, Balkans, and 
Iberian Peninsula), and Greenway came to the conclusion that 
it would be best to recognize only two subspecies : a blackish one 
(nominate cinclus), and a brown one (aquaticus) . He was aware 
that the range of the black form is discontinuous, and also that 
a more realistic treatment might include but a single subspecies. 
Black and brown forms are, however, separable at a glance. 

Vaurie, after studying the material in the European collec- 
tions, now shares the opinion of Greenway. "We consider that it 
is best to synonymize pyrenaicus, sapswortJii, atroventer, and 
amphitryon with nominate cinclus, and to synonymize meridion- 
alis, orientalis, olympicus, and the forms described by von Burg 
and Troller, with aquaticus. SapswortJii was described by Arri- 
goni in 1902 from Corsica but we find that good series in New 
York and Europe from Corsica and Sardinia are not separable 
from specimens from the Pyrenees. Olympicus Madarasz, 1903, 
type locality, Cyprus, and amphitryon Neumann and Paludan, 
1937, type locality, Lasistan, northeastern Turkey, will be dis- 
cussed below. 

The populations of Austria and Italy (meridionalis) , or from 
the Balkans (orientalis), vary too much individually to warrant 
nomenclatural recognition as about one third to one half or more 
of the specimens examined cannot be distinguished from typical 
aquaticus of Germany. Professor Stresemann, with whom Vaurie 
has discussed this question, is now also of the opinion that 
orientalis is not sufficiently constant and is best synonymized with 
aquaticus. 

The subspecific status of the birds of the Iberian Peninsula and 
Pyrenees is the least clear and has been the subject of most com- 
ment. The result of Vaurie 's examination of relevant material 
in European collections is set forth below ; that in New York was 
discussed in 1955. 

Eleven specimens from the Pyrenees support the statement 
made by Vaurie in 1955 after he had examined 25 birds that are 



4 BREVIORA No. 89 

virtual topotypes of pyrenaicus. As stated then, this popula- 
tion varies individually, but "in series . . . differs from nom- 
inate cinclus by being not so black, browner above and below 
including the crown and nape, which are distinctly paler, and 
in fresh plumage by having the gray edges of the feathers of the 
back better developed." Whistler and Harrison (1930, p. 467) 
stated that their three specimens from the Pyrenees ''cannot be 
separated from Swedish topotypes, ' ' but much larger series show 
that Witherby (1928) is correct when he states that the birds 
of the Pyrenees though "very much like" nominate cinclus are 
"not quite so black." Witherby recognized pyrenaicus but we 
believe it is best to synonymize it with nominate cinclus, as it is 
impossible to separate the birds of the Pyrenees from specimens 
which, in series, are even more similar to nominate cinclus. 

Nine specimens from the province of Santander are very 
similar above to the birds of the Pyrenees, including the prom- 
inent gray edges, but in series average a little more blackish, 
especially below. Three specimens from Riano, northern Leon, 
are still blacker. These three match four from the Serra do Gerez 
in northern Portugal (topotypes of atroventer), one from Serra 
da Estrela, also in northern Portugal, three from Candeleda, 
Sierra de Gredos, Avila, and twelve from Lagunilla, Be jar, Sala- 
manca. All these are very blackish below and similar to nominate 
cinclus but not quite identical. They average deeper black below 
but above they are more similar to the birds from Santander and 
the Pyrenees, the crown and nape averaging slightly more rufous 
and the gray edges of the feathers being, as a rule, better de- 
veloped than in nominate cinclus. 

However, individuals are found in northwestern or central 
Spain that are not similar to nominate cinclus. One specimen 
from Santo Domingo de Silos, Burgos, and also one from Can- 
deleda are browner. Of three specimens from San Ildefonso la 
Granja on the northern side of the Sierra de Guadarrama, one 
is similar to the black birds from Riaiio, though very slightly 
browner, and the other two are a little browner still. One from 
Cercedilla, on the southern side of the Sierra de Guadarrama, 
is distinctly browner than the foregoing and Witherby (1928) 



1958 REMARKS ON CINCLUS 5 

remarked that it is intermediate between nominate cinclus and 
aquaticus, adding; that one from nearby Escorial "is like 
aquatimt^." 

This last specimen is now in the collection of the American 
Museum of Natural History and was examined by Vaurie in 
1955 tog-ether with two from the Sierra Nevada in southern 
Spain. These three are identical and not separable from aquati- 
cus. Five other skins, examined in 1957, from the Sierra Nevada 
are also very brown and identical with aquaticus. 

In short, it appears that the population of southern Spain is 
constant and not separable from aquaticus. The birds of central 
Spain and the Pyrenees vary individually but, taken as a series, 
these populations are closer to nominate cinclus than they are to 
aquaticus, while in northwestern Spain and northern Portugal 
the populations seem constant again, and, though not quite iden- 
tical, are best called nominate cinclus as they are so similar to it. 

Two of the additional specimens from the Sierra Nevada were 
collected in 1955 by Dr. G. Niethammer and are in the collection 
of the Bonn Museum. This museum possesses also the type and 
paratypes of amphitryon. This series is identical with nominate 
cinclus in coloration, as stated by Neumann and Paludan (1937), 
who separated it from the latter on the basis that it had a ' ' some- 
what shorter wing and an almost imperceptibly thinner bill" 
(trans.). The bill shows a tendency to be more slender, but the 
difference is extremely slight and not of taxonomic importance, 
and the wing measurements show too much overlap to warrant 
the recognition of amphitryon. According to Vaurie, the wing 
length measures 93, 93, 94, 94 (+, molting), 95, 98, 98 (95) 
in the males from northeastern Turkey, as against 92, 94, 95, 95, 
95, 96, 96, 97, 98, 98, 99, 100 (96.3) in males from Scandinavia. 

Two other forms {olympicus from Cyprus, and uralensis 
Serebrovski, 1927, type locality, Urals) require brief comment 
though we are handicapped by the lack of adequate material. 

The material that we have seen from Cyprus consists of three 
rather worn and not too well prepared specimens collected in 
the spring of 1906. Vaurie (1955) stated he could match them 
with specimens of orientalis, but in Greenway's opinion they 
resemble caucasicus rather than Macedonian birds. Hartert 
(1910), who had the same specimens, stated that he "was not 



6 BREVIORA No. 89 

able" to separate them from caucasicus but cautioned that more 
and better prepared material was necessary from Cyprus. Never- 
theless, he sjaionymized olynipicus with caucasicus, though with 
a query. The name olympicus has remained a synonym of cau- 
casicus ever since but, unfortunately, olympicus was described 
in January 1903 as against December of the same year for 
caucasicus; it should replace the latter if indeed the populations 
of Cyprus and the Caucasus are not separable. Fresher and 
more abundant material from Cyprus is necessary, however, to 
establish beyond dispute that the two forms are not separable. 
We therefore recognize provisionally the validity of olympicus. 
Serebrovski (1927) has separated the population of the Urals 
as uralensis, stating that it differs from nominate cinclus by 
being paler and more brownish above and below, but is darker 
above and less rufous below than aquaticus. The validity of 
uralensis has been recognized by Hartert and Steinbacher (1935), 
Dementiev (1935), and Portenko (1937), but Sudilovskaya 
(1954) considers that uralensis is a synonym of nominate cin- 
clus. Her opinion, however, may be extreme as she also synony- 
mizes caucasicus (which in our opinion is perfectly valid) with 
nominate cinclus. The only specimen of uralensis that we have 
seen is not fully adult but shows the differences mentioned by 
Serebrovski when compared to specimens in the same plumage. 
The population of the Urals is very widely isolated from all 
the other populations of the species and may well be distinct. 

Cinclus pallasii 

As Vaurie (1951) has remarked, there are four specimens of 
the Brown Dipper, in the collection of the American Museum 
of Natural History, having gray rumps contrasting with brown 
backs. These were taken in Amurland and near Mukden. 

It would perhaps be possible to separate on the basis of such 
material the populations of extreme eastern Siberia and Japan, 
on the one hand, from those of China and Formosa, on the other. 
The probability is strong that this one difference in color of rump 
is due to a single gene, however, and a theory that a single popu- 
lation inhabits eastern Asia, Japan, Formosa, and northern 
Indochina is probably the most correct one. To be sure, there is 



1958 REMARKS ON CINCLUS 7 

a small size difference, as is shown in Table 1, below. That this 
is significant may be doubted. Of a series of 15 kinds, 6 are 
intermediate. Application of a coefficient of difference test shows 
a 20 per cent overlap. 







Table 1 






WING 


(in 


nun.) 




■i III aria 11(1 


China 






Fiinnii.sd 


94.0 1 


104.0 


1 






96.0 1 


102.0 1 


105.0 


1 






102.0 1 


103.0 1 


106.0 


1 






103.0 1 


106.0 1 


107.0 
108.0 


1 
1 






105.0 1 




109.0 


9 






Efx=: 406.0 




110.0 


3 






n = 4 




111.0 


1 






m = 101.50 




114.0 


1 






= 3.87 




120.0 


1 










121.0 


1 









Japanese populations are intermediate to some extent. Of a 
series of 13 specimens from Japan and Amurland, 5 are impos- 
sible to diagnose as belonging- to one population or the other. In 
other words, only 62 per cent can be recognized on the character 
of the gray versus brown rump. Certain Japanese specimens 
have a somewhat more reddish tinge than those from the Asiatic 
mainland, but only 65 per cent of the series at hand can be 
distinguished by this character. 

Vaurie (1955), recognized only two subspecies: nominate 
pallasii Temminck, 1820, type locality, "Crimea," but error for 
Okhota River, eastern Siberia, as shown by Stresemann (1948, 
pp. 115, 126) ; and tenuirostris Bonaparte, 1850, type locality, 
central Asia. Among the forms synonymized with nominate 
pallasii were wilderi La Touche, 1925, type locality, Hopeh, and 
dorjei Kinnear, 1937, type locality, eastern Bhutan. 



8 BREVIORA No. 89 

The two co-types of wilderi from the Eastern Tombs near 
Peking have brown rumps and do not appear to differ in any 
way from birds of western and southern China (see above). 

The affinities of dorjei were not made too clear by Kinnear (1937, 
p. 263), and Professor Stresemann kindly pointed out to Vaurie 
that the latter was probably wrong in considering that dorjei 
was not separable from nominate paUasii. In Stresemann 's 
opinion dorjei was probably a valid race, or, if not, a synonym 
of tenuirostris rather than of nominate pallasii. Examination of 
the original series shows that Stresemann is correct and that 
dorjei is valid. 

Its diagnosis and range, as given by Vaurie (1958) are: 
"Darker than tenuirostris in all plumages but paler than nom- 
inate pallasii. This race, though intermediate in coloration, is 
closer to tenuirostris. Range : Eastern Himalayas, east of ten- 
uirostris [which ranges eastward about as far as northern Sikkim 
where it grades into dorjei']., hills of Assam south of the Brah- 
maputra, and higher hills and mountains of Burma south to the 
Chin Hills in the west and the Shan States in the east, to north- 
ern Siam and probably to neighboring northern Yunnan though 
there are no records." 

CiNCLUS SCHULTZr 

Cincliis schultzi Cabanis of the mountains of northwestern 
Argentina has been treated as a distinct species by all authors. 
In view of the intraspecific variation in other parts of the world 
it would appear to reflect the probable biological truth better 
to treat it as a subspecies of leucocephaliis. Compare Cinclus 
cinclus leucogaster of Afghanistan with caucasicus and cash- 
meriensis, for example. 

The brown breast of schultzi replaces the white of the more 
northern form (leucocephalus) . The w^hite wing bar of schultzi 
is not consistently of greater extent. It is a grayer bird and a 
very distinct subspecies. 



1958 REMARKS ON CINCLUS 9 

ACKNOWLEDGMENTS 

Vaurie received much friendly cooperation during his recent 
visit to Europe and takes pleasure in thanking the curators of 
the various museums and their staffs for much help. Professor 
Stresemann kindly discussed several of the forms mentioned in 
this paper, giving him the benefit of his great experience, and 
Mr. Macdonald of the British Museum sent to New York the 
specimens of dorjei for further study. 

LITERATURE CITED 

Anonymous [= von Burg] 

1924. Die schweizerischen Wasseramseln. "Der Weidmann" (Biilach- 
Ziirich), 6(24): 7. 

Dembntiev, Georges P. 

1935. Systema A\dum Rossicarum. L'Oiseau, p. 456. 

Hartert, Ernst 

1910. Die Vogel der palaarktischen Fauna. 1(6): 788-795. 
1922. Op. cit., 3 -.2172. 

Hartert, Ernst and Friedrich Steinbacher 

1935. Die Vogel der palaarktischen Fauna. Erganzungsband, p. 344. 

Kinnear, N. B. 

1937. In Ludlow and Kinnear, The birds of Bhutan and adjacent 
territories of Sikkim and Tibet, pt. 2. Ibis, 249-293. 

Mayaud, Noel 

1953. Liste des oiseaux de France. Alauda, 21(l):l-63. 

Neumann, Oscar, and Knud Paludan 

1937. Zwei neue geographische Rassen aus Klein-Asien. Orn. Monatsb., 
45:15-16. 

Portenko, L. a. 

1937. Fauna Ptits Vnepoliar. Chasti Severnogo Urals. Moscow-Lenin- 
grad. Akad. Nauk SSSR, p. 164. 

Rokitansky, G. F. von 

1939. Zur Verbreitung der Wasseramseln, spezieU der Rasse Cinclus 
cinclus orientalis Stres. Ann. Naturhist. Mus, Wien, 49:282-294. 



10 BREVIORA No. 89 

Serebbovski, p. 

1927. New races of Palearctic birds. Doklady Akad. Nauk SSSE, 
1927 (A):325-326. 

Stresemann, Erwin 

1948. Dr. 0. H. Mercka ornithologischc Aufzeichnungen wahrend der 
Billingsschen Expedition von Oehotsk naeh Alaska (1787-1791). 
Zool. Jahrb., Abt. fiir Syst., 78(1) : 97-132. 

SUDILOVSKAYA, A. M. 

1954. In Dementiev and Gladkov (editors) ; Ptitsy Sovietskogo Soiuza. 
Mosco-w, 6:672. 

Troller, Julius 

1935. Die Wasseramsel, Cinclus cinclus L., und ihre Unterarten in 
der Schweiz. Areh. Suisses d'Ornith., 1(6) :193-224. 

Vaubie, Charles 

1951. Notes on tlie wrens and dippers of western Asia and India. 
Amer. Mus. Novitates, 1485:9-19. 

1955. Systematic notes on Palearctic birds, no. 16. Troglodytinae, 
Cinclidae, and Prunellidae. Amer. Mus. Novitates, 1751 : 12-17. 

1958 (in press). The birds of the Palearctic Fauna, part 1. Passeri- 
formes. London, H. F. and G. Witherby. 

Whistler, Hugh, and James M. Harrison 

1930. Some auturnn observations of the avifauna of the western and 
central Pyrenees. Ibis, 453-470. 

Witherby, H. F. 

1922. Kesults of a collecting trip in the Cantabrian Mountains, north- 
ern Spain. Ibis, pp. 323-345. 

1928. On the birds of central Spain with some notes on those of south- 
east Spain. Ibis, pp. 385-436, 587-663. 



BREVIORA 

Miuiseimm of Connparsitive Zoology 



Cambridge, Mass. July 30, 1958 Number 90 



A FOSSIL VAMPIRE BAT FROM CUBA 

By Karl F. Koopman 
Academy of Natural Sciences of Philadelphia, Philadelphia, Pa. 

Of the nine families of bats in the Western Hemisphere, all but 
three (Desmodoutidae, Fnripteridae, Thyropteridae) have been 
known from the West Indies for some time, althongh the Em- 
ballonuridae reach the West Indies only in Grenada. The precise 
ecology and food habits of the Thyropteridae and Fnripteridae 
are poorly known so that no explanation for tiieir absence othei 
than the presence of water barriers is apparent. The Desmodon- 
tidae, on the other hand, are the vampire bats, Avhose blood diet 
is well known. The absence of this family from the West Indies 
might, therefore, be explained by the paneity of native, non- 
flying, land mammals of which in the Greater Antilles, only five 
genera are known, none larger than a cat. Chicken-sized or larger 
ground birds are also absent. Thus it is possible that at the time 
of arrival of man in the West Indies, there were no animals 
present which could support a population of vampire bats. 

For a number of years, however, it has been known that during 
the late Pleistocene, several genera of large and medium-sized 
mammals existed in the Greater Antilles. Four genera of ground 
sloths are known from Cuba, while elsewhere in the Greater An- 
tilles, both ground sloths and large rodents are know^n. It there- 
fore appears that at this time a suitable habitat for vampire bats 
did exist. Until now, however, none of the fossil deposits in 
which these larger forms are found, at least in Cuba, has yielded 
bat remains. 



2 BREVIORA No. 90 

During: the past IS years, the Speleological Society of Cuba 
has made extensive excavatious in numerous caves, chiefly in 
western and central Cuba, uncovering abundant mammal re- 
mains, including a great deal of ground sloth material (Ar- 
redoudo 1955). One of the richest series of fossil deposits was 
in the L'lmas cave, a few miles west of Havana. The following 
statement was prepared for me by Sr. Oscar Arredondo of the 
Sociedad Espeleologica de Cuba, Havana, and Sr. Gilberto Silva 
Taboada, a student of recent bats, also of Havana. 

"The Lamas Cave is merely a rocky .shelter located in a land 
slope to the southwest of Santa Fe Beach, on the northern coast 
of Habana province. On January 21, 1951, an excavation was 
made at this cave by Oscar Arredondo, Director of the Paleonto- 
logical Section of the Speleological Society of Cuba, together 
with Dr. Manuel Rivero de la Calle, from Las Villas Central 
University. 

"According to the stratigraphical profile, the flrst level was a 
layer of very dark earth. 60 centimeters in depth, containing 
jilenty of shells and human bones pertaining to the oldest Indian 
culture of Cuba (the Cuanahatabeyes), together with skeletal 
remains of Capronnjs, Gcocaproiiiys, and Boroniy.s offella. The 
second level, extending about 40 centimeters in depth, was com- 
posed of a red earth completely lacking any bones or areheological 
remains. The third level was a 30 centimeter deep layer of a 
light yellow colored earth, containing bones of small extinct 
rodents, like Geocapromys. 

"The fourth level, having a depth of 1.40 meters and com- 
posed of a humid earth, reddish-yellow in color, Avas the richest 
fossil-bearing layer of the trench. Thousands of scattered frag- 
mentary bones and skulls, representing almost all the known 
extinct Cuban fauna, were unearthed from this level. Here was 
found a fragmentary bat skull in association with the extinct 
rodents Gcocapromxjs columbianus and Boromys torrei, the in- 
sectivore Nesophontes micrus, the ground sloths Megalocnus 
rodens and Mesocnus iorrei, an unknown bird allied to the ex- 
tinct Phororacos'^ of Argentina, and also Testudo cuhensis and 
the living Crocodylus rhomhifer." 

1 This iilentiflcation may be erroneous. 



l})r)8 FOSSIL VAMPIRE HAT KKOM CUBA 3 

AVe are licre eoneernod with the richest, deepest, and therefore 
oldest, of tile four layei's. This, it may l)e noted, consists entirely 
of extinct species, at least as far as the inanunals ai-e concerned. 
I can verify the identity of at least the Ncsuphoiitrs, Boroinys, 
Geocapronujs, and "Testudo" {Geochelone), since Sr. Arredondo 
has very generously turned ovei- to nie a small sample of the 
fossils from this layer. The assemblaoe probably indicates a 
truly Pleistocene level and tlie presence of <>round sloths 
[Megolocuus, Mesocnus) and of large ground birds (whatever 
their at^nities may be) indicate a very ditferent habitat, in 
terms of ability to support vampire bats, from that which existed 
in late pre-Columbian times. 

As mentioned above, among the numerous fossil bones of this 
deposit, there was found the rostral portion of a bat skull, which 
Sr. Gilberto Silva Taboada kindly presented to me. It is now 
No. 4737 in the Vertebrate Paleontology collection of the Mu- 
seum of Comparative Zoology. In this specimen, the very char- 
acteristic vampire dentition is immediately apparent, togethei- 
with the peculiar rhombic shape of the palate characteristic of 
this group of bats. Comparison with the three living species of 
the family Desmodontidae shows that the fossil is clearly refer- 
able to Desmodus rofiDidus, the commonest mainland species. 

Unfortunately the condition of the skull permits few measure- 
ments to l)e made on it (Plate). The following are the only 
standard ones I was able to make. In each case, the figures in 
parentheses refer to a series of seven specimens of Disuwdus 
rotKiidiis i)ntri)U(s from El Pachon, Tamaulipas. Measurements 
are in millimeters. 

Least interorbital width — 5.1 (4.5-5.5) 

Length from anterior edge of incisor to posterior edge of canine 
— 4.9 (4.6-4.9) 
Width across base of incisors — 3.9 (3.3-3.7) 

Except for the slightly greater width across the incisors, the 
Cuban fossil appears to be indistinguishal)le from the Middle 
American subspecies, DcsDiodus rofuiidits niuruius. 

We may assume that Desmodus reached Cuba after prey w^as 
available in the form of ground sloths and large ground birds, 
and that when these forms became extinct and thus the ecological 



4 BREVIORA No. 90 

niche for vampires disappeared, the latter also died out in Cuba. 
The introduction of domesticated animals in post-Columbian 
times has presumably reopened the niche, but, at least so far, 
vampires have failed to ^et acro.ss the fairly wide water bar- 
riers and hence do not form part of the living bat fauna of Cuba. 
In conclusion, I wish to acknowledge the help very graciously 
given me by the two above-mentioned Cuban naturalists, 8r. 
Oscar Arredondo and Sr. Gilberto Silva Taboada. 

LITERATURE CITED 

Arredondo, O. 

1955. Resumen de las Ac-tividades paleontologk-as realizada.s por la 
Seecion de Geologia y Paleontologia. Bol. Soc. Espeleologica dc 
Cuba. Alio I (Deeiembrc), pp. 3-31. 



tt 





Palatal views of J)csnw(hii-: skulls. Top, the Cuban fossil, MCZ (Verteliiato 
Paleontology) 4737. Bntfom, D. rotundus $, MCZ (Mammalogy) 10445 
from Bogaba, Panama. 



BREVIORA 

MmseiJiitTii of Coiraipsirative Zoology 



Cambridge, Mass. August 13, 1958 Number 91 



CONTRIBUTION TO A REVISION OF THE EARTH- 
WORM FAMILY LUMBRICIDAE. II. INDIAN SPECIES 

By G. E. Gates 

INTRODUCTION 

The domain of the ' ' recently evolved and dominant ' ' Inmbricid 
family, a group which has "almost completely conquered the 
Palaearctic" (Stephenson, 1930, p. 668), long has been thought 
to reach well into India. The southern boundary of that domain, 
on recent distributional maps (for example those of Tetry, 1938, 
and Julin, 1949), runs from the Mediterranean so as to include 
the Punjab portion of the Sind Valley and all of the Gangetic 
Valley to Calcutta from whence it is extrapolated, with question 
marks or predictions, across northern Burma and through China 
to the Pacific. 

Two routes from European centers of lumbricid evolution to 
the vicinity of Calcutta could have been available to the advancing 
lumbricids : through the Himalayas in an easterly direction and 
then down to the plains, or south along the Indus River and 
thence easterly through the Gangetic Valley. In either case, 
lumbricids could be expected throughout an eastern portion of 
the Gangetic plain as well as in the western part of the Himalayas. 
A survey of the earthworm fauna of the Allahabad sector (Gates, 
1945, 1947) and collecting in such Himalayan sites (Gates, 1951) 
as could be visited during wartime, provided data that made a 
review of the taxonomic status of the supposedly endemic species 
advisable. 

The results of that study are presented herewith. The text 
of the present contribution, except for this paragraph, has been 
excised from a chapter of an unpublished manuscript written 
several years ago during tenure of a John Simon Guggenheim 
fellowship. 



2 BREVIORA No. 91 

SYSTEMATICS 

Genus AlLOLOBOPHORA Eisen, 1874 
" Allolobophora caliginosa (Savigny, 1826) " 

A complex of four or more morphologically distinguishable 
species long has gone under Savigny 's name. The data aA'ailable 
in the literature rarely permit a decision as to which species was 
present in a particular collection. Indian localities from which 
"caliginosa" has been recorded are: Fern Hill and Ootacamund 
(at elevations of 6-8,000 feet in the Nilgiri Hills, South India), 
Mt. Abu (Rajputana), Lahore, Ferozepur, Kotla, Murree, 
Peshawar, Mardan, Chitral (Punjab and Northwestern Frontier 
Province), Gilgit, Gurez, Gundarbal, Anchar Lake (Kashmir), 
Almora, Mussoorie, Naini Tal, Simla, Kufri, Junga, Kasauli 
(western Himalayas). All except the Nilgiri localities are well 
within the temperate zone. None of the complex ever was found 
at Darjiling where exotic lumbricids now appear to be dominant 
(Gates, 1951). There are no records for Ceylon, Burma, the 
Andaman and Nicobar Islands, the Malay Peninsula, and other 
parts of southeast Asia, not even for hill stations where intro- 
duction might have been expected. 

The characterization "cosmopolitan" so frequently applied 
to "caliginosa" obviously implies more than has been warranted 
by the facts, even if the complex had been but a single species. 

AlLOLOBOPHORA jassyensis Michaclsen 1891 

This little known species has been reported from a single Indian 
locality well north of the thirty-second parallel in the Murree 
subdivision of the Punjab. Individuals of the caliginosa complex 
sometimes have been mistaken for jassyensis. Presence of the 
latter in India, if confirmed, presumably is due to importation 
by man. 

AlLOLOBOPHORA TRAPEZOTDES (DugCS, 1828) 

Helodrilus {Helodriliis) mariensis Stephenson 1917, Rec. Indian Mus., Cal- 
cutta, 13, p. 414, fig. 6. (Type locality, Murree. Types, several, pre- 
sumably in the Indian Museum.) 

Allolohophara (Eophila) mariensis, Stephenson, 1923, Oligochaeta, in Fauna 
of British India, Ceylon and Burma, London, p. 504. 



1958 REVISION OP INDIAN LUMBRICIDS 3 

Stephenson's species is known only from the original descrip- 
tion. Subsequent collections at the type locality provided only 
specimens that have been referred to the caliginosa complex. 

The types probably were brittle and strongly contracted, as 
female and spermathecal pores were unrecognizable. A number 
of differences, of a minor sort, apparently were thought to ob- 
viate consideration of caliginosa. The greenish coloration may 
have been an artifact. Shape of the posterior portion of the body 
(shown in fig. 6) may have resulted from unusually strong 
contraction at preservation and from dehydration in too strong 
alcohol. A similar condition of the posterior end has been seen 
in a number of specimens of trapezoides recently examined. 
Location of the first dorsal pore at 4/5 is of little importance 
taxonomically. A functional pore has been found in trapezoides 
as far forward as 6/7. Just which ones of a series of more or 
less pore-like markings at mD really are patent apertures through 
the body wall into the coelomic cavity often is undiscoverable in 
field-preserved material. Absence of mention of tubercula pu- 
bertatis in Stephenson's description presumably means that 
those structures (as often is the case in browned or other alcoholic 
museum material) were quite unrecognizable. The crypts or 
yellowish projections from the oesophagus in x, that were men- 
tioned by Stephenson, are, of course, calciferous sacs. The 
supposed hearts of xii are only the vertical portions of the extra- 
oesophageal trunlcs (cf. 0. cyaneum below). 

Spermatophores have not been found externally nor sperma- 
tozoa in the spermathecae of any of a considerable number of 
specimens of trapezoides that have been available from various 
parts of the world. The male funnels usually have shown no 
indications of presence of mature sperm. Male sterility requires 
that reproduction be parthenogenetic and that permits reduction 
in number as well as in size of the seminal vesicles. Most liable 
to early elimination after reproduction becomes asexual are the 
vesicles of x and ix. Absence of those vesicles certainly is no 
justification for assigning a species to Relodrilus or to Eophila, 
whatever those two genera may prove to be. In Allolohophora, 
where the species undoubtedly belongs, there is no good reason 
for specific distinction from trapezoides. 



4 BREVIORA No. 91 

Allolobophora tuberculata Eisen 1874 

Among specimens from Simla that were originally (Gates, 
1951) referred to caliginosa are a number that can now be placed 
in Eisen 's species. Synonyms are: similis Friend 1911, arnoldi 
Gates 1952, and caliginosa (various authors). Elsewhere, the 
species now can be recorded from the United States (northeast- 
ern portion), Canada, England, Chile, Denmark, Norway, 
Sweden and probably Germany. The distribution, even as now 
known, must have resulted in part from transportation. Obviously 
there can be no imputation of endemicity in India. 

Genus BiMASTOS Moore, 1893 
BiMASTOS PARVUS (Eiscu, 1874) 

Jalla, Patna, Bihar, March, 0-0-1, E. C. Lacy. 
Rawalpindi, Punjab, May, 0-0-6. F. G. Dickason. 

Forty -four specimens of this species were collected at Allahabad 
during a period of four years. More than half were found in 
dirt around roots of potted plants. The others were obtained from 
a single small area (sansiharicus habitat) by a drainage canal 
that was sampled at least once a month through most of the 
period (Gates, 1945, 1951). The sansiharicus habitats, in the 
region around Jubbulpore to the south of Allahabad were in- 
vestigated (Gates, 1956) on several occasions during that same 
period but yielded no specimens of parvus. 

Indian localities where parvus has been found are : Kodaikanal 
(Palni Hills, South India), Darjiling (eastern Himalayas), 
Patna (Bihar), Allahabad and Saharanpur (United Provinces), 
Partabgarh (Rajputana), Lahore, Lyallpur, Ferozepur, Pesha- 
war, Mardan, Chitral (Punjab and Northwestern Frontier 
Province), Gorai, Srinagar (Kashmir), Kasauli, Barogh, Naini 
Tal (western Himalayas). All are in the temperate zone except 
Kodaikanal which is a summer resort where elevations are 6000- 
7000 feet. The species was found in Burma, during twenty years 
of intensive collecting, only at elevations of more than 4000 feet, 
in and around four hill stations where European cultivated plants 
had been introduced. A hill station also provided the only record 
for the Malay Peninsula. B. parvus obviously is more widely 
distributed in southeast Asia than the supposedly cosmopolitan 



1958 REVISION OP INDIAN LUMBRICIDS 5 

"caliginosa." The species usually is assumed to have originated 
in America from whence, or indirectly through Europe, it was 
brought by man to India. 

The plants along with which parvus was brought to the orient 
were not taken only to the hill stations of Burma or to the 
Indian localities just mentioned. Such plants, potted or other- 
wise, are common throughout the Allahabad sector of the Gangetic 
Valley and also in the Jubbulpore region. The plants undoubt- 
edly have been taken on innumerable occasions into the tropical 
lowlands of India, Burma and the Malay Peninsula. Absence in 
those lowlands after a hundred years and more of such introduc- 
tions must have resulted from inherent inability of the species to 
adapt itself to a tropical climate. 

BiMASTOS EiSENi (Levinscn, 1883) 

This peregrine species, which probably does not belong in 
Bimastos, is known in India only from the region of Naini Tal 
in the western Himalayas. 

Genus DeNDEOBAENA Eisen 1874 
Dendrobaena octaedra (Savigny, 1826) 

This species, hitherto unknown from India, is now recorded 
from Kodaikanal (Palni Hills, South India) where it appeared 
to be quite common. This is another form to which the charac- 
terization of cosmopolitan has been erroneously applied. South 
of Siberia, the species has been found only at the Kodaikanal 
summer resort. The species is also absent from almost all of 
South America. 

D. octaedra is not usually geophagous and, accordingly, would 
not have been expected to be carried around the world as much 
as the earth-eating species. Nevertheless, specimens of this species 
have been intercepted at American ports during the last few 
years, in shipments of plants from Spain, Belgium, Holland, 
Germany, Denmark, Norway, Sweden, Italy, Czechoslovakia and 
Portugal. Importations into North America in days when 
quarantine restrictions were unenforced or lacking must have 
been much more numerous. The species obviously is exotic in 
India as well as in South and North America. 



6 BREVIORA No. 91 

Dendrobaena rubida (Savigny, 1826) 

Athecal Indian specimens that have been identified as Bimastos 
constrictus or Bimastos tenuis belong here. D. suhrubicunda also 
is a synonym. 

Indian localities from which the species is now known are : 
Kodaikanal, Fern Hill and Ootaeamund (Palni and Nilgiri Hills 
of South India, at elevations of 6000-8000 feet), Darjiling (east- 
ern Himalayas), Almora, Naini Tal, Simla and vicinity (western 
Himalayas). 

Genus ElSENIA Malm 1877 
EisENiA foetida (Savigny, 1826) 

Indian localities from which the species has been recorded are : 
Ponmudi, Travancore ; Kodaikanal and vicinitj^ (Palni Hills) 
where it is very common; Fern Hill, Coonoor (Nilgiri Hills) ; 
Calcutta; Darjiling and vicinity (eastern Himalayas) where it is 
also very common; Simla, Kasauli, Dharmsala (western Hima- 
layas). Absence of records for the Punjab, where Stephenson 
and his students worked, is noteworthy. 

The Travancore and Calcutta records never have been con- 
firmed. E. foetida was deliberately introduced, by the hundreds 
or thousands, on various occasions within the last twenty years, 
to lowland tropical areas of the Malayan and Indian peninsulas. 
All such introductions have been failures. Many more times the 
species must have been brought into the plains near the hill 
stations. Some such importation, most probably from Darjiling, 
may well have been responsible for the Calcutta record. 

EiSENiA HORTENSis (Michaelscu, 1890) 

Eisenia veneta f. hortensis, Gates, 1951, Proc. Nat. Acad. Sci. India, B, 
vol. 21, pp. 19, 21. 

This peregrine species has been found, in India, only at two 
Himalayan localities, Darjiling and Simla. 

Eisenia rosea (Savigny, 1826) 

Helodrilus (Bimastus) indicus Michaelsen 1907, Mitt. Nat. Miis. Hamburg, 
vol. 24, p. 188. (Type locality, Calcutta. Types, five, presumably in 



1958 REVISION OF INDIAN LUMBRICIDS 7 

the Indian Museum but probably valueless as they already were ' ' weak- 
ened" from maceration in 1907.) 
Helodrilus (Allolobophora) prashadi Stephenson 1922, Rec. Indian Mus., 
Calcutta, vol. 24, p. 440. (Type locality, Gundarbal, Kashmir. Types, 

several, in the British Museum.) 
Allolobophora (Allolobophora) prashadi + Allolobophora (Bimastus) indica, 

Stephenson, 1923, Oligochaeta, in Fauna of British India, Ceylon and 

Burma, London, pp. 501 and 506. 

Stephenson's species is known only from the original descrip- 
tion. Insofar as can be determined from that description the 
types differed from Savigny's rosea only by the absence of 
spermathecae. As various strains of 7'osea are now known to be 
athecal there is no longer any reason for recognition of prashadi. 

Michaelsen 's species is known only from the types the descrip- 
tions of which contain no indications that external characteristics 
or internal anatomy warrant distinction from athecal, quadri- 
vesiculate strains of E. rosea. Tubercula pubertatis, not men- 
tioned, may have been unrecognizable because of poor condition 
or they may have been lacking — they are disappearing in some 
of the athecal strains. The diameter, 6 mm., is 2 mm. greater than 
the maximum usually mentioned for rosea but the types may have 
been bloated (as are some recently examined specimens where 
thickness in the clitellar region is about 7 mm.) . 

The types of indicus could have been collected at the Botanical 
Gardens in Sibpur where there were secured types of at least 
one other earthworm species erroneously thought to be endemic 
in Bengal. Indeed, in absence of subsequent records to confirm 
domicile in the area, it now seems possible that the specimens in 
question may have been secured directly from earth around roots 
of imported plants. Certainly, there is now not the slightest 
reason for believing that any lumbricid is endemic in Bengal. 

All individuals of athecal xgtrains of rosea that have been 
studied by the writer w^ere male sterile. Tw^o spermatophores, 
each containing "two oval sperm masses" were, however, present 
on one of the types of indicus. Acquisition of ability to reproduce 
asexually often precedes male sterilization. As soon as reproduc- 
tion becomes parthenogenetic, important organs such as the 
spermathecae and male terminalia can be eliminated. In the genus 
Pheretima, for example, several instances are know^n in which 
extrusion to the exterior of sperm has become impossible though 



8 BREVIORA No. 91 

maturation still is profuse. Too little is known as yet about the 
consequences of becoming parthenogenetic, in the Lumbricidae, 
to permit attribution of much taxonomic value to the presence of 
spermatophores even if containing sperm. 

Savigny's species obviously does not belong in the genus 
Eisenia but may as well remain there until lumbricid genera can 
be defined by reference to structures that are less liable to rapid 
evolutionary change than are the organs of the reproductive 
system. 

E. rosea has been reported from Kashmir, Murree Subdivision 
of the Punjab, Chitral in the Northwestern Frontier Province, 
Simla (western Himalayas), Darjiling (eastern Himalayas), and 
to the south only from Kodaikanal (Palni Hills). The new 
synonymies require no additions to the distribution as domicile 
in the region of Calcutta is improbable. 

Genus LUMBEICUS L. 

L. rubellus Hoffmeister 1845 has been recorded from the 
Nicobar Islands. No confirmation has been provided in the last 
sixty years. Subsequent collections from those islands have con- 
tained no lumbricids and domicile is improbable. The specimens 
may have been found in imported earth. 

L. terrestris L. This species has been found only once in India 
and then at Simla in the western Himalayas. 

Genus OCTOLASIUM Orley 1885 
OcTOLAsiUM CYANEUM (Saviguy, 1826) 

Helodrilus (Dendrobaena) Tcempi Stephenson 1922, Rec. Indian Mus., Cal- 
cutta, vol. 24, p. 441. (Type locality, Kufri, Simla Hill states. Types, 
two, presumably in the Indian Museum.) 

Allolobopliora (Dendrobaena) kem-pi, Stephenson, 1923, Oligochaeta, in 
Fauna of British India, Ceylon and Burma, London, p. 502. 

Octolasium oyaneum, Cernosvitov, 1937, Rec. Indian Mus., Calcutta, vol. 39, 
p. 111. (Murree, Punjab.) 

Dendrobaena kempi, Gates, 1939, Vest. Cesk. Spol. Nauk, Praze, 1938-1939, 
p. 151. (Ootacamund, Nilgiri Hills, South India, at elevations of 
6700-8000 feet.) 

Octolasium cyaneum, Gates, 1951, Proc. Nat. Acad. Sci. India, B, vol. 21, 
p. 19. (Simla.) 



1958 REVISION OF INDIAN LUMBRICIDS 9 

Stephenson's species was distinguished from other Indian 
lumbricids, except mariensis, by presence of hearts in xii. The 
Ootacamiind specimens also appeared to have an extra pair. Junc- 
tions with the ventral blood vessel, which lateral hearts must 
have, were not found in specimens secured at Simla not far from 
the type locality of kenipi. However, the supposed hearts are 
only dorsally directed continuations of the extra-oesophageal 
trunks. These continuations, unfortunately, do look very much 
like hearts in a dissection from the dorsal side. There remain 
then no taxonomically important differences from cyaneiim. 

Reproduction probably is not biparental as no sperm have 
been found in the spermathecae (nor any spermatophores ex- 
ternally) of any of the specimens that have been available from 
various parts of the world. Uniparental reproduction, whether 
sexual or asexual, presumably would predispose to successful 
colonization after introduction. 

The species would appear to have been much more common at 
Ootacamund than elsewhere in India or indeed at any of the 
localities from which the writer has collected it. 

OcTOLAsiUM LACTEUM (Orley, 1881) 

Eophila liimalayana Cernosvitov 1937, Eec. Indiaai Mus., Calcutta, vol. 39, 
p. 109. (Type locality, Simla. Types, two, presumably in the Indian 
Museum.) 

Cernosvitov 's species is known only from his description. Sev- 
eral days' search at the type locality in 1944 by a good collector 
yielded a number of specimens of lacteum as well as representa- 
tives of two other peregrine forms that were previously unknown 
in India. 

Absence of seminal vesicles in segments ix and x is the only 
character available to distinguish liimalayana from lacteum. 
Undoubtedly the quadrivesiculate condition was the reason, and 
indeed the only one, for referring types of liimalayana to the 
genus Eophila. Unfortunately the condition of the material sub- 
mitted to Cernosvitov for study was poor, as the writer saw 
before it was mailed. Most of the specimens were more or less 
softened or brittle and alcohol-browned. In macerated individuals 
the smaller vesicles of the ix-x, especially when vestigial, may be 
indistinguishable from adjacent tissues. In brittle material the 



10 BREVIORA No. 91 

vesicles are apt to be broken off during dissection so as to leave no 
macroscopically recognizable traces of their former presence. 

Reproduction in lacteum probably is not biparental (though 
copulating couples have been seen, Gates, 1953), spermatophores 
never having been found externally nor sperm seen in the 
spermathecae of any of the specimens that have been available 
from various parts of the world. Some strains probably are male 
sterile but in others maturation of sperm appears to be so sparse 
as to warrant anticipation of parthenogenesis. With reproduc- 
tion now asexual, seminal vesicles no longer are necessary and 
can be eliminated without harmful results to the individual or the 
strain. Seminal vesicles of ix-x have been quite small to vestigial 
in some strains recently examined and apparently have been 
lacking in others. The pair in ix alone or that in x alone has been 
quite small, vestigial, or unrecognized in other strains. Occa- 
sionally even the vesicles of xi have been vestigial when those of 
ix or ix and x were better developed. As these reductions or 
eliminations take place within the limits of what must otherwise 
be considered a species, there is now no good reason for recog- 
nition of kimalayana. 

Indian localities at which lacteum has been found are Darjiling, 
Mussoorie, Simla (all Himalayan), Murree subdivision (Punjab). 
All are well within the temperate zone. 

DISCUSSION 

Fourteen lumbricid species, if the unallotted residue of the 
caliginosa complex is counted as one, are now known to be pres- 
ent in the Indian subcontinent. All of them obviously are exotic 
there. No evidence is available to indicate any endemicity in 
adjacent regions of Baluchistan, Afghanistan and Tibet, below 
the Hindu Kush and Karakorum ranges. Lumbricids never were 
found in northern Burma and no endemics were discovered by 
Chen (1933, 1946) in the Yangtze Valley and the interior prov- 
ince of Szechuan in China. There is now no reason to anticipate 
lumbricid endemicity south of the Tian Shan and Altai moun- 
tains, Mongolia and Manchuria. Korea and Japan appear to 
have one or more native forms but none is known from Man- 
churia. 

Twelve of the Indian lumbricids are present in the British 
Isles where no members of the family now seem to be endemic. 



1958 REVISION OF INDIAN LUMBRICIDS 11 

Twelve are widely distributed in North America. Many of them 
are present in South America, Africa, Australia, New Zealand 
as well as in various oceanic islands. One species, Bimastos par- 
vus, usually is assumed to have had an American origin. The 
others certainly are all from Europe, from whence the worms 
were ferried to other continents and to the oceanic islands. The 
only way earthworms are known to be transported for such 
distances is in ballast or in earth with those plants that are 
cultivated by man. Transportation of the plants is unquestion- 
able. The data now available (in MS) show that even a handful 
of the fertile, unsterilized earth surrounding the plant roots is 
liable to have contained cocoons or small juveniles and that larger 
samples often must have contained one or more adults. From 
the earliest transoceanic centers of successful colonization the 
exotic worms may have been transported by flood waters as well 
as by anglers, and, certainly, were distributed, on a geometri- 
callj' increasing scale, by florists and horticulturalists as well as 
by innumerable householders. The alien species accordingly are 
properly called peregrine, the original meaning of which is 
foreign or imported. 

In India, lumbricids are common in the Punjab and in the 
Northwestern Frontier Province. In the Himalayas, lumbricids 
never have been reported from Sikkim, Bhutan and Assam where 
there are no summer resorts, nor from Nepal where until very 
recently Europeans we/e excluded. Only in the region around 
Darjiling and in the sector west of Nepal are these worms known 
to be present. In the regions around the hill stations of both 
sectors lumbricids not only are common but also, in some part 
of the year at least, dominant. In the Gangetic Valley, which is 
well within the temperate zone, lumbricids are rare. Only 44 
individuals, all of one species {B. parvus), were included among 
the 30,000 earthworms identified during the course of a four-year 
survey of the fauna of the Allahabad sector. No lumbricids were 
present among some 15,000 thousand earthworms collected in a 
Jubbulpore sector of the Deccan below Allahabad. Farther down 
the peninsula lumbricids have not been found until well into the 
far south and then at elevations of 5000-8000 feet but only in 
and around summer resorts. 

Exotic plants are grown in pots as well as in the ground at 
all Indian, Burmese and Malayan localities where lumbricids are 



12 BREVIORA No. 91 

present. These plants were taken to the hills since 1800, as sum- 
mer resorts were developed by Europeans, but could have been 
introduced into many lowland areas of southeast Asia during the 
previous three centuries. Much earlier than 1500, during the 
period between Alexander's invasion and the downfall of the 
Graeco-Bactrian kingdoms, some cultivated plants may have been 
taken from Europe to the Punjab and Kashmir. 

The exotic plants with which European earthworms were taken 
to the hill resorts of southeast Asia are common in the western 
portion of the Gangetic plains, in the Jubbulpore region of the 
Deccan as well as in Ceylon where there are no lumbricids. The 
same plants also have been taken many times from the hills to 
the lowlands, probably most frequently to larger municipalities 
quickly reached from hill resorts, e.g., from Darjiling to Cal- 
cutta. Absence of all lumbricids in most of such plant trans- 
portations and presence of only B. parvus in the remainder is 
improbable. Deliberate mass introductions of lumbricids to cer- 
tain sections of India and the Malay Peninsula in the last twenty 
years have been no more successful. 

Failure of the very lumbricids that have successfully colonized 
in the Americas, Africa, Australia and New Zealand to establish 
themselves in so many parts of southeast Asia to which they Avere 
introduced does not now appear to be attributable to the rigor 
of any competition provided by native species. An American 
glossoscolecid earthworm, Pontoscolex corethrurus (Miiller, 
1856), possibly introduced more recently and much less fre- 
quently, is widely domiciled throughout southeast Asia as well 
as Malaysia and in some localities appears to have become the 
dominant species. An exotic ocnerodrilid and a peregrine 
megascolecid, as well as a south Indian species of Lampito, now 
are common throughout the Gangetic Valley. A score or more 
of other exotic earthworms are established in Ceylon. 

Moreover, the very numerous unsuccessful introductions that 
there must have been do not now seem to be attributable, as a 
rule, to absence of appropriate habitats. Most of the peregrine 
lumbricids seem to be tolerant of considerable variation in pH, 
moisture content and basic chemical constitution of the soil as 
well as in the amount of digestible organic matter. Nor has the 
reversal of seasons that exists below the equator prevented acqui- 
sition of domicile. 



1958 REVISION OF INDIAN LUMBRICIDS 13 

A first approximation to an explanation of the unsuccessful 
introductions probably could have been given long ago if distri- 
butional data had been particularized instead of being camou- 
flaged by geographical generalizations and erroneous adjectives 
such as cosmopolitan. The Lumbricidae, prior to human inter- 
vention in the situation, were confined to the temperate zone of 
the Northern Hemisphere and, possibly, to some adjacent Arctic 
portions of Eurasia. Transported to everj^ part of the world 
where Europeans have lived, lumbricids have acquired perma- 
nent domicile in the temperate zones and the tropics but in the 
latter only on the highlands. The elevations at which worms were 
collected usually are not stated in the literature but the tropical 
sites appear to be above the 5000 foot level. An inherent inability 
to survive in the less rigorous and more equable climate of the 
tropical lowlands seems to have been demonstrated. 

All lumbricids are able to withstand the rigors of a temperate 
zone winter. Forms such as Z>. octaedra, L. ruhellus and A. 
chlorotica (Savigny, 1826), that now live respectively in Nova 
Zemlya, northern Siberia and Greenland, must be able to with- 
stand freezing temperatures (and being frozen?) for long pe- 
riods. D. octaedra also has been able to colonize around Mexico 
City and Bogota (Colombia) as well as on the Palni Hills of 
south India. The other two species appear to be somewhat less 
adaptable as they are not known to be present anywhere in the 
tropics though more liable to transportation. A. trapezoides 
appears to be even less adaptable as it is certainly known only 
from areas with a climate more like that of the Mediterranean 
lands from which it presumably was taken. The "endemic" 
lumbricids occasionally must have been transported but without 
being able to take advantage of the opportunities for extending 
their range. Furthermore, the distributional records published 
in the last fifty years seem to hint that even in their own areas 
the natives are being replaced by peregrine forms that may have 
been introduced there too by man. Accordingly, the family, as a 
whole, seems now to be definitely limited in ability to ''ad- 
vance," by climatic factors (presumably thermal), and seems to 
lack the youthful characteristics implied by such words as 
' ' powerful " or " conquering. ' ' Even in the area that the lumbri- 
cids have "conquered" without human assistance, dominion is 
shared with two other families, the Hormogastridae and Criodrili- 
dae. 



14 BREVIORA No. 91 

Characteristics of youthful vigor and great evolutionary 
adaptability, it now appears, were too hastily imputed to the 
Lumbricidae as a whole because of the distributions of less than 
a score of peregrine forms. Reproduction in some of these forms 
is parthenogenetic which has been thought to predispose to 
successful colonization. Peregrine forms with obligatory bi- 
parental reproduction, however, are much more common in con- 
siderable areas of North America. More important then than 
parthenogenesis now seems to be an ability to tolerate human 
interference in the earthworm environment. Those species, the 
haemerophiles, which are able to endure the disturbances of the 
soil involved in agricultural and horticultural practices are pre- 
disposed, as it were, to transoceanic and subsequent intraregional 
carriage as well as to survival in new areas that are also culti- 
vated or otherwise influenced by man. One of the peregrine 
species, Eisenia foetida, has been thought to be haemerobiontic, 
i.e., dependent on culture, though that characterization, at least 
in the vicinity of Darjiling, seems exaggerated. Even D. oc- 
taedra, sometimes said to be haemerophobic, certainly lives so 
close to sites of human activity as to be frequently transported. 

Introduction of lumbricids, according to Stephenson (1930, 
p. 905), "frequently causes the disappearance of the endemic 
earthworm fauna." The evidence usually cited for that con- 
clusion is absence of native species in and around municipal 
areas of Chile and Australia that were investigated by Michaelsen 
during the present century. Whether endemics previously had 
been common at those sites is of course unknown, and the earth- 
worm habitats therein had been long subject to modification 
by man. Such little information as is now available hints that 
indigenes are haemerophobic. Culture, then, could have left 
areas formerly inhabited by earthworms open to rapid coloniza- 
tion by any of the preadapted species. These, of course, were 
most likely to be from Europe, as were the settlers. 

As the supposed competitive virility of the Lumbricidae also 
seems to be evidenced by little more than the predisposition of a 
few species to colonization in vacant but climatically restricted 
areas, there remains no reason for believing the family to be the 
"youngest" of the earthworms. Although India and China now 
must be excluded from the lumbricid domain, there still seems 
to be little reason to question a former range from the Mississippi 



1958 REVISION OF INDIAN LUMBRICIDS • 15 

River in North America across Europe and through Siberia to 
Japan. Most of the earthworm families of Michaelsen 's last classi- 
fication (1928), such as Sparganophilidae, Criodrilidae, Hormo- 
gastridae, Syngenodrilidae, Eudrilidae, Moniligastridae, are 
much more restricted geographically. If area is an indicator of 
family age (and there is at present little else from which to 
judge), the Lumbricidae are much older than has been thought, 
possibly as ancient as any of the four remaining families in 
Michaelsen 's system. 

Prior to the Pleistocene ice ages there may well have been, in 
the vast area between the Mississippi and Japan, manj'' more 
lumbricid species than are extant. Advance of ice sheets resulted 
in disappearance of earthworms from Greenland, Iceland, Can- 
ada, the northeastern part of the United States, northern Europe 
and perhaps also the British Isles. In North America a few spe- 
cies of two lumbricid genera survived but to this day they are 
largely confined to infrequently investigated habitats in the 
Appalachians. In Europe, the endemics, with a few exceptions 
that need further consideration, also have not advanced into the 
glaciated area. In that region the common species are those 
which seem most likely to have been living, during the glacial 
periods, just south of the ice cap. How closely they were able to 
follow the northward retreating ice remains to be learned. As 
none is known to be haemerophobic and many are haemerophilic 
the influence of man on their distribution in northern Europe 
must be determined first. That influence, hitherto underestimated 
for much of the world, enabled peregrine lumbricids, mainly 
from northern Europe, to acquire domicile throughout most of 
the settled regions of non-tropical North America in less than 
500 years. 

SUMMARY 

Soil around roots of plants has carried lumbricids to every 
part of the world where Europeans have lived. A high degree 
of tolerance for human disturbance of their habitats predisposes 
a score of species to transoceanic as well as subsequent intra- 
regional carriage and also colonization in vacant habitats where 
haemerophobic endemics have been eliminated by agriculture. 



16 BREVIORA No. 91 

Acquisition of domicile in the tropics only at elevations above 
5000 feet by otherwise successful colonizers shows an inherent 
inability to adjust to the milder and more equable climate of the 
lowlands. The size of the lumbricid domain, though India and 
China are excluded, indicates that the family is old. The Lumbri- 
cidae are conservative except as parthenogenesis permits more 
rapid accumulation of mutations. 

REFERENCES " 

Chen, Y. 

19.33. A preliminary survey of the earthworms of the lower Yangtze 
valley. Contr. Biol. Lab. Sci. Soc. China, Zool., vol. 9, pp. 177- 
296. 

1946. On the terrestrial Oligochaeta from Szechwan. III. Jour. West 
China Border Ees. Soc, B, vol. 16, pp. 83-141. 

Gates, G. E. 

1945. The earthworms of Allahabad. Proc. Nat. Aead. Sci. India, 
vol. 15, pp. 44-56. 

1947. Earthworms of the Allahabad sector of the Gangetic Plain. 
Ibid, vol. 17, pp. 117-128. 

1951. On the earthworms of Saharanpur, Dehra Dun and some 
Himalayan hill stations. Ibid., vol. 21, pp. 16-22. 

1953. On the earthworms of the Arnold Arboretum, Boston. Bull 
Mus. Comp. Zool., Harvard College, vol. 107, pp. 500-534. 

1956. Earthworms of the region south of the Allahabad sector of the 
Gangetic Plain. Proc. Nat. Acad. Sci. India, vol. 26, pp. 145-152. 

JULIN, E. 

1949. De Svenska Daggmaskarterna. Ark. Zool., vol. 42 (17), pp. 1-58. 

MiOHAELSEN, W. 

1928. Oligochaeta. In Kiikenthal and Krumbach, Handbuch der 
Zoologie, vol. II, fasc. 2, part 8, pp. 1-108, 103 text figs. Berlin 
and Leipzig. 

STia'HENSON, J. 

1930. The Oligochaeta. Oxford. 930 pp. 

Tetky, a. 

1938. Contribution k 1 'etude de la fauna de 1 'est de la France. Nancy. 



BREVIORA 

MiuseuntTi of Comparsitive Zoology 



Cambridge, Mass. September 17, 1958 Number 92 



A NEW GENUS OF ERETHIZONTID RODENTS FROM 
THE COLHUEHUAPIAN OF PATAGONIA 

By Bryan Patterson 

Completion of a monograph of the Deseadan rodents by Pro- 
fessor Albert E. Wood and myself, which includes incidental 
discussion of Colhuehuapian rodents, necessitates publication of 
this note. A full account of the new form will be given in a 
revision of the Colhuehuapian mammals now in preparation. 

Material of this rodent is contained in the collections of the 
Museo Argentino de Ciencias Naturales (MACN), the Museo 
Municipal de Mar del Plata, the Museum Nationale d'Histoire 
Naturelle, The American Museum of Natural History and the 
Chicago Natural History Museum. For the privilege of examina- 
tion and description, I am obliged to the authorities of these 
institutions, and in particular to Dr. J. -P. Lehman who was so 
kind as to forward specimens from Paris to Cambridge. For the 
opportunity of working in Buenos Aires and in Paris my thanks 
are due, respectively, to the John Simon Guggenheim Memorial 
Foundation and to the Carnegie Corporation. 

Seven species of erethizontids have been described from the 
Colhuehuapian : Eosteiromys homogenidens Ameghino, Steiro- 
mys pseudonectus Bordas, Steiromys segregatus Ameghino, 
Parasteiromys uniformis Ameghino, Steiromys axiculus Ame- 
ghino, Steiromys tahulatus Ameghino and Steiromys nectus 
Ameghino. The first three of these seem to be synonymous. 
Eosteiromys is a valid genus with a non-molariform Pi-, distinct 
postorbital processes, and nasals very strongly convex both 
anteroposteriorly and transversely. "Parasteiromys" uniformis. 
as Wood and I point out, appears to be referable to Steiromys, 
and is the only representative of this genus in the fauna. The 



BREVIORA 



No. 92 



last three species are indistinguishable, and S. tabulatus and 
S. nectiis are hereby placed in the synonymy of S. axiculus. This 
species, which is clearly not referable to Steiromys, is the type 
of the new genus. 





B 



Figure 1. Hypsosteiromys axiculus Ameghiiio. A, RP± — MJ.; B, LP-r- 



M: 



X4. 



Hypsosteiromys 1 gen. nov. 

Type species: — Steiromys axiculus Ameghino 1902. 

Distribution ;— Colhuehuapian, Patagonia. 

Diagnosis: — Cheek teeth with higher, more compressed lophs 
and lophids and deeper fossettes and fossettids than in other 
forms; Pi_not molariform, lacking metaloph; neoloph absent on 
M^^^, present on M-, rudimentary on P-i ; lophs and lophids 
showing marked tendency to irregular, very shallow notching; 



1 v^'^., on high, plus Steiromys, in allusion to the relative height of the molar 
crowns compared to those of other erethizontlds. 



1958 



NEW ERETIIIZONTID RODENT 



lower 1 small, short, extending posteriorly to end of M3 ; con- 
spicuous depression in maxillary anterior to tooth row and 
medial to root of zygomia. 



Hypsosteiromys axiculus Ameghino 

Steiromxjs axiculus Ameghino 1902, p. 109. 

Steiromys nectus Ameghino 1902, p. 109. 

Stciromys tahuJatus Ameghino 1902, pp. 109-110. 

Stichomys regius? Bordas (non Ameghino) 1939, p. 428 fig. 9. 

Type: — MACN no. A 52-171, fragment of right ramus with 
root of I, Mtij (Mj not fully erupted) . 

Type of 8. nectus: — MACN no. A 52-177, fragment of right 
ramus with root of I, dmj, Mp2 (^2 ^^^ fuHy erupted). 

Lectotype of S. tahulatus :—MAC^ no. A 52-172, fragment of 
left ramus with Fj — M3, crushed alveolus of I. Two other 
syntypes, an upper and a lower incisor from different indi- 
viduals (MACN no. A 52-180), are of very dubious reference. 

Horizons and localities: — All specimens but one are from the 
( 'olhue-Huapi formation in the Barranca south of Lake Col- 
hue-Huapi, Province of Chubut ; the exception (Bordas 1939, 
MACN no. 11247) is from the Trelew formation in the South 
Barranca of the Chubut Valley at Caiman, Province of Chubut. 

Diagnosis: — As for the genus. 

I append measurements of Ameghino 's types, and drawings by 
Mrs. Dorothy Marsh of two specimens in the Museum Nationale 
d'Histoire Naturelle collection. Measurements in millimeters. 





IT 


dm^" 


P4 


MT 


M2 


M3 




ap. 


tr. 


ap. 


tr. 


ap. 


tr. 


ap. 


tr. 


ap. 


tr. 


ap. 


tr. 


MACN no. 
A 52-171 


— 


3.0+ 










5.6 


4.4 


5.7 


4.8 






MACN 
A 52-177 


• — - 


2.4 


7.3 


3.5 






5.0 


4.3 










MACN 
A 52-172 










7.1 


5.2 


5.3 


4.9 


5.2 


5.2 


5.7 


4.6 



4 BREVIORA No. 92 

REFERENCES 

Ameghino, ¥ 

1902. Premiere contribution a la connaissance de la faune mam- 
malogique des couches a Colpodon. Bol. Acad. Nac. Cienc. 
Cordoba, 17:71-138. 

BOBDAS, A. F. 

1939. Diagnosis sobre algunos mamiferos de las capas con Colpodon. 
Physis, 14:413-433. 

Wood, A. E. and B. Patterson 

(In press.) The rodents of the Deseadan Oligocene of Patagonia and the 

beginnings of South American rodent evolution. Bull. Mus. 

Comp. Zool. 



BREVIORA 

MiiseiiMti of Comparative Zoology 



Cambridge, Mass. September 18, 1958 Number 98 



A NEW BARYLAMBDID PANT(3D0NT FROM THE 

LATE PALEOCENE 

By Bryan Patterson and Elwyn L. Simons 

A quarry discovered during the field season of 1939 by a Chi- 
cago Natural History Museum party working in the late Paleo- 
cene beds of western Colorado yielded numerous disarticulated 
remains of a large barylambdid pantodont. It was at once 
(^-ident, following preparation, that this animal was distinct 
generically from the large Barijlarnhda faberi, and certainly 
distinct specifically from the smaller Haplolamhda quinni (Pat- 
terson, 1939), both from the same horizon. The nature of the 
type and only specimen of H. quiinii (the anterior half of a 
skeleton, much of it pieced together and restored from weathered- 
out fragments) left some doubt as to the degree of relationship 
between this form and the new one, and description of the latter 
Avas accordingly postponed. During recent years much panto- 
dont material has been obtained from the Paleocene of the Polecat 
Bench formation of Wyoming by Princeton University parties 
under Professor Glenn L. Jepsen, supported by the William 
Berryman Scott Research Fund. This has now been prepared 
and is under study by Simons. Comparison of the Chicago and 
Princeton collections shows that Haplolamhda and the new form 
are represented in both and that they are beyond doubt generi- 
cally distinct. A specimen from Colorado in the Carnegie 
IMuseum also proves referable to the new form. 

Edinger (1950, 1956) has published some notes on the frontal 
sinus and endocranial cast of Colorado specimens of the new 
form under the name of Haplolamhda? sp. 

"We wish to express our sincere thanks to Professor Jepsen and 
to Dr. J. LeRoy Kay for the opportunity of studying specimens 
in the collections under their respective charges. 



BRF.VIORA 



No. 93 



PANTODONTA 
BARYLAMBDTDAE 



Leptolambda^ 



<i'on. nov 



Type species. L. schiiudii sp. nov. 

Distrihution. Late Paleocene, Tiffanian stage; Silver Coulee 
beds, Polecat Bench formation, Park and Bighorn counties, 
Wyoming; Plateau Valley beds, DeBe(iue formation, ^lesa 
County, Colorado. 

Diae/nosis. 1.^ CvP-i-M.^, no diastemata. Incisors above small, 
with lateral crest on each side ; lower incisors larger than up- 
lier with lateral crests more prominent, forming a spatulate 
tooth comparatively longer anteroposteriorly and shorter trans- 




Figure 1. LeptoJamhiln schmidti gen. et sp. nov. Upper dentition of 
right side, PU Xo. 14996 (unshaded teeth from PU Xo. 14680). x V... 



versely than in Baruhniibda. Canines smaller than in Bavii- 
lamhda, in supposed females incisiform. Premolars of typical 
barylambdid pattern; P| one rooted; P-1- usually broader trans- 
versely than in Bayylambda. Talonid bases of P 2:3- compara- 
tively larger than in Barylambda or HapJoJanihda, with serial 
homologue of crista obliqua directed anteriorly, not running 
obliquely anterointernally to long axis of tooth as in BaryJamhda. 
Upper molars differing from those of Haplolamhda and resem- 
bling those of Bar]ilamhda in M-=- larger than M— ; distinguished 



.. ,„ jlender ; with reference to the less robust post- 
ore slenrter construction of the long bones than in rtnru. 



I From the Greek ir/j/os. thin, sieiuier ; wun reiereiice to cne less rouust post- 
cranial skeleton and more slenrter construction of the long bones than in Bary- 
lambda , plus -lambda, referring to the Zanibda-shaped molar crests, and to agree 
with related genera. 



1958 NEW BARYLAMBDID PANTODONT 3 

from Bcn-i/ldDihdd by presence of heavier einjiula on M— , absence 
of backward rotation of protocoiies of AI— . jNIetacone of M— less 
reduced tlian in /Idplohnitlxla. Skull with wider, long'er frontal 
area than in Bdri/lainlxld and llaplolanibda ; head as a whole 
smaller in ])roportion to body size than in other barylambdids. 
Cervical centra and neural arches very short, weak; thoracic and 
lumbar vertebral centra comparatively lon<>'. Postcervical verte- 
brae with lower nein-al spines than in Barylaiiihda and Haplo- 
lamhda. Caudals without (or at most with small, anterior) 
haemapophyses ; caudal centra longer in i)roportion to width 
and height than in Barylamhda. Scapula not as broad antero- 
posteriorly, particularly in postscapular portion, as in Bary- 
lamhda, broader than in Haplolamhda ; spine proportionately 
narrower than in Baryhnitbda, lacking tuberosity. Clavicle more 
slender than in Barylauihda. Bones of forelimb shorter relative 
to hind than in Baryhnnhda; deltopectoral crest of humerus not 
as fiat, less projecting medially ; shaft of ulna less broad from 
front to back, strongly retrofiexed, olecranon relatively longer 
than in Barylamhda and Haplolamhda. Metacarpal I and 
tra]')ezium not fused as in HapJoJamhda. Metacarpal V much 
shorter than in Barylamhda, relatively shorter than in Haplo- 
la)iihda, with large lateral expansion in proximal half. Proximal 
and mesial phalanges differing from Barylamhda in much 
greater compression (or foreshortening) along axis of digit. 
Bones of hind limb less robust than in Barylamhda; femur with 
more slender, less flattened shaft, third trochanter nearly vesti- 
gial ; tibia with more slender sliaft, no pronounced cnemial crest. 
distal extremity proportionately less broad; proximal articula- 
tion for fii)ula less projecting laterally; tuber calcis of calcaneum 
very much shorter. 

/ Leptolambda schmidti- sp. nov. 

Figs. 1-3 

Type. CNHM^ No. P26075, incomplete skull, mandibles, numer- 
ous vertebrae and ribs, incomplete scapula and pelvis, various 
leg and foot bones. 

2 Named for Dr. Robert G. Schiiiiilt. who, with Dr. James H. Quinn, discovered 
the quarry from wliich nearly all the Plateau Valley material of this species was 
obtained. 

3 Ahl)reviations are as follows : CXIIM. Chicago Natural History Museum ; CM, 
Carnejrie Museum : PU, Princeton University. 



BREVIOBA 



No. 93 



Hypodigm. Type and CNHM Nos. P26076-7, incomplete sknll, 
various vertebrae and ribs, incomplete scapula, various leg and 
foot bones ; P15558, P15571, incomplete dentitions ; CM No. 11353, 
facial region of skull with dentition ; PU Nos. 14680, 14879, and 
14996, partial skeletons; PU Nos. 14681, 14990, 14992. mandibles. 

Horizon. DeBeque formation, Plateau Valley local fauna. 
Mesa Co., Colorado ; lower levels of the Silver Coulee beds. Pole- 
cat Bench formation. Park and Bighorn counties, Wyoming. 






B 



Figure 'l. A, ulna of Ihuylarnhda faheri Patterson, CNHM No. P26110, 
and B, ulna of Lcptolamhda .scJiiitiiIti gen. et sp. nov., PU No. 14879. x %. 



Localities. CNllM Xos. P26075-7 from one ([uarry apjiroxi- 
mately four miles 8SE. of DeBeque, Mesa Co., Colorado (due 
to complete lack of association in this quarry there is uncer- 
tainty as to the ])recise allocation of the individual bones to 
these numbers) ; Ci\I No. 11353, 2 to 3 miles west of DeBeque, 
Mesa Co., Colorado; VV Nos. 14680 and 14681 from the south 
side of Polecat Bench, Park Co., Wyoming; PU Nos. 14879. 
14990, 14992, nm\ 14996 from separate localities ah)ng a NU-SW 



1958 



NEW BARYLAMBDin PANTODONT 



line between the towns of Lovell and Greybnll in T 54 and 55 N, 
R 95 and !)(i AV, Bijiiiorn Connty, Wyoming. 

I)i.a(/H()sis. As for tlie genns ; size approximately as in Bary- 
lambda faberi or somewhat smaller, hut proportions clearly 
very different. Sjieeimens from Colorado and from Wyoming fall 





B 





D 



Figure .'5. Femora and til)iae of A, C, Barylamhda faheri Patterson, 
CNHM No. ri494-4, and B, I), of Leptolamhda schmidii gen. et sp. nov., 
CXHM Xos. P26075, type and P2(i077. x %. 



6 BREVIORA No. 93 

iuto two rather sharply defined size groups, one 20-30 per cent 
smaller than the other. The distinction is most apparent in the 
region of the canine, within the dentition, and in the postcranial 
skeleton. Differences of approximately this order of magnitude 
also occur in Bafyhniihda fahcri. We interpret them as indicat- 
ing a sexual difference in size. 

Table 1 
MEASUREMENTS OF THE DENTITION, IN MILLIMETERS 







si 


l-O 














^•5 




c 


11 


ap 
tr 




6.0 
5.2 








11 


ap 

tr 




8.0 
6.0 


8.8 
6.2 


8.0 
6.2 


12 


ap 
tr 






7.7 
6.3 






12 


ap 
tr 




10.0 
6.2 


11.0 
6.8 


10.5 

7.0 


I-"' 


np 




8.8 


9.1 






I-'^ 


ap 




11.5 


12.7 


10.5 




tr 




9.0 


6.8 








tr 




7.0 


7.7 


7.0 


c 


ap 


1-1.5 


13.4 


10.9 






C 


ap 




12.8 


15.7 


12.3 




tr 


15.0 


13.5 


8.8 








tr 




8.8 


11.4 


7.2 


pi 


ap 




15.0 


14.6 


13.8 




Pl 


ap 




13.5 




16.0 




tr 




14.0 


8.4 


8.0 






tr 




8.6 




8.6 


p- 


ap 


18.0 


16.0 






18.2 


P-" 


ap 




20.5 


21.5 


19.0 




tr 


25.0 


26.0 






25.5 




tr 




12.3 


15.7 


12.6 


p3 


ap 


20.5 


17.0 




17.0 


17.8 


p8 


ap 


22.5 


20.5 


20.5 


19.0 




tr 


27.2 


29.5 




25.0 


27.0 




tr 


18.0 


16.0 


18.3 


15.0 


P-l 


ap 




18.0 








I'-i 


ap 


22.0 


21.0 


21.6 


20.2 




tr 




32.0 










tr 


20.5 


18.0 


19.5 


17.0 


Ml 


ap 




26.0 


23.0 




24.5 


Ml 


ap 


2.1.0 


22.5 


23.2 


21.8 




tr 




36.0 


31.5 




31.6 


tr 


trig 


19..1 


17.0 


18.0 


18.2 


M^ 


ap 


26.0 


27.0 


23.5 


22.8 


25.3 


tr 


tal 


19.5 


17.8 


18.3 


16.0 




tr 


36.0 


39.0 


32.5 


34.1 


35.3 


Mo 


ap 


26.7 


23.0 


24.5 


23.3 


M3 


ap 


21.0 


21.11 


17.4 


18.1 


19.8 


tr 


trig 


21.0 


17.0 


18.2 


17.2 




tr 


34.5 




30.0 


31.2 


33.4 


tr 


tal 


17.0 


17.0 


18.3 


15.4 


4 


The a 


nteroposterior 


 diameters 


of the 


M;: 


ap 


31.5 


26.0 


30.4 


29.0 


upper cheek teeth of this sp 


lecimen ap- 


tr 


trig 


19.0 


16.3 


18.2 


17.5 


pea; 


r to h; 


five bee: 


11 somewhat 


redueed h\ 


tr 


tal 


13.5 


13.4 


14.4 


12.5 



wear. 



1958 NEW BARYLAMBDII) PANTODONT 7 

RcDKirhs. The Carnegie Museum specimen furnishes an in- 
teresting footnote to the history of Paleoeene exploration. The 
Plateau Valley fauna did not come to seientifie attention until 
the early 19;30's, following Edwin B. Faber's discovery of the 
tyiie and other specimens of Banjlambda faheri. CM No. 11353, 
h()W(>\('i-, was collected by Earl Douglass about 1909, was tenta- 
tively identified as Pantolamhda? sp., and the horizon recognized 
as Paleoeene. Having made and correctly interpreted this re- 
markabl(> find, Douglass went on to the discovery of the great 
Jensen dinosaur quarry, exploitation of which .so fully occupied 
the rest of his working life that he was never able to follow up 
the lead provided by this first identified Paleoeene mammal from 
western Colorado. As an additional item of interest, his specimen 
provides our only adequate knowledge of the facial region of 
Lcpfolamhda schmidti, a part otherwise represented only by 
isolated maxillaries in the Chicago Natural Historj- Museum 
material and by a much crushed skull in the Princeton collec- 
tions. 

The field evidence in Colorado suggests that Leptolamhdo 
schmidti differed from other Plateau Valley pantodonts as re- 
gards habitat. The previously known forms were all found in 
mud- or siltstones, whereas every specimen identifiable as L. 
schmidti was collected from the fillings of stream channels. The 
quarry that yielded the great bulk of the material w^as in such a 
situation, the disarticulated bones occurring at the junction of a 
channel fill with the underlying mudstone but clearly deposited 
in the channel. Leptotamhda presumably inhabited higher 
ground marginal to the basin in which DeBeque sedimentation 
was going on, wuth fragments and occasionally whole or partial 
carcasses being carried down into the area of deposition. The 
specimens of Leptolamhda collected in the Bighorn Basin of 
Wyoming neither confirm nor deny this conjecture. At least 
three specimens, PU Nos. 1-1680, 14681, and 14879 were collected 
from mudstones, whereas PIT Nos. 14990, 14992, and 14996 were 
recovered from sandstones. Nevertheless, the possibility that 
Barylamhda and Leptolamhda were adapted to rather different 
environments is suggested, in Wyoming, by the fact that the 
former genus has not been recovered from the Bighorn Basin, 
although Leptolamhda is comparatively common there for a 



8 BREVIORA No. 93 

pantodont, being represented by more than twenty-tive separate 
finds. Since these two genera occur at the same stratigraphic 
levels in Colorado it is clear they were co-existent in time and 
therefore the absence of Barylamhda in the Polecat Bench for- 
mation suggests that the environmental conditions under which 
it flourished did not exist in the region of the Bighorn Basin 
during the deposition of the Silver Coulee beds. 

REFEEENCES 

Edinger, T. 

1950. Frontal sinus evolution (particularly in the Equidae). Bull. 

Mus. Comp. Zool., 103:-409-496. 
]9.j(). (^l)jets et resultats de la Paleoneurologie. Ann. Palaeont., 

42:97-116. 

Patterson, B. 

1939. Xew Pantodonta and Diiiocerata from the upper Paleocene of 
western Colorado. Geol. Ser. Field Mus. Nat. Hist., 6:351-384. 
(And earlier references there cited.) 

Simons, E. L. 

MS. The Paleocene Pantodonta and their allies. Doctoral disserta- 
tion, Department of Geology, Princeton University (to be pub- 
lished). 



BREVIORA 

Miuseunra of Coimparative Zoology 



Cambridge, Mass. September 18, 1958 Number 94 

AFFINITIES OF THE PATAGONIAN FOSSIL MAMMAL 

NECROLESTES 

By Bryan Patterson 

Since its description by Ameghino in 1891, the curious little 
fossorial Necrolestes from the Santa Cruz, beds has never rested 
securely in the system. Ameghino himself always regarded it as 
an insectivore allied to the African chrysoehlorids. Scott (1905), 
in his description of excellent material collected by the Princeton 
Patagonian Expeditions, compared it with Chrysochloris and 
Notoryctes and concluded that it was an insectivore related to 
the African form : ' ' What does seem probable is that Necrolestes 
is an offshoot from some early member of the family which was 
at the same time the ancestor of Chrysochloris." Gregory (1910, 
p. 259) and Schlosser and Woodward in various editions of Zittel 
accepted Necrolestes as an insectivore, but others have queried 
or denied such a relationship. Leche pointed out that Necrolestes 
resembled Notoryctes about as much as it did Chrysochloris, that 
as a marsupial it would raise fewer zoogeographical difficulties 
than as an insectivore, and that its affinities were far from 
decided : " . . . die Beziehungen des Necrolestes zu den lebenden 
Saugern durch das bisher dargelegten material keineswegs als 
festgestellt betrachtet werden konnen" (1907, p. 143). Abel (wi 
Weber 1928, pp. 128-129), while discussing the genus under the 
Insectivora, expressed similar reservations. In current reference 
works, Necrolestei^ has been questionably included either among 
the marsupials (Homer 1945, p. 611) or among the insectivores 
(Simpson 1945, p. 53). Saban (1954, p. 422) has made the 
passing suggestion that it may be a palaeanodont edentate. One 
author who came out unequivocally for marsupial affinities was 
Winge (1941, pp. 79-80, 217-218), who regarded the incisor 



2 BREVIORA No. 94 

formula, reported by Scott to be;*, as proof of such relationship. 
He believed Necrolestes to be a didelphoid derivative, classifying 
it in a separate division, Necrolestini, of his family Cladosictididae 
( =Borhyaenidae ) . 

No first hand account of this little creature has appeared 
since Scott's study. His work was perforce done without benefit 
of the binocular microscope, and his conclusions as to affinities 
suffered from the fact that he evidently compared Necrolestes 
only with the equally highly specialized Notoryctes and the 
chrysochlorids, not at all with more generalized insectivores and 
marsupials. It had always seemed to me that a form represented 
by such excellent material • — with most of the skeleton preserved 
in one or another of the Princeton specimens it is actually among 
the better known fossil mammals — should reveal enough in 
the way of structure to permit at least its infraclass affinities to be 
firmly established. Recent examination and further preparation 
of the Princeton material has shown this to be indeed the case. 
Necrolestes is beyond any doubt a marsupial and, as regards its 
position within the order, Winge's suggestion of a remote rela- 
tionship with the borhyaenids is perhaps as reasonable a one as 
can now be offered. Scott's account is in general very good 
indeed. The brief description that follows is supplementary to 
his and corrects it in a few points. 

I wish to express my sincere thanks to Professor Glenn L. 
Jepsen and to Dr. Donald F. Baird, who very kindly took the 
photographs from which the drawings prepared by Mrs. Dorothy 
Marsh were made. 

MARSUPIALIA 
BORHYAENOIDEA 
NECROLESTIDAE 

Necrolestes patagonensis Ameghino 

Knowledge of the species, the only one of the family thus far 
discovered, rests essentially on three specimens in the Princeton 
collections : 

15065. Skull, crushed dorsoventrally and lacking parts of 
occiput and basicranium, mandible (now missing), various 
vertebrae and ribs, proximal end of humerus. Ten miles South of 
Coy Inlet, Santa Cruz; collected by 0. A. Peterson. 



1958 AFFINITIES OF NECROLESTES 3 

15384. Occiput and part of basicranium, mandible, some iso- 
lated upper teeth, numerous vertebrae from all parts of the 
column, pelvis, humerus, radius, ulna and various bones of the 
fore foot. Five miles South of Coy Inlet, Santa Cruz; collected 
by 0. A. Peterson. 

15699. Skull, incomplete posteriorly, mandible, right humerus, 
radius and ulna, left radius, incomplete ulna, femur, tibia, fibula, 
calcaneum, astragalus, cuboid. Killik Aike, Santa Cruz ; collected 
by J. B. Hatcher. 

DENTITION 

Dental formula. It can be definitely stated that 
this is I ^, Cf, postcanines -| (probably P|, M-|)- Recogni- 
tion of the correct number of upper incisors at this late date, 
more than half a century after the full account of the material, 
comes as somewhat of a surprise. The evidence is unmistakable, 
however (Fig. 1 A, B). In 15699, I^^ are preserved on the left 
side and the alveoli of these four teeth, together with I i, on the 
right. No. 15065 retains right and left I—, right I-^nd the alveoli 
of all other members of the series. Curiously enough, van Iterson, 
the artist who drew Scott's figures, indicated the correct number 
(Scott 1905, pi. 64, figs. 1, 2a). For 15699 he showed five in- 
cisors in right side view, reversing li^ from the left side, and 
for 15065 the right and left I- , right I— and the alveoli of 
right I^^ (the alveoli of left I-^ were at that time concealed 
by matrix). The figure of 15065 has right 1-5- erroneously 
labeled as I-^. Determination of the postcanine formula is un- 
certain. Believing that he was dealing with an insectivore, 
Scott gave it as Pf , Mg . Since we have a marsupial on our 
hands, either this formula could be correct, with M-f missing, or 
the first premolar could be the missing tooth, which would 
give P|, Mj. The latter possibility seems to me the more 
likely of the two. The last four cheek teeth above and below 
are uniform in structure, which suggests, although of course 
does not prove, that all are molars, while the second pair of 
teeth, especially the lower, although largelj^ molariform, do show 
some resemblances to the anterior pair. 



4 BREVIORA No. 94 

Upper teeth. The first four incisors are of similar struc- 
ture, with the crowns rectangular, compressed anteroposter- 
iorly, the labial face slightly convex in the dorsoventral 
diameter and bearing a faint median groove. I— is the 
largest, I— slightly smaller and I-^of equal size and slightly 
smaller than 1J-. I — Is the smallest of the series. It is a bluntly 
pointed tooth, worn obliquely on the posterior face by the 
anterior surface of the lower canine. 

Relative to the sizes of the other teeth, the single-rooted, tri- 
hedral canines, both upper and lower, are enormous. The upper 
is somewhat variable, that of 15699 being stouter, more triangular 
and with a larger anterior face than that of 15065. 

The first upper cheek tooth, P^as here interpreted, is double- 
rooted, implanted obliquely, and consists of a high, conical 
paracone, an anterior basal cusp and a shearing edge running 
from the apex of the paracone to the large but non-cuspidate 
metastylar area. The remaining teeth of the postcanine series 
are unilaterally hypsodont, curving outward toward their bases 
and tapering to small, bluntly pointed single roots (Fig. IC). 
The enamel extends nearly to the end of the root of the 
lingual surface and for only a short distance up the crown on 
the labial. M^^ are essentially narrow isosceles triangles as re- 
gards crown contour; Pyhas a convex anterior and M— a convex 
posterior border. ]\I— are subequal, M— is slightly smaller and M— 
reduced in size. The internal cusps are presumably paracones, 
being in line with the paracone of P—, and the greater part of the 
crowns is formed by the stylar area. There is no vestige of a 
protocone. These teeth are thus descriptively zalambdodont, but 
no structural details are apparent in the worn material available. 

Lower teeth. The incisors resemble those of the upper series and 
are not, as Scott stated, laterally compressed. ly is much larger 
than the others, which are subequal. The canine is narrower 
and less trihedral than the upper, with a smaller anterior face. 

Pyis double-rooted, elongate anteroposteriorly and bears three 
cusps, a high, acute protoconid and smaller, subequal an- 
terior and posterior basal cusps. The molars are alike in 
structure and are mirror images of the uppers, with no traces 
of talonids. The second postcanine, P-j- as I believe, differs 
from them in being less narrowly triangular and in the pos- 



1958 AFFINITIES OF NECROLESTES 5 

session of a vestigial anterior basal cusp serially homologous 
with that occurring on P ^ . The less worn of the lower molars 
available have heart-shaped crown contours caused by a shallow, 
ephemeral indentation between the bases of the paraconids and 
metaconids. No doubt P-3--M— are mirror images of the cor- 
responding upper teeth as regards unilateral hypsodonty, enamel 
distribution and root structure but no loose examples are 
available. Wear on all molars is limited to the crowns, where 
the abrasion surfaces are nearly plane. It is quite evident that 
in active occlusion the trigonids of the lowers worked across the 
trigons of the uppers. 

SKULL 

As is the case in other small fossorial forms, no suture is 
apparent in any specimen of Nccrolesfes. The chief peculiarity 
of the skull is of course the extraordinary upturned snout, which 
extends well above and in front of the incisors (Fig. 1 A, B). 
Scott was in some doubt as to whether this feature might not 
have been exaggerated by distortion in 15699, particularly as 
there is little trace of it in 15065. It appears to be natural, 
however. No. 15699 has undergone slight lateral compression, in 
the course of which the maxillaries and premaxillaries have been 
broken away from and slightly pinched in relative to the nasals, 
but this does not appear to have affected the dorsal profile of 
the facial region. No. 15065 has been badly crushed dorso- 
ventrally in this region and the upturned portion has been 
w^eathered off. The anterior narial openings (Fig. 1 A, B) extend 
forward to the tips of the nasals and do not end short of them, 
as Scott's figure shows them as doing. The median spine de- 
scribed by Scott as extending upward and forward from the 
mid-point of the premaxillaries evidently continued to the tips 
of the nasals as an ossification in the septum. It is partly broken 
away but remnants of it are still present. Between I- and the ca- 
nines, the premaxillaries are very deeply excavated for the 
reception of the lower canines, perhaps the most extreme develop- 
ment of this character to be seen in the Marsupialia. 

The upwardly arching z^^gomatic arch is well developed for 
so small an animal (Winge described it as slender, but this was 
in comparison with the arch of the borhyaenids) and quite simi- 



BREVIORA 



No. 94 



lar to that of small didelphids such as Marmosa. Extension of 
the jugal to the glenoid articulation is a marsupial character. 
Despite the obliteration of the sutures, there can be little doubt 
that the jugal of Necrolestes was so constructed. A prominent 




Fig. 1. Necrole.stcfi patagoneiisis Ameghino. A, B, riglit anr1 left sides 
of snout region of skull, X3 approx. (LI, left incisor) ; C, left M2, anterior 
view, X6 ; D, left innominate bone, lateral view, X2 approx. 



j)reglenoid process is present. The glenoid surface extends for- 
ward to this buttress and is in consequence deeply concave 
anteroposteriorl}'. Precisely similar conditions exist in other 
marsupials, in which the buttress is formed by the jugal. A post- 



1958 AFFINITIES OF NECROLESTES 7 

glenoid process is lacking;. The extent of the lachrymal cannot, 
of course, be seen. The lachrymal foramen, however, is so very 
similar to that of various didelphids as regards both position and 
size (far more so than to that of tlie chrysochlorids, with which 
Scott compared it) as to suggest that the bone itself was likewise 
comparable. Within the orbito-temporal region, the optic foramen 
appears to be confluent with the sphenorbital fissure. 

The long, slitlike incisive foramina are very didelphid-like, 
while the complete absence of palatal vacuities is a point of 
resemblance to the borhyaenids. The posterior border of the 
palate is moderatelj- thickened and the posterior nares are di- 
vided by a median bony septum. For a marsupial this is very 
unusual ; it may be an adaptation for providing an increased 
rigidity that would be useful to an animal using its upturned 
snout as an earth mover. 

Specimens 15065 and 15384 reveal a good deal of the structure 
of the basicranium and auditory region, much of which had 
hitherto been concealed by matrix. The basicranium is very 
short. The greater part of the right side of the basisphenoid is 
preserved in 15065 and this portion is perforated, at the level 
of the glenoid cavity, by a large foramen, which appears to be 
the carotid. The bone is almost flat and there is no indication of 
a tympanic wing. The basioccipital, of which only a small part 
is preserved in 15384, was clearly short and w4de. As noted by 
Scott, the condyles are relatively very large and wide ; as in 
Marmosa there is a conspicuous median notch between them 
that opens forward from the foramen magnum. The latter 
opening is large and faces posteriorly. There is no paroccipital 
process, the "broad and distinct" process so identified by Scott 
being the mastoid. That part of the alisphenoid external to the 
foramen ovale, of which the lateral rim is preserved, is present 
in 15065; it shows no trace of any development of an alisphenoid 
bulla. It is perforated by a small foramen, presumably vascular, 
situated midw^ay between the foramen ovale and the glenoid 
cavity. The periotic is short and robust, with a well rounded 
promontorium, which I take to have been the basis for Scott's 
incorrect statement that the "tympanic bulla" was "ossified 
and moderately inflated." Anteriorly, the bone terminates 
bluntly and, posteriorly, there is no trace of a tympanic wing. 



8 BREVIORA No. 94 

The fossa subarcuata is very large, and the anterior vertical 
semicircular canal is free from surrounding bone. The mastoid 
process appears to be large. 

, Immediately posterior to the glenoid cavity is a more or less 
triangular pocket, whose medial border is formed by the petrous 
portion of the periotic, the lateral by the squamosal and mastoid, 
which descend below the level of the glenoid and are concave 
on their medial surfaces, while the squamosal appears to be 
the main contributor to the roof, with the other elements par- 
ticipating to a lesser extent. A low transverse ridge crosses the 
roof of this pocket, partially dividing it into a large anterior 
portion, which is open below, and a smaller posterior portion, 
which is floored by the mastoid process. The fenestra ovalis and 
the apertura tympanici canalis facialis open into the pocket and 
the groove for the facial nerve is plainly visible in the lateral 
side of the periotic. This pocket, then, is the tympanic cavity, 
and the tympanic itself, of which no part remains, obviously 
provided a partial floor to the ventrally open anterior portion. 
There is no emargination of the squamosal marking the passage 
of the auditory meatus, which must have passed laterally below 
and behind the glenoid cavity. Within the pocket, in 15065, 
there is a minute foramen immediately behind the glenoid and 
a larger one just posterior to it. These I take to be the postzygo- 
matic and postglenoid foramina. The tympanohyal, descending 
from the crista facialis, is fused with a small medial projection 
from the mastoid process in 15065 but stands free in 15384. The 
"foramen" stylomastoideum primitivum is entirely open, merely 
the termination of the facial groove, as in various didelphids. 
Posterior to the tympanohyal is the opening of a short tubular 
recess, covered laterally by the mastoid and exoccipital, that runs 
back to the condyle. An almost identical recess occurs in small 
didelphids, e.g. Marmosa, and presumably houses M. stapedius. 
Of the preserved foramina not heretofore mentioned, the lacerum 
posterius is large and the condylar, due to the shortness of the 
basicranium, is very close to it. The latter, so far as I can detect, 
is single. There appears to be no subsquamosal foramen. 

The posterior portion of the mandible is greatly reduced 
and the angle consequently very small. Notwithstanding, this 
jn'ocess bears an internal ledge and tliis is inflected and concave 
dorsallv. 



1958 AFFINITIES OP NECROLESTES 9 

The fossorial specialization has resulted in a highly modified 
skeleton that shows numerous resemblances to other, similarly 
adapted forms, as Scott pointed out. Making due allowance for 
this, there is nothing that argues against a didelphoid ancestry 
for the Necrolestidae. The lack of an anticlinal vertebra, a point 
emphasized by Scott as a resemblance to the chrysochlorids is 
also true for the thoracic series of some of the smaller didelphids, 
and in these the neural spines of the lumbars do not incline 
forward to any appreciable extent. There is clear evidence of 
a well-developed tail, in agreement with didelphids and bor- 
hyaenids. 

The proportioiLs of the pelvis closely resemble those of the 
smaller didelphids and, as Scott pointed out, there w^as surely 
a well-developed pubic symphysis. The ilium is more rodlike 
than in the Didelphidae, a diiference possibly related to the 
fossorial adaptation of Nccrolestcs. In very close agreement with 
didelphids, there is a conspicuous pectineal prominence that 
stands well out from the anterior margin of the pubis. The rela- 
tively large epipubis of the smaller opossums extends up nearly 
to the pectineal prominence, and I suspect that the same w^as 
true of Necrolestes. On the right side of 15384 there is a small 
fragment of bone attached to this area of the pelvis by matrix, 
and it is just conceivable that this could be a remnant of the 
epipubis itself. 

DISCUSSION 

Winge was correct in his opinion that Necrolestes was a 
marsupial. The incisor formula is diagnostic for a large seg- 
ment of this order. There is nothing of a conflicting nature in the 
structure of the skull or postcranial skeleton ; on the contrary 
numerous features described above point so strongly in the same 
direction that there is no further need to elaborate on the ordinal 
position of the genus. Saban was quite right in seeking to exclude 
the Necrolestidae from the Insectivora, and his tentative sug- 
gestion of palaeanodont affinities was a reasonable one, given 
the information in Scott's account. The new facts do not support 
it, however. The Palaeanodonta resemble the Xenarthra in the 
possession of an ossified bulla composed of ecto- and entotympanic 
elements (Patterson and Segall, MS.) which Necrolestes does not 



10 BREVIORA No. 94 

have; the dentition and the pelvis are quite different, and so 
on. There is, of course, a general resemblance between Epoico- 
therium and Necrolestes but, like the resemblance between the 
latter and the chrysochlorids, this is due to convergence.^ 

There remains for discussion the problem of where, within 
the Marsupialia, Necrolestes should be placed. The isolated posi- 
tion of the genus — its lack of any ancestors combined with its 
great specialization — prevents any satisfactory conclusion, al- 
though some approximation is possible. The facts of zoogeography 
would seem to limit the field to the known South American 
groups. One of these, the Caenolestoidea, may at once be ex- 
cluded from consideration. No known caenolestoid approaches 
Necrolestes in any significant way ; the dentitions of the earliest 
forms are specialized in completely different directions. The 
remaining two groups, the Didelphoidea and the Borhyaenoidea, 
on the contrarj^ do show resemblances of a basic sort to the Necro- 
lestidae. Some of these have been mentioned in the description. 
As regards the dentition, the incisors of didelphids — and pre- 
sumably of the earliest borhyaenids — could readily have given 
rise to those of Necrolestes; the canines are large in the two, and 
the lower is received into a socket in the upper jaw ; the zalamb- 
dodont molar type of the necrolestids could readily have evolved 
from tribosphenic molars of the sort that were no doubt present 
in the early members of both groups. The skull and postcranial 
skeleton of Necrolestes reveal, beneath the facade of fossorial 
specialization, unmistakable borhyaenid and didelphid resem- 
blances. 

If, as seems virtually assured, the ancestry of the Necrolestidae 
is to be sought near the base of the didelphoid-borhyaenoid com- 
plex, then the question arises as to whether it is nearer to one 
or to the other of these superfamilies, or is equally distinct from 
both. Necrolestes is one of the most specialized of American 
marsupials, specialized in a manner totally unlike anything seen 
in the Didelphoidea or in the Borhyaenoidea. To seize upon this 
as an excuse for erecting a superfamily for the reception of one 



1 Since this was written, McDowell (Bull. Amer. Mus. Nat. Hist., 115, 173-4, 
l'J58) lias asserted that A'ccro/c.y/e.s- is a member of the suborder Xeuarthra. In 
addition to the characters of the auflitory region just cited that oppose reference 
to the suborder I'alaeanodonta, the presence of rooted, enamel-covered teeth and 
the at)sence of the diainiostic xenarthran vertebral and pelvic structures prevent 
acceptance of this opinion. 



1958 AFFINITIES OF NECROLESTES 11 

genus whose phylogeny is totally unknown, would be the height 
of irresponsibility, however. Furthermore, there are one or two 
characters that hint at a more acceptable possibility. As will be 
evident from the descriptive notes, there are more resemblances 
to the didelphids than to the borhyaenids in the anatomy of 
Necrolestes. This is not surprising : the didelphids include the 
least specialized of marsupials and no doubt retain a good deal 
of the ancestral structure relatively unmodified. The adequately 
known borhyaenids are relatively much more specialized and 
would therefore be less likely to display basic resemblances to 
Necrolestes. However, in such characters as the absence of 
palatal fenestrae and the lack of an alisphenoid bulla, Necro- 
lestes is closer to all or some of the borhyaenids, which conceiv- 
ably may indicate that the family arose from the borhyaenoid 
stock shortly after its divergence from the didelphoid. Such evi- 
dence, of course, is of the slenderest yet I nevertheless clutch 
at these straws and, in accord with Winge, very tentatively place 
the Necrolestidae in the Borhyaenoidea.^ As a marsupial, Necro- 
lestes adds one more example to the long series of convergences 
and parallelisms for which the extinct South American mam- 
malian fauna is justly famous: one of convergence toward an- 
other order (Insectivora, Chrysoehloridae), one of parallelism 
within the order (Notorijctes). 

AVith the removal of Necrolestes from the Insectivora, an order 
of mammals — one widespread elsewhere at an early date — has 
definitely been eliminated from the early Tertiary South Ameri- 
can scene. As I have briefly stated (1957, p. 57), it is becoming 
evident that the earliest South American mammalian fauna must 
have been an unbalanced one, of the sort encountered on islands. 
If, as has usually been supposed in recent years, the Americas 
were united by land in the late Cretaceous and early Paleocene 
it is very difficult to understand the absence from the southern 
continent of any traces of multituberculates, insectivores, taenio- 
donts, ereodonts, fissipeds and pantodonts, to name only the 
groups that are known, or may with confidence be inferred, to 

^I prefer Simpson's classilicatlon of the marsupials into six groups (1945) to 
tliose classilit-ations that unite Didelphiilae, Borhyaeniclae and Dasyuridae into 
one. Admittedly, these families are closer to each other morpholnKically than any 
one of thi'iu is to other marsupial .i:,'r(Hips, l)ut keepinjc them separate (or ar least 
keeping the Australian and South American groups separate) accords better with 
the zoogeosraphical <lata, which, when adequately understood, have an importance 
for mammalian taxonomy and phylogeny that can hardly be overestimated. 



12 BREVIORA No. 94 

have been in existence in the north during all or part of this 
time.^ That none of these would have made its way over a land 
connection had such then been in existence is improbable in the 
extreme. It appears to me much more likely that no such connec- 
tion existed. Projecting backward in time from the known early 
Tertiary mammalian faunas of South America, it is possible to 
visualize the relatively few major groups represented as having 
evolved from a small number of successful colonizations by 
ancestors that survived the hazards of overseas transportation 
from North America by natural rafts. A minimum of three such 
colonizations would have sufficed, but a more plausible total 
would be four, perhaps even more. I am in full agreement with 
Darlington (1957, p. 364) on this point. 

One successful landfall by a breeding pair or by a gravid 
female could have started the marsupial radiation, and another 
the edentate. The various ungulate groups might conceivably 
have resulted from a single colonization, but at least two seems 
more probable. From small, generalized condylarths could have 
evolved the litopterns, astrapotheres, xenungulates and pyro- 
theres. The notoungulates, by far the largest and most varied 
group of South American hoofed mammals, were, I suspect, the 
descendants of ancestors that were already members of this order 
by the time they made their landfall; they may have arrived 
earlier than the condylarths. The really extraordinary abund- 
ance and diversity of notoungulates, especially notable by con- 
trast with the other ungulate orders, would be at least partially 
explicable under this view. It is implied, of course, that the 
Notoungulata were of northern origin and that the known 
Holarctic forms — Palaeostylops of the late Paleocene of Mon- 
golia and Arctostylo'ps of the early Eocene of North America — 
were lingering survivors in that region. It is possible to think 
of these northern forms as immigrants from the south, but I am 
encouraged to doubt this by the structure of their teeth, which 
seems to me sufficiently^ different in detail from anything known 
among South American notoungulates as to indicate some inde- 

■^ Primates had almost surely come into existence by tlie early Paleocene. They 
make their first appearance in Kouth America in the ('ullHiehuapian. whicli sug- 
gests that they nuiy not have reached the continent until after the Mustersan, 
perhaps at abcuit the same time as the rodents. With a uroup so rare in the 
fossil record tliere is always a huT'e element of d(pul>t as to the reality of 
absence, and I therefore omit them from the list, which is quite Imposing enough 
without them. 



1958 AFFINITIES OF NECROLESTES 13 

pendent evolution. 

From the haphazard, unbalanced beginning thus visualized, 
the South American mammalian fauna in time achieved balance 
of a sort peculiarly its own. The early immigrants had by chance 
drifted to an area whose size, physical diversity and multitude 
of available niches provided the space essential for such a de- 
velopment. 

REFERENCES 

Ameghino, F. 

1891. Nuevos restos de mamiferos fosiles descubiertos por Carlos 
Ameghino en el Eoceno inferior de la Patagonia austral — 
especies nuevas, adiciones y eorreciones. Rev. Argent. Hist. Nat., 
1: 289-328. 

Darlington, P. J. 

1957. Zoogeography: the geographical distribution of animals. New 
York: John Wiley and Sons, i-xiii, 1-675. 

Gregory, W. K. 

1910. The orders of manmials. Bull. Amer. Mus. Nat. Hist., 27: 1-524. 

Leche, W. 

1907. Zur Entwicklungsgeschichte des Zahn-Systems der Saiigetiere, 
zugleich ein Beitrag zur Stanimesgeschiohte dieser Tiergruppe. 
Zweite Teil : Phylogenie. Zweites Heft : Die Familien der 
Centetidae, Solenodontidae und Chrysocliloridae. Zoologica, 20 
(49): 1-158. 

Patterson, B. 

1957. Mammalian phylogeny. In Premier symposium sur la speciticite 
parasitaire des parasites des vertebres. Neuchatel: Paul At- 
tinger, 15-48. 

Romer, a. S. 

1945. Vertebrate paleontology. Chicago : The University of Chicago 
Press, i-ix, 1-687. 

Saban, R. 

1954. Phylogenie des insectivores. Bull. Mus. Nat. d'Hist. Nat. Paris, 
(2) 26: 419-432. 



14 BREVIORA No. 94 

Scott, W. B. 

1905. Palaeontology. Part II. Insectivora. Repts. Princeton Univ. 
Expeds. Patagonia, 5: 365-383. 

Simpson, G. G. 

1945. The principles of classification and a classification of mammals. 
Bull. Amer. Mus. Nat. Hist., 85: i-xvi, 1-350. 

Weber, M. 

1928. Die Saiigeticre. Einfiihrung in die Anatomie und Systematik 
der recenten und fossilen Mammalia. Band II. Systematischer 
Teil. Jena: Gustav Fischer: i-xxiv, 1-898. 

WiNGE, H. 

1941. The interrelationships of the mammalian genera. Volume 1. 
Monotremata, Marsupialia, Insectivora, Chiroptera, Edentata. 
ILerbenhavn: C. A. Reitzel: i-xii, 1-412. 



BREVIORA 

Mnaseiainti of Comparative Zoology 



Cambridge, Mass. September 19, 1958 Number 95 



A NEW BOLIVIAN LAND SNAIL OF THE GENUS 

DRYMAEUS 

By Juan Jose Parodiz 

Acting Curator of Invertebrates, Carnegie Museum 
I'ittsliurgh, Pa. 

Drymaeus pereirai sp. nov. 

Description. Shell subcorneal, elongated. Whorls, seven, in- 
creasing gradually, the last being approximately % of the total 
length with its major diameter scarcely larger than one half of 
the same length. Suture ^vhite, marginated below and somewhat 
shouldered above, a character which is more evident at the base 
of the third post-embryonic whorl. Surface sulcate with very 
fine and irregular axial riblets, more distinct on the middle por- 
tion of the shell. The interspaces between these riblets are 
variable in width and there are 3, 2 or occasionally only 1 riblet 
per mm., with .some microscopical spiral liratioiis, more noticeable 
on the zone of the last whorl opposite the aperture. The riblets 
start on the suture, forming a very distinct marginated area. 
Color (in the holotype) white, with traces of brown bands broken 
into points and divided by three white zones. This spotted 
coloration is limited to the intercostal spaces and is not present 
on the high part of the riblets which remains white, giving a 
diffused silky aspect. Aperture oval, with rounded base, shiny 
white, enameled inside and somewhat iridescent; the superior 
part of the columellar lip a very pale pink ; peristome thin and 
not reflexed. The protocouch has the same pale pink-lilac shade 
of the peristome. 



BREVIORA 



No. 95 



Measurements in mm. : 

Shell 
Length Width 

Ilolotvpe 30 12.5 10.5 

Paratype 10 9 



Aperture 
Leno-th Width 



13.5 
12 



i.O 

6.5 



Last Whorl 
Length 

19.5 
16.5 




Drymaeus iwreirai, sp. nov. M.C.Z. no. 168843, El Carmen, Prov. Chi- 
quitos, Depto. Santa Cruz, Eepubliea de Bolivia. Hno. F. S. Pereira and 
Carl Gans, c-ollectors. (Holotype X "IV-i.) 

Locality. Estacion El Carmen (between Pahnito and Santa 
Ana) on the railroad Corumba-Santa Cruz, Prov. Chiquitos, 
Depto. Santa Cruz in the oriental plains of southeastern Bolivia. 
Holotype No. 168843 in Museum of Comparative Zoology and 
paratype in Carnegie Museum Aces. 16,073. Collectors : Hno. 
F. S. Vereira and Carl Gans, 1953. 

The paratype (which has the early whorls decollated) is evi- 
dently a younger specimen as shown not only by its smaller 
dimensions but by some traces of carination in the last whorl, and 



1958 DRYMAEUS PEREIRAI X. SP. 3 

fresher colors. The very pale spotted coloration in the paratype 
deserves an explanation, since it is very possible that other speci- 
mens with intermediate patterns will be found. The three 
brownish bands are more distinct, with the dividing white zones 
also very conspicuous ; the medium, broader band shows darker 
or clearer triangular areas ; the umbilical zone is completely 
white ; the lower spiral white zone continues into the aperture 
and the upper one, on the suture ; the dark bands are visible 
inside the aperture ; as in the paratype, the oblique axial riblets 
remain white. 

The marginated suture, the irregular axial-oblique riblets, and 
the very pale pink-lilac tint on the columellar lip, are the more 
important features of this new species. Two other species of 
Drymacus are known which have a roughened surface of pseudo- 
riblets: />. sulcosus (Pfeilt'er) from Mexico, which is a larger 
and stronger species with a thick and refiexed peristome, and 
some young specimens of the Central American D. jonasi (Pfeif- 
fer) — such as those figured by Pilsbry in 1899 (Manual of 
Conchology, (2) 13, p. 54, pi. 10, fig. 64) — which is different in 
many other characters such as the angulate l)ase of aperture and 
its broader shape. 

The region where this new species lias been found should have 
an extensive exploration from the malacological point of view. 
We know better a number of characteristic species from the 
western side of Santa Cruz, on the subandean zone and oriental 
Cordillera of Bolivia, as well as many others along the Paraguay 
River, from Corumba to the south. The locality El Carmen is in 
an intermediate region, more than 200 miles wide, between the 
Llanos of Izozo and the Paraguay River, being a northern ex- 
tension of the Boreal Chaco. 

The species is named for Hno. F. S. Pereira who obtained the 
specimens while collecting with Dr. Carl Gans along the zone 
Corumba-Santa Cruz in 1953. I wish to express my appreciation 
to Dr. J. Bequaert and Dr. W. J. Clench of the Museum of 
Comparative Zoology, for giving me the opportunity to report 
upon this new species. 



BREVIORA 

Miaseiiiimi of Comparsitive Zoology 



Cambridge, Mass. September 19, 1958 Number 9(i 

A NEW DICHOBUNID ARTIODACTYL FROM TPIE 

UINTA EOCENEi 

By C. Lewis Gazin 

Smithsonian Institution 

INTRODUCTION 

Recently, Mr. Bryan Patterson turned over to me for study a 
lower jaw belonging to an Antiacodon-\ike dichobunid from the 
Uinta upper Eocene. The specimen was discovered by him among 
some materials secured by the Museum of Comparative Zoology 
in an exchange with the Cleveland Museum of Natural History. 
The label accompanying the specimen reveals that the jaw was 
collected by Mr. D. C. Roberts near the entrance to Myton Pocket, 
a well-known collecting locality for upper Eocene mammals in 
the Uinta Basin of northeastern Utah. 

The specimen is particularly significant in showing that the 
Antiacodon line did not become extinct in Bridger middle Eocene 
time but survived at least until the Uinta C horizon of the upper 
Eocene. The form represented, while clearly related to Antia- 
codon, shows certain distinctive modifications rather beyond the 
limits that may be ascribed to that genus. Moreover, the ex- 
tended range in time of this line, together with the very distinc- 
tive and divergent characteristics brought to light (separating it 
from the better-known Homacodontinae), weakens my earlier 
objection (1952, p. 24) to applying separate subfamily recogni- 
tion to the antiacodonts. The name Antiacodontinae might well 
designate this phyletic sequence. 

The first antiacodont to be placed on record was the Bridger 
form, described by Cope (1872, p. 2) in his first " Palaeontological 

1 Published by permission of the Secretary of the Smithsonian Institution. 



2 BREVIORA No. 96 

Bulletin" as " Lophiotherium" pygmaeum. The same form was 
described scarcely a month later by Marsh (1872, p. 210) as 
Antiacodon venustus. Cope early recognized Marsh's genus but 
later felt that it was not sufficiently well characterized and in 
1875 (p. 256) proposed the name Sarcolemur for the species he 
had named Antiacodon fiircatiis. These I have shown (1955, 
p. 23) are synonymous, so that the genotypic Bridger species is 
Antiacodon pygmaeus. The first recognizable illustration of the 
lower jaw of Antiacodon (as Sarcolemur) was furnished b}' 
Osborn (1902) but, following Cope, he regarded it as a primate. 
Matthew in 1909 was evidently the first to realize that An- 
tiacodon was an artiodactyl, although earlier in the same year, 
while including "Sarcolemur" in the Artiodactyla, he believed 
Antiacodon to be a leptictid. Evidently the earliest form showing 
a relationship to the antiacodonts is the Wasatchian genus Hexa- 
codus (Gazin, 1952, p. 73), a form that might be regarded as an 
antiacodont, but that also includes within its range of character- 
istics certain indications of an ancestr}^ for Microsus, a typical 
homacodont. 

DICHOBUNIDAE Gill 
ANTIACODONTINAE, new subfamily 

AUXONTODON", new genus 
Type. Auxontodon pattersoni, new species. 

Generic characters. Small incisor-like canine. Much enlarged 
and probably caniniform Pi. Second to fourth premolars antero- 
posteriorly elongate. Marked diastema between Pi and P2. Little 
or no diastema between P2 and Ps. Cheek teeth resembling those 
of Antiacodon. Inferior margin of lower jaw strongly convex in 
anteroposterior profile. 

Comparison. The three lower incisors of Auxontodon, as indi- 
cated by the root portions, were small, about equal and decidedly 
procumbent. The root of the lower canine, which is also pro- 
cumbent and closely parallel to those of the incisors, is only 
slightly larger. The root of Pi, on the other hand, is much larger 

- From Greek, Auxontos, iucreasing : and odon, tooth ; in allusion to the in- 
creased size of the first lower premolar. 



1958 NEW MCHOBUNID ARTIODACTYL 3 

than that of the canine and appears to be more accurate, so that 
Avhile the root appears to extend well back in the jaw, the crown 
may have been more nearly erect. There is a marked diastema 
betw^een the alveolus for Pi and those for P2, and evidently only 
a very slight diastema separating P2 from P3. The diastema be- 
tween P2 and P3 in Antiacodon is variable and evidently some- 
what greater than in Auxontodon. In A.M.N.H. No. 12697 it is 
about 2.5 mm. whereas in U.S.N.M. No. 18000 it is 3.9 mm. 

P2 is not preserved in the Auxontodon specimen, but Pa is seen 
to be an elongate tooth with a distinctly high primary cusp or 
protoconid and a very well-defined metaconid almost posterior 
to it. There would appear to be no talonid on Ps but there is a 
low, distinct paraconid. P4 has a more distinctly developed 
talonid with a conspicuous entoconid, better developed and more 
widely spaced from the crest extending down the posterior slope 
of the protoconid than in Antiacodon. The metaconid is pro- 
minent as in Antiacodon but the anterior extremity of this tooth, 
though damaged, seems more forward-protruding than in the 
earlier genus. Mi shows a characteristic antiacodont form with 
a high paraconid and with the crista obliqua extending to the 
apex of the more worn metaconid. The hypoconulid is posterior 
to the crest between the hypoconid and entoconid, but appears 
less conical than in Antiacodon — more as a prominent upward 
arching of the posterior cingulum. 

The jaw of Auxontodon shows a decidedly convex lower mar- 
gin, viewed from the side. It is more nearly straight in Antiaco- 
don. A mental foramen of moderate size is noted beneath the 
posterior root of P3. It is in this position or slightly more forward 
in Antiacodon. However, in Auxontodon there is a larger fora- 
men below a point just posterior to the alveolus for the Pi, open- 
ing lateral to the root and extending backward parallel to it. 
In A.M.N.H. No. 12697 of Antiacodon pygniaeus the larger and 
more anterior mental foramen is just anterior to the posterior 
root of P2. 

Discussion. Perhaps the most significant feature observed in 
Auxontodon is the much-enlarged root of Pi. In homacodonts 
in general, although details in this respect are not known in all 
genera, there is no marked tendency toward differentiation of a 
stronglv caniniform tooth. Nevertheless, it was noted that in 



4 BREVIORA No. 96 

such more advanced liomacodonts as Mytononieryx and Pentace- 
mylus the root portion of Pi is distinctly larger than that of the 
canine. Moreover, it is more robust than in either of the roots 
for P2. However, in these genera the crown of Pi is premolar-like 
and forms with the incisors, canine and following premolars a 
nearly gradational series in size. In the more-selenodont Eocene 
artiodactyls, such as the agriochoerids, leptotragulids and hyper- 
tragulids. Pi early becomes caniniform. 

In none of the known material of Antiacodon pygmaeus is the 
anterior portion of the lower jaw sufficiently preserved to indi- 
cate with any certainty the extent to which Pi was developed. 
In A.M.N.H. No. 12697 of this species the jaw is preserved to 
about 1.7 mm. in advance of the roots for P2 but this portion 
of the lower jaw is relatively slender and there is no evidence 
of a root for Pi extending back in the jaw. Nevertheless, Antia- 
codon and Auxontodon show a rather strong tendency toward 
selenodonty and it is not surprising to find this line within the 
Dichobunidae showing tendencies paralleling the more highly 
selenodont families. 

AUXONTODOX PATTERSONI^, new species 
Figure la 

Type. Left ramus of mandible with Ps-Mi and part of M;:, 
M.C.Z. No. 9316. 

Horizon and locality. Uinta C, near entrance to Myton Pocket, 
about 7 miles east of Myton, Duchesne County, Utah. 

Specific characters. Size of lower jaw much larger than in 
Antiacodon pyg^naeus. Other characteristics of the species have 
not been distinguished from those cited for the genus. 

Discussion. The lower jaw, M.C.Z. No. 9316, is the only spec- 
imen known of Auxontodon pattersoni. Evidently this species 
was a very rare element of the fauna, as represented in the Uinta 
Basin, particularly considering the extensive collecting that has 
been carried on in the Uinta area and the relative abundance of 
artiodactyls encountered, both bunodont and selenodont. 

:i Named f(ir Mr. r.ryaii r;itlrrsi;ii. who first recoffiiiz(Ml the true relatioiishiiis of 
Auwontodon. 



1958 NEW DICHOBUNID ABTIODACTYL 5 

Measurements in millimeters of lower jaw and dentition in type 
specimen of Auxontodon pattersoni, M.C.Z. No. 9316 

Deptli of lower jaw at diastema between Pi and \'-i 8.0 

Depth of lower jaw beneath M^, lingually 12.1 

Ijengtli of cheek tooth series from anterior margin of 

alveolus for Pi to posterior margin of Ms 46.5 

Length of diastema between Pi and Pa 5.0* 

Length of c-heek tooth series from anterior margin of 

first alveolus of P2 to posterior margin of Ms 37.2 

Length of lower molar series, Mi-Ms, inclusive 19.2 
Pi, anteroposterior diameter of root portion at alveolus 4.0" 

Pi, transvers-e diameter of root ])orti()n at alv(»olus 2. .5 

Ps, length: greatest width 6.3^:3.0 

P4, length: greatest width 6.8'':3.6 

Ml, length: greatest width 6.1:4.4 

''Av>proxiniat'' 

References 

Cope, Edward D. 

1872. Description of some new Vertebrata from the Bridger group of 

the Eocene. Palaeont. Bull. No. 1, pp. 1-6 (also, Proc. Anier. 

Philos. Soc, vol. 12, pp. 460-465, 1873). 
1875. On fossil lemurs and dogs. Proc. Acad. Nat. Sei., Philadelphia, 

pp. 255-256. 

Gazin, C. Lewis 

1952. The lower Eocene Knight formation of western Wyoming and 

its mammalian faunas. Smithsonian Misc. Coll., vol. 117, No. 18, 

pp. 1-82, figs. 1-6, pis. 1-11. 
1955. A review of the upper Eocene Artiodactyla of North America. 

Smithsonian Misc. Coll., vol. 128, No. 8, pp. 1-98, pis. 1-18, 

charts 1-2. 

Marsh, Othniel C. 

1872. Preliminary description of new Tertiary mammals, Parts 1-lV. 
Amer. Journ. Sci. and Arts, vol. 4, pp. 122-128, 202-224. 

Matthew, William D. 

1909. The Carnivora and Insectivora of the Bridger Basin, middle 
Eocene. Mem. Amer. Mus. Nat. Hist., vol. 9, pt. 6, pp. 291-567, 
figs. 1-118, pis. 42-52. 

OsBORN, Henry F. 

1902. American Eocene primates, and the supposed family Mixodec- 
tidae. Bull. Amer. Mus. Nat. Hist., vol. 16, art. 17, pp. 169-214, 
figs. 1-40. 



BREVIORA 



No. 96 





Figure 1. (a) Auxontodon pattersoni, new genus and species, occlusal and 
lateral view of left ramus of mandible, M.C.Z. No. 9316, with (b) lateral 
view of left ramus of mandible of Antiacodon pygmaeus, U.S.N.M. No. 
18000. All natural size. Drawing prepared by Lawrence B. Isham. 



L-/ 



BREVIORA 



Musceiuim of Coeiparative Zoology 



Cambridge, Mass. October 27, 1958 Number 97 

FUSION OF CERVICAL VERTEBRAE IN THE 
ERETHIZONTIDAE AND DINOMYIDAE 

BY Clayton E. Ray 

Comparative postcraiiial osteology has been a curiously neg- 
lected source of evidence in the study of relationships among 
caviomorph and hystricomorph rodents though Landry, 1957, 
pp. 5-30, has made some use of postcranial osteology on the 
subordinal level. It seems useful, therefore, to call attention to a 
peculiarity observed in the cervical vertebrae of the living erethi- 
zontine (Fig. 1) and dinomyid (Fig. 2) caviomorphs. The second 
(axis) and third cervicals are firmly fused together in Erethizon, 
Coendou, Echinoprocta (the postcranial skeleton of Chaetomys is 
apparently unknown), and Dinomys. This condition was noted 
by Giebel (1877, p. 273) for Coendou {=Cercolahes) only and 
has apparently not been recorded in the literature since, having 
been overlooked, for example, by Scott (1905, p. 422) in con- 
trasting the skeletons of Sciamys and Erethizon, by Anthony 
(1926, pp. 124, 126) in his detailed comparison of the skeletons 
of Elastnodontomys, Erethizon, and Dinomys, and by Swena and 
Ashley in their recent (1956, pp. 8-10) osteology of Erethizon. 
The first four cervicals are missing from the type (in part) 
skeleton of Dinomys figured by Peters (1873, PI. 3, fig. 4). Lan- 
dry (1957, p. 46) and Fields (1957, pp. 337, 341, 384) have 
mentioned but not described the tendency toward fusion of cervi- 
cal vertebrae in Dinomys and in Olenopsis.'^ 

Cervical vertebrae of the following caviomorph and hystrico- 
morph genera have been available for examination or are ade- 

1 I'atterson (personal conamuiicatioii) (juestions Fields' (1957, p. o23) assiem- 
ment of his Colombian material to the genus Olenopsis. Thus, even though the 
animal describert is a ilinomyifl, it is not necessarily true that Olenopsis is a 
dinomyid. 



2 BRKVTOKA No. 97 

(juately described in the literature : Acanthion, Thecurus, 
Hysfri.r, Athcriirus, Triclnjs, Erethizon, Coendou, Echinoprocta, 
Kocardia (Scott, 1905, p. 465), Schistomys (Scott, 1905, p. 482), 
Cavia, Kcrodon, Galea, PrndoUchotis (Fields, 1957, p. 373), 
Dolchotu, Tlydrochoerus, T>in()my>i, " Ohnopsis'" (the material 
from the La Veiita of Colombia, not the Santa Vvwz of Pata- 
li'onia), I'Jlastnodonfomys, Amhlyrhiza, Cunicnlus, Dasyprocta, 
Myoprocta, Periniys (Scott, 1905, p. 437), Lagostomus, Lagidium, 
ChinchiUa, Ncoreomys (Scott, 1905, p. 395), Capromys, Geocap- 
vomys, Plagiodontia, Myoeasior, Sciamys (Scott, 1905, p. 422). 
Ctenomys, Ahrocoma, Isolohodon, ProecJwnys, Hoplomys, Iso- 
thrix, Echimys, Dactylomys, Thryonomys, and Bathyergus. 
Included in this list is at least one representative from every 
family of caviomorph and hystricomorph rodents (as given by 
Wood, 1955) excepting the Petromuridae. It is not likely that 
the condition of the cervicals in this monotypic x^.frican family 
would alter the general picture in any way. In none of the genera 
listed, save the three erethizontids and Dinomys, was cervical 
fusion noted. 

ERETHIZONTINAE 

The fusion in Erethizon is typical of the .Erethizontinae and 
will serve as the basis for description. Some two dozen individuals 
were examined. The neural spine of the third cervical is here 
relatively higher than in mammals generally and is enveloped on 
either side by the broad lateral lamellae of the neural spine of the 
axis. The two neural spines become more broadly fused to one 
another with increasing age (though in part distinguishable in 
all specimens seen) . In immature individuals the lateral lamellae 
of the axis are closely applied to the neural spine of the third 
cervical but are not fused to it, and in one individual (MCZ- 811 "i 
diverge posterodorsally to form a bifid process. The area of most 
profound fusion is between the postzygapophyses of the axis and 
the prezygapophyses of the third cervical, the trace of the articu- 
lar surface becoming obliterated with age. Although the centra 
become tightly fused, particularly near the ventral midline, their 

2 Throufrhout tliP discussion MCZ reCcis lo Muscuiii of < '(iiii|)iir;iti\ i' Zoolo^'y. 
AMXII to .\inoritaii .Museum of Natural History, CXIIM to Cliicaffo Xatural His 
lory Musouni, ami I'CMl* to Univorsity of California Museuni of I'alcoutoloKy. 



190(S FUSION Ol' ('K1J\I(AI>S IN ( AViOAiOKl'l IS 3 

union is in part (liscciniblc in all individuals. The pleurapophyses 
of the axis approach those of the third eervical quite closely, but 
attain contact in only one individual (MCZ 81!)), in which the 
distal extremity of the rifjfht plenrapophysis (only) of the axis 
touches and is narrowly fused to tiie dorsolateral surface of the 
correspond in<i- pleurapoi)hysis of the third cervical. In none of 
the specimens examined was there any sugjyestion of fusion be- 
tween the third and fourth cervicals. 

DINOMYIDAE 

Dinomys is represented by two subadult skeletons '^ (CNHM 
66891 and 57186), by a young adult skeleton (CNHM 69593), 
and by an adult skeleton in part pathological (AMNII 70354). 
The nature of the fusion between the axis and third cervical is 
very similar to that in the Erethizontinae. Already in the im- 
mature individuals the neural spines are closely appressed and 
largely fused, whereas the mutual zygapophyseal articulations 
are completely and indistinguishably fused. As in some individ- 
uals of Erethizon, the lateral lamellae of the axial neural spine 
in all four specimens of Dinomys diverge posterodorsally to form 
a bifid process. These lamellae are broadly separated in CNHM 
69593, and the intervening space is occupied by the neural spine 
of cervical three and by a connecting network of cancellous bone. 
In CNHM 57186 (Fig. 2A) and in AMNII 70354 a slit-like 
foramen situated on either side just medial to the zygapophyses 
passes into the neural canal between the neural arches of the axis 
and third cervical. This foramen may be interpreted as the 
vestige (preserved here for the passage of nerves and blood ves- 
sels) of the open space normally present between the neural 
arches of successive vertebrae. It is not present in CNIIM 66891, 
and is but inconsistently and weakly developed in the erethizon- 
tids examined. There is on the articular facet of each post- 
zygapophysis in CNHM 69593 a small foramen with a well-defined 
channel extending from it to the posterior (right) and postero- 

3 Not adults as stated by Fields (19.57, p. 320), but suluulults judging from the 
slight wear undergone by P"r. the open basisphenoid-basioceipital sutures, and 
the open epiphyses throughout the skeleton. The young adult is characterized 
by P -^ in full use, closed basisphenoid-basioceipital suture, oiion vertebral epi- 
physes, and tight but discernible limb epiph.vses. 



4 BREVIORA No. 97 

medial (left) edge of the facet. I suspect that this foramen is 
homologous to the one discussed above, and that the functionally 
single postzygapophysis is in fact compounded from a postero- 
lateral contribution from the axial postzygapophysis and an 
anteromedial contribution from the third cervical postzyga- 
pophysis. This suspicion is reinforced by the presence of comple- 
mentary subdivisions in the prezygapophysis of cervical four 
(particularly the left), by the enveloping of the third cervical by 
the axis to a greater extent than in any other specimen of Dino- 
mys, by the juxtaposition of the supposed axial postzygapophyses 
and the lamellae of the axial neural arch, and by the complete 
absence of the line of fusion as seen in lateral view in the other 
Dinomys specimens examined (Fig. 2A). Although the vertebral 
epiphyses (and second intercentrum at the base of the odontoid 
process) remain open throughout the column in the subadult 
and young adult specimens, the epiphyses between axis and third 
cervical are fused together (Fig. 2B). As in the erethizontines 
the pleurapophyses on either side approach closely but do not 
touch except in the youngest individual (CNHM 66891) in 
which the distal extremity of the left axial pleurapophysis is 
fused narrowly to the dorsal surface of the pleurapophysis of 
the third cervical. The pleurapophyses of the axis in CNHM 
69593 are peculiarly developed into broad lamellae which extend 
posterodorsally lateral to the prezygapophyses of cervical four. 
Neither in the two subadult individuals nor in the young adult 
are cervicals three and four fused to one another. However, in 
the older of the two subadults (CNHM 57186) and in the young 
adult (CNHM 69593), roughened areas are present immediately 
anterior to the postzygapophyses of cervical three, posterior to 
the prezygapophyses of cervical four, and in the area of contact 
between the neural spines, suggesting the possibility of fusion at 
these points, with increased age. The contact surface between 
centra three and four in CNHM 69593 is not at all planar but 
curves anteriorly at both dorsolateral corners. Clearly verj- little 
movement could have occurred between cervicals three and four 
in this individual. AMNH 70354 presents a special problem owing 
to its pathological nature and to the lack of a series of normal 
adult skeletons from which to determine the average condition. 
Here the centra of cervicals two through seven are fused as are 



1958 FUSION OF CERVICALS IN CAVIOMORPHS 5 

the neural spines and zygapophj^ses of cervicals two through five. 
The fusion of the centra between cervicals five and six and 
betw^een six and seven is effected by gross ventral exostoses. 
Otherwise cervicals five through seven are independent of one 
another just as they are in the three younger individuals at hand 
and in the subadult figured by Peters (1873, PI. 3, fig. 4), all of 
which strongly suggests that these vertebrae are not ordinarily 
fused in Dinomys. The fusion of centra two through five may 
be enhanced by lesser exostoses. All post-atlantal cervicals are 
more or less marked by the rugose, spongy, asymmetrical growth 
indicative of pathology, as are many of the other vertebrae 
(notably the posterior thoracic and anterior lumbar). However, 
the exostoses are not so pronounced as to definitely indicate that 
cervicals four and five are here abnormally included in the fused 
series. That cervical five was not fused to cervical four in the 
type skeleton is clearly shown by the accidental loss of the fourth 
and the preservation of the fifth (Peters, 1873, p. 231). On the 
basis of the condition in the subadult and young adult animals 
(and its analogy to the erethizontines) and the pathology of 
AMNH 70354, it seems reasonable to conclude tentatively that 
only cervicals two and three are normally fused in Dinomys, 
and that cervical four may be added in older animals. The final 
solution awaits more specimens. 

Thus Dinomys and the erethizontines are very similar in the 
fusion of cervical vertebrae, wath the reservation that fusion 
seems to be more variable in nature and extent in Dinomys than 
in the erethizontines (judging from the considerable diversity 
observed among only four individuals of Di^iomys). Certain dif- 
ferences may be noted in the morphology of the vertebrae in the 
two groups. The anterior articular facets of the axis in erethizon- 
tines are entirely convex and situated on low pedestals serving 
to separate them from the peglike odontoid process, whereas in 
Dinomys the facets are concave in frontal section in such a way 
that they form a curved surface continuous medially with the 
bluntly conical odontoid process (cf. Figs. IB, 2B). In the 
erethizontines the centra of cervicals two and three are relatively 
deep dorsoventrally, Avith a prominent anteroposterior mid-ven- 
tral ridge (especially in Ereihizon and Coendou) ; in Dinomys, 
relatively shallow dorsoventrallv with a flattened venter. In 



6 BRF.VIOIfA No. 97 

Erefhizon the articular facets of tlie postzygapophyses of cervical 
three are situated relatively higli on the neural arch, are about 
cjually latei-ally aud ventrally directed, planar, and somewhat 
isoJatoil from the body of the neural arch ; in Dinoiuys the articu- 
lar facets are relatively hj-w, dominantly ventrally directed, 
concave in transverse section, and an integral ]iart of the neural 
arch (cf. Figs. IC, 2C). 

Of extinct erethizontids and dinomyids onlj' the dinomyid from 
the La Venta Miocene of Colombia, described by Fields (1957, 
p. 323, et sqq.) under the name (Jlcnopsis { = Drytonwm}fs) 
aequaforialis (Anthony) 1922, includes cervical vertebrae. The 
specimens preserved are both articulated series of sub.adult ver- 
tebrae, the one set (UCMP 39969) including a fragment of the 
li^it postzygapophysis of the axis and cervical three through 
thoracic one, all perfectly preserved excepting the distal ex- 
tremities of pleurapophyses ; the other set (UCMP 11636) 
including most of the neural arch and spine of the axis, complete 
(excepting pleurapophyses) cervicals three, four, and five, and 
fragmentary cei'vicals six and seven lacking portions of the 
neural arches. I am very sorry to be unable to agree with Fields' 
characterization (1957, p. 337) of the cervical vertebrae as 
"showing tendency toward fusion of anterior part of series, as 
in Dinomys." 1 can discover no certain evidence for fusion 
between any of the vertebrae preserved. The prezygapophyses 
and anterior end of the centrum of cervical three in UCMP 
39969 bear well-preserved, smooth articular facets as in normal 
diarthroses. The fragment of the left postzyga])ophysis of the 
axis adhering to tlie third cervical appears to be merely ce- 
mented by matrix as are the articulations in all cases. The 
anterior articular facet of cervical three is exposed and smooth 
also in UCMP 41636. The neural arch of the axis in this speci- 
men is slightly skcAved upon that of cervical three in a clockwise 
direction as viewed from above with the result that the two 
vertebrae are not perfectly articulated but are separated by a 
thin layer oi' cementing matrix. The two neural spines are 
nowhere in contact, that of cervical three being (juite low as in 
the succeeding cervicals and imlike the tall spine of the third 
cervical in Dinoniijs. That the absence of fusion in the fossils 
is not due mei-ely to immaturity is shown l)y the youngest speci- 



1958 FUSION OF CERVICAIiS IN CAVIOMORPHS 7 

men of Dinomys ((^NIIM 66891) the dentition of whicli is com- 
parable ill stage of wear to that of the dentigerous elements 
associated with eaeh of the fossils (Fields, 1957, Fig's. 15, 16 A, 
16B). The possibility remains that fusion could have occurred 
later in the ontogeny of the extinct form, but even granting this 
possibility the tendency could have been by no means as strong 
as in Dinomys. 

TAXONOMIC DISCUSSION 

Evidence from myology (Parsons, 1894, pp. 295-296; Wood 
and White, 1950, p. 592), external characteristics (Pocock, 1922, 
p. 422), parasitology (Vanzolini and Guimaraes, 1955. p. 43), 
dentition (Wood, 1950, p. 95), serology (Moody and Doniger, 
1956, p. 54), and paleontology and paleogeography (AVood, 1955. 
p. 180) has clearly indicated that erethizontids and hystricids 
show no particular resemblances to one another beyond the 
possession of quills,"* structures shared also with hedgehogs, 
echidnas, and some tenrecs as well as with other rodents. This 
distinctness of New and Old World porcupines was recognized by 
some workers many years ago (e.g. Tullberg, 1899, p. 108), but 
only recently has the opinion been expressed that erethizontids 
are isolated from other caviomorphs. Simpson (1945, p. 94) 
followed by Wood (1955, p. 182) has accorded superfamilial rank 
to the New World porcupines, whereas Moody and Doniger 
(1956, p. 53) have asked, "Should the erethizontids be accorded 
their own suborder ? " In support of this suggested isolation are 
the findings that Eretliizon is serologically as distinct from Cavia 
and Dasyprocta as from Hysfrix (Moody and Doniger, 1956, p. 
52), that erethizoiitid Mallophaga fall into one isolated grou]") 
and those of other caviomorphs into another (Vanzolini and 
(Juimaraes, 1955, pp. 30, 31), and that "Deseadan caviomorphs 
are all very closely related, except for the erethizontids, which 
were already distinct" (Wood, 1955, p. 182). It should be 

* In piiiiit of fiict. the quills are structurally rather dissinillar in the two groups. 
For exainijle, the distal portion of the quill in all three cenera of Erethizontinae 
is furnished with a coverinK of imhricnte, proximallv directed harhs (desrrihed 
and illustrated for r.r€thi::on by Loweg. lilOO. p. 85.-?, PI. 28. fig. f> and by Shadle 
and Po-Chedley, l!t4'.l. p. 17."., fig. 1). whereas I am able tn find such barbs in 
none of the five genera of Ilystricidae (Loweg, 1900, p. 85.3 notes their absence in 
Hystrix crintdtn and none are described liy Lochte. 19.t7 in his detailed study of 
the quills in Hifi^trir hirKiitiroxtris) . 



8 BREVIORA No. 97 

pointed out that the serological, parasitological, and (Deseadan) 
paleontological information is unknown for dinomyids, which 
therefore cannot on these grounds be summarily lumped with 
"other caviomorphs" as opposed to erethizontids. Aside from 
the similarity in cervical fusion demonstrated here, Pocock 
(1926, p. 228) has noted a "tolerably close resemblance" be- 
tween the feet of Dinomys and of Erethizon and Coendou, and 
Fields (1957, p. 348) has pointed out the similarity of the audi- 
tory ossicles in Erethizon, Dinomys, and Olenopsis acquatorialis. 
Attractive as this evidence is, it furnishes an inadequate basis for 
asserting any special relationship between erethizontids and 
dinomyids. Indeed contrary evidence from dentition and cranial 
osteology seems at present to be more compelling. In this con- 
nection the relationship between the axis and third cervical in 
Chaetomys and in the Deseadan erethizontids will, when known, 
be of great interest in helping to determine, respectively, the 
affinities of Chaetomys, and the antiquity of fusion in the erethi- 
zontids. If fusion had already taken place in the Deseadan 
(Oligocene) erethizontids then it might reasonably be concluded 
that the fusion in dinomyids is independent since it apparently 
had not occurred by late Miocene time. Furthermore, one would 
expect fusion to occur in Chaetomys if it is truly an erethizontid."'' 
In conclusion, present evidence indicates that the constant and 
exclusive fusion of cervical vertebrae two and three, together 
with the morphological features described above, furnishes a 
valid taxonomic character for the living Erethizontinae. Ap- 
praisal of the possible broader value of the character awaits 
evidence from Chaetomys, normal adult Dinomys, and fossil 
erethizontids (and additional fossil dinomyids). 

5 A cautionary, if confusing, example is provided by tlie Pedetidae, in whicli 
cervical fusion may prove to be irrej^ularly distributed. Hatt (1932, pp. 640, 715) 
found cervicals two and tliree to be fused in Pedetes surdastcr but not in P. cajir 
(as Hatt was well aware, this character should be checked on more specimens 



1958 FUSION OF CERVICALS IN CAVIOMORPHS d 

ACKNOWLEDGMENTS 

1 wish to thank the following people for assistance during the 
preparation of tliis paper : Professor Bryan Patterson for read- 
ing an early stage of the manuscript and for examining certain 
caviomorphs and hystricomorphs in the U.S. National Museum, 
Dr. Ernest E. Williams for reading the manuscript and for 
encouraging the undertaking of the study, Professor Albert E. 
Wood for information on Platypittamys, Dr. D. Dwight Davis 
for observations on Ahrocoma in the Chicago Natural History 
Museum and for the loan of Dinomys skeletons, Dr. Donald E. 
Savage for the loan of the cervical vertebrae of Olenopsis aequa- 
torialis in the University of California Museum of Paleontology, 
Drs. Stuart 0. Landry, Jr., and Robert W. Fields for useful 
information via correspondence, and Miss Barbara Lawrence and 
Dr. Charles P. Lyman for free access to the collections in their 
care at the Museum of Comparative Zoology. The illustrations 
were made by Mrs. Patricia Chaudhuri. The work was carried 
out while on the tenure of a National Science Foundation pre- 
doctoral fellowship. 

LITERATURE CITED 

Anthony, H. E. 

192(i. Mammals of Porto Rico, living and extinct — Rodentia and Eden- 
tata. New York Acad. Sci., Scientific Survey of Porto Rico and 
the Virgin Islands, vol. 9, pt. 2, pp. 97-241. 

Fields, R. W. 

1957. Hystricomorph rodents from the late Miocene of Colombia, South 
America. California Univ. Geol. Sci. Pub., vol. 32, no. 5, pp. 
273-404. 

GlEBEL, C. G. 

1877. 771 H. G. Bronn's Klassen und Ordnungen des Thier-reichs, vol. 
6, sect. .T, nos. 15, 16, pp. 257-304. 

Hatt, R. T. 

1932. The vertebral columns of ricochetal rodents. Bull. Am. Mus. Nat. 
Hist., vol. 63, art. 6, pp. 599-738. 

Landry, S. O., Jr. 

1957. The interrelationships of the New and Old World hystricomorph 
rodents. California Univ. Zool. Pub., vol. 56, no. 1, pp. 1-118. 



10 BREVIORA No. 97 

LOCHTE, T. 

1957. Die Haare, Borsten und Stacheln des Stachelschweines (Hystri.r 
Jiirsutirostris) . Der Zoologisehe Garten, Leipzig, vol. 23, no. 
1/3, pp. 145-162. 

LOWEG, T. 

11)00. Studien iiber das Integument des Erethizon dorsatus {Erethizon 
dorsatum Cuvier). Jenaische Zeitschrift, vol. 34, no. 4, pp. 833- 
866 (n.s., vol. 27). 

Macinnes, D. G. 

1957. A new Miocene rodent from East Africa. Brit. Mus. (Nat. 
Hist.). Fossil Mammals of Africa, no. 12, pp. 1-35. 

Moody, P. A. and D. E. Doniger 

1956. Serological light on porcupine relationships. Evolution, vol. 10, 
no. 1, pp. 47-55. 

Parsons, F. G. 

1894. On the myology of the sciuromorphine and hystricomorphinc 
rodents. Proc. Zool. Soc. London, pp. 251-296. 

Peters, W. C. H. 

1873. Ueber Dinomys, eine merkwiirdige neue Gattung von Nagethieren 
aus Peru. Festschrift zur feier des Hundert-Jiihrigen Bestehens 
der Gesellschaft Naturforschender Freunde zu Berlin, pp. 227- 
234. 

POCOCK, R. 1. 

1922. On the external characters of some hystricomorph rodents. 

Proc. Zool. Soe. London, 1922, pp. 365-427. 
1926. The external characters of a young female Dinomys hranickii 

exhibited in the Society's gardens. Proc. Zool. Soc. London, 

1926, pp. 221-230. 

Scott, W. B. 

1905. Mammalia of the Santa Cruz beds, III. Glires. Rept. Princeton 
Univ. Exped. Patagonia, 1896-1899, vol. 5, pp. 384-489. 

Shadle, A. R. and D. Po-Chedley 

1949. Rate of penetration of a porcupine spine. Jour. Mammalogy, 
vol. 30, no. 2, pp. 172-173. 

Simpson, G. G. 

1945. The principles of classification and a classification of mniumnls. 
Bull. Am. Mus. Nat. Hist., vol. S5, pp. 1-349. 



1958 FUSION OF CERVICALS IN CAVIOMORPHS 11 

Stromer, E. 

1926. Restf Land- und Siisswasser-bewohnendcr Wirbelticre aus den 
Diamantenfeldern Deutsch-Siidwestafrikas. Chap. XXI, pp. 107- 
153 in Kaiser, E., 1926, Die Diamantenwiiste Siidwestafrikas, 
II: Berlin, 535 pp. 

SwENA, B. and L. M. Ashley 

1956. Osteology of the common porcupine. Dept. Biol. Sci. and Biol. 
Station, Walla Walla College, Washington, no. 18, pp. 1-26. 

TULLBERG, T. 

1899. IJber das System der Nagethiere, eine phylogenetische Studie. 
Nova Acta Reg. Soc. Sci. Upsaliensis, vol. 18, sect. 2, no. 1, 
pp. 1-514. 

Vanzolini, p. E. and L. R. Guimaraes 

19.')5. Lice and the history of South American land mammals. Rev. 
Brasileira de Entomologia, vol. 3, p]). 13-46. 

Wood, A. E. 

1950. Porcupines, paleogeography and parallelism. Evolution, vol. 4, 

no. 1, pp. 87-98. 
lit.!;"). A revised classification of the rodents. Jour. Mammalogy, vol. 

36, no. 2, pp. 165-187. 

Wood, A. E. and R. R. White, III 

1950. The myology of the chinchilla. Jour. Morphology, vol. SO, no. 3, 
pp. 547-598. 




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BREVIORA 

Museiiinn of Comparative Zoology 

Cambridge, Mass. December 12, 1958 Number 98 

TWO NEW SPECIES OF BATHYLAGUS FROM THE 

WESTERN NORTH ATLANTIC WITH NOTES 

ON OTHER SPECIES 

By Daniel M. Cohen 

Ichthyological Laboratory, U.S. Fish and Wildlife Service 
U.S. National Museum, Washington, D.C. 

While reviewing the Atlantic bathylagids for the forthcoming 
volume of "Fishes of the Western North Atlantic," two appar- 
ently unnamed species were encountered and are herein described 
as new. In addition, notes on several related bathylagids are 
presented. 

Bathylagus compsus, sp. nov. 

Figure 1 
Bathylagus glacialis Beebe {-non Re^an), 1933, j). 114. 

Holotype. U.S. National Museum 171755, 44.4 mm. in stand- 
ard length; collected by Richard H. Backus on board the "Blue 
Dolphin," haul RHB 457, July 17-18, 1953, Isaacs-Kidd mid- 
water trawl. Trawl down at 2255 hours, up at 0210 hours. Depth 
69.5 m. Locality, within a circle 12 miles in diameter with the 
center at 39° 45' N., 71° 08' W. Paratype: USNM 171754 (1), 
38 mm. in standard length ; data as for the holotype. 

Diagnosis. A Bathylagus with an extensive gill opening, reach- 
ing almost halfway up the side of the body; an anal fin base 
longer than the length of the caudal peduncle ; and dark pigment 
present on the margins of the scale pockets. 

Counts and measurements. Measurements given as per cent of 
standard length, holotype first, followed by the paratype in 
parenthesis. Dorsal 11 (10), anal 19 (20), pectoral 9 (9), ven- 
tral 9 (10), branchiostegals 2 ( — ) ; preanal 75.9 (76.8), preven- 



BREV'IORA 



No. 98 



tral 52.5 (52.1), predorsal 47.3 (50.0), prepectoral 27.0 (— ). 
head length 25.0 (26.3), snout 5.6 (5.3), eye 10.0 (9.2), greatest 
body depth 16.9 (15.3), least depth of caudal peduncle 6.3 (6.6), 
dorsal fin base 8.8 (6.8). anal fin base 16.9 (15.0), interorbital 
9.0(— ). 

Description. A graceful, elongate fish with the greatest depth 
a short distance behind the head. The dorsal profile of the head 
descends from the nape to the interorbital in almost a straight 
line. It curves downward near the anterior edge of the orbit and 




Figure 1. USNM 171755. Holotype of Baihylagus compsus. 



descends to a point above the nares where another inflection 
occurs causing the profile to drop more steeply to the upper lip. 
The dorsal rim of the orbit does not enter the dorsal profile in 
the larger specimen but does in the smaller. The ventral profile 
of the head rises more gently than the dorsal profile descends. 
The interorbital area is broadly concave between the supraorbital 
canals. The maxillary extends back to a vertical from a point 
slightly in front of the anterior edge of the orbit. Minute papillae 
are present on the ventral margins of the maxillary and premax- 
illary. The jaws are rounded and subequal. 

The eyes are directed laterally and lack adipose eyelids. A 
crescent of white tissue is present over the iris of the posterior 
half of the eye. 

The height of the gill opening is slightly less than half the 
greatest height of the body and extends beyond the dorsal edge 
of the pectoral fin base by a distance equal to two and one-half 
times the length of the pectoral fin base. In his description of 
post-larvae of this species, Beebe (1933, p. 119) stated ''gill 
openings characteristically small"; however, he gave no quanti- 
tative data, nor did he discuss this character in larger specimens. 



1958 NEW SPECIES OF BATHYLAGUS 3 

The pectoral fin inserts a short distance behind the head on 
the ventrolateral contours of the body. The dorsal fin orip:inates 
close to the midpoint of tlie body and has its base raised above the 
body. The ventral fins insert on the ventral surface of the body 
under the posterior part of the dorsal fin base. The anal fin base 
is set off from the body and is longer than the dorsal fin base and 
longer than the length of the caudal peduncle. The adipose fin 
is placed over the posterior end of the anal fin base. The fins are 
all broken off short. 

The pigmentation of specimens in alcohol is striking and 
serves as an important diagnostic character, at least in adolescent 
specimens. The ground color is a light yellow-brown. The muzzle, 
the opercular apparatus and a ring around the orbit are dark 
brown. The opercle is overlaid with an iridescent sheen and is 
probably silvery in life. The iris is iridescent gun-metal blue 
and may also be silvery in life. The dark lining of the peritoneal 
cavity shows through the body wall and outlines the body cavity. 
A single row of large, dark chromatophores runs the entire length 
of the body slightly below the lateral line. A second and less 
distinct line parallels the first line a short distance above the lat- 
eral line. The right side of the paratype shows a striking resem- 
blance to Beebe's (1933, p. 118) figure of an adolescent specimen. 
The left side of the paratype and both sides of the holotype vary 
in that the anterior portion of the upper line is lost in a scatter- 
ing of smaller, lighter chromatophores. The venter is lightly pep- 
pered with small, dark chromatophores which are more densely 
distributed on the dorsum. Brown pigment cells are also present 
on the dorsal surface of the head behind and before the interor- 
bital area and on the cheeks. The caudal, dorsal and adipose fins 
have scattered brown chromatophores, while the anal, ventral and 
pectoral fins are immaculate or bear only a very few pigment cells. 
In both specimens the posterior half of the body has the ragged 
remnants of scale pockets colored by large, dark chromatophores 
(not shown in Fig. 1). 

Relationships. Bathylagus compsus appears most closely re- 
lated to B. nigrigenys and B. longirostris, the only other known 
species of Bathylagus with extensive gill openings and elongate 
anal fin bases. The types of B. compsus differ from specimens of 
B. nigrigenys of comparable size in having more anal rays, 19 



4 BREVIORA No. 98 

to 20 in conipsus, 18 to 17 in nigrigenys; a shorter preventral 
distance, 52.1 to 52.5 in conipsus, 56.7 to 61.4 in nigrigenys; in 
having- a less deep body, 15.3 to 16.9 in compsiis, 18.9 to 22.6 in 
nigrigenys. In addition, adolescent nigrigenys apparently lack 
the striking- pigment pattern found in compsus. Other less 
trenchant differences are also present. 

I have examined neither adolescent specimens of B. longiros- 
tris nor adult specimens of B. cumpsus and so hesitate to com- 
pare proportional measurements ; however, B. Inngirostris differs 
markedly from conipsus in its complete lack of pigment on the 
scale pockets. 

Discussion. In his studies on Bermuda Bathylagus, Beebe 
(1933) identified one of the forms upon which he reported as 
Bathylagus glacialis Regan, a species originally described from 
the Antarctic. More recent studies by Norman (1937) and 
Cohen (MS.) strongly suggest that B. glacialis Regan is a syno- 
nym of B. antarcticus. I have examined a series of B. anfarcticus 
from southern waters, and I find they differ from Beebe 's de- 
scriptions of his Bermuda material. Unfortunately, I have not 
been able to obtain Bermuda specimens for direct comparison ; 
however, the two specimens at hand differ so markedly from B. 
antarcticus (which is probably restricted to far southern seas) 
and agree so well with Beebe 's description that I feel no qualms 
at referring Beebe 's glacialis to Bathylagus compsus. 

Beebe (1933) also presented an osteological study of this 
species, and there seem to be a number of discrepancies between 
his findings and those of Chapman (19-43), who presented an 
osteological study of Bathylagus pacificus. The differences ap- 
pear so marked that they could be used to separate genera or 
even higher taxa. I was fortunate in having at my disposal the 
identical cleared and stained specimens upon which Beebe based 
his descriptions (Stanford University 45302, from Bermuda) 
and I was able to note several points which require comment. 
These are briefly noted since they are of some importance in the 
classification of the bathylagids. 1. The frontals : Beebe states 
that the frontals are fused and he also figures them as such. I 
find that the frontals are not fused but instead are slightly sep- 
arated in the midline. The underlying cartilage has taken up 
some of the stain and the edges of the bone are difficult to see 



1958 NEW SPECIES OF BATHYLAGUS 5 

unless they are gently lifted. 2. The sphenotie : The bone which 
Beebe labels the pterotic in his dorsal view of the skull (Fig. 38) 
is actually the splienotic and is so labeled in the lateral view 
(Fig. 3G). ■I. Supraniaxillary : Beebe shows supraniaxillaries 
in Figure 36, although he does not mention them in the text. I 
find no supraniaxillaries, but instead a more heavily stained 
ridge which may have been mistaken for a joint. 4. Branchio- 
stegals : Although he shows two branehiostegals in all of his 
figures and I find two branehiostegals in his specimen, Beebe 
gives three or four branehiostegals as a generic character. I 
know of no species of recent Bathylagus with other than two 
branchiostegal rays. Hubbs (1919) notes a single branchiostegal 
ray in Bathylagus pacijicus; however, I have found two in this 
species. Chapman (1943) also reports two. 

With regard to general shape and placement of bones, I find 
Bathylagus compsus in good agreement with Chapman's account 
of B. pacificus. Since Beebe did not dissect his specimen, his 
account of the deeper bones of the skull is necessarily sketchy. 

The name compsus is from the Greek kompsos, elegant or 
pretty, and refers to the general appearance of this attractive 
sjjecies. 

Bathylagus geeyae, sp. nov. 

Figure 2 

Holotype. Chicago Natural History Museum 49730, 71 mm. 
in standard length; collected by the "Caryn," haul 38, August 
(5, 1948, 35 foot otter trawl. Trawd in at 2245 hours, out at 0200 
hours. Depth of haul 500-550 m., 1500 m. wire out, bottom depth 
2000 to 2560 m. Locality, Bermuda, 32° 13.5' N., 64° 32.5' W. 

Paratypc. Stanford University 50974 (1), 35.5 mm. in stand- 
ard length, collected by the Bermuda Oceanographic Expedi- 
tions of the New York Zoological Society, 1929-1930, net 35, 
April 24, 1929. Net down at 9 :54 A.M., duration of haul 3 hours, 
36 minutes. Depth of haul 1646 m. to surface. Locality, Ber- 
muda, an 8 mile circle with its center at 32° 12' N., 64° 36' AV. 

Diagnosis. A Bathylagus with an extensive gill opening, reach- 
ing more than halfway up the side of the body ; an anal fin base 
equal to or shorter in length than the length of the caudal 
peduncle ; and dark pigment present on the margins of the scale 
pockets. 



6 BREVIORA No. 98 

Counts and Measurements. Measurements given as per cent 
of standard length, holotype first, followed by the paratype in 
parenthesis. Dorsal 11 (13), Anal 13 (13), pectoral 12 (13), 
ventral 10 (11), branchiostegals 2 ( — ), gill rakers on lower arm 
of first arch in holotype 16, preanal 81.0 (79.1), preventral 58.4 
(61.1), predorsal 48.2 (50.7), prepectoral 28.9 (32.7), head 
length 25.3 (29.6), snout 5.9 (7.0), eye 9.4 (see description for 
comments on eye of paratype), greatest body depth 13.4 (16.9), 
least depth of caudal peduncle 8.4 (8.4), dorsal fin base 12.0 
(11.8), anal fin base 9.7 (9.9), lateral line scales probably be- 
tween 45 and 50 on both specimens. 

Description. An elongate and much laterally compressed fish 
with the greatest depth behind the head, tapering very little to 
the caudal peduncle. The dorsal profile of the head slopes gently 
downward in almost a straight line from a point behind the eye 
to a point near the nostrils, from whence it descends more 
steeply. The dorsal rim of the orbit projects into the dorsal pro- 
file of the head. The interorbital area between the supraorbital 
canals is broadly rounded. A well developed pineal apparatus 









Figure 2. CNHM 49730. Holotype of Bathylagus greyac (this figure is 
a reconstruction of the somewhat distorted holotype and should not be used 
for measurements). Lower figure is of the opercle of Bathylagus ochoien.'^i.^. 

is present and is visible through a clear window in the deeper 
pigmentation of the braincase. A broad ring of dense pigment 
about the window gives the appearance of an iris. The posterior 
edge of the maxillary does not reach a vertical from the anterior 
edge of the orbit. Miimte papillae are present on the epidermal 
covering of the maxillary, premaxillary and dentary. The jaws 
are rounded and equal. 



1958 NEW SPECIES OF BATHYLAGUS 7 

The mouth of the holotype is damaged, making it difficult to 
discern any details of dentition; however, the single row of 
upper teeth are conical and apparently restricted to the head of 
the vomer. The compressed dentary teeth are smaller than the 
vomerine teeth with the exception of a pair of small, spike-like 
teeth which project at the symphysis of the dentaries. 

The eyes are directed laterally and lack adipose eyelids. A 
crescent of white material is present over portions of the iris. 
The left eye of the paratype presents certain peculiarities. It 
has a horizontal diameter of 3.5 mm. compared to 4.2 mm. for 
the right eye. In addition, the left eye has two small lenses 
visible through the pupil. 

The height of the gill opening is more than half the greatest 
height of the body and extends beyond the dorsal edge of the 
pectoral fin base by a distance equal to about two and one-half 
times the length of the pectoral fin base in the holotype and about 
one and one-half times in the paratype. 

The pectoral fin inserts a short distance behind the head on 
the ventro-lateral contours of the body. The dorsal fin originates 
close to the mid-point of the body. Both the dorsal and anal fin 
bases are set off from the body. The dorsal fin base is slightly 
longer than the anal fin base. The anal fin base is equal in 
length to the length of the caudal peduncle. The ventral fins 
insert on the ventral surface of the body under the posterior 
part of the dorsal fin base. The dorsal adipose fin is placed over 
the center of the anal fin base. A weak ventral adipose fin is 
present directly anterior to the vent of the paratype. 

The ventral half of the holotype has a light, straw-colored 
ground color. The dorsal half of the fish is brown, while the 
operele and the snout are dusky. Small, dark-brown ehromato- 
phores are scattered at random over the sides of the fish on a 
field of still smaller, more densely distributed, light-brown 
chromatophores. The sides of the head and all of the fins are 
lightly peppered with dark-brown pigment cells. Dark pigment 
is also thickly- distributed on the margins of the scale pockets. 
Some of the scale pockets bear remnants of a purplish, trans- 
lucent material which is common to a number of Bathylagus 
species. Tlie paratype has faded badly, and little in the way of 
pigmentation remains. 



8 BREVIORA No. 98 

Relationships. B. greyae is closest to Bathylagus ocJiotensis 
(which has B. nakazaivai as a synonym, Cohen, MS.). It differs 
in having a greater preanal distance, 81.0 in the type of greyae, 
73.7 and 77.6 in two specimens of ochotensis of comparable size; 
in having a longer preventral distance, 58.4 to 61.1 in greyae, 
51.3 to 57.1 in 17 specimens of ochotensis; in having the anal fin 
base slightly shorter than the dorsal fin base, while in ocJiotensis 
the anal fin base is longer than the dorsal fin base. In addition, 
the postero-ventral margin of the opercle is smooth in greyae, 
while in ochotensis it is drawn out into five, finger-like projec- 
tions (Fig. 2). 

This species is named for Mrs. Marion Grey of the Chicago 
Natural History Museum in recognition of her contributions to 
the study of deep-sea fishes. 

Bathylagus longirostris Maul 

Until the present time this species has been known only from 
the type, apparently taken at Madeira. I have examined a single 
specimen from the Gulf of Mexico, south of Pensacola, Florida, 
collected by the U.S. Fish and Wildlife Service vessel "Oregon" 
and deposited in the Chicago Natural History Museum. In addi- 
tion, two specimens collected in the Central Pacific (30° 16.5' N., 
179° 54' E. and 29° 54' N., 168° 32' W.) by the Pacific Oceanic 
Fishery Investigations (POFI) are apparently referable to B. 
longirostris. Descriptions of the above specimens will be included 
in a projected revision of the Bathylagidae. 

Bathylagus bericoides (Borodin) 

This species has been previously recorded three times, each 
from the Western North Atlantic. I have examined an unre- 
corded specimen from Bermuda (SU 42715) and a specimen 
from "Oregon" st. 1028 (Chicago Natural History Museum). 
The latter was provisionally identified as Bathylagus micro- 
cepha.lus by Springer and Bullis (1956). B. bericoides is also 
present in POFI collections from the central Pacific (39° 31' N., 
178° 54' W.; 39° 08' N., 164° 51' W., and 42° 16' N, 179° 52' 
W.). These specimens will be treated at greater length in the 
future. 



1958 NEW SPECIES OF BATHYLAGUS 9 

ACKNOWLEDGMENTS 

1 am indebted to Dr. Richard Backus, Mrs. Marion Grey, Dr. 
George Myers, Miss Margaret Storey, Dr. Donald Strasburg, 
Dr. Denys Tucker, and Mr. Loren Woods for the loan of speci- 
mens. I thank Mr. G. E. Maul for information on the type of 
B. longirostris and Miss Esther Google for preparing the figures. 

LITERATURE CITED 

Beebe, William 

1933. Deep-sea fishes of the Bermuda Oceanographic Expeditions. 
Family Argentinidae. Zoologica (New York), 16: 97-147. 

Chapman, Wilbert M. 

1943. The osteology and relationships of the l)athypelagie fishes of 
the genus Bailiylagus Gunther with notes on the systematic 
position of Leuroglossus stilbius Gilbert and Therobromus 
caUorhinus Lucas. Journ. Washington Acad. Sci., 33: 147-160. 

HuBBS, Carl L. 

li)19. A comparative study of the bones forming the opercular series 
of fishes. Journ. Morph., 33: 66-71. 

Norman, John R. 

1937. Fishes. Rep. Br. Aust. N.Z. Antarct. Res. Expedn. 1929-31, 
Adelaide, (B) 1 (2): 49-88. 

Springer, Stewart and Harvey Bullis 

19.36. Collections by the Oregon in the Gulf of Mexico. Dept. of the 
Interior. U.S. Fish and Wildlife Service. Spec. Scient. Repts. 
— Fisheries, 196: 1-134. 

Taaning, Aaqe V. 

1931. Postlarval stages of Bathylagus from the North Atlantic. Vi- 
densk. Medd. fra Dansk Naturh. Foren., 92: 269-274. 



BREVIORA 

Miasemm of Compsirsitive Zoology 



Cambridge, Mass. December 19, 195S Number 99 

A NEW SUBSPECIES OF CHAMAELEO JACKSONI 

BOULENGER AND A KEY TO THE SPECIES OF 

THREE-IIORNED CIIAMAELEONS 

By a. Stanley Rand 

Biological Laboratories, Harvard University 

A small collection of reptiles collected on Mt. Meru, Tangan- 
yika Territory, in August 1956, by Lt. Col. J. Minnery was sent 
to the Museum of Comparative Zoology by Mr. C. J. P. lonides. 
It included three male chamaeleons that Mr. Loveridge believed 
to represent a new form but did not describe since he desired to 
confirm the characters on a large series. Fourteen additional 
specimens were obtained through Mr. lonides in 1957 and seven 
in 1958 after Mr. Loveridge had retired as Curator of Herpetol- 
ogy. Five of these last have been sent to the British Museum 
(Natural History). These twenty-four specimens, together with 
an additional one already in the collection of the Museum of Com- 
parative Zoology, form the basis on which the following form is 
described. (The name has been suggested hy Mr. Loveridge.) 

ChAMAELEO JACKSONI MERUMONTANA SUbsp. nov. 

Type. A male, Museum of Comparative Zoology No. 56152 
(collector's No. 7651), apparently adult, collected at Laikinoi, 
near Arusha, Mt. Meru, Arusha District, Northern Province, 
Tanganyika Territory, at 7500 ft. alt., August 1957 by Lt. Col. 
J. Minnery. 

Paratypes. Twenty-four. MCZ 44327, male, Mt. Meru, east at 
9000 ft. alt., 1938, Cooper collector ; MCZ 54790-92, males, same 
locality as type, August 1956, Lt. Col. J. Minnery collector ; MCZ 
56153-59, males and 56151, 56160-64 females, same data as type; 



2 BREVIORA No. 99 

MCZ 56989 male and 56990 female, same locality as type, Janu- 
ary 1958, Lt. Col. J. Minnery, collector; BM (5 specimens), same 
locality as type, January 1958, Lt. Col. J. Minnery, collector. 

Diagnosis. A chamaeleon with three annulated horns in the 
male, closest to Chamaeleo jacksoni Boulenger, but differing from 
it in smaller size, in having the occipital and postorbital areas 
covered with smooth or weakly convex scales (rather than 
strongly convex scales) and in having fewer and larger scales 
between the preocular horns. 

Description. Three annulated horns, one rostral and two pre- 
ocular in position and subequal in length. (In males only, females 
with only a rostral horn, proportionately shorter and with two 
sharp horn scales instead of the preocular horns.) Preocular 
horns arising from the anterior end of the raised supraorbital 
ridges and separated by a mosaic of enlarged flattened scales, a 
few of them with antero-posteriorly oriented ridges. (This ridg- 
ing is more marked in some of the paratypes than in the type.) 
The minimum number of scales counted between the preocular 
horns (males) or horn scales (females), 7 in the type, 6 or 7 in 
the paratypes (see table below). 

The supraorbital ridges continued posteriorly as low ridges to 
the posterior border of the head, separating the posterior dorsal 
and lateral head scales into postorbital and occipital areas. These 
areas are covered with enlarged scales, some of which are flat and 
others weakly convex. (The number of completely flat scales in 
these regions varies in the paratypes but never are there many 
strongly convex scales present.) 

A moderately low curved profile to the casque ; a parietal crest 
present, two or three scales high, forked anteriorly to form low 
ridges that extend antero Jaterally toward the supraocular ridges, 
but do not meet them. These branches of the parietal crest sep- 
arate the large flat occipital scales from an area of small strongly 
convex scales that are distinct from both the occipital scales and 
the interorbital scales anterior and lateral to them. 

No occipital lobes; a very low ridge marking the posterior 
border of the casque and separating the head scales from the 
smaller nape scales. 

Small, soft, triangular tubercles on the upper portion of the 
eyelid. 



1958 NEW SUBSPECIES OF CHAMAELEON 3 

A median dorsal crest of enlarged scales extending from the 
shoulder region nearly to the base of the tail, these trianguloid 
scales largest and most pointed anteriorly, becoming smaller and 
more rounded posteriorly, separated by one to four scales. In 
some cases the small scale immediately in front of the large one 
is distinctly larger than the others but it never approaches the 
largest in size or shape. No crest on the tail. 

The scales on the rest of the dorsal and lateral surfaces 
markedly heterogeneous in size on both body and tail, with an 
irregular arrangement of large flat scales among small granular 
ones and intermediates in size and convexity. The dorsal surfaces 
of the limbs similarly scaled but with large plates predominating, 
particularly on the forelegs. 

The ventral surface covered by uniform granules with some 
very tiny ones between them, except for the sides of the throat 
where there are elongate patches of the very small granules. 
Xo gular or ventral crest evident. 

Color (in alcohol) : The horns, the interorbital and snout scales, 
the supraorbital ridges, and the anterior enlarged dorsal crest 
scales light brown. The rest of the animal blue, lighter dorsally 
and on the head, limbs, and tail and darker on the lips, sides of 
throat and sides and venter of the body. (All the male paratypes 
show this distribution of light yellow brown but vary in the 
intensity of blue on the rest of the body. The females are blue 
with the rostral horn dark brownish.) 

Measurements: See Tables 2 and 3. 

Discussion. The Mt. Meru specimens described above are most 
similar to Chamaeleo jacksoni Boulenger which was described 
from Nairobi. However, they differ in a number of respects. 
The comparison below is based on eighteen adult specimens of the 
typical race from various localities in Kenya Colony. 

The most striking difference is the small size of the Mt. Meru 
series. The largest male measures 91 mm. in snout-vent length, 
the largest female 86 mm. In the Kenya series of eighteen, eleven 
measure 97 mm. or over, snout-vent length. The largest male is 
167 mm., the largest female 140 mm. 

In merumontana the occipital and postorbital scales are flat or 
a few of them weakly convex, giving a smooth appearance to the 
occipital and postorbital areas. In the Kenya specimens many 



4 BREVIORA No. 99 

of these scales are strongly convex giving a bumpy appearance 
to these areas. 

In the Mt. Meru series, the scales between the preocular horns 
(interorbital scales) are larger and fewer in number than in the 
Kenya specimens. The table below shows the specimens arranged 
according to the least number of scales counted betw^een the 
horns. 

Table 1 
SCALES BETWEEN PEEOCULAE HOENS OE HOBN SCALES 



"lumbers of 
















scales 


6 


7 


8 


9 


10 


11 


12 


s'uinljc'i- of JNIt. i\Ieiu 


S 


17 

















specimens 
















with each 
















scale count Kenya 





1 


3 


8 


5 





1 



In the Mt. Meru series the large fiat interorbital scales (some of 
them sometimes having a median keel) differ sharply from the 
mure posterior small strongly convex scales between them and the 
branches of the parietal crest. In the Kenya specimens the inter- 
orbital scales do not differ markedly from the scales behind them. 

In the Kenya specimens the dorsal crest is formed by groups 
of two scales, the first of which is smaller than the second, but 
both are larger than the scales between groups, and both are 
pointed — at least in part. The enlargement of the anterior 
scale is much less evident in some of the smaller specimens. In 
the Mt. Meru specimens the anterior scale in each group is rela- 
tively smaller. It is usually but little larger than the surrounding 
scales, so that the crest seems to be composed of single, isolated, 
enlarged scales. 

The ridges on the head of the Kenya form are more distinct 
than those of the Mt. Meru series. This is particularly true of 
the ridge along the posterior border of the casque and the anterior 
branches of the parietal crest. The more marked parietal crest 
gives a noticeably higher profile to the casques of the large males 
from Kenya. 



1958 NEW SUBSPECIES OK CHAM AELEON 5 

The body scales of all of the specimens are markedly hetero- 
geneous. The two forms differ only in that the large scales tend 
to be flatter in tlie Mt. Mern specimens than in the Kenya speci- 
mens. 

In seven of the eight Mt. Meru females the rostral horn is well 
developed, although shorter than that of a male of equivalent 
size. The eighth female has a very small rostral horn, and none 
has preocular horns. In the six large Kenya females, one has 
both rostral and preocular horns developed. In the other five 
the preocular horns are 2 nnn. or less. Three of these have a 
rostral horn and two have only a sharp rostral scale. 

Loveridge, 1957, gives the range of Chamacleo jacksoni as 
"Highlands of Kenya Colony and Tanganyika Territory." Ap- 
parently his inclusion of Tanganyika was based on the MCZ 
specimen No. 44327 from Mt. Meru, listed here as a paratype 
of C. j. merumoniana. No other specimens or records of C. jack- 
soni from Tanganyika have been located. Therefore, the range of 
C. j. jacksoni can be restricted to the highlands of Kenya Colony, 
and that of C. j. mcrumontana can be given as Mt. Meru, between 
7500 and 9000 ft. alt. 

Of the type and those paratypes collected by Lt. Col. J. Min- 
nery, C. J. P. lonides writes: "They are found on bushes and in 
low small trees. Laikinoi is a farm on the very edge of the rain 
forest." (Letter dated 5 December 1957.) 

Three-horned chamacleo7is. In the course of describing this 
form, specimens of the five other species of three-horned chamael- 
eons were examined. A few notes and a key for the identification 
of these species are included below. 

The six species characterized by the possession of three annu- 
lated horns in the male do not seem particularly closely related. 
The other morphological characters that they have in common 
are negative rather than positive : lack of spurs in either sex ; 
lack of axillary j^ockets. 

Among them, C. jacksoni seems closest to C. johnstoni Boulen- 
ger, again primarily on negative characters : absence of occipital 
lobes, absence of a gular crest, absence of a dorsal fin, but also in 
the possession of heterogeneous body scales. 

C. wcrneri Tornier and C. fullehorni Tornier seem quite closely 
related to eacli other, though not to either C. jacksoni or C. 



6 BREVIORA No. 99 

johnstoni. The only other close relationship involving these 
species seems to be between C. johnstoni and C. ituriensis 
Schmidt, the latter a hornless chamaeleon of the eastern Belgian 
Congo forest. 

C. oweni Gray is widespread throughout the lowland rain 
forest in West Africa extending into eastern Belgian Congo. All 
the others are mountain forest forms in East Africa : C. fulle- 
horni from the Ngosi Volcano, Poroto Mountains, Ukonde, Tan- 
ganyika Territory; C werneri from the Uluguru and Uzungwe 
Mountains, Tanganyika Territory; C. johnstoni from the virgin 
forest of Uganda, Belgian Ruanda-IJrundi and the adjacent 
Belgian Congo ; C. deremensis Matschie from the Usambara and 
Uluguru Mountains, Tanganyika Territory; C. jacksoni from 
the highlands of Kenya Colony and Mt. Meru, Tanganyika Terri- 
tory. 

It is noteworthy that except in the Uluguru Mountains no two 
species of three-horned chamaeleons are known to occur together. 
This is evidence supporting the suggestion that the three horns 
serve these chamaeleons as a means of species recognition. The 
case of the two species occurring together on the Uluguru Moun- 
tains supports rather than contradicts this view. These two 
chamaeleons are very different in appearance. C deremensis is 
large, laterally compressed and with a high dorsal fin ; C. werneri 
is smaller and stocky with no dorsal fin. Even with the presence 
of three horns in both there could be little opportunity for 
confusion. In eastern Belgian Congo where a close relative of 
the three-horned C. johnstoni, C. ituriensis, occurs with the three- 
horned C. oweni, both species are similar in size and proportions, 
but C. ituriensis is hornless. 

Key to the three-horned chamaeleons 

1. High sail like dorsal fin C. deremensis 

No high sail like dorsal fin 2. 

2. Body scales equal in size or nearly so C. oweni 

Body scales differing markedly in size 3. 

3. Well developed occipital lobes 4. 

No occipital lobes 5. 

4. Occipital lobes continuous across back of casque as 

a flap C. iverneri 

Occipital lobes separated by a gap in the middorsal 

region C. fulleborni 



1958 



NEW SUBSPECIES OF CHAMAELEON 



5. Dorsal crest of much enlarged scales in groups of 

one or two 6. 

No dorsal crest of much enlarged scales 7. 

G. Occipital and postorbital scales strongly convex, scales 
between preocular horns smaller and more numer- 
ous (7-12), size larger C. j. jacksoni 

Occipital and postorbital scales flatter, scales between 

preocular horns larger and fewer (6-7), size smaller C. j. merumontana 

7. * Dorsal profile of body smooth or undulating C. j. jo'hnstoni 

Dorsal profile of body crenulated C. j. crenulata 

* C. johnstoni crenulata not seen, this section of the key taken from Laurent, 
1951. 

Table 2 

MEASUREMENTS OF MT. MERU SPECIMENS 

w 

a a 

— a u 

a ^ c ** ® 

-• o . It' tc -—^ 4-1 

= t. > ^ Sr S"" = -^^ 

s= >< 5= i-r ^^ 2§" «S 

Sz 32 x^:; 5J^j CL(^ S^o wa 

MCZ 44327 male 68 13 11 22 7 

54790 '' 88 19 18 28 6 

54791 " 76 16 14 24 7 

54792 <' 60 7 6 19 7 
*56152 " 91 19 18 29 7 

56153 " 73 18 18 24 6 

56154 " 78 15 13 26 6 

56155 " 75 16 15 25 7 

56156 " 73 17 16 24 7 

56157 " 90 20 16 29 7 

56158 " 71 11 10 24 6 

56159 " 64 10 8 20 7 

56989 " 86 23 22 26 6 
BM " 78 17 15 26 7 

" 82 16 15 24 7 

"57 9 7 19 7 

" 79 14 13 25 7 

56151 female 74 7 — 23 7 

56160 " 77 7 — 23 6 

56161 "80 8 — 22 6 

56162 "81 8 — 25 7 

56163 "74 8 — 23 7 

56164 " 86 6 — 26 7 

56990 "78 8 — 24 6 
BM "79 2 — 24 7 

* Type of C. j. merumontana 



8 



B[iEVIORA 



No. 99 



Table 3 
MEASUREMENTS OF KENYA SPECIMENS 



u 
i) 

s 

N 






a 

5a 


a 

o 
.a 

C3 - 

o a 


o s 


Head length 
(snout to end 
of casque) 


it: a 

a, o 

"3 ? 
II 


Embu Dist 


lict 














7842 




male 


110 


15 


15 


35 


9 


7842 




female 


140 


— 


— 


41 


12 


Meru 
















11467 




male 


167 


35 


33 


52 


9 


29454 




I < 


118 


22 


22 


41 


10 


Lukenya 
















34995 




female 


99 


7 


— 


31 


10 


34996 




male 


111 


16 


16 


36 


9 


Parklands, 


Nairobi 












18365 




male 


98 


15 


13 


30 


9 


18366 




female 


123 


13 


6 


37 


9 


18367 




It 


97 


5 


2 


29 


8 


31365 




t i 


103 


3 


2 


32 


9 


31366 




( ( 


105 


— 


_i_ 


31 


8 



LITERATUEE CITED 

BOULENGER, G. A. 

1896. Description of a new chamaeleon from Uganda. Ann. Mag. Nat. 
Hist., (6)17: 376. 

Laurent, R. 

1951. Deux reptiles et onze Imtraciens nouveaux d'Afrique central. 
Rev. Zool. Bot. Afr., 44: 360-381. 

LOVERIDGE, A. 

1957. Checklist of the reptiles and amphibians of East Africa. Bull. 
Mus. Comp. Zool., 117: (2): 153-362, i-xxxvi. 



BREVIORA 

MmseiLiini of Comparative Zoology 



Cambridgk, Mass. December 23, 1958 Number 100 

OX THE PINEAL ORGAN OF THE TUNA, 
THYNNUS THYNNUS L. 

By Uno Holmgren 

ZooJogical Institute, Uppsala, Sweden and 
Biological Laboratories, Harvard University, Cambridge 

A conspicuous piueal spot has been found in many seombrid 
fishes, e.g-. Thynnus, Germo, Parathynnus and Neothynnus and 
the related genera Auxis, Katsuivonus, Enihynnus and Sarda. 
Underlying- the pineal spot a fontanelle or "pineal window" 
(Rivas, 1952) is present in the last four mentioned genera and 
also in Scomber and Scomheromorus. Rivas investigated the 
pineal apparatus of the tuna, Thynnus thynnus L. and claimed 
a photoreceptory role of the pineal area, but he "failed to reveal 
the presence of a pineal organ, which is apparently absent or 
greatly reduced" (Rivas, 1952, p. 174). My researches show, 
however, that a jiineal organ is present in the tuna. 

A supply of the bluefin tuna, Thynnus thynnus L. was obtained 
from an expedition on board M/S "Delaware," belonging to 
U.S. Fish and AVildlife Service. Five fishes were dissected, and 
the pineal organ of each was sectioned and stained, using 
Bodian's protargol method (Romeis, 1948, p. 421) to determine 
the presence of nerve elements, and Heidenhain's azan stain 
(Romeis, 1948, p. 344) for the general histology. The sections 
were cut at 8 microns, using paraffin technique. 

Dissections of the Bouin-preserved material of the tuna re- 
vealed, as had previously been pointed out by Kishinuya (1923), 
Gregory (1933), Godsil and Byera (1944).^ and Rivas (1952), 
the presence of a fontanelle, underlying a conspicuous white 
pineal spot 15 x 30 mm. in size, in a specimen weighing 350 lbs. 
(Fig. 2). The spot is situated slightly caudally from an assumed 



2 I3REVIORA Xo. lUO 

line eoniiecting tlu^ centers of the eyes. The dermal layer over 
the pineal fontanelle was found to be transhieent, transmittin<i 
about 20 i)er cent of the liji'ht, measured Avitli a ])h()tometer. 

The fontauelle in the frontal bone, below the dermal layer 
(Fi«i'. '■]). was filled with a jelly-like translucent substance, and 
measured slightly less than the overlying white pineal spot, about 
10 X 30 mm. in the specimen mentioned above. 

About .") mm. below the ed^c of the fi-ontal bone, the jelly-like 
material pa'^sed into a more cartila<>inous tyi)e of tissue. A 
tubular excavation around the pineal organ was formed below 
the pineal fontanelle, leading down to the brain at an angle of 
60 or 70 degrees (Fig. 1). The excavation had a diameter of 
10 mm. The end-vesicle of the pineal organ was situated immedi- 
ately below a dome-shaped cartilage (Figs. 1, 4). The vesicle 
was approximately 10 mm. in diametci- and connected to the 
diencephalon by a stalk. \'entrally the stalk had a white cord 
which distally seemed divided into two branches ; in the specimen 
photographed the stalk was 30-35 unn. in length (Fig. 1). 

The histological examination of the end-vesicle showed a struc- 
ture similar to that of most other teleost fishes. The lumen, 
hoAvever, was larger and the walls were thinner, consisting of 
only one or two layers of cells (Figs. 4, 5). Because the pineal 
cells were secretory and had cytoplasmic processes extending into 
the lumen of the vesicle, the cells of the end -vesicle and the stalk 
(Figs. 4, 5) were similar to the primary sensory cells in teleosts 
described by N. Holmgren (1920) and *Friedric'h-Freksa (1932). 
There were also present su])porting cells ("8tiitz/,ellen" Fried- 
rich-Freksa. 1932) which had more hyperchromatic nuclei and 
more indistinct cytoplasm than the sensory cells. 

A cross-section of the pineal stalk revealed, especially in the 
distal part, folded walls. Connective tissue, stained blue with 
lleidenhain's stain, formed a thin layer inside and outside the 
wall of the stalk. Silver impregnation did not give any evidence 
of axons eitlier in the pineal vesicle or in the stalk. A tractus 
pinealis is ])resent, at least in the ])i'()ximal |)ortion of tlic ])ineal 
stalk, in many teleost fishes. Apparentlv the tuna is one of the 
exceptions. 



1958 I'INEAL ORGAN OK THE TINA 3 

Tlic pineal structures in the tuna were t'ound to i)e very vuscil- 
lar, osj)e('ially below tiie eiul-vesiele of tiie pineal organ ( Fi<i'. 4) 
and in its walls. There, many small blood vessels filled with 
blood corpuscles were found. 

Kivas (1952) suggested positive |)hot()tropism in tunas usinji: 
the experiments of Hsiao (1952) to support his idea. The 
latter found that both the Pacific yellowfin tuna {Neoihynnus 
inacroptcrKs) and the Pacifie little tunny {Enfhynnns yaito) 
showed phototaxis. In these experiments the pineal area was 
considered to be partly responsible for phototaxis. Breder and 
Kasquin (1947, 1950) also have shown a correlation in certain 
teleost fishes between the degree to which the pineal area is 
exposed to light and the amount of positive phototaxis. Grune- 
wald-Lowenstein (1956) demonstrated in the teleost Astyarwx 
that prolonged exposure to light or darkness had an effect on 
the glycogen content of the pineal. Rasquin (1958) reported 
that the pineal secretion in teleosts w^as affected by light ; this 
has also been found to occui- in higher animals (Quay, 1956). 
While there is evidence that light in some way stimulates the 
pineal's secretory activity, it is also known that the diencephalon 
in some fishes is light sensitive (Scharrer, 1928; Young, 1935). 
This had already been indicated by von Prisch (1911) w^ho 
demonstrated some continuing sensitivity to light by the 
teleost Phoxinus after extirpation of the pineal. Since von Frisch 
gives us reason to believe that the lining of the ventricle of the 
diencephalon has light-sensitive cells, and the work of Scharrer 
and Young corroborate this, the whole i)ineal area nuist be in- 
cluded in any discussion on the location of the light-sensitive 
apparatus. While the histological examination of the tuna's 
pineal organ showed thin walls which could possibly transmit 
light, the interior cells did not show any pronounced secretory 
activity. Therefore, taking into consideration the preceding evi- 
dence presented in this paper, one can conclude that while there 
are morphological indications of the possible light sensitivity of 
the pineal area in the tuna, there is not j^et enough physiological 
evidence to make a firm commitment as to the possible ecological 
or hormonal control by this area, 



4 BREVIORA No. 100 

REFERENCES 

Breder, C. y\. and I'. R.vsQi'ix 

1947. Comparative studies in the light sensitivity of blind ehaiaeius 

from a series of Mexican caves. Bull. Amer. Mus. Nat. Hist., 

89(5): 325-51. 
1950. A preliminary report on the role of the pineal organ in the 

control of pigment cells and light reactions in recent teleost 

fishes. Science, 111(2071): 10-12. 

Friedrich-Freksa, H. 

1932. Entwicklung, Bau and Bedeutung der Parietalgegend bei Tele- 
ostiern. Zeitschr. wiss. Zool., 141: 52-1-42. 

Frisch, K. v. 

1911. Das Parietalorgan der Fische als functionierendes Organ. Sitz- 
ber. Gesell. Morph. Physiol., 27: Ki-lS. 

GoDSiL, 11. C. and R. D. Byers 

1944. A systematic study of the Pacific tunas. Calif. ])iv. Fish and 
Game Fish Bull. Sacramento, 60: 1-131. 

Grexjory, W. K. 

1933. Fish skulls: a study of the evolution of natural mechanisms. 
Trans. Amer. Philos. Soc, (n.s.) 23 (2) : 75-481. 

Grunewald-Lowenstein, M. 

1956. Influence of light and darkness on the pineal body in Astyaiiti.i- 
mexicanus (Filippi). Zoologica, New York, 41: 119-128. 

Holmgren', N. 

1920. Zur Anatomie und Histologie des Vorder- und Zwischenhirns 
der Knochenfishe. Acta Zool., 1: 137-315. 

Hsiao, S. C. 

1952. Observations on the reaction of the tuna to artificial light. 
Spec. Sci. R^-p. U. S. Fish Wildlife Serv., 91: 36-58. 

KiSHINOUYE, K. 

1923. Contrilnitions to the comparative study of the so-called scom- 
broid fishes. Jour. Coll. Agrir. Tokyo, 8: 21)3-475. 



1958 PINEAL ORGAN OK THE TI'NA 5 

QiAV, W. B. 

]i>r)(i. Vohiiiu'tiic jind rytologic vaiiat icDi in the piiR'al IkkIv of Pcro- 
myscu^ leucopu,^ (Rodentia) with respeit to sex, captivity and 
day-length. Jour. Morph.. 98: 471-498. 

R.ASQUIN, P. 

1958. Studies in the control of pigment cells and light reactions in 
recent teleost fishes. Bull. Amer. Mus. Nat. Hist., 115: 1-68. 

RiVAS, L. R. 

19r)2. The pineal apparatus of tunas and related sconibrid fishes as a 
possible light receptor controlling phototactic movements. Bull. 
Mar. Sci. Gulf and Caribbean, 3: 169-180. 

ROMEIS, W. 

1948. Mikroskopische Technik. l.'tth ed. Munich. 

SCHARRER, E. 

1928. Die Lichtempfindlichkeit blinder Elritzen. (Untersuchungen 
iiber das Zwisehenhirn der Fisehe). Zeitschr. vergleich. Physiol., 
7: 1-38. 

Young, J. Z. 

1935. The photoreceptors of lampreys: II. The functions of the pineal 
complex. Jour. Exper. Biol., 12: 254-270. 



Fig. 1. ]Jissec-ted .spei-iir.eii showing tlic end -vesicle of tlie jjineal organ 
and the i)ineal stalk, ending in tlie diencephalon. Tliis pieture also shows 
tlie tulielike excavation extendintj- from tlie iipi>er edges of the frontal lione 
to the brain. Photo 3.") x. 



Fig. 2. The pineal spot of Thynnus iliynnus. Specimen of 350 lbs. 
Natural size. 

Fig. ?y. The pineal fontanelle in the frontal bone. A jelly-like material 
is present in the fontanelle. 

Fig-. 4. Part of the pineal organ situated below the cartilaginous tissue 
mentioned above. Note the conspicuous lumen of the organ and the presence 
of blood vessels below the end-vesicle. Fixation Bouin 's solution. Heiden- 
hain 's azan stain. 150 x. 

Fig. 5. Section of the wall of the end-vesicle. Cytoplasmic processes 
are extending into the lumen of the pineal organ (arrow). Fixation Bouin 's 
solution. Heidenhain's azan stain. 1200 x. 

Fig. 6. Cross-section through the distal part of tlie pineal stalk, showing 
its folded walls. Fixation Bouin 's solution, Heidenhain's azan stain. 300 x. 



BREVIORA 

Museiiim of Comparative Zoology 



Cambrid(;e, Mass. March 2, 1959 Number 101 



CERVICAL RIBS IN TURTLES 
By Ernest E. Williams 



The existence of cervical ribs in Recent turtles is frequently 
denied. In fact, however, cervical ribs in turtles have been dis- 
covered and rediscovered several times over. They are rudi- 
mentary in all post-Triassic forms and the nature of these highly 
peculiar rudiments is evident only in embryos or young speci- 
mens. Tlie only description with any pretensions to fullness is 
that of C. K. Hoffman (1879, repeated in Bronn's Thierreich 
1890). There is brief mention of their occurrence in embryos by 
Ruckes (1929), Naef (1929) and Emeliaiiov (1937). None of 
these previous observations has been sufficiently detailed to make 
the nature and relationships of the rib elements quite clear, and, 
except for ratber crude and incomplete sketches in Hoffman, no 
figures have been given. 

I have myself seen cervical ribs and rib rudiments in botb 
adults and embryos (Emydinae) or in adults only (Testudininae, 
Chelydridae, Dermochelyidae, Chelyidae and Pelomedusidae). 
Embryos of Emys orhicidaris given me by Dr. B. AV. Kunkel and 
of Pseudemys sp., loaned me by Dr. H. Ruckes, and of Chrysemys 
)iiorgi)iafa and Chelydra .serpentina in the Minot Collection, 
Harvard University, have provided me with the opportunity to 
study the embryonic condition and developmental history of the 
cervical rib rudiments, while I have been able to examine adult 
conditions from specimens in the osteological collections of the 
Museum of Comparative Zoology (MCZ), the American Museum 
of Natural History (AMNH), the United States National Mu- 
seum fUSNM), the British Museum (Natural History) (BM), 



BREVIORA 



No. 101 



the Institut des ►Scieiiees Naturelles in Brussels, the Senekenberii' 
Miiseuni in Frankfort, the Zoolop'ische Staatssammlung- in Munich 
and the Laboratoire d'Anatomie Comparee of the Museum 
National d'Histoire Naturelle de Paris (LAC). 

In tiie course of tlie present study a wide variety of conditions 
lias been observed, iiu'hulin<i' (in adults) considerable intra- 




Juber- 
culum 



Corpus 
costae 



Capitulum 



Fif^'. 1. Diasiaiu to show the lili region in prechondrnl and cnrtilago 
stages of tlie cervical vertebrae of Eini/.s orhiculdris. 



specific \ariatiuii. hi many iustaiuH's the ribs are fragiiieiited 
or broken into little ossicles, which lia\-e lieen many times rc- 
portcil but ijircly correctly iiiter|)rete(l. In yet otiicr instances 



If),")!) C'lOKVICAIi KIHS IN TUK'l'Li;s 3 

tlio ribs ai'o still nioi'e i-udiinentary, i'e])i'es('iite(l in the adults only 
by cryptic fused eleiiicnts. It would, in fact, be vci-y difficult lo 
understand tlie i-ibs of Kecent adult forms if it were not for llic 
evidence of embryos, and (as \vc sball see) of the Triassic fossils. 

The embryonic cei-vical i-ihs of emydine turtles are I ir()-li((i<l((l 
witli a dorsal tubercular attachment at the reo'ion of the neuro- 
central suture and a ventral attachment to the anterior inter- 
central ossicle, when that is present, oi" to the anterior centrum 
when the intercentrum is absent. These ribs may first be dis- 
tinguished in precartilage stages (Fig. 1). They are then not very 
different from the precartilage cervical ribs of other reptiles. 

With the onset of chondrification, however, these ribs undergo 
a partial degeneration. The dorsalmost portion chondrifies at 
its place of attachment to the neurocentral sutui-e, but no joint 
forms here, and the separateness of the element is indicated at 
most by a suture line in cartilage (a line of small densely-packed, 
poorly-ditferenticited cartilage cells like those which constitute the 
cartilage neurocentral suture). 

From the dorsal rib rudiment at the neurocentral suture a 
band of prechondral (later ligamentous) tissue extends ventro- 
anteriorly toward the anterior intervertebral region. In this 
band there is embedded at the middle of its length a se]mrate 
small cartilage nodule. The prechondral or ligamentous band 
then continues forward to attach anteriorly to a separate antero- 
ventral nodule alongside a median intercentrum or to this element 
fused onto the intercentrum as a lateral parapophysial projec- 
tion. 

In the Emydinae, therefore, the bicipital rib of the membranous 
stage has fragmented into three distinct chondral elements. The 
morphological interpretation of these elements will be discussed 
a little later ; for the present it will be Avell to have neutral terms 
for them, avoiding questions of homology. I suggest, therefore, 
calling the dorsalmost element the alpha rib element, the middle 
one the beta, and the ventralmost the gamma element. 

I have seen this fragmented condition of the cartilage ribs in 
both Emys and Pseudemys embryos. I have not had the material 
to follow these structures into ossification stages, but the same 
elements are readily recognized in adult Kdij/s and Pseudemys 
and in other Recent emvdines. 



BREVIORA 



No. 101 



In all liviiio' cryptodires the alpha element of the embryonic 
ribs is apparently represented by the low indistinct transverse 
processes fonnd anteriorly at the junction of neural arch and 
centrum on each cervical. C. K. Ilotfman (1890) has reported 
that this element ossifies separately in tortoises but remains 
cartilaginous in other forms. The second statement appears to be 
erroneous (except for Derynochehis) ; the first I cannot confirm 
from personal observation, but the statement has been repeated 
(whether copied or verified) by .J. S. Thomson (1932). 

I have three times observed separate ossification and hyper- 
trophy of this element, in all cases as individual variations and 
always on the eighth vertebra — once asymmetrically (on the 




Fig. 2. Pyxis arachnoidcs. Anterior view of alniorninl 8th vertebra. Sym- 
metrical hypertrophied alpha elements. 



left side only) in a spt^ciiiieii of I'si iidi inijs scripla (hydiis (La- 
fayette College), once symmetrically in (iopln rus Ixrlandieri 
(A. M.N. 11. 91916), and ouce symmetrically in I'lfxis anivh noidcs 
(L.A.C. 1885-34) (Fig. 2). Siebenrock (liKHJ) has reported the 
same condition in the last form. These conspicuous abnormal 
examples apart, the alpha element appears in all Recent adult 
types to be a cryptic fused element. 

The ventral rib elements, when j)reseiit. arc less frequently 
rryptic, but extreme intraspecific variability makes their dc^sci-ip- 
tion somewhat difficult. 

Jn emydines, the gamma clement mav he |)resen1 on ail the 
eervicals, usualh' fused with (lie intefeent fa on tlie anterior 



1959 CERVICAL KIBS IX TITKTLES 5 

vertebrae, free on tlic middle vertebrae, and fused afraiii but 
with the anterior end of the eentrum on the last eervieal. I have 
se(ni free <2:aninia ossieU's in: HafiKjur haska, Kaelnifia frcfm}), 
Orlltia horncrnsis, Malaclcniys terrapin, I'scudcmys scripta, I'scu- 
(iemifs fioridana, Pseudcmys nihriventris, Pscudemys terrapen, 
Chi-ysemys picta, Graptemys geographica, Graptemya oculifera, 
Clemmys iiisculpta, Clcmmys nnihienhergi, Clenimys guttata, 
Clemniys leprosa, Clcmmys marmorata, Emys orbicularis, Emy- 
doidca htandingi, Deirochclys rcticularia, Tcrrapene Carolina, 
Geocmyda trijuga, Gcoemyda punctularia, Cuora amhoinensis 
and Cyclemys dcntata. When it is considered that these are 
small elements which, if free, are readily lost by the usual 
methods of skeletal preparation, it becomes very probable that 
<>annna ossicles are typically present in the Emydinae. Indeed, 
ill one instance they are probal)ly invariably present: the antero- 





Fig. 3. Tcnapene caroUiui. A. Ventral view of joint between 5th and 
Gth cei'vical centra showing separate beta and gamma rib elements. B. 
Anterior view of 6th vertebra in the same specimen. Compare Fig. 5. 

ventrolateral processes lyinw alongside the hypapophysis of the 
eijjrlith centrum in emydines are apparently fused gannna ossicles. 
Similar fusion of g'amma ossicles occurs rather frequently at the 
anterior end of the seventh centrum. 

The beta element, on the other hand, is unknown in adults 
except on the middle cervieals of a few individuals of certain 
emydine species. Here a^rain, however, the probability of loss 
of these very small and loosely-attached elements must be taken 
into account as a possilile explanation of the apparent rarity of 
these elements in adults. 



6 BREVIORA No. 101 

All instance in whidi hotli ventral rib elements are well de- 
veloped is seen in Figure 3A. The joint between centra 5 and 6 
of a specimen of Terrapenc Carolina from my own collection is 
shown in ventral view. The g'amma elements are relatively large 
flattened ossicles attached to the ventral side of the centrnm 
(centrum 5) just below the paired, convex, posterior central 
articular surfaces. These ossicles extend posteriorly and some- 
what laterally to lie ])artly underneath the anterolateral diminu- 
tive transverse processes (alpha rib elements) of the posterior 
vertebra (vertebra 6). (lamma and alpha rib elements are con- 
nected on each side by ligaments in which are embedded small 
rounded ossicles which are evidently the persisting beta elements. 
Figure 8B shows the same elements in anterolateral view. (Corn- 




Fig. 4. Pseutlcmy.i sp. cml)iyo. Seetion tlnouKli liglit side of n middle 
cervical vertebra in early cartilage stages showing all three lil) elements. 
Comiiaie Fig. 3B. 

pare Fig. 4.) The only element of the morphological chain which 
we saw in the eml)ryo that is missing in tliis case is the median 
intercentrum ; this is consistently absent on the posterior cervicals 
of turtles. 

Museum specimens in wliicii 1 have seen comparable condi- 
tions (i.e. both beta and gamma ossicles ))resent) are: Clenintys 
muhlcnhcrgi U.S.N.M. 2922S, Clemrnys insciilpta F.S.N.M. 
6308!) and rhrysrmys picta F.S.X.M. 29238, as well as Emjfs 
urbicularis l\M. 1!)2()-1 -20-2248 and other specimens of this 



]!)5!) Clil{Vl(Ah i;iBS L\ TUHTLES 7 

sperics at the British Muscuin. I should emphasize tliat tlicsc 
specimens are not the only ones by any means in wliieli 1 have 
observed beta and ji'anima ossicles together; these are only certain 
readily accessible specimens upon which my observations may 
be confirmed. Museum specimens are not likely to have preserved 
these elements; careful preparation of specimens is required to 
reveal them. 

As mentioned above, on anterior cervicals the gamma ossicles 
may be fused with the true median intercentra (or with the 
anterior of the two centra between which they lie), but occasional 
si)ecimens show them separate from and lying alongside the 
intercentra, presenting in such cases the appearance of tripartite 
intercentra. 1 have seen this condition in Chrysemys picta 
U.S.N.M. 29233, Pseudemys floridana U.S.N.M. 60492, and Orlitia 
horneensis U.S.N.M. 37788 — in the first case only at the odon- 
toid-second centrum articulation, in the two latter also at the 
joint between centra 2 and 3. I have seen such apparent triple 
intercentra also in Chehjdra serpentina (a specimen at Lafayette 
College, A.M.N.H. 28942, and specimens 18308 and 18361 in the 
Natural History Museum of the University of Kansas). 

Siebenrock (1907) mentions gamma ossicles in Plotysfernon 
megacephoIi())i, regarding them, howevcn-, as rudimentary ventral 
arches. He says : ' ' Rudimentare unteren Bogen, Hypapophysen, 
kommen so wie bei Macrodemmys Gray and bei vielen Emydidae 
am ersten bis f iinften Halswirbel vor ; sie f ehlen dagegen bei 
Chelydra Schw., wie ich mich an drei darauf untersuchten Skelet- 
ten iiberzeugen konnte." 

As Siebenrock states, gamma ossicles appear to be usually 
absent in Chelydra except at the odontoid-second centrum articu- 
lation, but one specimen of this genus at the Senckenberg Museum 
has gamma ossicles on centra 3 to 5, as does another specimen 
in the Staatssammlung at Munich. 1 have not myself seen any 
gamma ossicles in Maeroclemys, though Siebenrock mentions 
them above, nor in Staurotypus or in Kinosiernon. Beta ossicles 
have not been seen in any of the Chelydridae sensK lato. 

I have seen neither of the ventral rib ossicles in Dermatemys, 
and I have seen gamma ossicles only rarely in the Testudininae 
(e.g. at the odontoid-second centrum joint in Gopher us herlan- 
dieri A.M.N.H. 91916). l)ut Siebenrock (1906) records a series 



8 



15REVI0RA 



No. 101 



of gamma ossicles in Acinixys planicauda, and Thomson (1932) 
reports them on the fourth, fifth and sixth vertebrae of Toiindo 
ihera and T. graeca. 

In the Cheloniidae and Dermoehelyidae one ventral rib element 
is usually present on most or all the eervieals. It is cartilaginous 
in Dermochelys (e.g. U.S.N.M. 29492, see also the figure in Volker 
1913), bony in Caretta caretta (e.g. U.S.N.M. 29372) (Fig. 5) 
and Eretmochelys imhricafa (e.g. U.S.N.M. 59866). A special 




Fig. 5. Caretta caretta. A. Anterior view of 6th vertebra. B. Ventral view 
of joint l)etween 6th and 7th centra in the same specimen. 




Fig. 6. Oiclonia mydas. A. Anterior view of 7th cervical vertelira. B. 
Anterior view of right side of the 7th vertelira of another s])ecimen. 



condition is seen in Chclonia jiit/das (A.M.N.H. 46837) (Pig. 
6A) in which on vertebra 7 the rib rudiment is seen to be fused 
to the right anterior side of the vertebra in such a way as to 
enclose a small "vertebrarteriar" canal. The " vertebrarterial" 
canal is incomplete laterally on the left side of the same vertebra. 
In another specimen of the same species (A.M.N.H. r)912) (Pig. 
6B) the " vertebrarteriar " canal is completed lat(M-aIly liy car- 
tilage on both sides of the seventh vci'tebra. 



1!);")!) CERVICAL KIIJS IN TURTLES 9 

I have s(MMi ossified fjamma ossit'les also in the Eocene sea 
turtle Eospliargis (jiuas in the magnificently preserved specimen 
in the Rruskels Museum. 

Beta ossicles are unknown in the ("heloniidae, and they may in 
fact be merged with the rather larnrer apparent e(}uivalents of the 
jijunma ossicles of emydines. 

1 have seen no trace of rib rudiments except the transverse 
])rocesses in adult trionychids or in Careftochelys, but gamma 
ossicle ecpiivalents exist in at least ChcJus and Er]imnoehely!< of 
the Pleurodira. 

These peculiar conditions of the rib elements in modern turtles 
are fortunately traceable to a more primitive condition in Triassic 
turtles. Jaekel (1916) has described and figured the situation 
in Trinssochelys dux {-=Proganochelys qucnstedfi), and 1 can 
confirm his description in all essentials, having examined the 
magnificent skeletons of Profjanochelya in the possession of the 
Stuttsi'art Museum, which are to be described bv Dr. K. Staesche. 




Fig. 7. Proganochelys quenstedti (=Triassochelys dux). Anterior view 
of 7tli ccrvieal vertebra showing liieipital lili. (After Jaekel.) 

Figure 7 presents an anterior view of the seventh cervical 
vertebra in Proganochelys (after Jaekel). It will be seen that 
these are ribs of a normal bicipital type, the tuberculum lying 
across the neurocentral suture, the capituhnn articulating more 
ventrally with the side of the centrnm. Ribs of this type were 
present on cervicals 2 to 7. The eighth rib mucii resembled the 
abnormal vertebrae of Pseudcniys, Pyjris, etc. described above. 
It is single-headed rather than bicipital, elongate rather than 



.10 BREVTORA No. 101 

short. It curves downward distally to a degree appropriate to its 
leugrth. Its single head lies across the neuroeentral suture in the 
position of the tuberculum. Probably the eighth and the ribs 
behind it retain only the tuberculum of the tAvo original rib heads. 

The congruence of the bicipital ribs in Proganochelys with 
the ribs that would be expected if the embryonic rib elements of 
Recent turtles were fully developed is very complete. With this 
evidence before us, the interpretation of the otherwise very singu- 
lar, even anomalous rudimental rib ossicles of Recent turtles can 
hardly be questioned. 

I have seen similar small ossicles that are apparently rib rudi- 
ments in large lizards, e.g. Iguana iguana A.M.N. H. 1269 (Com- 
parative Anatomy collection). The phenomenon of rib frag- 
mentation is thus not confined to the neck of turtles. Indeed, 
there is a considerable literature on anomalous rib conditions in 
Amniota (Baur 1886, Howes and Swinnerton 1!)()1 — Sphcno- 
ilon; Xoble 1921 — Kphaerodactylus; Schinz and Zangerl 19371) 
— Gallus; Virchow 1929 — Ornithorhyncus; Albrecht 1882, 
Hayek 1928, de Burlet 1917, Federow 1910, Abel 1909 — various 
mannnals.) 

These special conditions have, I feel, implications of greater 
morphological moment than appears on the surface. They may 
require a careful re-examination of the origin and morphology 
of ribs throughout the Amniota. 

Rib fragmentation in tetrapods occurs typically in regions of 
the column in which special movements are taking place (neck of 
turtles, lizards, etc., tail of amniotes generally), and in which, 
tlicrefore, the thoracic rib pattern is not useful, or in acjuatic 
forms in which the thoracic pattern itself may prove unstable. 
It is a phenomenon, therefore, of degeneration — of breakdown 
of the two-headed ribs })rimitively present in tetrapods, ])re- 
sumahly throughout the cohnnn. However, it is precisely the 
origin and evolution of the two-headed ribs of tetrapods that 
is still a puzzle — a puzzle to which, 1 believe, these special condi- 
tions along with the ontogeny of less modified ribs have something 
to contribute. Sjiecifically, the (piestion of the possil)le origin of 
two-headed tetrapod ribs from a union of the dorsal and ventral 
I'ilis of fishes still needs a definitive ves or no answer. 



]})5!) CKKA'ICAL HIBS I X 'ITHTLKS H 

LITKRATURE (MTKD 

Abel, O. 

1909. Cotaceenstudioii. I Mitt(>ilnn{j. Dns Sk(>lc'tt von Kiiriiinodrlphis 
coclieteuxi aiis den 01)0iniioziin voii Aiitweipcii. SitzlitM-. Akiid. 
Wiss. Wien, Math.-Nat. Kl., 118 (1): 241-253. 

Al.HKlX'llT, I'. 

1882. Note sur uno sixienie costoide cervical clicz lui jcuiu' Hippopota- 
iiiiis tinipliihiiis. Bull. .\rus. Roy. Hist. Nat. linuxellcs, 1: 197-202. 

B.\uR, G. 

1886. The ril).s of Hphenodnn (Hofferia) . Anier. Nat., 20: 979-202. 

i)F Ht'RLBn', H. M. 

I!n7. BeitrJige ziii- Eiitwieklungsgeschiclitc dcr Wiilielsiiulc der Ceta- 
ceen. :\rori>h. .Talnl)., 50: .",73-402. 

I'^jrKLIANOV, S. M. 

1937. Die Morpliologie der T('tiaitndciiii])iieii. Zool. .T.-ihib. (A))t. 
Anat.), 62: 173-274. 

Fkderow, Y. 

1910. Zwei FJille der .selteiieii Bildiing vou Qiierfortsjitzen des ersten 
Brustwirl)els. Anat. Anz., 36: 556-500. 

Hayek, H. 

1928. Uber Querfortsatze und Rippenrudiinente in der Hals luid J.en- 
densegmenten. ]Mori)h. .Tahrb., 60: 371-416. 

Hoffman, C. K. 

1879. Over het voorkonien van lialriblien by d(> s(diild])adden. Yersl. 

Medel. kon. Akad. Wet., 14: .12-67. 
1890. Seliildkroteu. In II. G. Bronn — Klassen und Oi-dnungen des 
Thierreichs. Leipzig, 442 pp. 

Howes, G. B. and H. H. Bwinnerton 

1901. On the development of the skeleton of the tuatara, Sphcnodon 
pimctdtus, with remarks on the egg, on the hatching and on 
hatched yonng. Trans. Zool. Soe. London, 16: l-7(i. 

Jaekel, O. 

1916. Die \\'irbeltiere aus dem Kenper von Hall)erstadt. Serie II. 
Testndinata. Palaeont. Zeitsehr., 2: 88-214. 



12 BREVIORA No. 101 

Naef, a. 

19'2':K Notizen zur Moiphologie unci Stamniesgesehiehte der Wirbeltiere. 
15. Dreissig Thesen ii))cr Wii-lielsiiiile und Rippen iiisbesondeiT 
bei den Tetrapoden. Zool. .Tahil). (Abt. Anat.), 50: 581-6n0. 

Noble, G. K. 

1921. TIk' bony structure and ]ih.\ietic relations of Sphaerodactylus 
and allied lacertilian genera with a desci'iption of a new genus. 
Anier. ]\[us. Novitates, 4: 1-1 (i. 

RUCKES, H. 

1929. Studies in elielonian osteology. Part II. The morphological rela- 
tionships lietween the girdles, ribs and carapace. Ann. New York 
Acad. Sci., 31: 81-120. 

HcHixz, 11. R. AXi) R. Zangerl 

1937a. Uber die Osteogene des Bkelettes beini Haushulin, liei der 
Haustaube und beim llaubensteissfuss. ^forph. .Tahrb., 80: 
()20-628. 
19371;. Beitriige zur (Jsteogenese des Knochensystenis beini Haushuhn, 
bei der Haustaube, und l)eini llaubensteissfuss. Eine ver- 
gleichende osteologische Studie. Denkschr. Schweiz. Naturf. Ges., 
72 (Al)h. 2): 117-16.-1. 

SlEBENROCK, F. 

190(i. Schildkroten von Ostafrika und Madagascar. In Voeltzkow, 
Heise in Ostafrika in den Jahren 1903-1'90.5, 2: 1 -lU. 

1907. Uber einige zum Teil selteue Schildkroten aus Siidchina. Sitzber. 
Akad. Wiss. Wien, Math-Nat. Kb, 116 (1): 1741-1775. 

Thomson, J. S. 

1932. The anatomy of the tortoise. Sci. Proc. Roy. Dublin Soc, 20: 
359-461. 

VlRCHOW, II. 

1929. Die tiefen Riickenmuskeln des ()niilli<irlnn\cii.^. ^birpli. .lahrb., 
60: 481-559. 

V(")LKER, H. 

1913. Uber das Stannn-, (iliedmassen- und Ilaut- skelet von Dermo- 
cliiljis coridCKi I.. Zool. .lahrb. (Alit. Anat.), 33: 431-552. 





fc % 




jLiiJ^* 



''WW.'' 



PLATE 



Eiin/s orhictihiri.s embryo. A section tlirouyli tlie anterior end of tlie 2nd 
■ei'vical vevteliiTi in an advanced cartilage stag'c. Compare Fig. 1. 



BREVIORA 

Mmsenann of Comparsitive Zoology 



Cambridge, JVIasc;. April 9, 1959 Number 102 



A XEW JAMAICAN GALLIWASP (SAURIA, ANGUIDAE) 

By Garth Underwood 

Department of Zoology, University College of the West Indies, Jamaica 

The aiiguid lizards whose Jamaican vernacular name, "galli- 
wasp, " has gained some currency, are generally distributed in 
Central America and the Greater Antilles. Five distinct species 
are already known from Jamaica (Grant 1940, Cousens 1956). 
In this paper I describe the sixth Jamaican species. As a resident 
in this island I regret that 1 am obliged to describe this species 
on the basis of a single specimen ; however, after several years 
no further material has been secured. As the species is not 
merely very distinct but also represents a species group new to 
Jamaica 1 delay its description no longer as it widens the scope 
of known galliwasp radiation as well as reminding us of the 
gaps which remain in our knowledge. 

In May 1952 Mt. R. P. Bengry and Mr. G. R. Proctor, 3rd. of 
the Institute of Jamaica, were engaged in general collecting, 
zoological and botanical, on the north coast of Jamaica in the 
parish of St. Mary. In turning over a pile of coconut trash at 
Boscobel they captured a number of specimens of the common 
galliwasp of the lowlands, Diploglossus crusculus; Avith these 
galliwasps was the specimen here described. A number of at- 
tempts to find further specimens in the same area have failed ; 
1 can only suppose that this individual had wandered from its 
customary habitat, which remains unknown. The coast here backs 
onto limestone hills which are covered by dry scrub forest. 

A description l)asL'd upon a single specimen unavoidably con- 
tains tedious detail. By making comparisons with its nearest 
relatives I have endeavoured to recognise features significant in 



Z BBEVIORA 

(lefiiiint>' the species. With more material it will probably be 
possible to simplify and improve the description. 

I do not follow Burt and Burt (1931) and Taylor (1956) in 
maintaining i\ distinction between the genera Diploglossus and 
Celestus. After examining material of 22 different species I 
cannot see good grounds for supposing that there are two stocks 
of galliwasps which merit generic rank, and I therefore use 
the older name Diploglossus. 

As the generic name is compounded from Greek roots, I form 
the specific name likewise. To draw attention to what I believe 
to be its most striking single feature I call it : 

Diploglossus microblepharis sp. nov. 

Type and only known specimen. M.C.Z. 55764; collected by 
R. P. Bengry and G. R. Proctor, 3rd. May 1952, Boscobel, St. 
Mary, Jamaica, West Indies, 18°24'N, 76°58'W. 

Diagnosis. A plump, short-limbed galliwasp with small palpe- 
bral aperture, nasal shield excluded from rostral, frontal broader 
than long, very pronounced median keels on proximal caudal 
scales, few scale-organs on dorsal scales (less than one per scale). 

Description. I believe this form to be most closely related 
to D. delasagra of Cuba and D. pleeii of Puerto Rico ; the de- 
scription has been prepared with specimens of these species 
before me. A number of the features of the head squamation 
appear to be related to the reduction of the size of the palpebral 
aperture; these are nuirked with an asterisk; in some cases modi- 
fication of the squamation is asymmetrical. To make the descrip- 
tion easier to follow T have italicised those characters in respect 
of which the new species differs from both of the related species. 
Where a count can be made on two sides T have separated the 
left hand count from the right hand count by a comma. Plead 
broad; width: length = 0.7!) :1. Rostral twice as broad as high, 
in contact with supranasals and 1st labial. Three supranasals 
(asymmetrical), right meets frontonasal, 1st loreal, nasal, rostral 
and in midline left supranasal (also in j)Jccii) ; the two left 
supranasals together apparently represent a divided scale similar 
to the right supranasal ; together the two scales make the same 
contacts as the right supranasal. Nasal meets 1st and 2nd labials, 



NEW JAMAICAN UALLIWASI' 



;j 



1st loreal and suprauasal. Prefrontal about as long as fronto- 
nasals, contact with 2nd loreal and 1st supraorbital, ah art contact 
with 3rd loreal. \'ery broad frontal, width greater than lengtii, 
broad contact between frontal and parietals (also in pleeii). 
Four supraorbitals* , 1st much the largest. Third loreal meets 
1st supraorbital, not separated by intercalary scale*. Preocular 




Fig. 1. Dorsal view, bond of Biplofflnssnn microhlppJiaris type. 



meets 1st supraciliary, 1st supraorliital, ^rd loreal, 5th supra- 
labial and subocular*. Subocular meets 5th and 6th labials (also 
in pleeii). Three supraciliaries. Postorhitals 3,2*. Postocular ap- 
pears as last member of partly concealed row below supra- 
ciliaries. Three temporals between f ronto-parietal and labials : 
upper temporal separated from parietal ; lower temporal meets 
labials 6 and 7 and nearly enters margin of orbit between sub- 
ocular and lower postorbital*. Single nasal between supranasal, 
loreal 1 and labials 1 and 2. Three loreals; 1st low, elongated, 
between supranasal, nasal, labials 2 and 3 and loreal 2 (also in 
pleeii) ; 2nd loreal scpiarish, between frontonasal, prefrontal, 



4 BREVIORA 

loreals 1 and 3 and labials 3 and 4; 3rd loreal squarish, between 
2nd loreal, prefrontal, supraorbital 1*, preocular and labials 4 
and 5. No scale between 3rd loreal and prefrontal*. Eight supra- 
labials (also in pleeii). Small mental, half width of rostral; 7 
infralabials, prominent postmental in contact with labials 1 and 
2, followed by 4 pairs of chinshields (also in pleeii). 1st pair 
meeting in midline. 





Fig. '1. A. Scales aiuund left eye of Diploglo.s,su.s inicroblepharin type. 
B. Scales around right eye of Diploglossus pleeii. 



Head length (snout to posterior margin of earhole) 14 mm., 
width 11 mm. Palpebral aperture about 1 mm. long, ear-opening 
about i/> mm. in diameter. Palpebral marginal scales reduced, 
lower palpebral scales very thin. Pupil circular. 

Middorsal scales with 16-20 parallel striae and a j^ronounced 
central keel, caudal end of most striae broken. Accentuation of 
central keel fades out and interruption of sti-iae increases on 
lateral doi-sal scales; striae fade out altogetlier on Hanks, the 
vcnti-al scales being smooth and fiat. Towards shouhlers central 



NEW JAMAICAN GALLIWASP 5 

keels fade, and on neek striae fade out; towards rump central 
keels become very pronounced, and keels become so pronounced 
on proximal balf of tail as to fj:ive it, dorsally, the appearance 
of a polygonal prism. Towards tip of tail keels fade out. \'entral 
caudal scales smooth and flat. 

Scale organs on head extend back dorsally about as far as 
occipital scale, ventrally about as far as 15th postmental scale ; 
scale organs very few on back (less than 1 per scale), absent on 
flank save before groin, absent on belly ; a few scale-organs on 
outer face of thigh; scale-organs present on back and sides of 
proximal tapering portion of tail, aggregated on the mesial 
(dorsal) side of the central keel of each scale. Fourteen scales 
from a middorsal row gave the following scale-organ counts : 
011010010 3 1100 (mean 0.64) ; 4 groin scales gave 
the counts : 2 8 1 1. Scale-rows at midbody 43, mental to vent 
109 scales, occipital to point above vent 105, from anal to chest 
(on line joining axillae) 72, some union of dorsal scale-rows over 
rump, at level of ankles 20 scale-rows round tail. 

Third digit of forelimb slightly longer than 4th ; lamellae, 3rd 
digit 7, 8, 4th digit 7, 7. Scales on outer face of upper arm with 
faint striae (no central keels), scales of forearm quite smooth, 
scales of inner face of forelimb with striae. Ilindlimb : length of 
4th toe to length of 3rd toe as 1.4 to 1 ; 4th toe lamellae 11, 12 ; 
scales of outer face of thigh with striae ; scales of shank also 
with central keels; scales of inner face of hindlimb with faint 
striae but no keels. 

Cieneral colouration medium brown. Top of head uniform 
brown ; upper labials white with brown spots, lower labials with 
a few spots only ; throat white ; neck and back speckled wdtli 
brown, speckles on back loosely arranged in about 20 backwardly 
pointing chevrons ; speckling fades out fairly abruptl.y on sides, 
above ear-hole, on flanks and above groin ; speckling less regular 
on tail ; upper side of forelimb and outer face of forearm and 
hand closely speckled ; upper side of liindlimb and outer face 
of shank and foot speckled, inner face of thigh darker brown, 
some brown onto sole of foot. 

Dimensions. Snout -vent 87 mm., tail (tii) regenerated) 83 
mm., forelimb 13 mm., hindlimb 18 mm., axilla-groin 58 ram. 



b BBEVIORA 

Discussion. Burt and Biirt (19;U) make an explicit distinetion 
between Diploglossus and Celestus. Their discussion begs the 
(juestion as to wliether or not there are two natural groups which 
might properly be regarded as genera; they seek only a formal 
basis for their discrimination. They overlook the fact that two 
West Indian species of '' Celestus" have claw sheaths and there- 
fore belong to Diploglossus as they define it. This mistake ap- 
pears to be due to the fact that Boulenger (1885) did not have 
material of delasagra or plceii. He implied that they do not have 






Fig. 3. A. Lateral view of 4111 toe of Dijyloglo.ssus nticrohlcphuiit: type 
B. Lateral view of ith toe of Diploglossus plceii. (Claw is coin-enlcd liy 
sheath.) C. Lateral view of 4th toe of Diploglossus Jicicarrlii. 



slieathed claws by ])lacing them with the other island forms in 
section 11 of his key : "No large ungual sheath ; frontal anteriorly 
in contact with a single shield." 1 am very doubtful of the state- 
ment by Burt and Burt that the claws of the forms with a sheatii 
are "semi-retractile to retractile"; I see no indication of mobil- 
ity. 1 checked a series of species in respect of : claw sheaths, 
number of prefrontal shields, nasal-rostral contact and scale- 



NEW JAMAICAN (iAI.Ll VVASl' 7 

organs on tlie dorsal scales. The results are summarised in the 
accompanying table; I have included the Ilispaniolan endemic 
genera Saurcsia and Wetmorena. 

We see that there are four Antillean forms which have 
sheathed claws and conversely two mainland forms with naked 
claws. As is recognised by Burt and Burt and by Taylor the 
number of prefrontals is not correlated with the claw-sheath 
condition. The character of three prefrontals is however confined 
to the mainland (and Malpelo) species, with the interesting ex- 
ception of darlingtoni from the mountains of Hispaniola. Of the 
three M.C.Z. specimens, one, the type, has 3 prefrontals, the 
other two have only one (Cochran, 1941, states that the type 
has only one). Contact between nasal and rostral scales is shown 
by all the mainland forms but also by two Antillean forms 
{delasagra and pleeii). 

The scale-organs of the trunk call for more detailed comment. 
These organs do not appear to have been noticed in the Angui- 
dae; McDowell and Bogert (1954) state that they are not pres- 
ent; Scortecci (1940) did not find them. I have, however, found 
them, at least on the head, on every anguid examined. They were 
checked by prying off the horny scales with a needle and mount- 
ing in polyvinyl alcohol. The preparations are in the Museum 
of Comparative Zoology. I am not yet in a position to offer a 
critical appraisal of this character. A cursory comparison of the 
scale organs on various parts of the body of several species indi- 
cated some striking differences in respect of their occurrence on 
the dorsal scales ; representatives of the available species were 
therefore checked for this character. The scale-organs are visible 
as thin refracting spots in the horny scales. On the trunk and 
tail scales they are localised in two areas on either side of 
the midline and nearer to the posterior margin. Commonly there 
are scale-organs on both sides of the midline of a scale. Counting 
the scale-organs separately for each side of a longitudinal row 
there is sometimes a significant asymmetry ; the higher number 
of scale-organs is in such cases found on the side nearer to the 
mid-dorsal line, i.e. the mesial side. The scales of the flanks and 
the sides of the tail usually have all the scale-organs on their 
dorsal sides, i.e. the sides nearer the dorsal midline. I have not 
found scale-organs on belly scales nor on ventral tail scales. 



8 



BREVIORA 



Most of the dorsal scale-organ counts were made on a row of 
about ten dorsal scales taken from near the dorsal midline. To 
form an idea of the reliability of such counts I mounted pieces 
of dorsal slough from tAvo Jamaican D. criisculus. Each lateral 






Fig. 4. Scales and scale-organs in Diploglossus microblepharis. A. Mid- 
dorsal scale to show interrupted striae. B. Scale from right side of tail near 
base to show size and position of scale-organs (reversed under microscope). 
C. Right fronto-nasal to show distribution of scale-organs. 



dorsal scale-row is clearly indicated in this species by a break in 
the colour pattern. Including the lateral dorsal rows one slough 
had a complete block of 196 scales (14 rows of 14) and the 
other of 403 scales (13 rows of 31). Counts of scale-organs were 



NEW JAMAICAN GALLIWASP 9 

made on all of these. On tlic outer three rows of scales 97.8 
per cent of the scale-organs lay on the inner side. The scale- 
organs were less asymmetrically distrilmted on the inner rows of 
scales; there were however more on the inner side of each scale- 
row than on the outer side. The reversal of the asymmetry did 
not take place about the midline, however, but between the 2nd 
and 3rd rows to the left of the midline. The lateral dorsal scale- 
rows had rather high scale-organ counts ; the other rows differed 
from one another in apparently random fashion, but a difference 
between two rows would be maintained throughout their lengths 
(not scale by scale but for groups of scales large enough to 
smooth out random variations). One specimen had the highest 
count (3.0 organs per scale) for the 5th row, right of centre, and 
tiie lowest counts (2.2 organs/scale) for the 2nd, 3rd and 4th 
rows, right of centre. The other s})ecimen had the highest count 
(5.6 organs/scale) for the centre row and the lowest (4.2 
organs/scale) for the 3rd row, right of centre. Discounting the 
lateral dorsal scale-rows it was possible to make a count of 10 
consecutive scales in 60 different ways for one specimen ; the 
lowest total was 20, the highest 33, mean 26.25, standard varia- 
tion 11.0 per cent of mean. There were some irregularities in 
the squamation of the other specimen; avoiding these, 222 such 
counts were possible; the lowest total w^as 31, the highest 67, 
the mean 51.49, standard variation 15.7 per cent of mean. 1 tried 
some counts of 10 scales in a transverse row and 10 scales 
diagonally but the results did not suggest that these would be 
more reliable. 1 have not eliminated the possibility that the 
counts may be correlated with size or with sex. 

All of the mainland species lacked dorsal scale-organs save 
monolyopis (23 scales from two specimens of the latter averaged 
4.] organs/scale). A group of Antillean species (dclasagra, 
pleeii and darlingto7ii) likewise lacks dorsal scale-organs; micro- 
blepharis is intermediate in that many scales are without organs. 
It is noteworthy that SSauresia and W etynorena both are in agree- 
ment with the majority of insular forms in the possession of 
dorsal scale -organs. 

Preliminary examination of a number of forms suggests that 
these scale-organs may have value as a systematic character at the 
species level. There may be differences in the number of organs 



10 BREVIORA 

per dorsal scale ; when the organs are absent from the back they 
may or may not also be absent from tlie flanks. The number of 
organs on the scales of the groin may also be useful. For 
example, the type of Panolopus cost at us Cope has a dorsal 
average of 2.2 organs/scale (10 scales), the type of Diploglossus 
stenurus Cope (listed as a synonym of costatus by Cochran) 
has a dorsal average of 5.0 organs/scale (16 scales) ; to this is 
added a mean of 7.4 organs/groin scale for costatus (5 scales) 
and 17.4 organs/groin scale for stenurus (5 scales). When it is 
noted also that the adpressed limbs of costatus fail to meet by 
about 15 scales whereas those of stenurus overlap by about 5 
scales, there seems to be little question that stenurus is a distinct 
form. D. eymeagrammus is noteworthy for the fact that each 
groin scale bears a single very large organ. Differences were 
also noted in the distribution of the scale-organs on the occipital 
scale. For example, the Jamaican D. hewardi has scale-organs 
scattered around the jiosterior margin of the occipital whereas 
D. crusculus and D. harbour i have the scale-organs generally 
scattered over the occipital. 

Purveying the distribution of variant characters amongst the 
different forms, 1 cannot arrive at any firm ideas concerning the 
species groups. However, to provide a basis for discussion I put 
forward a scheme which accords reasonably well with the evi- 
dence. I propose to regard monotropis as primitive in respect of : 
the presence of dorsal scale-organs ; the presence of sheathed 
claws; the contact between nasal and i-ostral scales; the presence 
of 3 prefrontal scales and the stoutness of the limbs. The general 
occurrence of scale-organs on the trunk of lizards of various 
families suggests that this is a i)rimitive condition. As with 
the teeth, however, we cannot assume that the onlj^ changes in 
phylogeny have been reductions in number, so we must be pre- 
pared to allow re-extension of scale-organs onto the back if other 
evidence requires it. Conditions in lizards generally suggest 
that many small scales are more primitive than few large scales ; 
on these grounds I suppose three prefrontals to be more primitive 
than one. In the case of claw-sheaths we have either to suppose 
that they have arisen within the genus several times or that they 
have been lost several times; as a working hypothesis 1 prefer 
the second proposition. As regards the pi'oposition that the 



NEW JAMAICAN GALLIWASP 11 

nasal-rostral oontact is primitive, 1 can only ai-<jru(' that this 
liypothesis ai)i)ears to tit the otlier assumptions better. That stout 
limbs are more i)riniitive than weak limbs is not likely to be 
(juestioiied. In South America fasciatus and resplendens re- 
semble monotropis but lack the dorsal scale-organs; Ophiodes 
nuiy be derived from this branch. On Malpelo, )nill( punctatus 
likewise resembles >no)i()t)-opi,s but has the claw-sheath reduced 
as well as lacking- dorsal scale-organs. In northern Central 
America enneagrammus and vijatiochloris superficially resemble 
such Antillean forms as crusadus and costatus; however tliev 
retain a nasal-rostral contact and lack dorsal scale-organs. The 
three species delasagra, plecii and microhlepharis do appear to 
represent a natural group ; even here we have some complexity 
since uiicroMepharis, presumably primitive in letaining a few 
dorsal scale-organs, has lost the nasal-rostral contact and partly 
lost the claw-sheath retained l)y the other two species. It seems 
clear therefore that microhlepharis cannot be derived from 
delasagra or pleeii, nor can the converse view be entertained. 
The delasagra-pleeii and microhlepharis stocks may be separately 
derived from a Central American ancestral stock. On the evi- 
dence of the distribution of mammals, Darlington (lf)57, p. 510) 
suggests that Cuba and Jamaica represent two ports of entry for 
Central American stocks in the Am'illes. This group must be 
surely represented by an undiscovered torm in Ilispaniola. The 
remaining Antillean species pei-hajis form a natural group : 
darlingtoni is the most isolated form. On the grounds that they 
retain claw-sheaths I suppose Sauresia and Wetmorena to be 
early offshoots of the Antillean stock. [ cannot see good grounds 
for recognising the genus SaiD'esia as distinct from Diploglossus. 
The disappearance of one digit, small in a number of other forms, 
is a trivial feature Avhich cannot be of much adaptive sig- 
nificance; the species is not set otf in other respects. Wetinorena, 
on the other hand, is clearly set apart by the loss of the ear- 
opening, a feature of manifest adaptive significance in a burrow- 
ing animal. 

The forms so far discussed all belong to McDowell and Bogert's 
snbfamily Diploglossinae. All of them have numerous scale, 
organs on the head although they ma}- be i-educed on the trunk. 
A few representatives of their subfamily Gerrhonotinae were 



12 BREVIORA 

cursorily examined. No scale-organs at all were found on the 
trunk and only a fe^v on the anterior part of head. A loreal and 
3rd supralabial scale -were exaniijied of one specimen each of : 
Gerrhonotus liocephahis, Bariasia imhricutu, Elgaria mtdtica- 
rinata, Ahronia deppii and Ophisaurus apus. Elgaria most 
closely approached the Diploglossinae in the partially sheathed 
claws, short nasal-rostral contact and the possession of more 
loreal scale-organs than the others. The three prefrontals of 
Ahronia resemble those of DiploglossKs. The loreal and labial of 
Gerrhonotus had only one scale-organ each. The Barissia had 
none on the loreal and four on the labial. Ophisaurus resembled 
the others in the absence of scale-organs from the trunk and 
posterior part of head and in the small number on the anterior 
part of head. Angnis fragilis is similar to Ophisaurus in the 
possession of a few scale-organs on the anterior head only. 

I am indebted to Mr. C. Bernard Lewis, Director of the Insti- 
tute of Jamaica, who generously turned this specimen over to 
me for study. I am grateful to Dr. E. B. Williams for access to 
the collection of the Museum of Comparative Zoology and for 
the benefit of discussion. Dr. R. Zweifel kindly allowed me to 
study some additional specimens in the American Museum of 
Natural History. 

LITERATURE CITED 

BOULENGEK, E. G. 

1885. Catalogue of the lizards in the British Museum. London. 

Burt, C. E. and M. D. Burt 

1931. South American lizards in the collection of the American 
Museum of Natural History. Bull. Amer. ]\Ius. Nat. Hist., 61: 
227-395. 

Cochran, D. M. 

1941. The herpetology of Ilispaniola. Bull. U. S. Nat. Mus., 177:1-398. 

CousENS, p. N. 

1956. Notes on Jamaican and Cayman Island lizards of the genus 
Celestiis. Breviora, Mus. Comp. Zool., No. 56:1-6. 

Darlington, P. J. 

1957. The geographical distribution of animals. New York, 675 pp. 



NEW JAMAICAN GALLIWASP 



13 



Grant, C. 

1040. The reptiles in Lynn nnd Grant: Tlie licrpetology of .T.-iiiiiiica. 
Bull. Inst. Jamaica, Sci. Ser., 1:1-6"). 

.McDowell, S. B. and C. M. Bogert 

1954. The systematic })osition of Ldntlionotus and tlic alUiiitics of the 
anguinomorphan lizards. Bull. Anier. Mus. Nat. Tlist., 105:1-142. 

SCORTECCI, G. 

1940. Eecettori dcKli iKuauidi o di altri Sauri. Atti Roc. Ital. Sci. 
Nat., 79:1-10. 

Taylor, E. II. 

1956. A review of the lizards of Costa Rica. Sci. Bull. Univ. Kansas, 
38:1-322. 



Table 

Claw-sheath Prefrontal Xasal-rostral Scale-organs 

contact on back 



DipJoglossus 










fasciatus 


+ 


3 


+ 


— 


resplendens 


+ 


3 


+ 


— 


monotropis 


+ 


3 


+ 


+ 


millepunctatus 


% 


3 


+ 


— 


cyanochloris 


— 




+ 


— 


enneagrammus 


— 




+ 


— 


microhle pilaris 


¥2 




— 


few 


delasagra 


+ 




+ 


— 


pleeii 


+ 




+ 


— 


darlingtani 


— 


1 or 3 


— 


— 


hewardi 


— 




— 


+ 


ocdduu^ 


— 




— 


many 


duquesnayi 


— 




— 


+ 


crusculus 


— 




— 


+ 


barbouri 


— 




— 


+ 


hadius 


— 




— 


+ 


costat-u.s 


— 




— 


+ 


rugosus 


— 




— 


+ 


ciirtissi 


— 




— 


+ 


stenurus 


— 




— 


+ 


Sauresia 










sepsoides 


+ 




— 


+ 


W'ctmorena 










haetiana 


+ 




— 


+ 



BREVIORA 

MiLaseniim of Compsirsitive Zoology 



Camhridgk, Mass. April 10, 1959 Ximber 103 



TWO NEW SPECIP]S OF ELEUTHERODACTYLUS 
FROM PFERTO RICO 

By Juan A. Rivero 

Institute of Marine Biology and Department of Biology 

University of Puerto Eico 

Mayag'iiez, Puerto Eico 

Some years ago Dr. J. A. Ramos of the Biology Department 
of the University of Pnerto Rico collected, at the Oamhalache 
Forests, a small frog that differed strikingly from the Elcnthcro- 
dactylus crampioni collected at the same locality and date, by 
the very short length of the fingers and relatively short, thick 
liiiid limbs. The specimen was said to have been collected in 
bromeliads l)nt no voice record was taken, so that the description 
of the apparently new species was postponed nntil more para- 
types could be obtained and the voice recorded. So far, four 
more trips have been made to the same locality, at night and 
during the daytime, but all of them have been unsuccessful, 
although a strange voice that could have been of frog or insect 
was heard on several occasions. The Cambalache Forests are 
of the semiarid, deciduous type, and spiny trees and bushes are 
common, so that catching a calling frog at night is not an easy 
matter. 

More recently, another apparently new frog was collected at 
Dona Juana Forests and, although this first collection consisted 
of 40 individuals, the animal was not heard to produce any voice 
in spite of the fact that all the specimens were caught at night 
and enough time was given each one to call. The description of 
this form was likewise postponed and, in the month following 
the first collection, 16 other individuals of the same species were 
collected at El Yunque, on the other end of the island. No voice 



2 BREVIORA No. 103 

was heard to come from any of these animals, so that it was tenta- 
tively concluded that the species is nuite. Since well developed 
gonads are fonnd in both sexes, the lack of voice cannot be at- 
tributed to immaturity, especially if it is considered that frogs 
of the genus Eleuthcrodactylus seem to call throughout the year 
in Puerto Rico. 

Descriptions of both species have been prepared in the belief 
that they should be recorded and that their jniblication may 
be of help in determining distribution and habits. 

The author wishes to express his appreciation to Mr. Jorge 
Rivera Lopez, to Dr. Ruth Turner, and to his wife, Mrs. Eneida 
B. Rivero, who helped him collect in the field. 

Eleutherodactylus ramosi sp. n. 

Type. Museum of Comparative Zoology No. 30428, a 9 from 
Cambalache Forests, n. Puerto Rico, Coll. J. A. Ramos, Aug. 
1952. 

Diagnosis. A small Eleutherodactylus with uniform colora- 
tion, very short fingers and short limbs, the heel and elbow of 
which do not meet when adpressed along the side of the body. 

Description. Head broader than long ; snout subovoid ; tongue 
large, % free and nicked behind ; vomerine odontoids in two 
strong oblique series behind and between the choanae ; eyes of 
moderate size, their diameter equal to distance between eye and 
nostril ; interorbital space broader than an upper eyelid ; canthus 
rounded ; loreal little inclined, not concave ; tympanum moder- 
ately distinct, covered above by a supratympanic fold, V2 the 
eye diameter ; fingers strikingly short, the longest not longer 
than the orbital diameter ; no definite palmar tubercles ; larger 
disks smaller than the tympanum ; no apparent metatarsal 
tubercles ; toes free, the first well separated from the others ; 
disks of toes smaller than those of fingers ; heel of the adpressed 
hind limb extends to the posterior margin of the tympanum ; 
heels overlap when legs are placed at right angles to the body; 
heel and elbow do not meet when adpressed along the flank. 

Skin above, smooth. Ventral surfaces smooth except on the 
posterior part of the belly, where slight rugosities or granula- 
tions can be seen under a lens ; posteroventral aspect of thighs, 
granular. 



VJdU 



NEW FROGS FROM PUERTO RICO 



Color. Above, unifonn brown, with iniimte dots (melano- 
phores) iindor a lens; no ai)parent. eantlial or supratympanic 
streaks. Below, infuseated white, the nielan()i)liores being scat- 
tered and sparser than on the npper surfaces. 

MiasurcDunts. (nun.) $ snout-vent 22; head length 7.3; 
head breadth 8.3 ; femur 8.2 ; tibia 9.6. 




Fig. 1. Eleutherodactyhis ramosi Type, MCZ 30428 (M. Estey del). 



Remarks. E. ramosi is most closely related to E. cramptoni 
with which it occurs at the Cambalache Forests. It differs from 
that species in its verj- short fingers, smaller disks, broader head 
and shorter and thicker hind limbs, the heel of which does not 
even approximate the elbow when both anterior and posterior 
limbs are adpressed alongside the flank. It was the very short 
fingers and thick hind limbs that made the author describe this 
species on the basis of only one specimen. 



4 BREVIOKA No. 103 

BlEUTHERODACTYLUS ENEIDAE Sp. 11. 

Type. Museum of Comparative Zoology Xo. 30-129, a 9 from 
Dona Juaiia Forests, \'illalba, Puerto Kico. Coll. J. A. and E. B. 
Rivero, August 1956. 

Diagnosis. A medium sized Eleutherodactylus with rounded 
canthus rostralis, interorbital space as broad as an upper eyelid, 
tubercular dorsum, barely overlapping heels, mottled and 
variegated dorsal coloration and immaculate ventral surfaces. 

Description. Head slightly l)roader than long; snout short 
subovoid ; tongue oval, ^ ^o free and nicked behind ; vomerine 
odontoids in two oblicpie series behind the small, oval choanae, 
their external extremities commencing at level with the vertical 
of the center of the latter ; canthus rounded ; loreal sloping ; inter- 
orbital space as broad as an upper eyelid ; eye diameter greater 
than distance between eye and nostril, almost as long as the 
snout ; tympanum small, not too distinct, ^ the eye diameter ; 
first finger shorter than second, which is longer than last ; three 
metacarpal tubercles and two smaller ones in line behind the 
wrist, just in back of the central metacarpal tubercle ; disks fan 
shaped, broader than the t^'iiipanum ; two small metatarsal 
tubercles ; subarticular tubercles distinct ; toes free, their disks 
slightly smaller than those of the fingers : heel of the adpressed 
hind limb extends to the posterior corner of the eye ; heels barely 
overlap when tibiae are placed at right angles to the body. 

Skin above, studded with small, evenly distributed tubercles; 
eyelids tubercular ; a small tubercle at the heel. Below, granular 
on the belly and thighs, very slightly granular on the throat; a 
ventral discoidal fold. Male with a slight fold on each side of 
the throat. 

Color. Above, light gray with blotches and mottles of darker 
gray or l)rownish gray ; eyelids darker than the body color; loreal 
area whitish gray, witli small darker spots along the upper lip; 
limbs light gray, with l)i-owiiish gray striations and mottling; 
loins with a golden brown stain ; posterior aspect of the thighs 
with a diffuse, irregularly margined brownish marking on the 
distal end. ^"entral surfaces immaculate, powdered with brown, 
except for a thin gray rim along the lower margin of the li]). 

Measurements, (mm.) 9 snout-vent 26.2; head length 10.5; 
head breadth 10.5; femur 11.9; til)ia 12.6. 



1IJ5!> NEW FROtJS FKOM I'l'EKIO RRO 5 

Re)it(n-hs. Paratypos include M!) specimens from Dona . I nana 
cloud forests, with tiie same data as the type (IMMv. (Jll 12. 
614-51) and 16 from El Yiuuiiie (U.P.K. 5!):)-6l(). El Ynn(iue, 
sixth i-ain shelter to Pinnacles, 1 Sept. Ifi56). All were collected 
at iiig:ht on the floor or on the leaves or trunks of busiies, palms 
or tree ferns at not more than 5 or 6 feet from the ground. 

The largest of the 54 paratypes is only 2 nnn. larger than 
the average sized type, an indication of a very uniform size for 
fully grown individuals. This is contrary to the situation existing 
in Elcuthcrodacff/l us porioriccuxis where considerable variabil- 
ity is found in any collection. 

The following variations are found among the paratypes of 
FA( >iih( rodaciyhiH encidac : 

1. In poorly preserved specimens, a slight canthus may be 
apparent, but ordinarily the snout is rounded in cross-section and 
there is no angle at the canthal margin. This character easily 
differentiates this species from E. portoricensis, its closest Puerto 
Rican ally. 

2. The heel of the adpressed hind limb may extend to any 
point on the eye (posterior corner, middle or anterior corner). 

3. The golden brown stain of tlie loins is present in most but 
not all individuals. 

4. A small, black spot or short streak is generally found above 
the tympanum. 

5. The dorsum is always mottled and variegated but the 
ground color may be lighter or darker than in the type. 

Eleuihcrodaciylus cneidae is most similar to E. portoricensis 
from which it can be distinguished by its more uniform, adult 
size, rounded canthus not bordered by a white line (as in most 
E. portoricensis) , absence in most specimens of a well defined 
interorbital bar ; smaller tympanum ; dark eyelids ; narrower 
interorbital space (much broader than the upper eyelids in E. 
portoricensis), more tubercular dorsum, different coloration and 
])r()l able absence of voice. 

Some examples of E. porioricensis are also mottled above, but 
the mottling is usually brown, and the well defined canthus, 
broader interorbital space and persistent interorbital bar make 
the species easily distinguishable. It can also be said that if a 
specimen is more than 80 nun. snout-vent length, it is not 
E. eneidae. 



6 BREVIORA No. 103 

EleutkerodactyUis cneidae is quite common in the cloud forests 
of Doiia Juana and El Yunque and it is very strange that the 
species had not been named before. It appears possible, however, 
that collections of E. portoricensis may include this species. 
This may be the case, for example, with Schmidt's (1928:46) 
Figure lie, but this figure may just as well represent a color 
phase of E. portoricensis. 

E. eneidae is somewhat similar in coloration to E. flavescens 
from the Dominican Eepublic, but it differs from that species in 
many important details. It also looks somewhat like Schmidt's 
(op. cit.-.od) figure of E. locustus but Dr. R. G. Zweifel of the 
American Museum compared it with the type of that species and 
found that three specimens of E. cneidae "are very different in 
size and general appearance from locustus.'" Furthermore, 
Zweifel found very little difference between cramptoni and 
locnstus, and suggested that they might be synonyms. Males 
of E. crampto)ii have vocal sacs (described as absent by Schmidt) 
and the proportion of tibia and femur (heels cross in locustus, 
meet in cramptoni) , mentioned as a difference between the two 
species, does not, according to Zweifel, seem to exist. On the 
other hand, Schmidt described the voice of E. locustus as the 
"most distinctive of any noted in Puerto Rico," so that it is 
})erhaps better to keep the two forms separate until more field 
studies can be made. 

EEFEEENCES CITED 

Schmidt, K. P. 

1920. Contriljutions to the herpetology of Porto Kico. Ann. Xcw 
York Acad. Sei., vol. 28, pp. 167-200. 

1927. A new Eleutherodacti/lus from Porto Rico. Auier. Mus. 
Novitate.s, No. 279, pp. 1-3. 

1928. Amphibians and land reptiles of Porto Rico, with a list of 
those reported from the Virgin Islands. Sci. Survey Porto Rico 
and the Virgin Islands. (New York Acad. Sei.) Vol. 10, part 1, 
pp. 1-160. 



"^d^ 




FJeutherodactylus eneidae , Type, MCZ 30429 (J. A. Eivero phot.). 



BREVIORA 

MiaseiLinn of Comparative Zoology 



('A:\inKiDGE, i\lAss. April 13, 1959 Number 104 



STUDIES OX FISHES OF THE FAMILY OPHIDIIDAE. III. 
A XEW SPECIES OF LIJPOPHIDIUM FROM BARBADOS^ 

By C. Richard Robins 

The Marine Laboratory, University of Miami 

Anionof the eolleetions of The Museum of Comparative Zoology, 
Harvard University, are two eiisk eels collected off Barbados, 
British AVest Indies, during the operations of the Steamer 
"Blake." They represent a new species of Lepophidium which 
is described below. 

Lepophidium kallion, sp. nov. 
(Figure 1) 

Holotype. Museum of Comparative Zoology 28057 A, a female, 
117 mm. in standard length; collected in 200 fathoms "off' Bar- 
bados," "Blake" station 291, presumably in February or March, 
1879 (see Pierce and Patterson. 1879:15). 

Paratype. MCZ 28057B (1), a male, 162 mm. in standard 
length ; data as for holotype. 

Diagnosis. Dorsal rays 133, anal rays 108; pectoral rays 23 
or 24, gill rakers 10, vertebrae 74 (15 precaudal and 59 caudal 
vertebrae), pyloric caeca 6 (1 dorsally, 5 ventrally). Tapper 
sides with dark brown markings in two series, the upper form- 
ing a longitudinal stripe on either side of the dorsal-fin base. 
Snout tip fringed with numerous fleshy projections. 

Description. A healed injury at the caudal end of the para- 
type has resulted in the loss of about eight caudal vertebrae 
including the hypural plate. Counts of vertebrae and of rays in 
the vertical fins, listed in the diagnosis, are therefore based 

iCuntiilnitioii Xn. 221 from The Marine Laboratory. University of Miami. 



2 BREVIORA No. 104 

solely on the liolotype. Pectoral rays number 23-23 (holotype) 
or 23-24 (paratype), the caudal rays 9. Three rudimentary gill 
rakers are on the upper arm of the tirst arch of botli spi^'imens 
and 6 or 7 developed rakers are on the lower arm. The holotype 
has an additional rudimentary raker on the lower arm ; both 
specimens liavc a total of 10 gill rakers on the first arch. 
Branchiostegal rays number 7. The pyloric caeca are blunt, and 
number 6 (1 dorsally and 5 ventrally). 

L. kallion possesses the general characteristics of Lcpophidiinn 
(see Robins, 1959:361). 

Morphometric data follow (in each instance the figure in 
parenthesis refers to the paratype) : standard length, mm., ]]7 
(162-damaged) ; head length 23.0(34.6) ; tip of snout to origin 
of dorsal fin, 27.8 (39.3) ; tip of snout to origin of anal fin, 42.8 
(67.2) ; tij) of snout to occiput, 15.5(22.5) ; tip of snout to pos- 
terior end of maxillary, 11.1 (15.8) ; depth of body, at occiput. 
13.7 (23.2); at origin of dorsal tin, 13.0 (24.6), at origin of anal 
fin, 13.1 (20.7); length of pectoral fin. 11.4 (14.5); length of 
pelvic rays, 8.6 and 6.4 (12.4, 9.5) ; length of caudal fin, 7.1 
(damaged) ; tip of snout to posterior end of lateral line, 112 
(150) ; horizontal diameter of bony orbit, 3.5 (5.1) ; post-orbital 
length of head, 12.7 (19.5) ; snout length (to margin of orbit), 
4.2 (6.5) ; bony interorbital, 3.5 (5.1). 

Despite their age the distinctive color pattern is readily seen 
on the type specimens. The dorsal, caudal and anal fins are pale 
with scattered melanophores and with a well defined dark mar- 
gin. The pectoral fin is clear with some melanophores on its 
basal third. 

The body is almost straw-colored in preservative although the 
scattered melanophores lend a dusky appearance to the belly. 
Two series of dark brown markings on the upper sides dominate 
the body coloration. The uppermost series starts as a collar just 
behind the occiput, runs laterally to a point above the opercle 
and then extends as a dark stripe, just above the lateral line, 
most of the distance to the caudal-fin base (Fig. 1). It is broken 
posteriorly into a series of dashes. In the holotype the stripe is 
less continuous than in the paraty])e. The lower series of dark 
brown markings extends posteriorly from the shoulder as a series 
of elongate blotches, just below the lateral line. Caudally, the 
lower series crosses the lateral line and merges with the dorsal 



1 !*'")!» NEW SPECIES OF LEPOPIIIDIUM 3 

series. Two dark spots occur on the nape between tlic dorsal- 
fin origin and the occipital coHar. Tlic head is nfenerally dusky. 
A sliort dark bar extends from the upper end of the gill slit, 
reai'ward along the dorsal niai-gin of the operculum. The snout 
is not spotted but a few dai-k si)ots may have been present on 
top of the head, behind the interorbital region. As seen from 
above, the dark margin of the dorsal fin appears to be a mid- 
(h)rsal stripe. 

The floor of the mouth and the gill bars are pale and the gill 
chamber silvery with numerous melanophores. The roof of the 
mouth especially toward the pharynx is dusky but not black. 
The gastrointestinal ti-act is sooty black except for the stomach. 



Fisu:e 1. Laternl view of the paratype of Lepophidiiim l-allio-n, MCZ 
i;S0o7B, 162 nun. in standard length, eollected off Barbados, "Blake" 
station 291. 

As in other species of Lepophidium, the swimbladder is similar 
in both sexes, a rather thin-walled sac, with no posterior opening 
or tube. It occupies the dorsal portion of the body cavity. 

Numerous short, pointed, non-depressible teeth are present on 
the premaxillary, dentary, vomer, palatines, and pharyngo- 
branchials. Those on the outer row of the premaxillary and 
dentary are test developed. The dentigerous areas are papillose, 
especially the lips. A prominent spine is easily detected under 
the skin on the snout and on the upper posterior margin of the 
opercle. 

Scalation is typical of the genus. The scales are cycloid, imbri- 
cate and in regular rows on the body. The postorbit^l portion 
of the head is also scaled. In L. kallion the scales are imbedded 



BREVIORA 



No. 104 



and the skin must be scraped away to determine the number of 
scale rows. About 8 and 25 rows are present between the lateral 
line and the dorsal- and anal-fin origins, respeotively. Some 
165-170 rows cross the lateral line between the posterior edge of 
tile hypural plate and the point of opercular attachment. So 
many other characters are available to the systematist that the 
labor involved in approximate enumeration of scale rows in 
ophidiids seems unjustified. 





2mm 



lOmm 




d / 

0/ 



Figure 2. (a, b). Diagram of lateralis system on head of Lepopliidiuni 
kallion (see text for explanation), (e). Development of fleshy tal)S on tiji 
of snout of L. kalluin. (d). Tip of snout of L. mdrmnratum (Goode and 
Bean). 



The lower rim of the snout is incised anterior to the nostrils. 
Four fleshy tabs project from this margin (Fig. 2 c). The actual 
shajic of these tabs apparently varies, foi- tlie ])aratype is less 



1!)")!) M;\V SPECIES OF LEPOPHIDIIM 5 

(Iccoi-atcd tliaii the Iiolotype. Tavo rows of ficsliy tabs i-uii dor- 
sally from ill fi-oiit of the aiitoi"ioi' nostril an.l join just aiitcrioi' 
to the rostral spine { Fi<>'. 2 v) . 

IJohlke and Kobins (in j)ress ) have demons! i-atcd specific dif- 
ferences in the lateralis system on the head. In L. hallioti this 
system may l)e divided into the lateral, supratemporal, supra- 
orbital, infraorbital and })reopereulomandibnlar canals. Ilubbs 
and Camion (1935, pi. 2) provided illustrations of these canals 
in several darters and Robins and Miller (1957:216-217) simi- 
larly treated Cuttus. 

A sehematic presentation of the lateralis system of L. kallion 
is given in Figure 2 (a, b). In the lateral view the eanals are 
>hown in black. A single pore occurs on the lateral canal just 
anterior to the attachment of the gill membrane. Three pores 
(one median) are on short caudally-projecting side canals ex- 
tending from the supratemporal canal. Each supraorbital canal 
consists of six pores (one above and slightly behind the eye, one 
median coronal pore, two over the anterodorsal margin of the 
eye, one behind and above the anterior nostril and one opening 
caudally into the snbnasal depression. Each infraorbital canal 
consists of eight pores (one behind the eye, one behind and below 
the eye, one at the posterior end of the suborbital rim, four on 
the sul)orbital rim above the jaw% and one opening anteriorly 
into the subnasal depression. The preopereulomandibular canal 
is more difficult to trace on the type material of L. kallion, espe- 
cially on the sides of the head. Apparently it opens through 6 
pores. The four pores along the ramus of the lower jaw" are 
large and easily seen (Fig. 2 b). One tiny pore opens oft" a side 
canal just below the junction of the preopereulomandibular 
canal with the lateral canal. Another pore open., oft' a long side 
canal on the lower third of the operculum. In at least one other 
species of Lepophidium a second pore occurs below this one but 
it could not be found in L. kallion. 

The posterior nostril is nearly on the anterior rim of the orbit 
and is small and circular. The larger anterior nostril opens 
through a short tube, above the subnasal depression, near the 
anterior end of the snout. 



6 BREVIORA No. 104 

The name is from the CTi-eelv haJlion, meaning- more beautiful, 
and refers to the distinctive pattern of this speeies in contrast 
to the more drab species of Lcpophidiuni. 

Relatio)} ships. Serious consideration need be accorded only 
/.. nwrworaium (Goode and Bean). Other Atlantic species were 
l)riefly mentioned by Robins (1959:366-367). L. )nan))()rat}(iii 
differs from L. kail ion in many features. 1) The upper sides are 
spotted instead of striped. 2) The upper half of the head and 
the snout are spotted. 3) It possesses more gill rakers, usually 
11, occasionally 12 or 13. 4) The longer division of the pelvic 
ray is considerably longer than the pectoral fin. 5) Probably, 
there are fewer dorsal rays ( 121-129) and, 6) fewer pyloric caeca 
(3-5). 7) The snout is smooth, its lower rim entire (Fig. 2 d) 
without any of the numerous fleshy tabs of hallion. 8) A sub- 
nasal pore is present on the infraorbital canal just below the 
posterior nostril. 

L. kaJlion and L. marmoraium are intimate species within 
the framework of the genus. The color patterns of the two al- 
though quite different are of a similar nature, quite unlike that 
of L. staurophor Robins, J., jcannae Fowler or L. ccn'iiiuni 
(Goode and Bean). The pigmentation of the gut, peritoneum, 
and oral and branchial chambers are the same, a distinctive 
combination. The gill rakers are similar, those on the lower arm 
diminishing gradually in size instead of shifting from equally 
developed rakers to rudiments. 

More detailed consideration of morynordinin is beyond the 
scope of this paper. 

ACKNOWLEDGMENTS 

Thanks are due to the Museum of Comparative Zoology for the 
loan of specimens and to Dr. Henry B. Bigelow, Mr. William C. 
Schrocder and Mrs. Myvanwy Dick for courtesies extended to the 
writer during his visit to the Museum. Dr. (Jiles W. Mead of 
the Tnited States P^ish and AVildlife Service kindly provided 
x-rays of the type specimens of L. kail ion. Mr. Walter R. Courte- 
nay photographed the i)aratype. This study has been supported 
by a grant-in-aid from the National Science Foundation (NSF- 
CJ-7116) which 1 gi-atefnlly acknowledge. 



I!*.")!) XKW ST'KCIFS OF T.Ki'oiM nniT'^r 7 

LITERATURE CITED 

BoHLKE, James E. and C. RiciiARn Robins 

(in ]iress). Studies on fishes of the family Ophidiidae. II. Three new 
species from the Bahamas. Proc. Acad. Nat. Sci. Philadelphia. 

HuBBS, Carl L. and Mott Dwight Cannon 

1935. The darters of the genera Eololcpis and Villora. Misc. Pnhl. 
Univ. Mich. Mus. Zool., 30:1-93, 3 pis. 

Pierce, B. and C. P. Patterson 

1879. List of dredging stations occupied by the United States Coast 
Survey Steamers "Bibb," "Hassler," and "Blake," from 
1867 to 1879. Bull. Mus. Comp. Zool., 6(1):1-15. 

Robins, C. Richard. 

1959. Studies on fishes of the family Ophidiidae. I. A new species 
of Lepophidium from the Caribbean Sea. Bull. Mar. Sci. Gulf 
and Carib., 8(4) :360-3G8, 2 figs. (1958). 

Robins, C. Richard and Robert R. Miller 

1957. Classification, variation, and distribution of the sculpins, genus 
Coitus, inhabiting Pacific slope waters in California and south- 
ern Oregon, with a key to the species. California Fish and 
Game, 43(3) :213-233, 3 figs. 



^~ <- 



o^^^u 



BREVIORA 

MuseiiintTi of Comparative Zoology 



Cambridge, Mass. April 14, 1959 XrMHKK 105 

BVFO GIJNDLACHI, A NEW SPECIES OF 
CUBAN TOAD 

By Rodolfo Ruibal 

Division of Life Sciences 
University of California, Riverside 

INTRODUCTION 

The discovery of a new species of Cuban Bufo is surprising- 
considering the amount of herpetological colleetino- that has 
previously been done in Cuba, the apparent abundance and 
widespread distribution of the new species, and the fact that 
Cuba already possesses three species of Bufo. Bufo peltacephalus 
Tschudi, B. cmpusus Cope and B. longinasus Stejneger have 
Ion": l)een known from the island, the last species having- been 
described in 1905. In contrast, the other Greater Antillean 
islands, Hispaniola and Puerto Rico, possess but a single species 
each of Bufo: B. lemur in Puerto Rico and B. guniheri on His- 
paniola. 

The new species is named in honor of Juan Gundlach, the 19th 
century Cuban naturalist and collector. Gundlach in his ''Con- 
tribucion a la er])etologia cubana,"' published in 1880, actually 
recognized the existence of the species here described as new. 
lie mentions collecting a small species of toad 33 millimeters in 
length, near H(d)ana. The toads had a distinctive call and some 
were in amplexus. (4undlach sent some specimens of this toad 
to Wilhelm Peters at the Berlin Museum who declined to recog- 
nize the specimens as a new species because he thought that they 
might represent juveniles of Bufo peltacephalus. Gundlach 
justly chided Peters for failing to realize that a juvenile would 
not be calling, in amplexus and laying eggs. It has taken nearly 
80 yeai-s to prove Gundlach correct. 



2 



BREVTORA 



No. 105 



BUFO GUNDLACHI, sp. IIOV. 

Typr. M.C.Z. W^nil, male, collected about 14 kilometers N.E. 
of the city of Cama<>iiey, Provincia de Camaguey, Cuba, by 
Kamon Molina and Kodolfo Kuibal on August 22, 1!)57. 

Paratypes. M.C.Z. 30552-81, all with the same data as the type. 

Diagnosis. A small toad differing from Bufo longinasns, the 
only other known small toad of Cuba, by the following charac- 
ters: shorter hind limbs (see Table 1); high supraorbital and 
supratympanic crests that are lacking in B. longinasxis; the skin 




Figure 1. Dorsal view of the head of B. gundlachi, MCZ 305.")2. The pis- 
ment patterns have been excluded from the drawing. 



of the males more spinous than in B. longinasus males; and in 
Iiaving the parotid gland small and latei-al rather than large and 
dorsolateral as in B. Jonginasus (see Figures 1-4 for compari- 
sons). Readily distinguished from juvenile specimens of B. 
pcltacephalus by the structure of the suprii()rl)ital and supra- 
tympanic crests as well as by the i)resence of a thin white 
mid-dorsal stripe in B. giiii<llaclii, absent in juvenile B. pclta- 
cephalus. Readily separable from adult B. pfltac< phalus by size 
(see Table 2). Separable from B. cinpnsiis by the fact the 



1950 



NEW SPECIES OF CUBAN TOAD 



tnbei'cles on the doi-sum of B. niipiisxs are not spinous, as they 
are in B. (jiiixlldcJii, tliat in B. onpiisus the tynipaniiiii is pro- 
])ortionately much smaller than in B. (jiindlachi, and that the 
metatarsal tubercle of B. oiipusus is lar<ie and keratinized while 
that of B. [jiiudlachi is smaller and not keratinized. 

Description of ttjpc and parafifpes. A small, short-legged toad 
(see Table 1 and Table 2). Head with high supraorbital crests 
that are continuous with the supratympanic crests (see Figure 
1). The posterioi- end of the supratympanic crest is usually 
thicker than the crest proper. The area between the two supra- 
tympanic crests is marked by a number of w'ell-developed pro- 




Figure '2. Dorsal view of the head of B. longi7uisus dun)ii, AMKII 
60802. The pigment patterns have been excluded from the drawing. 



tuberances or bosses. A low canthal crest is often present. The 
rest of the head surface may show smaller isolated rugosities and 
ridges (see Figures 1 and 3). There is considerable variation 
in the size and extent of the rugosities in the various specimens. 
Eye with a horizontal elliptical pupil. Tympanum small and 
not sharply demarcated. Parotid gland spinous, lateral in posi- 
tion, irregular in outline, and not clearly demarcated in many 
specimens. Skin of the dorsum, tubercular, with some of the 



BREVrORA 



No. 105 



tubercles bearing a spinous termination. Ventral skin roug:h and 
<?ranular. 

Manus with large tarsal tubercle and inner toe of males with 
a darkly pigmented patch on median surface. Third finger the 
longest, then fourth, second, and first (in that order). A large 
flat metacarpal tubercle. Pes with a pair of metatarsal tubercles 
and a tar.sal tubercle. Toes partially webbed. 

Color. The new species is very variable in general pattern as 
well as color. In life (type and paratypes) the dorsal area 
varied in color from brown to dark green. A distinct, thin, white 
middorsal line was evident in all specimens. The most common 
pattern w^as one of a dark green or brown dorsal area containing 




Figure 3. Lateral view of the head of B. gundlachi, MCZ 30552. 



some darker blotches and having a reddish brown band border- 
ing it on each side. In turn this reddish Iji-owu band was bor- 
dered ventrally by a darker brown l)an(l. The dorsal anterior 
surface of the head was usually lighter in color than the dorsal 
body area (see Figure 5). The iiuiles differ from the females 
in having a darkly pigmented throat. 

After preservation in alcohol or formalin the same general 
color and pattern is visil)le except that no green is evident and 
the general pattern is often obscured and darkened. The dorsal 
area may be dark and patternless or show three pairs of dark 
blotches, one pair above the level of the forelimbs, another pair 
at the middle of the body, and the third pair at the posterior tip 
of the urostyle. This dorsal area is in many specimens bordered 
by a lighter band (the reddisli bi-owu l)an(l of tlie living animal). 



1!)5!) 



NEW Sl'KCIEK OF CUHAX TOAD 



The darker band that liorders this hciiul vciitrally originates on 
the tympanum (see Fi<>ure 3), covers most of the parotid jrland 
(except for the ventral edge of the gUiiid which is white), and 
extends along the ventrolateral margin of the l)ody. TTsually 
there is evident a darker stripe anterior to the insertion of the 
forelimb and in some cases a short thin dark stripe is present on 
the midline of the chest. Otherwise (with the exception of the 
dark throat of males) most specimens have a spotless ventral 
surface, though a few specimens show various degrees of dark 
mottling on the throat, chest and/or belly. 

Other spccime)is. Specimens of the new species have also been 
collected in Pinar del Rio, Habana, Isla de Pinos, Las Villas, 
and at other localities in Camague3\ Most of these specimens 
were collected bv Dr. Albert Schwartz of Albright College, who 




Figure 4. Lateral view of the head of B. Innginasus dnnni. AMNH 
00802. 



very generously made his material and notes available to me. 
Morphologically, all of the populations are similar. The only 
specimens that may be sufficiently distinct from the tyjies to 
I'epresent a ditt'erent subspecies are from Isla de Pinos. These 
specimens aj)pear to be small, immature individuals; however, 
they differ from other specimens in being much less rugose and 
spinovis and in having a more shari)ly contrasting color pattern. 
Since only two specimens are Hvailable from Isla de Pinos it 
would appear best to reserve any subs])eciiic designation of the 
forms until more specimens are available. 

The only other noticeable differences observed were that the 
Contramaestre, C'amaguey, specimens had swollen and enlarged 
parotids as compared to other populations. Also the specimens 



6 BREVIORA No. 105 

from near llerradura. ]'inar del Kio, were characterized by the 
("act that males were more spinous than the females. 

List of other specinicus examined. Pinar del Rio: 2.8-2.8 ini. 
E. of La Mulata, AMXII 60821-28; 2.9 mi. E. of Isabel Kul)i() 
(formerly Mendoza), AMNH 60829: 2 mi. S.E. of llerradura. 
AMNII 60880-56; near llerradura. MCZ 2833 (3 spec.) ; 5.5 mi. 
E. of Candelaria, AMNII 60890-918. Ilabana : Prov. of Habana, 
MCZ 19394; Isla de Pinos, AMNH 61222-23. Las Villas: 28 
km. E. of Trinidad. AMNII 60884-89. Cama«-uey : Embarca- 
dero de Moron, AMNII 60857; 9.4 mi. S. of Contramaestre, 
AMNH 60858-83. 

Ecological data. The type and paratypes were collected at 
two sites in the open serpentine savanna between Camaguey and 
the Sierra de Cubitas. The toads were found in m<^dium size 
clioruses in the afternoon after a heavy and intense rainstorm. 
All the low, poorly-drained areas of the savanna were flooded 
and the congregations of toads were iu these shallow, grass- 
covered temporary pools. The toads were in the water when 
calling and were wary, submerging at the approach of the col- 
lectors. The call was loud for such a small animal and might 
be reproduced as CKEE-CKEE-CKEE. Also calling at this time 
was a small species of EleutJierodactt/lus. A single specimen of 
B. empusus was found near the pools. No egg-laying had yet 
occurred when the specimens were collected. 

The many specimens collected by Albert Schwartz were col- 
lected in similar shallow ponds in open fields. In this respect 
the new species differs from B. loii<jinasi<s, which so far has 
always been collected along streams in wooded areas (Stejneger, 
1905; Barbour, 1914 and 1926). 

B. gundlacJii has been collected from the provinces of Pinar 
del Kio, Ilabana, Las Villas, and Canuiguey, and from Isla de 
Pinos. It is, therefore, a widespread form in Cuba and will 
probably be found to inhabit also Matanzas and the lowland 
areas of Oriente. All of the other species of Cuban toads are 
characterized by a similar islaudwide distribution : B. empusus 
and B. peltaccpludns have been collected in all the provinces and 
on Isla de Pinos, while B. loiiginasus has been found in what 
appear to be isolated po])ulations in Pinar del Rio, Las Villas 
and Oriente. 



1 <»;')!) 



NEW SPECIES OF CUBAN rOAI) 



Since it seemed unlikely that the American museums had 
missed collecting B. gnndlachi until 1957, ii search M'as made of 
the collections of B. cmpusus in the Museum of Comparative 
Zoology and in the American Museum of Natural History with 
the idea that specimens of the new species had been collected 
but had mistakenly been identified as juvenile B. cmpusns (the 
juveniles of B. empusus are rare in collections and thus are 
poorly known, while the young- of B. peltaccphalus are relatively 
common and known). This proved to be the case and a number 




HINDLIMB 
LENGTH 



riguie 5. Dorsal view of B. (/inidladti, MCZ 305.")2. Measuieinent of the 
liind limb length is indicated. 



of specimens of the new species were uncovered in this fashion. 
The relative scarcity of the new species from collections and the 
fact that all of the sjiecimens collected by Albert Schwartz and 
myself were from choruses Avould indicate that B. gnndlachi is 
a subterranean species and thus difficult to locate outside of the 
breeding season. 



BREVIORA 



No. 105 



DISCUSSION 

111 Table 1, a suiiiinary of the (iuantitativt' data expre.s.siu<>- 
the difference in the lengtli of the hind limb of B. (jiindlachi as 
compared to the other species of West Indian toads is provided. 
In the case of B. oxpusiis and B. pdtaccphahis it was felt to be 
more meaniiio'fid to measnre only jnvenile specimens since only 
the small s])('cimeiis of these two species conld be confu.sed with 
B. (jundlachi. The measnremeiit of the body lent>th was done 
Avith Vernier calipers and actnally is the measurement from 
the tip of the snout to tlie posterior tip of the urostyle. When 
measnrin<»- the specimen, it was held so as to flatten the body. 
The len<>t]i of the hind limb is considered to ho the distance from 




Figure 6. Lateral view of the head of B. empu.sns, MCZ 3733. 



the vent to the tarso-metatarsal joint measured with a ruler 
(see Figure 5). The data on Table 1 do not demonstrate any 
apparent ditference between the various populations of B. gnnd- 
lachi in regard to hind limb length. The data do demonstrate 
a significant difference between B. (jundlachi and B. longinasns 
dunni (and probably the other subspecies of B. longinasiia as 
well) in this re^jard. The dnta for the juvenile specimens of B. 
peltacephaliis overlaj) the lower range of B. gundlavhi while the 
juvenile specimens of B. eurpusits over-lap the upper range of the 
new .species. 



11).')!) 



NEW yi'LClES OF CUBAN TOAD 



A comparison of th(> external characters of llic new species 
with those of the otlier species of Cuhan toads indicates that B. 
(inndlachi resembles B. ()iipiisiis ;iiid B. peUdccpJinJus more than 
it does B. lo)ighiasus. 



/>. iiiiiju.siis, B. pcltavcphdUis 
and B. (jundlaclii 

1. I'aiotid gl.'iiui short and lateral 

in position. 
'1. Various degrees of supratyni- 

panie. and sii])raorl)ital cranial 

crests. 



/.'. loiKjinana.s 

1. I'arotid t;land elong:ate and ex- 
] landed dorsally. 

2. Absence of well developed cranial 
crests. 




Figure 7. Lateral view of the head of B. pcltacepholus, MCZ 292S7. 



As a consequence of the i)eciiliarly ossified tympanic re<>ion, 
B. empusHs would appear morphologically to stand out from B. 
(jundlaclii and B. pcJtacepJialus. HoAvever, all three species ap- 
pear to be more closely related to the otlier West Indian species, 
B. lemur and B. guniheri, than to the Cuban B. longinasus. In 
addition to the characters listed above, B. longinasus is dis- 
tinctive in the pointed snout of adult specimens, a feature which 
is, however, less obvious in the youn<i-. Schmidt (1928) rightly 
pointed out that the Puerto IJican and Hispaniolan species are 
more similar to each other than to the Cuban species. Never- 
theless, B. lonur and B. gunihcri are characterized by short 



10 



BREVIORA 



No. 105 



lateral parotids and well-developed cranial crests and are tlius 
similar to the peltacepkalus-empusus-gundlachi group. B. longi- 
iiasn.s, as Barbour (1914) observed, stands out as a unique West 
Indian species unrelated to any of the other known forms. 

A word is in ordei- here concerning Bufo turpis Barbour from 
Virgin Gorda near Puerto Rico. I have compared the type (MCZ 
4099) of this species with a number of specimens of B. lemnr 
from Puerto Rico (MCZ 30614-5 and AMNH 10148 to 10151) 




Figure 8. Dorsal view of the head of B. guntheri, MCZ 3104. 



and can find no valid reason for considering B. turpis a distinct 
species. The differences in the shai)e of the dorsal crests cited by 
Barbour (1917) are insignificant and clearly within the ex- 
pected range of variation of a species. Nor does the single 
specimen of B. turpis give any evidence of being siibspecifically 
distinct. Consetpiently B. turpis should be placed in the 
svnonvmv of B. lemur. 



id:)!) 



NEW SPECIES Ol' CUBAN TOAD 



11 



The key to tlie West Indian sj)ecies of Btifo here i)resente(l 
does not inchule the subspeeics of B. Jonginasus. H. longinafius 
ramsdcni described by Barbour in 1914 is known only from the 
two not too well preserved orijiinal speeimens from Oriente, and 
B. luiujinasiis is only known from Stejneger's original type from 
Pinar del Kio. B. lo)igiiiasus dunni described l)y Barbour in 
1926 is the only foi-iu of which sei-ies of specimens are available. 
Until further si)ecimens of />. /. longinasus and B. I. ramsdeni 
are available it will lie impossible to critically determine the 




Figure 9. Lateral view of the head of B. g)tiii]icri, MCZ 3104. 



validity of the three named forms. Also not included in the 
key is tlie introduced B. nun'iinis. This form has been success- 
fully introduced into the Lesser Antilles and Puerto Rico, Ilis- 
paniola, and Jamaica. It has been introduced into Cuba at least 
once (Scaramozza, 1947) but interestinoly enouprh there is no 
evidence that it has established itself on Cuba. It is an attractive 
hypothesis to attribute the failure of B. marinu:< in Cuba to the 
existence of B. pcltacephalus. This native species is almost as 
large as B. mariiius and successfully inhabits tlie cultivated 
areas of Cuba and thus represents a form that has adapted itself 
to a situation in which much of the original habitat has been 
destroyed. 



12 



BREVIORA 



No. 105 



Kfi) to flic )iafirc \V(.st Indian species of BuFO 

]. Tyinpaniun not evident, covered by i)ustulous skin (rigure 4); parotid 

elongate, the length greater than the width (Figure 2) 

B. lonfjina.su.s 

Tynipaniini not covered by pustulous skin; parotid 7iot elongate '2 

'2. Tympanum small, completelj' encircled by bone (Figure 6); sharply in- 
clined profile to snout (Figure (i) . B. empu.sus 
Tympanum not encircled liy bone; profile not sharply inclined ;^ 

'A. Hind limbs short, the body length/hind limb ratio ranging from 1.57 to 
1.93; adult size small (maximum body length, 36 mm.); a thin median 

dorsal white line (Figure 5) B. gundlachi 

Hind limbs longer, the body length/hind limb ratio ranging from 1.27 
to 1.6o; adult size larger (body length as great as 170 mm.) ; no median 
dorsal white stripe 4 




Figure 10. Lateral view of the head of /.'. li-m>ir, M("Z 30615. 



4. .\ cauthal knob (Figure 7 ) ; orbital crests indistinct B. pcliaccpltahix 
Xo canthal knob; well developed supraorbital and supratympanii- 
crests 5 

5. Median supraorbital crest aud posterior supraorbital crest forming a 
right angle (Figure 8) ; preorbital crest e(|ual to su])raorl)ital crest 

in thickness B. gunthrri 

Supraorbital crests confluent and forming an aic; preorbital cresi thiii- 
ucr aud less jirououuced than su|)ra()rbital cicst (Figure 10) U. liiiiiir 



1 f)r)f) 



XEW SPECIES OK CUBAN TOAD 



13 



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14 BREVIORA No. 105 

Table 2 

Maxiiuum body length of native "West Indian spreios of Biifo 

Maximum Body 
Speciefs Scir Leriffth (in mm.) 

Kimdlachi $ 36 

longinasus 9 34 

empusus 9 74 

peltacephalus $ 170 

5?untheri 9 80 

lemur 9 105 

ACKNOWLEDGMENTS 

The new species of Ciil)an toad -was collected while doing field 
work during the summer of 1957 in Camaguey with the finan- 
cial assistance of a grant from the Penrose Fund of the American 
Philosophical Society. T am grateful to the Society for this 
assistance. I Avould also like to thank Miss Dereth Bogert for 
her excellent drawings, and Dr. Doris Cochran for measuring 
the type of B. longinasus longinasus for me. 

LITERATURE CITED 

B.\RBOUR, T. 

1914. A contribution to the zoogeography of the West Indies. Mem. 

Mils. Comp. Zoo]., 44 (2^: 209-346. 
1917. Notes on the hcrpetology of the Virgin Islainls. Pioc. Biol. 

Soc. Washington, 30: 97-104. 
192(i. Xew amphibia. Oct-. Papers Boston So.-. Xnt. Tli.st., 5: 19M94. 

B.\RBOUR, T. .\ND C. T. RaMSDEN 

1919. Ilerpetology of Cuba. Mem. Mus. Comp. Zooi., 47 (2): 71-21:;. 

(irXDLACII, J. 

1880. Contrilnu-ion a la erjietologia cubann. C. Montiel y ('in., ll;ili;ni;i, 
pp. 1-99. 
SCARAMOZZA, L. C. 

1947. Correspondi'iu-i.-i. Mem. Soc. Cubanu IJist. X;il., 19 (li: 112 
113. 
Schmidt, K. V. 

1928. Amphibians and huid reptiles of Porto Hied, with a list of 
those reported from the Virgin Islands. Sci. Survey Porto 
Rico and Virgin Isl., 10: 1-1 (id. 
Stkj.nkgek, L. 

1904. Herpetology of Poito liicu. Ann. liept. l'. S. .\;it. Mus. fi.i- 

1902, pp. .349-724. 
190."). r)escii]»tion of a new tiiad from Cuba. Proc. T'. S. Nat. .Mus., 
28 (1406): 765-767. 



( jsJbOU 



BREVIORA 

Mmseiim of Coimiparative Zoology 



Cambridge, Mass. April 28, 1959 Number 106 



THE OCCIPITO-VERTEBRAL JOINT IN THE BURROW- 
ING SNAKES OF THE FAMILY UROPELTIDAE 

By Ernest E. Williams 

In 1939, R. Hoffstetter described an amazing condition of the 
head joint in the peculiar burrowing snakes of South India and 
Ceylon — the Uropeltidae. Quite in contrast with the condition 
found in all other amniotes, the first vertebral centrum was said 
to be fused with the occipital condyle, while the second vertebral 
centrum was unmodified and precocious like those behind it. 
According to Hoffstetter, also, the atlas neural arches, united 
ventrally by a ligament, embraced the neck of the occipital con- 
dyle. The species he personally examined was Rhinophis hlythii. 

The head joint of Rhinophis planiceps (^ R. philippinus), as 
Hoffstetter was well aware, had previously been described by 
Baumeister (1908, pp. 499-502, fig. 22). He, like Hoffstetter, 
had found the first vertebral centrum (= odontoid process) to 
be absent as a distinct structure. His account differed, however, 
from that given more than thirty years later by Hoffstetter in 
one very important detail. He did not find the atlas embracing 
the neck of the occipital condyle ; instead it was in its normal 
place posterior to and articulating with the head of the condyle. 
Baumeister also interpreted the situation very differently, be- 
lieving the first vertebral centrum to have been reduced and for 
the most part fused with and forming the major element in the 
anterior articular surface of the second vertebral centrum. 

Whether as described by Hoffstetter or by Baumeister, it is 
sufficiently evident that the head joint in the uropeltids exam- 
ined by them is profoundly different from the head joint in 
any other amniotes. In the course, therefore, of a general study 
of the head joint in reptiles I have found it necessary to give 



2 BREVIORA No. 106 

special attention to this feature in this family and in related 
groups. The simple anatomy of the articulation needed to be 
carefully restudied, its taxonomic incidence established, and its 
morphological interpretation reassessed. 

Since a detailed discussion of these points would be a lengthy 
digression in any more general paper, I publish this account 
separately. 

Skeletons of the following members of the family Uropeltidae 
have been specifically prepared for this study. In each case the 
head joint was examined at the moment of separation of head 
and vertebral column and before further preparation. 

Melanophidium wyandense 
Plat y 2)1 efrurus madurensis 
Plectrurus perroteti 
PseudotypJdops philippinus 
Rhinophis hlythii 

philippinus 

sanguineus 
Teretrurus rhodogaster 
Uropeltis ceylanicus 

ocellatus 

ptdneyensis 

ruhrolineatiis 

This list includes at least one member of every genus currently 
recognized in the family and several species of two of these 
genera, including both the species for which the head joint has 
previously been described. 

In this suite of specimens differences of a specific or generic 
character are indeed observable, but they involve only minor 
details. The general pattern is remarkably uniform. 

In every case the second vertebra articulates directly and by 
a simple concavity with the ball -like convexity of the occipital 
condyle. There is no intervening odontoid process. The condylar 
articular surface is never notched dorsally or indented medially. 

An atlas neural arch is always present. It is never placed 
over the neck of the condyle, as reported by Hoffstetter (Fig. 
IB), but instead is, as Baumeister stated, always in articulation 
posteriorly with the neural arch of the second vertebra, and 
ventroanteriorly with the ball-like convexity of the occipital 



1959 



UROPELTID HEAD JOINT 



condyle. In its articulation with the occipital condyle it fur- 
nishes a dorsolateral supplementary rim to the cup-like anterior 
concavity of the centrum of the second vertebra. 

An atlas ventral arch may or may not be present. It is present 
as a distinct element only in Vropeltis pulneyensis (Figs. 3A 
and B). In some of the forms examined it may have been fused 




Fig. 1 A. Head joint of Typhlops steinhmisi (after Hoffstetter) . B. 
Head joint of Ehinophis Uythii (after Hoffstetter). C. Real condition of 
the head joint of Ehinophis blythii. Abbreviations: Ci = centrum 1, the 
odontoid ; C2 = centrum 2, the axis centrum. lo = second intercentrum ; 
I3 = third intercentrum. O = occiput. 



into the ventral margin of the anterior concavity of the second 
centrum. The presence of a slight anteroventral projection — 
absent in U. pulneyensis — is suggestive of this (Figs. 2B and D ; 
Fig. 3D). 

The contribution of the pedicels of the atlas neural arch- 
halves or of the ventral arch (when present) to the concavity 
which receives the occipital condyle, is in all cases minor. The 



BREVIORA 



No. 106 



major articular surface is always provided by the second cen- 
trum. 

The uropeltid head joint ma^' be usefully contrasted with that 
of a more primitive snake, Cylindrnphis rufus, which is quite 
typical of the more usual squamate condition. 






Fig. 2 A. Anterior view, atlas and axis of Bhinophis sanguineus. B. Side 
view, atlas and axis of Ehinophis sanguineus. C. Anterior view, atlas and 
axis of Platyplectrurus madurensis. D. Side view, atlas and axis of 
Platyplectrurus madurensis. 



l!)5i) ITROPKLTID IlKAD .JOINT 5 

111 Cyiindi'ophis rnfiis the occipital condyle bears definite 
traces, in well-marked grooves, of its origin by the union of 
three bones and is not hemispherical but distinctly indented on 
its dorsal surface. This indentation is the fovea dent is (Gaupp, 
1908, p. 518), and is the place of attachment of the ligament urn 
apicis dcntis. In many forms this ligament chondrifies or ossifies 
in ■whole or in part as a small conical projection from the odon- 
toid process which it is then convenient to call an apex dentis 
(new term). In C. ruftis the apex dentis is a small cartilage cone. 

The occipital condyle is received into a concavity formed by 
the ventral portions of the atlas ring, the lateral margins of 
which are formed by large articular surfaces on the pedicels of 
the atlas neural arch-halves and the articular surface of the 
ventral arch. 

The dorsal margin of the concavity is furnished by the liga- 
nientum transversum atlantis, while the outer portion of the 
bottom of the cup is the septum interarticulare (Gaupp, 1908, 
p. 528). Beneath the ligamentum transversum atlantis and per- 
forating the sept inn interarticulare by means of the foramen 
septi the apex dentis projects forward into the fovea dentis of 
the condyle. 

The contact of the occipital condyle in C. rufus is thus pri- 
marily with the atlas ring and its associated ligaments, only to 
a minor degree with the odontoid process, mostly by its apex 
dentis, and not at all with the centrum of the second vertebra. 

The atlas ring in C. rufus, as in other typical squamates, forms 
posteriorly a cup for the reception of the odontoid process, en- 
tirely similar to that described for the occipital condyle. The 
same parts participate in a substantially similar way. 

The odontoid process in C. rufus projects forward from the 
second vertebra. It is not a conical, distinctly tooth-shaped proc- 
ess, as in most mammals. In C. rufus it is an essentially wedge- 
shaped element, flat dorsally, posteriorly with a flat surface that 
is suturally united with the true centrum of the second vertebra. 
Its anterior surface slopes sharply backward. Medially, just 
below the point of juncture of anterior and dorsal surfaces the 
small cartilaginous cone that is the apex dentis arises so abruptly 
as to seem a separate element; it has frequently, though prob- 
ably incorrectly, been regarded as representing the vestigial 
centrum of a proatl antic vertebra. 



6 



BREVIORA 



No. 106 



Ventral and somewhat posterior to the odontoid process is the 
second or axis intercentrum, an element much larger than the 
odontoid process; it is suturally united to the ventroanterior 
surface of the true centrum of the second vertebra and projects 







rig. 3 A. Anterior view, atlas and axis of Uropeltis pulneyensis. B. Side 
view, atlas and axis of U. pulneyensis. C. Anterior view, atlas and axis of 
Uropeltis ocellatus. D. Side view, atlas and axis of V . ocellatus. 



1959 IIROPELTID HEAD JOINT 7 

obliquely backward as the first of two ventral spines on the axis 
vertebra. Anteriorly this element articulates with the ventral 
arch of the atlas. 

With minor differences in details of shape and size this de- 
scription would serve for the head joint of any typical squamate 
or of Sphenodon (cf. Typhlops, Fig. lA), and with no serious 
modifications except in regard to relative size of the odontoid 
process and second intercentrum, it will serve for any crocodile 
or most turtles. 

The essential differences between the uropeltid head joint and 
that typified by Cylindrophis rufus are : 

1. The condyle of the uropeltids lacks a fovea dentis. 

2. There is neither a ligamentum apicis dentis nor an apex 
dentis. 

3. The articular surfaces on the atlas neural arch pedicels 
are reduced. 

4. The atlas ventral arch is reduced as a whole or more fre- 
quently has disappeared altogether as a distinct element. 

5. There is no odontoid process. 

Somewhat aberrant among squamates is the head joint of 
Cylindrophis maculatus of Ceylon (Fig. 4). The occipital con- 
dyle is not notched dorsally, nor indented mediall3^ The grooves 
indicating the place of union of the three participating bones, 
are, however, present. There was apparently no ligamentum 
apicis dentis or apex dentis. The basal portions of the atlas ring 
— including the neural arch pedicels, but especiallj^ the ventral 
arch — are highly developed. The two neural arch pedicels 
almost meet mid-dorsally, reducing the ligamentum transversum 
atlantis to an extremely short connection between the two. The 
area which could have been filled by a septum interarticulare is 
very small ; perhaps there was none. The odontoid process itself 
seems somewhat reduced and truncate in front. 

In certain ways in which uropeltids differ from typical squa- 
mates, C. maculatus appears to be modified in a direction oppo- 
site to that seen in the uropeltids. The basal parts of the atlas 
are enlarged in C. maculatus, reduced in uropeltids. 

In certain other ways C. macidatus approaches the uropel- 
tids: in the absence of the fovea dentis and of the correlated 
ligamentum apicis dentis or apex dentis. 



8 



BRE\aORA 



No. 106 



Even the enlargement of the atlas basal elements can, how- 
ever, be interpreted as tending in the direction of the nropeltids. 
The typical squamate and other reptilian condition is complex, 
essentially two joints, one within the other. On the outside 
there is a major condylar convexity fitting into an atlantal cup ; 
inside this is a joint oriented in the opposite direction — a con- 
vexity arising from the odontoid (atlas centrum), fitting into a 
small concavity within the condjde. See diagram Figure 5A. 

The nropeltids and C. maculatus have simplified this arrange- 
ment by abolishing the internal odontoid joint and perfecting a 





Fig. 4. Cyli)uh-op]tis tiianilatits. A. Anterior view of atlas. B. Side view 
of axis. 



ball and socket joint entirely comparable to the joints between 
vertebrae, posteriorly. (Pig. 5B.) 

If this be the correct interpretation of the results achieved by 
the uropeltid - C. maculatus modification, it may reasonably be 
wondered why the extreme uropeltid condition was found 
adaptively necessary. C. maculatus has achieved a reasonably 
good ball and socket joint without losing the odontoid process 
entirely. A hypothesis may be offered here, for future mechani- 
cal and functional analysis to test. The axis of C. maculatus 
even with some reduction of the odontoid is much longer than 



1!)59 ITROPELTID HEAD JOINT 9 

the vertebrae behind it. The uropeltid axis on the contrary is 
ff.s short or shorter than the vertebrae behind it. We may assume 
that in tlie still unanalyzed functioning of the uropeltid head 
joint this shortening- of the axis^ has been mechanically and thus 
selectively valuable. That this shortening has been accomplished 
by reduction of the odontoid to essential nullity we may believe 
to have been an accident of evolution in this specialized group 
— or we may see in it the culmination of a trend begun in an 
ancestral form which must have been similar to C. maculatus. 

It must be explained that although C. mamdatiis occurs in 
the same area as the uropeltids, shows in several respects special 
morphological resemblances to them, and may w^ell have had a 
long history independent of other members of the group, part of 
the resemblances between maculatus and the uropeltids may well 





B. 

Fig. 5. Diagrammatic horizontal section of the head joint region. A. in 
most reptiles. B. in uropeltids. 

be parallelism. At least, as Bellairs and Underwood (1951, p. 
231) have indicated, the presence of two ventral transverse scale 
rows to each body segment in uropeltids rather than one as in 
Cylindrophis is possibly primitive and prompts the inference 
that the uropeltids have not been directly derived from Cylin- 
drophis or its immediate relatives. 

Thus, while it is interesting to suggest that the uropeltid head 
joint may in the course of its evolution have gone through a stage 
directly comparable to that in C. r)iaculatus — a stage in which 
the basal parts of the atlas were enlarged rather than reduced 
— there is no compelling reason derived from any demonstrated 
direct phyletic relation of C. maculatus to the uropeltids to ac- 
cept such a stage as a necessary intermediate. It would appear 

1 Most uropeltids have also elongated the pedicel of the occipital condyle, but 
this is not true in M elan o phi dium which seems in skull and scutellatlon the most 
primitive member of the family. 



10 BREVIORA No. 106 

simpler to suppose tliat in the iiropeltids the modification of the 
head joint was accomplished from the first by strong reduction 
of the ventral parts of the atlantal ring and of the odontoid 
simultaneously. The question is one. however, on which critical 
evidence is unavailable. 

If the course of evolution of the uropeltid head joint is likely 
to remain for some time unknowii, more hope may be offered 
that its ontogeny can be examined. The uropeltids are ovo- 
viviparous ; it will therefore be worthwhile to examine the col- 
lections already in existence for pregnant females. Though this 
would be unlikely to provide a stage by stage view of the on- 
togeny of the head joint region, in the present state of our 
knowledge any glimpse of the development (as of the function 
or the evolution) of this region in uropeltids would put us sub- 
stantially farther ahead. 

LITEEATURE CITED 

Baumeister, K. L. 

1908. Beitrage zur Anatoinie und Physiologie der Ehinophiden. 
Integument, Driisen der Mundhohle, Augen und Skeletsystem. 
Zool. Jahrb. (Abt. Anat.), 26: 423-526. 

Bellairs, A. d'A. and G. Underwood 

1951. The origin of snakes. Biol. Review, 26: 193-237. 

Gatjpp, E. 

1908. Ueber Entwicklung und Bau der beiden ersten Wirbel und der 
Kopfgelenke von Echidna aculeata nebst allgemeinen Bemer- 
kungen liber die Kopfgelenke der Amnioten. Jenaisciie Denk- 
schriften, 6: 483-538. 

HOFFSTBTTER, R. 

1939. Sur 1 'articulation oceipito-vertebrale des Uropeltidae (ophidiens 
fouisseurs). Bull. Mus. Hist. Nat. Paris, (2) 11: 426-433. 



BREVIORA 



MitasemiM of Connparsitive Zoology 

Cambridge, Mass. May 6, 1959 Number 107 



A REVISION OF THE DACETINE ANT GENUS 

NEOSTRUMA 

By William L. Brown, Jr. 

Museum of Comparative Zoology, Harvard University 

INTRODUCTION 

The genus Neostruma Brown includes six known neotropical 
species forming a compact and distinctive group in subtribe 
Strumigeniti of tribe Dacetini. This genus evidently was derived 
within the New World tropics from the gundlachi group ("sub- 
genus Pyramica") of the large tropicopolitan genus Strumi- 
genys Fr. Smith. 

The >S^. gundlachi group is also neotropical in distribution. It 
contains a graded series of species, linked to the more "typical" 
Strumigenys through the S. connectens group (Kempf, 1958, 
Rev. Brasil. Ent., 8:59-68), that foreshadow Neostruma in head 
shape, development of serial denticulation of the mandibles, 
lengthening of labral lobes, reduction of pilositA" and spongiform 
appendages, and other characters. 

Neostruma is set off from these by its very long labral lobes 
with their short trigger hairs, and by the distinctive form and 
armament of the mandibles, featuring very short apical fork 
teeth and a series of denticles on each side of a submedian (pre- 
apical) tooth (Fig. 5). With the exception of N. myllorhapha 
new species (Fig. 4), a "countercurrent" specialist with sec- 
ondarily elongate mandibles, the Neostruma species show a co- 
adaptive complex of characters converging toward those of 
Smithistruma Brown, but not reaching the same degree of com- 
pleteness (see Brown and Wilson, in press). Among the charac- 
ters involved in this complex are the following: 

1. Shortening of mandibles. 

2. Reduction of apical fork. 



BREVIORA 



No. 107 



3. Development of serial mandibular deuticulation. 

4. Transformation of basal tooth into a perpendicular "basal 
lamella." 

5. Slight lengthening, depression and narrowing of head, with 
some lessening in prominence of occipital lobes and occipital 
excision. 

6. Shortening and incrassation of antennal scapes. 

7. Elongation of labral lobes. 





Figs. 1-4. Neostruma spp., dorsal views of heads, mandibles and antennae. 
Fig. 1. N. zeteTci n. sp., paratype worker. Fig. 2. N. metopia n. sp., holotype 
female, funiculus omitted. Fig 3. N. crassicornis (Mayr), left antennal 



scape of worker. Fig. 4. N. myllorhapha n. sp., holotype worker, 
drawn to same scale. 



All figures 



8. Reduction in length of labral trigger hairs. 

9. Inability to open mandibles widely (limit is much less than 
180°). 

10. Hunting behavior is of "strike -hold -sting" type (as based 
on observation of N. mustelina; see below). 

11. Foraging less active and less frequent {N. mustelina) 
than in several Strumigenys species studied, but much as in 
Sntithistruma spp. 



1959 REVISION OF ANT GENUS NEOSTRUMA 3 

The species form small colonies, chiefly in the leaf litter of 
I'ain forest or tropical ever<ii-een forest, and nests occupy cavities 
in rottinsT twin's, ])ieces of bark or similar forest-floor vegetable 
debris so far as we know. Pour of the species (hrevicornis, myl- 
lorhapha, metopia, zeteJci) occur in Central America, one (mus- 
tclina) is widespread in tropical Mexico, and one (crassicornis) 
is known only from southeastern Brazil. It seems strange that 
no records of the genus are available for northern or Am.azonian 
South America, but then collecting in these regions has scarcely 
begun so far as the Dacetini are concerned. All of the records 
we now have are from mainland localities; the genus apparently 
is absent from the West Indies. In the areas where it occurs, the 
genus is usually not scarce, but exists in fair numbers along with 
species of Sfrumigcnys, particularly of the gundlachi group, 
and many other cryptobiotically foraging ants. The food, judg- 
ing from what we know of Neostruma mustelina, consists pri- 
marily of small entomobryomorph Collembola and possibly some 
other minute terrestrial arthropods as well. Hunting behavior 
is like that of Smitliistruma rather than like the Strumigenys so 
far studied. 

The material used in this study has come from various sources 
(see Acknowledgements at end of paper), but the principal 
places of deposition are the United States National Museum, 
Washington (USNM), and the Museum of Comparative Zoology 
at Harvard College, Cambridge, Massachusetts (MCZ). 

The measurements and proportions used, and their abbrevia- 
tions, are as in my other papers on dacetine ants ; see especially 
Brown, 1953, Amer. Midi. Nat., 50 : 7-15, or Brown, 1953, Jour. 
New York Ent. Soc, 61 : 53, 101. 

Neostruma Brown 

=: Neostruma Brown, 1948, Trans. Amer. Ent. Soc, 74:111. Type species: 
Strvmigenys crassicornis Mayr, 1887, by original designation (+ 3 spp.). 
<^ Strumigenys, auct. (+ 3 spp.). 

Worker. Small to minute, resembling members of the Strumi- 
genys gundlachi group (from which Neostruma appears to have 
been derived; see Brown and AVilson, in press). Mandibles in- 
serted rather far apart, linear, tapering to an apical fork of 
two very short teeth separated by 2-4 minute intercalary den- 



4 BREVIORA Xo. 107 

tides. Basal laiuella present, separated from basal condyle by 
a deep notch, perpendicular to shaft, broad and flat, subtruncate 
or rounded at apex, normally hidden beneath clypeus when 
mandible is closed. Inner borders of mandibles armed Avith a 
row of small teeth or denticles on the apical half or tAvo-thirds, 
arranf?ed as follows : a preapical series of 3-8 denticles, imme- 
diately proximad of apical fork ; a single larger submedian 
tooth; proximad of this, a short medial series of indistinct den- 
ticles, beyond which the shaft is unarmed to the basal lamella 
(Fig. 5). Labrum with a broad basal part and greatly elongate, 
parallel tapered lobes, extending half or more the exposed length 
of the closed mandibles. At their apices, the labral lobes each 
bear 2-3 short trigger hairs; one or two additional pairs of fine 
hairs on their upper surfaces before the tips. When the man- 



Fig. 5. Neostrxima zeteM n. sp., greatly magnified dorsal view of left 
mandible as dissected out of the head. 

dibles are closed, the labrum is tilted ventrad, but when they are 
open, it is raised to point straight forward between them. Ap- 
parently the same "springtrap" arrangement holds in this 
genus as in Sfrumigenys. Palpi segmented 1, 1, as in Strumi- 
genys. 

Head rather elongate, depressed, with moderately expanded 
occipital lobes, narrowed anterior portion, and moderately well- 
developed elliptical compound eyes set well forward of midlength. 
but only partly or not at all visible from above. There is a tend- 
ency, best seen in hrevicornis, zeteki and especially metopia, for 
the head to be impressed or concave in front of the vertex. An- 
tennae 6-segmented, as in Strumigenys, but scape shortened and 
incrassate, with curved (and sometimes lobulate) anterior mar- 



gin. 



1J)5!) RFVISIOX OF AXT GENUS NEOSTRUMA 5 

Alitrunk (.-ompact ; {)ronotuin depressed and nearly plane 
above, bounded by a distinct anterior and less distinct lateral 
margins, meeting at blunt humeral angles. In profile, the dorsal 
outline is convex from pronotum to propodeum, with a depres- 
sion (or flat area) in the region of the mesonotum. Promeso- 
notal suture obsolete ; iiietanotal groove weak to absent. 
Propodeal teeth laterally compressed, acute, their apices directed 
slightly dorsad, their infradental lamellae narrow and concave. 
Petiole pedunculate, its rounded node with a feebly developed 
posterodorsal collar of spongiform lamella ; no ventral append- 
ages. Postpetiole subreniform, much broader than petiole, with 
weak lateral and ventral spongiform appendages. Gaster with 
a narrow basal spongiform margin and a band of short basal 
costulae ; rest of surface smooth and shining. 

Head, alitrunk, legs, and both nodes finely and evenly reticu- 
late-punctulate and opaque, except for a smooth, shining patch 
on the mesokatepisternum. 

Mandibles with oblique pointed and linear-spatulate hairs di- 
rected mesad from dorsal surfaces, especially near insertions. 
Clypeal and anterior scape margins each with a series of border- 
ing spatulate hairs; one erect truncate, clavate or remiform 
erect hair on the posterior edge of each occipital lobe, one on 
each lateral occipital border (Figs. 1 and 4), one on each hu- 
merus, and one on each side of the mesonotum (these hairs are 
often small and fragile, and are often missing in part) ; two on 
the petiolar node, four on the postpetiole, and 16-20 on the 
gastric dorsum. Ground pilosity moderately developed to obso- 
lescent except in N. crassicornis, consisting of sparse, fine, recli- 
nate or subreclinate hairs on the head, legs and antennae, often 
invisible on head, shorter on clypeus. 

Color varying from ferruginous yellow to dark brown. The 
gaster is often darker than the rest of the bodv. 

Female like worker, but with larger eyes and fully developed 
flight apparatus, and otherwise differing from the workers as 
Struniigenys workers difl^er from their females. Forewing as in 
Wheeler's figure 11 D in his book, "Ants," p. 25, except that in 
Neostruma the apical section of R (labeled "f" by Wheeler) is 
lacking. 

Males known only for N. mustelina; these are basically like 
males of the smaller species of Struniigenys. 



6 BREVIORA No. 107 

SPECIES EVOLUTION AND RELATIONSHIPS 

The six species recognized here represent six morphologically 
distingnishable samples; of these, crassicornis is distinct in a 
number of good characters from the other five samples, and there 
is every reason to maintain it as a good species. Its range in 
southern Brazil and northern Argentina, so far as tlie present 
incomplete collections shoAv, is separated by the bulk of the 
South American continent, or over 3,000 miles, from the nearest 
known locality of any of the other species. In Panama, we find 
two sj^mpatric species, zeteki and metopia, which are very dif- 
ferent from one another (Figs. 1 and 2). In zeteJci, the head is 
long and exceedingly flattened, and the mandibles are fairly 
long, with many denticles, while metopia has a deep, short, broad 
head and very short mandibles. So far, our knowledge of the 
distribution of both of these species is limited to knowing that 
they both have been taken on Barro Colorado Island in the 
Canal Zone. 

Two of the remaining three species (hrevicornis and mustc- 
lina) are very similar to one another and to zeteki, and mustelina 
also approaches metopia. In fact, these four species can be ar- 
ranged as the morphocline zeteki ^ hrevicornis ^ uiustc- 
lina ^ - metopia. In their distribution, however, zeteki and 
metopia occur together in Panama, hrevicornis is in Costa Rica 
and Honduras (and presumably also in between, in Nicaragua), 
and mustelina is widespread in tropical Mexico. Thus we see 
that the morphocline is partly also a geographical cline running 
from Panama to Mexico: zeteki ^ hrevicornis ^ muste- 
lina. This situation suggests that zeteki, hrevicornis and mus- 
telina could amount to one clinally varying species occujiyiug 
much of Central America and tropical Mexico, and that the 
zeteki population may represent extreme character displacement 
in the presence of metopia, which is itself displaced toward the 
other extreme. An alternative possibility should not yet be 
excluded : metopia could belong with hrevicornis and mustelina, 
and could represent a reversal of cline where zeteki is present 
and displaces it. Despite the ijiterest of this possible character 
displacement situation (see Brown and Wilson, 1956, Syst. Zool., 
5: 49-64), we unfortunately lack the material needed to confirm 
it at the present time. We have no samples from Guatemala 



1959 REVISION OF ANT GENUS NEOSTRUMA 7 

(^ between the ranges of mustelina and hreuicornis) or from 
Avestern Panama (between hrevicornis and zeteki with metopia). 
A second difficulty is that the available material of hreuicornis, 
and especially of mustelina, each representing a wide and varied 
range in Central America and Mexico respectively, shows no 
obvious infraspecific clinal trends corresponding to that seen 
among the several species. Instead, allowing for individual or 
local nest-series variation in size, color and other minor charac- 
ters, the constancy of the "diagnostic" characters holds very 
well, allowing a complete separation of four morphological spe- 
cies. In view of all of the facts outlined, and the various in- 
terpretations possible, I am treating mustelina, hrevicornis, 
zeteki and metopia provisionally as separate species without 
strong prejudice as to their eventual status. 

The last species to be discussed is the remarkable iV^. myU 
lorhapJia, known only from the single type worker from Costa 
Rica. As Figure 4 shows, this species is quite distinct from all 
its congeners. It represents an apparent reversal of the trend 
in Neostruma toward shortening of the mandibles. That this is 
a true reversal is attested by the great elongation of the labral 
lobes. The Neostruma ancester of N. myllorhapha obviously had 
the shortened mandibles, reduced trigger hairs, and elongate 
labral lobes characteristic of all the existing species of the genus. 
When the mandibles started to elongate in the line leading to 
myllorhapha, the trigger hairs did not elongate again, but in- 
stead the labral lobes bearing them were drawn out to an extreme 
length, in this way compensating for the unrecovered loss of 
length of the hairs needed to activate the snap -closure of such 
long mandibles. 



'o 



Neostruma mustelina (Weber) 

Sirumigenys Tnufitclina Weber, 1934, Eevista de Ent., 4:34, fig:. 11, female. 

Type locality: Jicoltepee, Veracruz. Syntypes in MCZ. 

Worker (8 specimens measured from 3 nest series). TL 1.6- 
1.8, HL 0.41-0.46, ML 0.15-0.16, WL 0.43-0.46 mm.; CI 79-81, 
MI 35-39, ICD 48-52. Habitus intermediate between Figure 1 
and Figure 2. The dimensions and proportions overlap those of 
N. hrevicornis, but the tendency in N. mustelina to smaller size, 
relatively broader and deeper head, shorter mandibles, and 



8 BREVIORA No. 107 

deeper promesonotum is evident on comparison of series. The 
space between the apical fork and the submedian tooth is shorter 
than in hrevicornis, and bears only 4 preapical denticles, a 
number quite constant in 55 worker and female specimens exam- 
ined, representing 13 separate nests. These preapical denticulae 
are longer and more acute than is usual in hrevicornis, and the 
third one (counting from the apex toward the submedian tooth) 
is a little larger than its fellows. It is the constancy of the pre- 
apical denticular number in Mexican series which, above all 
other considerations, leads me to recognize mustclina as a species 
distinct from hrevicornis. (Before the recent acquisition of sev- 
eral good Mexican series, I had considered N. niustelina to be 
synonjanous with N. hrevicornis, and had sent out specimens 
labeled accordingly. All such material labeled as "N. hrevi- 
cornis" from Mexican localities should now be considered as 
belonging to N. tnustelina.) 

The color varies from light yellowish-ferruginous to dark 
brown. Other characters as in hrevicornis. 

Female (3 specimens measured from 3 different localities) : 
TL 1.9-2.3, HL 0.48-0.52, ML 0.18-0.20, WL 0.51-0.55, forewing 
L 1.6-1.8 mm. ; CI 83-84, MI 36-38, ICD 49-51. 

Male (1 specimen reared from nest taken at Las Hamacas, 
Veracruz) : TL 1.7, IIL 0.38, HW across eyes 0.38, WL 0.52 mm. 

Habitus much like that of other small strumigenite males ; 
head rather large compared to rest of body. Mandibles simple, 
acute, too small to be opposable. Labrum broad, divided, not 
elongate. Strong longitudinal frontal groove in front of median 
ocellus. Antennae not clavate. Notauli present, moderately dis- 
tinct. Angle of propodeum obtusely rounded; teeth reduced to 
low carinae, rounded above. Petiole low, node rounded ; both 
petiolar and postpetiolar nodes smooth and shining, like entire 
gaster and lower posterior sides of alitrunk. Rest of body reticu- 
late-punctulate, appendages finely punctulate, opaque. Erect 
hairs few and fine, positioned more or less like the clavate hairs 
of the female. Sparse fine reclinate ground hairs on both sur- 
faces of head and on appendages. Color brown, head blackish, 
appendages tan. 

Material examined. 13 nest series, half of them extensive, all 
from Mexico : tamaulipas: Aldama (F. Bonet leg.), veracruz: 



1959 REVISION OF ANT GENUS NEOSTRUMA 9 

Jicoltepec, syntypes of mustelina, collector unknown. Pueblo 
Xuevo, near Tetzonapa, and Las llamaeas, about 17 km. north of 
Santiago Tuxtla (E. 0. AVilson leg. j see notes below). Volean 
San Martin, dark brow^n series, ca. 1100 m., forest debris (C. J. 
Goodnight leg.). Guerrero : Oaxtepec (F. Bonet leg.), ciiiapas : 
Ocosingo and Finca El Real, Ocosingo Valley, leaf mold in hill- 
side (C. and M. Goodnight and L. J. Stannard leg.). 

Several colonies of this ant were taken by Wilson during 
August, 1953, at Pueblo Nuevo and Las Hamacas, Veracruz, in 
tropical evergreen forest. The ants were found foraging in the 
leaf litter and nesting in small rotting bits of wood in the litter. 
One nest (No. 235) contained 24 workers, 1 queen, 4 worker 
pupae and 5 larvae. Another (No. 237) held 26 w^orkers, 1 
queen, 18 worker pupae, 13 larvae of various sizes, and a number 
of eggs. In one natural nest, an entomobryid was found with 
larvae feeding on it. Nests in captivity readily captured ento- 
mobryid Collembola, including one relies ell a-\\\^e species. The 
ants are sluggish hunters, and often show immobile ''ambush" 
behavior, with mandibles held open at about 30° -40° (as in 
Strumigenys gundlachi). When the ant strikes a springtail, the 
springtail almost always jumps, often several times in rapid 
succession. The ant hangs on, bulldog-fashion, and immediately 
curls in a rapid attempt to sting. On one occasion, the collem- 
bolan prey jumped and left behind a leg in the jaws of the ant, 
which bore it back to the nest. Small symphylans, millipedes, 
campodeids, a mosquito, mites and other arthropods offered to 
the Neostruma were ignored or avoided. One small symphylan 
was caught, but later rejected. One nest was maintained in the 
laboratory for two years, during which time it produced two 
separate sets of males and winged females (sexes in about equal 
numbers). From this nest, placed under a bell jar in the sun- 
light, the males and winged females flew out, but remained in- 
active for days on the ceiling and upper walls of the jar, 
apparently without mating. They appeared to try to move to- 
ward the light. During the two years w^e kept this colony in a 
small plaster nest, it was given nothing to eat but entomobryid 
and isotomid collembolaus, a diet which allowed it to maintain 
itself in a flourishing condition. 

One Las Ilamacas colony produced an anomalous individual, 
probably a gynandromorph, in which the right half of the head 



10 BREVIORA No. 107 

and both mandibles are worker, while the left half of the head is 
fully male. 

Neostruma brevicornis (Mann) 

Strumigenys hrevicornis Mann, 1922, Proc. U. S. Nat. Mus. 61(13): 38, 
worker, female. Type locality: La Ceiba, Honduras. Syutypes in ITSNIM, 

MCZ. 

Worker (10 specimens measured from 4 nest series). TL 1.8- 
2.2, HL 0.45-0.53, ML 0.17-0.22, WL 0.44-0.53 mm.; CI 75-80. 
Ml 37-42, ICD 46-50. Habitus intermediate between Figure 1 
and Figure 2. Although its dimensions and proportions overlap 
those of N. mustelina, N. brevicornis is generally larger and has 
a slightly narrower head and longer mandibles, and the head 
and promesonotum are usually a little more strongly depressed. 
The space between the apical fork and submediau tooth is rela- 
tively a little longer and usually bears 5 minute, subequal pre- 
apical denticulae, these appearing smaller than the preapical 
denticulae of N . mustelina. Very rarely, 6 denticulae are pres- 
ent, and one specimen was seen with 5 on one side and 4 on 
the other. 

Medial denticulae 3-5 in number, small and irregular. 

Color varying widely from yellowish-ferruginous to dark 
brown. The dark brown specimens (Tablazo) apparently come 
from a highland area in Costa Rica that has produced melanic 
variants of many wide-ranging dacetine species. 

Female (2 specimens from 2 nests) : TL 2.1-2.3, HL 0.48-0.52, 
ML 0.19-0.20, WL 0.58 mm.; CI 77-81, MI 38-40, ICD 50. 

Color as in corresponding workers or a little darker. 

Material studied: Honduras: La Ceiba (W. M. Mann leg.; 
syntypes). costa rica : Bataan, Abaca Plantation, "in rhizome" 
(C. II. Batchelder leg.). "Costa Rica," without further locality, 
2 separate series (F. Nevermann leg.). Turrucares (A. Bierig 
leg.) Tablazo, dark brown series (A. Bierig leg.). Peralta Sta. 
(P.P.Calvert). 

Neostruma zeteki new species 
(Figs. 1 and 5) 

Holotype worker. TL 2.0, HL 0.53, ML 0.20, WL 0.52 mm., 
CI 68, MI 38, ICD 38. Paratypes : TL 1.8-2.1, HL 0.50-0.56, 
ML 0.19-0.22, WL 0.47-0.53 mm., CI 67-70, MI 37-40, ICD 37-39. 



1959 REVISION OF ANT GENUS NEOSTRUMA 11 

This species is distinguished from brevicornis by its relativelj^ 
narrow head and by the very strong depression of the head and 
alitrunk. The mandibles show 6 to 8 subapieal denticles and a 
quite distinctly enlarged submedian tooth. The medial denticu- 
lar series varies from 4-7 units. The coloration is light to me- 
dium ferruginous, gaster darker brown. The average size is also 
larger than that of hrevicornis, but these two species are very 
similar in all other respects. 

Dealate female. TL 2.5, HL 0.58, ML 0.22, WL 0.63, CI 71, 
MI 38, ICD 41. Male unknown. 

Ilolotype (USNM) : Barro Colorado Island, Panama Canal 
Zone (James Zetek). Paratypes (USNM, MCZ, etc.) Several 
small series, received mixed with other dacetine and basicerotine 
species, mainly collected by Mr. Zetek on Barro Colorado during 
1941-1943. Known only from the type locality. 

Several colonies are represented in the type material, and 
judging from these the species seems very constant and dis- 
tinctive. 

Neostruma metopia ncAv species 

(Fig. 2) 

ITolotype female, dealate. TL 2.2, HL 0.50, WL 0.56, Cf 78, 
MI 32, ICD ca. 50. 

Very close to brevicornis, and similar to the female of that 
species except in the much shorter mandibles and the form of the 
vertico-occipital region of the head. In metopia, the ocellar 
triangle is raised on a blunt ridge or prominence, the latter al- 
most rectangular in lateral view. The ocelli are much closer 
together than in brevicornis; the distance between the anterior 
and each posterior ocellus being 2-21/4 ocellar diameters, while 
in brevicornis, this same distance equals 4-41/2 ocellar diameters. 
The vertex in front of the raised ocellar area falls away rather 
sharply and is here decidedly concave. In full face view the 
prominence appears as a curved, blunt and poorly defined trans- 
verse ridge centering at the ocellar triangle. The small space 
between the posterior ocelli is shining and partly smooth, and 
much of the region inside the triangle is blackened. A single 
small, suberect, spatulate occipital hair rises posterior to, and 
to each side of the ocellar triangle, but there are no other spe- 
cialized erect or suberect hairs on the vertieocciput. The brcvi- 



12 BBEVIOBA No. 107 

cornis female usually has an additional pair of hairs placed even 
with or a bit anterior to the ocelli. This character is, however, 
subject to damage, and should not be trusted on the basis of 
one specimen. 

The preapical denticles are 3 in number on both mandibles. 
Medial denticles 3-4, uneven. Submedian tooth slightly but dis- 
tinctly larger than the largest of the denticles. 

Color medium-light ferruginous. Color sculpture and otlier 
characters as in hrevicornis. 

Holotype (Weber Coll.). Barro Colorado Island, Panama 
Canal Zone (N. A. Weber leg., 1938, no. 871). 

Neostruma crassicornis (Mavr) 
(Fig. 3) 

Strumigenys crassicornis Mayr, 1887, Verb, zool.-bot. Ges. Wien, 37:577, 

worker. Type locality: "St. Catharina." Types in Naturhistorisehes 

Museum, Vienna. 

Worker. TL 2.0-2.3, HL 0.52-0.57, ML 0.18-0.20, AVL 0.47-0.53 
mm., CI 70-75, MI 34-37, ICD 48-50. This species, which ranges 
in the Parana Basin and eastward, is distinguished from the 
four Central American species by its slightly larger size, by the 
peculiar, sublobately incrassate antennal scapes, by its more 
swollen, reniform postpetiole, by the extreme reduction of its 
spongiform appendages, and by the more broadly spatulate 
ground pilosity of the head. 

The preapical mandibular series of denticles varies in number 
of units from 3-7, and the submedian tooth is relatively large 
and distinct (contrary to Emery's figure, otherwise a fairly good 
representation). Color light to medium ferruginous. 

Female (one specimen measured). TL 2.7, HL 0.58, ML 0.22, 
WL 0.62 mm., CI 78, MI 38, ICD ca. 50. 

Material examined. Southeastern brazil : Parecj^ Novo, 2 se- 
ries (Hansen leg., Rambo leg.). Nova Teutonia, S. Catariua (F. 
Plaumann leg.). Argentina: Posadas, Misiones (F. Silvestri 
leg.). 

Neostruma myllorhapiia sp. nov. 
(Fig. 4) 

Holotype worker. TL 2.4, HL 0.58, ML 0.37, WL 0.62. scape 
L 0.23, funiculus L 0.52 mm. ; CI 74, MI 64, ICD 45. 



195!) REVISION OF ANT GENUS NEOSTRUMA 13 

Like hrcvicornis, but differino- in the strikingly elongate, 
coarsely denticulate mandibles and the correspondingly long 
labral lobes and antennal funiculi. Head gently and evenly con- 
vex above, without a distinct impressed area in the center. Ali- 
trunk rather evenly convex in lateral-view })rofile, mesonotal 
area slightly flattened, but not impressed. Sculpture and erect 
hairs as in drevicornis, the hairs rather fine, truncate. 

Color light ferruginous, mandibles and appendages more yel- 
lowish, gaster brown. 

Holotype and sole known specimen labeled simply, "Costa 
Rica" (F. Nevermann leg.), from the collection of Father 
Thomas Borgmeier of Jaearepagua, Brazil. 

ACKNOWLEDGEMENTS 

Thanks are due to Father Thomas Borgmeier, Dr. M. R. Smith 
and Dr. Neal A. Weber for the opportunity to study material 
under their care. Dr. E. 0. Wilson furnished valuable notes on 
the behavior and ecology of N. mnstelina in its natural habitat, 
and furnished captive live nests of this species for laboratory 
study ; he also criticized the manuscript before publication. The 
figures were mostly drawn by Mrs. Nancy Buffler. 



BREVIORA 

Mnasemim of Coimpsiraitive Zoology 



Cambridge, Mass. May 7, 1959 Number 108 



SOME NEW SPECIES OP" DACETINE ANTS 
By William L. Brown, Jr. 

Museum of Comparative Zoology, Harvard University 

The five species of ants described here include two in Colo- 
hosiruma Wheeler (which o-enns I currently consider to include 
Alistruma Brown and Clarkistruma Brown), one in the gund- 
lachi group of Sfruynigenys Fr. Smith, one in Smithistruma 
Brown, and one in Codiomyrmex Wheeler. These descriptions 
are offered in order to validate names used in a general paper 
on dacetine evolution bv E. 0. Wilson and mvself, which we 
hope to publish shortly. The paratype variation and other data 
pertaining to the two Colohostriuna and the S. gundlachi group 
species are to be discussed in revisions of these groups now in 
preparation. Abbreviations for measurements and proportions 
used are given in Brown, 1953, Amer. Midi. Nat., 50 : 7-15 ; and 
Brown, 1953, Jour. New York Ent. Soc, 61: 53 and 101. The 
initials [MCZ] refer to deposition of types in the Museum of 
Comparative Zoology, Harvard University, Cambridge, Massa- 
chusetts. 

COLOBOSTRUMA CERORNATA liew spCcicS 

(Figs. 1, 2) 

Holotype worker. TL 2.5, HL 0.59, HW 0.52 (CI 88), ML 
0.17 (MI 29), WL 0.71, scape L 0.31, greatest diameter of eye 
0.17 mm. 

Shape of head as in Figure 1 (drawn from a paratype) ; seen 
in side view, deep, convex above, with occiput rounded. Man- 
dibles with a stout apical tooth and lesser preapical tooth ; inner 
margins straight, edentate (though appearing falsely denticulate 



BREVIOIJA 



No. 108 



due to integumental microstructui-e) ; beneath the border, how- 
ever, are fine, sharp teeth arranged serially, A'isible only from 
below or when mandibles are open (Pig. 1). 

Eyes very large, with 80-85 coarse facets, set in shallow de- 
pressions or scrobes below cariniform dorsolateral margins of 
head. Antennae with curved scapes to fit around eyes when 
retracted ; funiculus with 3 segments, the second of these long, 
representing the fused segments II, III, and IV of the related 
6-segmented species. Labrum large, covering under-mouthparts 
when retracted, bipartite, tlie two rounded lobes separated by a 
shallow notch. 





Figs. 1 and 2. Colobostruma cerornata new species, worker paratype. 

Fig. 1. Head; full-face (dorsal) view, right antenna and pilosity omitted. 
Fig. 2. Alitrunk, nodes and base of gaster, side view, pilosity omitted. 



Alitrunk as in Figure 2 (drawn from a paratype). Prome- 
sonotum gently convex, weakly submarginate along dorsolateral 
borders; humeral angles prominent, subacute, margined with 
white waxlike material, also edging the anterior pronotal border 
except in the middle. No promesonotal suture ; alitrunk con- 
stricted at distinct metanotal groove. Propodeal lamellae (no 
teeth distinguishable) waxlike, white, thick, soft and irregular, 
readily bent or distorted (as seen in paratypes). Petiole and 
postpetiole with waxlike borders, simulating modest spongiform 
appendages (as in Strumigenys) . Gaster both above and below 
with coarse longitudinal costulae, waxlike at base, the dorsal 
costulae extending more than 1/3 the length of the basal seg- 
ment. Gaster otherwise loosely shagreened, weakly shining, as 
is also the posterior portion of the postpetiolar disc. 



1959 NEW DACETINE ANTS 3 

Head dorsally covered with shallow indistinct foveolae, mostly 
contiguous; the boundaries between these form fine longitudinal 
rugules or eostules. This coarser (but not conspicuous) sculpture 
is overlain by dense, opaque punetulation which also extends to 
the rest of the head ; gula also with loose longitudinal costula- 
tion. Alitrunk, petiolar node, and anterior part of postpetiolar 
disc distinctly longitudinall}' costulate, the costulae often whitish 
and waxlike in appearance, especially on sides of alitruiik and 
extending onto propodoal lamellae. Intercostular spaces, an- 
tennae, legs and mandibles chiefly very finely and shallowly, but 
densely, punctulate-granulose, opaque. 

Head and scapes covered evenly above by numerous whitish, 
suborbicular, subappressed squamose hairs of nearly equal size 
throughout. Hairs on alitrunk dorsum narrower, more cochlear 
and more erect, less abundant, not conspicuous. Numerous short, 
spatulate erect hairs on both nodes and both gastric surfaces, 
curved caudad. Mandibles and legs with short, sparse appressed 
hairs. 

Color dull yellow ; legs, mandibles and antennae light yellow. 

Holotype [MCZ] a worker taken foraging on Dempster Head 
(also called Telegraph Hill), at Esperance, Western Australia, 
February 4-6, 1955 (E. 0. AVilson leg., journal no. 428). Demp- 
ster Head is a large, rounded gneissic-outcrop hill with low 
heath-like shrubs and some dwarf wattles and mallee. Paratypes 
were taken at the type locality and on the sandplain about 8 
km. northeast of the old (abandoned) Thomas River Station, 
about 65 miles (airline) east of Esperance, near the end of 
January, 1955, by E. 0. Wilson and C. P. Ilaskins. All speci- 
mens were taken at night by sweeping the low heath-like vegeta- 
tion. Variation is mostly in size, and this is not particularly 
marked. 

This species is very distinct from all other Coloho struma in 
its very large eyes, small size, and particularly in its peculiar 
pilosity (recalling that of certain Strumigenys species), and its 
waxlike costulation and pedicellar appendages. The 3-segmented 
funiculi are also found in two or three other Coloho struma spe- 
cies. This is one of a pecidiar large-eyed, depigmented fauna 
of ants, including three species of Coloho struma, found foraging 
at night on low heath-like vegetation of the coastal plain of this 
isolated part of Western Australia. 



4 BREVIORA No. 108 

CoLOBOSTRUiMA AUSTRALis iiew species 
(Fig. 3) 

Ilolotype ^vorker. TL 3.2, IIL 0.73, HW 0.66 (CI 90), ML 
0.19 (MI 27), WL 0.85, scape L 0.37, greatest diameter of com- 
pound eye 0.17 mm. 

Head like that of C. aliiiodis (Forel), but shorter; evenly con- 
vex above; lateral lobes behind eyes evenly rounded. Clypeus 
broadly triangular, flat, rounded l)ehind, with slightly convex 
anterior (free) margin mesad. Mandibles triangular, flattened 
above, with slightly down-curved apices ; masticatory margins 
very finely denticulate (as seen when mandibles are fully 
closed) , but ventral to this margin is a row of fine, acute, oblique 
teeth, visible from above only on opened mandible. Apex of 
mandible with one stout tooth and one smaller (preapical) 



Fig. o. Colobofit rama australis new species, worker paratype (Kallista), 
petiole, postpetiole and gastric base as seen from above. 

triangular tooth. Antennae with sigmoidally-curved, incrassate 
(but not lobed) scapes; funiculus witli 5 distinctly separated 
segments. Labrum large, tongue-like, with rounded, entire mar- 
gin. 

Alitrunk with a depressed, nearly flat pronotum having blunt, 
almost rounded humeral angles, not marginate. Mesonotum dis- 
tinct, convex, rising above pronotum and propodeum. Promeso- 
notal suture and metanotal grooves marked by shallow sulci. 
Propodeum downsloi)ing caudad, convex between large, broad 
lamellae, each forming a large acute tooth above, concave below, 
with a ventral projection. 



1959 NEW DACETINE ANTS 5 

Petiolar node subcuboidal as seen from side and above, with 
short, downsloping peduncle, lateral alae, and a short, oblique 
anteroventral spine trailing a narrow, areolate keel. Postpetiole 
shorter and lower in side view, but broader in dorsal view (Fig. 
3). Basal costulae extending about 1/8 the length of the basal 
segment. Gaster otherwise smooth and shining. Head, alitrunk 
and both nodes coarsely but shallowly foveolate, the foveolae 
centrally punctate, most distinct and mostly contiguous on head, 
becoming smaller, less distinct and sparser caudad and on 
clypeus. 

Tnterfoveolar spaces vary from nearly smooth, shining (on 
occiput) to finely shagreened and opaque (nodes, clypeus). Man- 
dibles, antennae, legs, petiolar peduncle finally punctulate- 
granulose, opaque. Lower pleura indefinitely rugulose, opaque. 

Pilosity confined to four stiff truncate and a few smaller 
flexuous hairs at the gastric apex. Legs, mandibles, and antennae 
with very fine, short, sparse reclinate pubescence. Color ferrugi- 
nous yellow, with dark lirown markings, one on vertex, one on 
center of clypeus, extending to frontal triangle, a large diamond- 
shaped spot in the center of the first gastric tergum, flanked on 
each side by a similar spot. This "three diamond" pattern 
occurs also in some other species of the genus. 

Holotype worker [MCZ] one of a nest series taken in a large 
red-rotten log in wet Eucalyptus reg7ians-tree fern forest at 
Kallista, in the Dandenong Range, Victoria, Australia (J. Clark 
and W. L. Brown leg., September, 1950). Several paratype 
series, including males and females, were examined from Kallista 
(J. Clark leg.), Narbethong (J. J. McAreavey leg.), and Fern- 
tree Gully (J. Clark leg.. Brown leg.) in Victoria, and from 
Moss Vale (T. Greaves leg.) and Dorrigo (AV. Heron leg.) in 
New South Wales. This species is obviously widespread in moist 
forests in southeastern Australia. Its nests are normally in rot- 
ting wood. It is distinguished from the other Colohostrimm 
species, described and undescribed, by means of its 6 -segmented 
antennae with non-lobate scape, its "normal" head, with rounded 
lobes in back of the eyes, its partly shining sculpture and its 
predominantly yellow color. 



6 BREVIORA No. 108 

Strumigenys jamaicensis new species 

Holotype worker. TL 2.6, HL 0.54, HW 0.44 (CI 81), ML 0.41 
(MI 76), WL 0.60, scape L 0.37 mm. (SI 84). Resembles 8. 
gundlachi, but larger in size and with relatively much longer 
mandibles and antennae. Mandibles with gently convex outer 
borders and straight inner (masticatory) borders, the latter 
bearing 8 denticles on the left and 7 on the right ; denticles strong 
and acute, occupying about the apical 2/3 of the free inner bor- 
der, roughly decreasing in size from apex toward base of man- 
dible. 

Alitrunk relatively longer than in gundlachi, forming two 
separate convexities divided by the rather deeply impressed 
metanotal groove. Petiole, postpetiole and gaster also a little 
longer and more slender than in gundlachi. Sculpture and pilos- 
ity much as in gundlachi, although the ground hairs tend to be 
less conspicuous in the new species, and the erect hairs larger ; 
gundlachi has more but shorter erect hairs on the gastric dor- 
sum. Color blackish -brown ; mandibles and legs medium brown. 
The color is considerably darker than is usual for gundlachi, but 
samples of gundlachi from Jamaica are also rather dark — 
nearly as dark as jamaicensis. 

Holotype (Collection of the Illinois Natural History Survey, 
Urbana, Illinois) a worker from a series taken at Corn Puss 
Gap (about 2000 feet), St. Thomas Parish, eastern Jamaica 
(H. B. Mills leg.), INHS accession no. 49627, Mills no. J-18. 
Paratypes from the type collection and in two series from Hard- 
war Gap (about 4000 feet), Portland Parish, Jamaica (Mills 
leg.), INHS accession nos. 49613 and 49616, Mills nos. J-4 and 
J-7, deposited in INHS, MCZ and elsewhere. Paratypes are 
similar to holotype ; principal variation is in mandibular teeth, 
varying from 6-8 in number. 

The large size, long mandibles, long scapes and slender alitrunk 
will distinguish this species at once from the other two members 
of the S. gundlachi group on Jamaica, *S'. gundlachi Roger and 
S. eggersi Emery. The long scapes and blackish color, among 
other characters, distinguish S. jamaicensis from the Cuban and 
mainland species of the group. 



1959 NEW DACETINE ANTS 7 

Smithistruma webeki new species 
(Fig. 4) 

Ilolotype worker. TL 2.3, HL (omitting cervical flange) 0.61, 
IIW 0.39 (CI 64), ML (estimated from open mandibles) 0.08, 
WL 0.58, scape L 0.27 mm. 

Head and mandibles as shown in Figure 4; dorsum of head 
gently convex behind in both directions, in side view its dorsal 
outline nearly straight from vertex to clypeus, central or 
"frontal" part of head gently convex from side to side. Clypeus 
with a broad and very shallow transverse impression near mid- 
length ; tumulus of clypeus broad, indicated by dashed lines in 
Figure 4. Eyes fairly large and convex. Mandibles small, with 
basal lamella like that of S. iruncatidens Brown, but more slen- 
der and with broader apical expansion; diastema distinct; first 
tooth of principal series very small, second tooth largest; all 
teeth acute. Scapes incrassate, broadest near the middle, only 
sliglitly bent near their bases. 

Pronotum depressed, broad, gently convex and forming a 
single profile convexity with mesonotum; promesonotal suture 
present but inconspicuous. Seen from above, promesonotum 
broadly oval, separated in center front by a short transverse 
margin from the cervix, but not marginate on sides ; no humeral 
angles. Metanotal groove distinct but shallow; propodeal dor- 
sum slightly convex (straight in profile), terminating in a pair 
of very small triangular teeth subtended by narrow, nearly 
straight infradental lamellae. Petiole with a peduncle about as 
long as its node; node high, rounded, with a flat, smooth, steep 
anterior face defined by weak carinae on both sides and above. 
Free portion of nodal summit seen from above trapezoidal, a 
little broader than long. Postpetiolar node with a convex, smooth 
and shining disc about twice as wide as the petiolar node. Both 
nodes, as well as the anterodorsal gastric margin (but not the 
antero ventral face of the gaster), provided with unusually volu- 
minous spongiform appendages in all the usual positions. Base 
of first gastric segment with coarse costulae extending about 1/3 
the free length of the segment. Gaster otherwise smooth and 
shining. 

Head, alitrunk and petiolar node generally punctate-rugulose, 
this sculpture much coarser and more irregular than in other 



8 



BREVIORA 



No. 108 



Smithistrunia species. Pronotal rugules more longitudinal. The 
mesepisterna and propodeal dorsum are densely punctulate, and 
the metapleura, propodeal declivity, anterior face of petiolar 
node and tumulus of clypeus are mostly smooth and shining. 
Antennae, scrobes, exposed parts of mandibles and legs finely 
punctulate-granulose . 












Fig. 4. Smithifitruma v:chcri lU'w species, holutype worker, full-face view 
of head, showing principal fringing pilosity only. Fig. n. Codiomyrmex 
semicomptus new species, paratype worker with rather narrow head, full- 
face view of head. 



The dorsal surfaces of head, thorax proper, both nodes and 
both upper and lower sides of gaster, and also the anterior bor- 
der of clypeus and scajaes, with long, fine curved hairs. On the 
dorsal surface of the head proper, the 50-60 hairs are erect, 
mesally curved, and have narrowly flattened-clavate apices. On 
the clypeus, there are an additional 80 or more hairs of various 
lengths, predominantly directed anterolaterad, but with their 
apices mostly recurved dorsad and more or less posteromesad. 
The central clypeal hairs have slightly enlarged apices. Hairs, 
more peripherally situated on cranium, scapes and clypeus, are 
finer and tend to have apices merely truncate, rather than en- 
larged (Fig. 4). The long hairs ou the rest of the body become 
progressively finer caudad, and many of them show "furled 



I!).")!) NEW DACETINE ANTS 9 

coaehwhip" looping, or else arch back to the surface. Arched 
and looped hairs are abundant on the o;astric dorsum. Ap- 
pendages clothed with fine arched-reclinate hairs. 

Color light reddish-ferruginous, extremities more yellowish. 

Ilolotype a unique worker from Ango, Belgian Congo, 1948 
(N. A. Weber leg., no. 2170), in the Weber Collection. Several 
years ago I saw another worker of this species from northern 
Angola (Machado leg.) in Dr. F. Bernard's collection; because 
at that time I thought he might eventually describe this species, 
T did not record the details. 

CODIOMYRMEX SEMICOMPTUS UCW SpeciCS 

(Fig. 5) 

Ilolotype worker. TL 2.1, HL 0.56, HW 0.42 (CI 75), ML 
0.10 (MI 18), AVL 0.52, scape L 0.25 mm. Shape of head and 
mandibles as in Figure 5, drawn from a paratype, but slightly 
wider across occiput. Dorsal surface of cranium strongly con- 
vex in the center, sloping toward occiput and anteriad (toward 
clypeus), so that the occipital lobes are narrowly- rounded api- 
cally as seen from side. Mandibles strongly convex, rising above 
the anterior clj'^peal border (as is usual in Codiomyrmex) ; with 
five strong, acute conical teeth, the first (nearest base) and 
fourth a little smaller than the second and third, the apical 
(fifth) tooth smallest. The basal lamella is normally hidden be- 
neath the clypeus at full closure ; it is set at a lower level than 
the principal teeth and oblique to them, and is separated from 
them by a brief diastema; in shape and size it resembles the 
lamella of Smithistruma weheri (Fig. 4), but is sharply truncate 
without being broadened at the apex. Clypeus almost perfectly 
plane. 

Alitrunk narrow, only slightly more than half as wide as the 
head, its dorsum almost flat from side to side and defined b,y 
lateral margins, but forming one gentle convexity from pro- 
notum to propodeal teeth, with only a slight dip at the posterior 
mesonotum. Seen from above, the pronotum is evenly rounded 
and marginate, without humeral angles and only slightly wider 
than the propodeum; promesonotum with a faint median longi- 
tudinal carinula. Propodeum curving evenly down into the 
concave declivity between the dorsolateral margins and the 



10 BREVIORA No. 108 

propodeal teeth ; teeth with horizontal dorsal borders, completely 
involved in the broad infradental lamellae, which are briefly 
concave below the short acnte dorsal tips, then below this broadly 
convex. 

Petiole large, its pednncle rapidly enlarging caudad, bnt dis- 
tinctly differentiated from the large node. Node high in front, 
with a strong, l)lnntly raised anterodorsal margin and steep 
bicarinate anterior face, the dorsal surface then strongly convex 
and sloping candad : node distinctly longer than high, and, seen 
from above, longer than broad, Avith rounded sides and truncate 
anteriorly. Postpetiolar disc broadly subelliptical, broader than 
long and about 1 1/3 times as broad as petiolar node. A full 
complement of voluminous, areolate spongiform appendages 
present in all the usual positions on both nodes and at the base 
of the gaster. G aster depressed, much broader than deep. Basi- 
gastric costulae distinct, crowded, effaced mesally, extending 
about 1/5 the length of the first segment. 

Gaster otherwise, both nodes, entire alitrunk, most of legs, 
anterior frontal area of head, and clypeus smooth and shining 
except for scattered piligerous punctures. Scrobes, upper sides 
of hind coxae and petiolar peduncle densely reticulate-punctate. 
Mandibles, antennae and small parts of legs finely punctulate- 
granulose, subopaque to opaque. Cranium, both dorsal and 
ventral surfaces, loosely and rather coarsely rugose or costulate, 
the rugae running more or less longitudinally. Between the 
rugae are various coarse and fine punctures, rendering the sur- 
face here opaque (more shining near the midline). 

Occiput and rest of dorsal surfaces of body, including the 
entire legs, with numerous long (mostly 0.07-0.10 mm.) fine 
hairs with tapered or truncate tips, becoming more spatulate on 
center of head, and on anterior half of head proper represented 
by shorter, appressed linear-spatulate hairs directed mesad, this 
appressed pilosity repeated in miniature on clypeus. Scapes each 
with a row of short remiform hairs, inclined apicad. Mandibles, 
antennae, gula, and tarsi with fine reclinate hairs. 

Color deep reddish-brown, head and gaster darkest ; clypeus, 
mandibles, antennae and legs yellowish-ferruginous. 

Holotype [MCZ] a worker, chosen from a series of 18 workers 
taken together in heavy, dense eucalypt forest adjoining rain 



1950 NEW DACETINE ANTS 11 

forest at Shipton's Flat, about 20-25 miles soutli of Cooktown, 
Queensland, Australia (P. V. Dar]in<i'ton leg.). The 17 para- 
types are very similar to the holotype, and vary hardly at all in 
measurements ; several are subteneral : these will be placed in the 
MCZ and with other ant eollections. 

This species greatly resembles the larger Sumatran ant 
Wehoistnmia jacohsoni (Menozzi) in shape of head and thorax 
proper, as well as in sculpture and pilosity, but many details 
are different, including the diagnostic shape of the petiole 
(clavate, with only feebly differentiated node in all Wederi- 
struma). From the two other known Codiomyrmex species, C. 
thaxtcri Wheeler (Trinidad) and C. loveridgei Brown (Nyasa- 
land), the new species is distinct in many characters, but it fits 
this genus in formal diagnostic characters and is obviously simi- 
lar in overall habitus. It is becoming clearer with each new 
find that the genera Codiomyrmex, Weheristruma, Glamyro- 
myrmex and their relatives are interrelated. Although each 
species in this relict group is very distinct, the generic limits 
are at present not very satisfactory. 

This capture is a remarkable one, regardless of eventual 
generic placement, because it represents the first short-mandibu- 
late species of subtribe Strumigeniti found on the Australian 
continent. 



BREVIORA 



Musemiinni of Coinipar,ative Zoology 



('AMRRinoE, Mass. May S, 1959 Xitmber 1(H) 



OX THE PIXEAL AREA AXD ADJACENT STRUCTURER 

OF THE HRAIN OF THE DIPNOAN FISH, 

PROTOPTERUS ANNECTEN8 (OWEN) 

By Uno Holmgren^ 

Zoological Institute, Uppsala, Sweden, and 
Biological Laboratories, Harvard University 

INTRODUCTION 

The morphology of the pineal area of the dipnoan fish, Protop- 
terus anticctens, so far has not been described. In connection with 
an investigation of the central nervous system of Protopterns 
atinectens, however, Burckhardt (1890, 1892) gave a brief de- 
scription of tlie dorsal region of the brain, indicating the pres- 
ence of a pineal organ and, interior to that organ, a conspicuous 
structure which he named coronarium. Because of the taxonomi- 
cal position of Proioptcrns, and the considerable variation in 
the morphology of the pineal structures, especially among teleost 
fishes (Studnicka 1905, Tilney and Warren 1919, Friedrich- 
Freska 1982, Rasquin 1958 and others), it seemed of interest 
to describe tlie dorsal differentiations of the brain of this dipnoan 
fish, with special reference to the pineal area, and to compare 
them with the corresponding structures of teleosts. 

Material aud nieihods. P^our specimens of Protopterns annec- 
tens fixed in Bouin's solution, and one specimen in formol-alcohol, 
were investigated. Two heads were sectioned, one transversely 
and one sagittally, at 50 microns, using celloidin technique. In 
the others, the brain was dissected out and cut in paraffin at 
7 microns. The celloidin sections were stained in Mallory's 

1 Fellow of the Commonwealth Fund. 



2 BREVIORA No. 109 

phosphotungstic acid haematoxvlin (Romeis 1948, p. 163), and 
the paraffin sections were stained in Gomori's aldehyde fnehsin 
(Gabe 1953) and in Malloi-y's triple stain. 

Terminologi). Stndnicka (1905, p. 4) defined the pineal area as 
the dorsal parts of the diencephalon from the "commissnra pos- 
terior to the paraphysis." This definition has been adopted 
in this study. Structures situated anterior of the velum trans- 
versum are <>ene]'ally called paraphysis. The latter belongs 
morj^holofiically to tlie telencephalon (Kappers 1957, ]). 52). In 
his description of the central nervous system of Protopterus 
annectens, Burckhardt (1892) called the conspicuous structure 
anterior of the velum transversum, the coronarium, probably 
because of its vascular pattern. However, this structure should 
be called paraphysis (Stndnicka 1905, p. 4). The present study 
has confirmed Studnicka's opinion, for the structure satisfies 
all the criteria of a true paraphysis cerebri, mainly because of its 
position in front of the saccus dorsalis and the velum transver- 
sum but also because of its general structure. The terminology 
is in agreement with that discussed by Kappers (1956). 

DESCRIPTION 

No pineal spot is associated with the pineal area as found in 
certain teleosts, amphibians and lizards. The general morphology 
of the pineal brain area is shown in Figure 1. It includes a 
number of well-defined structures, namely : an organon subcom- 
missuralis, underlying a conspicuous commissnra posterior, a 
commissnra hal)enulae, a pineal organ, and a saccus dorsalis. 

The organon subcommissuralis (Fig. 5) consists of columnar, 
ciliated, secretory cells containing granular Gomori-positive ma- 
terial as in teleosts and other vertebrate classes (Wingstraiid 
1953). The pineal organ, arising from the commissnra posterior 
area, consists distally of an cnd-vesieh^ wliich is attached to the 
brain by means of a proximally divided stalk. The pineal organ 
extends forward and overlies both the saccus dorsalis and the 
paraphysis and is situated close to the sknll roof (Figs. 1, 2). 
The end- vesicle did not penetrate the skull in my material (cf. 
Wiedersheim 1893). 

The histology of the vascularized pineal organ does not differ 
markedly from that found in tclcost fishes, in which mainly two 



1959 



nxKAh aki;a of i'kotopterijs 



types of cells have been observed: the sensory cells and the su])- 
])()rtiiiji- cells. The primary sensory cells ( N. Holin<>ren 1918, 
1!»2U, Friedrich-Freksa 1!>.'}2, Rascinin 1958, and others) in 
Pi-otoj)f(ri(s, as ill most teleosts, can be dilt'erentiated from the 
snpportin<i' cells ( "Stiitzzellen," Friedrich-Freksa 19;}2) which 
show a stron<i' attinity t'oi- the acid fuchsiii of Mallory's stain 
especially the hyperchromatic nnclei of the cells. The cyto|)lasm 
of the sujiportinji' cells is reduced and the cell membranes seem 
almost to enclose nuclei. lIoAvever, compared to teleost fishes, 



Pineal organ 



Commissura posterior 




Org, subcommissuraiis 
O.O 0.^ mm. 



Fig. 1. St-hematical picture of the pineal area and adjacent stnictures 
of the dipnoan fish, Prntopterus annericn.'i. The conspicuous paraphysis 
(Pa) is situated anterior to the velum transversuni (Ve). 



there are very few supporting cells present in the pineal organ of 
Prntopterus. The primary sensory cell-type has a well defined 
nucleus with nucleoli, and the cytoplasm had a distal process simi- 
lar to that of the ami^liibian sensory cell-type (N. Holmgren 
1918, Bargmann 1943). The sensory cells in teleosts have been 
found to be secretory in nature (X. Holmgren. 1920). Since the 
pineal sensory cells resemble both those of the teleosts and the 
amphibians and there is a significant decrease in supporting 



4 BREVIORA No. 109 

glial elements of the pineal organ, one is tempted to regard the 
pineal of Protoptcrus as a type intermediate between that of 
teleosts and amphibians. 

No parapineal organ or rudiment of it eould l)e found in any 
of the five specimens. A parapineal rudiment present in th(^ 
adult has so far not been described in teleost fishes. A parapineal 
organ is, however, ahvays present in the teleost embryo, l)nt it 
degenerates in the adult. 

Anterior to the pineal organ, the commissura habenulae is 
present in the dorsal roof of the diencephalon (Fig. 5), marking 
the limit between the parts of the diencephalon which posteriorly 
give origin to the pineal- and parapineal complex, and the parts 
which anteriorly form the saecus dorsalis and velum transversum. 

In the eonnnissura liabenulae area, it is observed that the right 
nucleus habenulae is larger than the left one in two specimens, 
and of about the same size in one specimen. Similar variation 
has been observed in teleosts. Gierse (1904, p. 618) and Handrick 
(1901, p. 7) reported asynnnetry between the habenular nuclei 
in certain deep sea teleosts. Such individual variation within 
one species was frequently found in a number of teleosts (N. 
Holmgren 1920) ; this fact already had been pointed out by 
Gierse (1904), who found that sometimes the nucleus habemilae 
dexter, sometimes the nucleus habenulae sinister was the larger. 
The significance of this asymmetry has been extensively dis- 
cussed in the literature (Gaskell 1890. Priedrich-Freksa 1982. 
and others). The saecus dorsalis which surrounds the pineal 
stalk consists of neuroependym. wliicli corresponds to the obser- 
vations in teleosts (Studnieka 1905, Friedrich-Freksa 1932). The 
anterior limit between the saecus dorsalis and the paraphysis i> 
formed by a not very well defined velum transversum (Fig. 1 ). 
which indicates the anterior end of the pineal area. 

The paraphysis, shown in Figures 3 and 6, is very nnicii 
folded and vascularized. Vessels with blood corpuscles are 
observed between the folds (Fig. (i ) of the paraphysis, where 
also aldehydefuchsin-positive material was observed (8). 

Although the specificity of the aldehydefuchsin stain is not 
well established so far, it uuiy be of interest to note that similar 
staining reactions were observed in the same slides in the sub- 
commissural organ and also in the nucleus jireopticus secretion 
of the h.vpothalamus. 



1959 PINEAL AREA OF PROTOPTERUS 5 

The descriptions and comparisons presented in this paper 
have shown that there is no considerable difference between 
the pineal morphology of Protopterus and that of teleost fishes. 
The pineal organ of Protopterus resembles that of primitive 
teleosts, especially that of the Clupeidae and Salmonidae. The 
pineal region of Protopteriis also resembles the pineal area of 
other dipnoans such as Ccratodus (Huxley 1876, Studnicka 1905, 
and N. Holmgren and v. der Horst 1925). A well defined pa- 
raphysis cerebri has previously been observed in another dipnoan, 
Ceratodus, and in the chondrostean Acipeyiser (Studnicka 1905). 
In selachians, on the other hand, the paraphysis, although present 
during the early ontogeny, is not found in the adult (Kappers 
1957). In teleosts the conditions are essentially similar, although 
the paraphysis is present in a number of species (U. Holmgren 
1959). 

The resemblance in the histology of the pineal (as described 
above) between Protopterus and the amphibian is not surprising, 
considering the taxonomic position of the lungfish. The presence 
of aldehydefuchsin-positive material in the paraphysis has been 
dealt with by Scharrer (1951), who described the presence of 
such material in the paraphysis of certain snakes. This secretory 
material of the snake arose from the nucleus paraventricularis 
area of the diencephalon. The described aldehydefuchsin-positive 
granules in Protopterus were 2-3 times larger than red blood cells 
and appeared in close association with blood vessels. Their struc- 
ture did not resemble that of blood elements. The origin and 
nature of the aldehydefuchsin-positive material could not be de- 
termined on the limited material at hand. 

SUMMARY 

The pineal area of Protopterus annectens was studied in five 
specimens. Its morphology resembles that of primitive teleost 
fishes. The histology of the pineal cells, on the other hand, is 
similar to that of amphiliians. 

ACKNOWLEDGEMENTS 

Material for this study was provided by Prof. K. (1. Wing- 
strand, Institute of Comparative Anatomy, Copenhagen, who 
also provided laboratory space during the early stage of this 



6 BREVIORA No. 109 

study. I am also much obliged to Mrs. ]\I. Dit-k of the ^Museum 
of Comparative Zoology, Cambridge, for providing additional 
material from the collections of the museum. Dr. Tilly Edinger 
of the Museum of Com])arative Zoology, and Dr. A. B. Dawson 
of the Biological Laboratories, Harvard University, offered valu- 
able suggestions when the manuscript was being prepared. 

REFERP]NCE8 
Bargmann, W. 

1943. Die Epipliysis cerebri. In Haiidb. der mikr. Aiiat. des Menseh, 
6(4): 307-500. 

BURCKHABDT, R. 

1890. Die Zirbel von IcJitJitioplii.s f/liitimisu.s uiid Protnptenis nnnecten.t. 

Anat. Anz., 6:348-49. 
1892. Centrabieivensystems von I'rotdptrrns (iniicctcns. Beiliii. 

Eriedrich-Freksa, a. 

1932. Entwickhing, Bau luid Bcdcutunj;- tier I'arietalfjegend liei Tele- 
ostiern. Zeitsdir. wiss. Zool., 141: .i2-]42. 

(J ABE, M. 

1903. Sur (]uekiue applications de la ct)loration par la fiu-hsin para- 
aldehyde. Bull. :Mici-. Ap])l., 3: ir)3-l()2. 

Gaskell, W. H. 

189(1. On the origin of vertebrates from a crlistacean-like ancestor. 
Quart. Jour. Micr. ScL, 31: 379-443. 

GlERSE, A. 

1904. I'ntersutdiungen iilier das Gehirn and Kopfner\('n von Cyclo- 
flioin (ircli iilih U.S. In (iegenbauer 's Mori)h. Jaliil'., 32: (i()2-687. 

GOMORI, G. 

1941. Observations with dil'feient stains on luinian islets of l.anger- 
hans. Ainer. Jour, i'atli., 17: 39.1-4n(i. 

Handrick, K. 

1901. Zur Kenntnis des Xervensystenis und die LeuchtorKane des 
ArfiyropclccHs hcinuiymnuti. Zoologica, 32: 1-63. 

Holmgren, N. 

1918. Zur KenntiiLs di'r Bai-iet;ilorgane von Hhiki hinponiria. Ark. 

Zool., 11(24) : 1-13. 
192(1. Zur .Anatomic mid Histologie des A'ordcr- nnd Zwiscdiniliirns 

der Kiioriicufisc-hc. Acta Zool.. 1: 137-.31.). 



lf>5f) PINEAL AREA OF PROTOPTERITS 7 

Holmgren, N. and C. J. v. TTokst 

1925. Contribution to tho niorpholofjy of tlio brain of Crratodius. 
Acta Zool., 6: 59-165. 

Holmgren, U. 

1959. On the structure of the pineal area of teleost fishes with special 
reference to a few deep sea fishes. Goteborgs Kungl. Vetenskaps- 
och Vitterhets-Saiuhiilles TIandl. Sjiitte foljden. Ser. B, Bd. 8. 
N:o 3. (Meddel, C^ijteborgs Mus. Zool. Avd., 128). In press. 

Huxley, T. H. 

187(i. On Ceratodus forsteri. Proc. Zool. Hoc. London, 1876:24-51. 

Kappers, a. 

1956. On the development, structure and function of tlie paraphysis. 
In: Progress in Neurobiology'. Proe. 1st interna t. meet, of 
neurobiologists, 1 955 : 130-145. 

1957. On the problem of the presence of a paraphysis cerebri in 
selachians. Pubbl. Staz. zool. Xapoli, 29: 41-67. 

Kasquin, p. 

1958. Studies in the control of pigment cells and light reactions in 
recent teleost fishes. I. Morphology of the pineal region. Bull. 
Amer. Mus. Nat. Hist., 115: 1-68. 

Romeis, B. 

1948. Alikroskopisehe Technik. 15th ed., Munich. 

SCHARRER, E. 

1951. Neurosecretion. X. A relationship between the parai)hysis and 
the paraventricular nucleus in the garter snake, TItamnophis. 
Biol. Bull., 101: 99-106. 

Studnicka, F. K. 

1905. Die Parietalorgane. /n Oppel's Lehrbuch der vergleichenen 
Anatomie der Wirbeltiere, Vol. V., pp. 254. 

TiLNEY, F. AND L. F. WARRBN 

1919. The morphology and evolutionary significance of the pineal 
l)ody. Amer. Anat. Mem., 9: 1-257. 

WiEDERSHEIM, W. 

1893. Grundriss der vergleichenen Anatomie der Wirbolthiere. 3rd 
ed., p. 255. 

Wixgstrand, K. G. 

1958. Neurosecretion and antidiuretic effect with remarks on the sub- 
commissural organ. Ark. Zool., 2: 47-67. 



PLATE 1 

Fig. -. Median section uf the pineal area showing the pineal organ. 
The pineal stalk is sunoundeil on each side by the saecus dorsalis and the 
paraphysis. Fixation Boiiin, ^lallory's jihosphotungstic acid stain, HO 
niierons. 25 x. 

Fig. 3. Cross-section of the brain at the level of the paraphysis. Note 
that the paraphysis (Fa) forms an epiphysis-like structure. Fixation 
Bouin, Mallory's phosphotungstic acid, 50 microns. 30 x. 





I'LATK I 



PLATE 2 

Fig. 4. Cross-SLH-tion of tho diencephulon showing the pineal stalk {P). 
Fixation Bouin, Goniori's aldehyde fuchsin. 100 x. 

Fig. 5. Sagittal section of the commissura posterior area. Note the large 
eommissura habenulae compared to the coininissura posterior. The pineal 
stalk has been removed. Fixation Bouin, Goniori's aldehyde fuchsin. 100 x. 

Fig. 6. A part of the paraphysis showing aldehydefuchsin-positive ma- 
terial (S, arrows). Fixation Bouin Goniori's aldehyde fuchsin. .300 x. 




Organon subcommissuralis 






PLATE 2 



BREVIORA 

Mnasenim of Compsirative Zoology 



Cambridge, Mass. June 16, 1959 Number 110 

THE SPIDEK GENUS C0LE080MA (ABANEAE, 

THERIDIIDAE) 

By Herbert W. Levi 

This paper is one of a series describing American spiders of 
the family Theridiidae and redefining theridiid genera. The late 
Miss Elizabeth Bryant revised the genus Coleosoma (1944, 
Psyche, vol. 51, pp. 51-58), and my paper is intended to comple- 
ment hers by re-illnstrating the species and giving their distri- 
l)ntions. 

I should like to thank Dr. AV. J. Gertsch of the American 
Museum of Natural History, Dr. R. V. Chamberlin, University 
of Utah, and Dr. A. M. Cliickering, Albion College, for making 
collections available to me. The work vas done with the help of a 
National Institutes of Health grant (E-1944). A National Sci- 
ence Foundation grant (G-4317) made possible the examination 
of types in European museums. I extend my thanks also to Prof. 
G. C. Varley for his hospitality at Oxford University, and to 
Prof. T. Jaczewski and A. Riedel of the Polish Academy of Sci- 
ences, Warsaw, for the loan of valuable specimens. 

Coleosoma 0. P. Cambridge 



!^^ 



Coleosoma O. P. Cajiibridge, 1882, Proe. Zool. Soc. Loiidou, p. 426. Type 
species by monotypy: Coleosoma l>la7idum O. P. Cambridge. 
Small (less than 3 mm. total length) theridiid spiders. Eyes 
\er\' small, eyes of posterior row one to two diameters apart. 
Clypeus projecting. Chelicerae without teeth on anterior margin 
or with one tooth. First leg longest ; in females fourth leg next in 
length, in male second leg. Tarsal comb present but indistinct. 
Abdomen sometimes modified in female (Fig. 6), always modi- 
fied in male. Colulus absent. 



2 BREVIORA No. 110 

Epioynum very weakly selerotized, openings indistinct, quite 
variable. Palpus with a functional median apopylisis (M in Fig. 
16) ; a selerotized radix (R), a weakly selerotized conductor (C) 
and a thread-shaped embolus (E). 

The males of Coleosoma have a selerotized ring around the 
anterior of the abdomen ; the ring continues as a ventral shield 
and has a pair of lobes on the dorsum (Figs, -l, 9, 10, 15). In 
some species the abdomen may be constricted (Figs. 9, 10). These 
characters separate the males of Coleosoma from both Chrysso 
and Theridion. Unfortunately I know of no way to separate fe- 
males from Theridion. Coleosoma is not close to Lithyphantes 
[=^Steatoda] as Miss Bryant thought. 

Simon incorrectly synonymized Coleosoma with llieridion, 
causing confusion. The catalogers, Petrunkevitch, Roewer, and 
Bonnet, have unfortunately followed Simon. 

It is of interest that although the few species known to belong 
to Coleosoma are very closely related, and the males are difficult 
to separate, the species C. foridanum has the epigynum (Fig. 14) 
and internal genitalia (Fig. 13) entirely different from the other 
two. While C. acutivenier and C. normale have similar genitalia 
in the female as well as in the male, thev differ greatlv in the 
abdominal shape of the female. The abdominal shape is usually 
very conservative and often a good generic character in Theri- 
diidae. This situation emphasizes the importance of having both 
males and females for correct generic placement, and of using 
more than one character for generic diagnosis. 

Besides C. hlandum of Ceylon, there are three additional species 
known, all occurring in America. One of these, C. fioridanurn. 
is cosmotropical and has been given numerous names. 

As Miss Bryant (1944) made ade([uate descriptions of species, 
this has been omitted here. 

Key to American Species of Coleosoma 

Males 

la. Abdomen with a distinct iiu'dian constriction (Figs. 9, IUk Palpal 

embolus of median length (Fig. 11) acutiventer 

llj. Abdomen Avithout median constriction (Figs. 4, 15) 2 

-a. Palpus narrow, enil)olus sliort, radix projecting (Fig. 17) fiorulanuni 
2b. Palpus wide, embolus long, radix not projecting (Fig. 5) jwrmalc 



1959 SI'IOER OENTS ('OLKOSOi\[.\ 3 

Females 

la. Abdomen with an extension hc.yond the spinnerets, all dark gray or 
dorsum dark gray in color (Fig. 6) acutiventer 

II). Abdomen shape not modified, usually light in color except for dorsal 
stripes 2 

2a. Epigynum an indistinct white transverse depression divided into three 
portions (Fig. 14) floridanum 

2h. Epigynum with two indistinct longitudinal marks (Fig. 3) . nnrmale 



CoLEOSOMA BLAXDUM 0. P. Cambridge 
Fig. 18 

Coleosovia hlandum O. P. Cambridge, 1882, Proe. Zool. Soc. London, p. 427, 
pi. 29, fig. 3. $ type from Ceylon, in the Hope Department of Entomol- 
ogy, Oxford University, examined. 

Theridion vituherabile Petrunkevitch, 1911, Bull. Amer. Mus. Xat. Hist., vol. 
29, p. 210. New name for CoJeosmna hlandmn thought to lie preoccujiied 
by Theridion bland wm Hentz. 
This species is known only from the type specimen. 

CoLEOSOMA NORMALS Bryant 
Figures 1-5; Map 1 

Coleosoma normale Bryant, 1944, Psyche, vol. 51, p. 56, figs. 2, 5, 8, 10, $, 
$. S type from Fort Myers, Florida, in the Museum of Comparative 
Zoology, examined. 

The embolus of a male frojn Arizona is shorter than that from 
Florida specimens. 

Hahiis. On shrubs; in dune grass (North Carolina). 

Records. North Carolina. Carteret Co.: Bogue Bank (K. D. 
Barnes) ; Beaufort (R. D. Barnes) ; Carrot Isl. (R. D. Barnes). 
Florida. Alachua Co. (H. K. Wallace). Dade Co.: Kendall (A. 
M. Nadler). Highlands Co.: 3 mi. S. of Lk. Istokpoga (A. M. 
Nadler). Indian River Co. : Sebastian (G. Nelson). Pinellas Co. : 
Pass-a-Grille (A. M. Nadler). Sarasota Co.: Miakka Riv. State 
Pk. (W. J. Gertsch). Arizona. Virgin Narrows (W. Ivie). 
Yuma Co.: Mittry Lk. (V. Roth). Mexico. Colima: Cuyutlan 
(F. Bonet). Nayarit: San Bias (C. and M. Goodnight), p'uehla: 
Tlacotepec (V. Roth, W. J. Gertsch). Guerrero: Acapulco (L. I. 
Davis). British West Indies: Great Bahama Isl., Pine Ridge 
(E. B. Hayden) ; South Bimiiii (A. M. Nadler). Venezuela: 
Rancho Grande (A. M. Nadler). 



4 BREVIORA No. 110 

CoLEOSOMA ACUTivENTER (Keyserling) 
Figures 6-11 ; Map 1 

Achaea acutiventer Keyserling, 1884, Die Spinnen Anierikas, Theiidiidae, 
pt. 1, p. 113, fig. 74, 2 . 9 type from ' ' Maragnioe, ' ' Peru, in the 
Polish Academy of Sciences, Warsaw, examined. 

Coleosoma flavipes O. P. Cambridge, 1895, Biologia Centrali Americana, 
Araneidea, vol. 1, p. 154, pi. 19, fig. 12, 5 . $ syntypes from Teapa 
[Tabasco, Mexico], in the British Museum, INTatural History. Bryant, 
1944, Psyche, vol. 51, p. 52, figs. 1, 4, 7, 9, 5 , $. NEW SYNONYMY. 

Argyrodes floridana Banks, 1900, Canadian Ent., vol. 32, p. 98. $ syntypes 
from Punta Gorda, Florida, in the Museum of Comparative Zoology, 
examined. NEW SYNONYMY. Not Coleosoma floridana Banks, 1900, 
described on same page. 

Chrysso nigripalpis Banks, 1929, Bull. Mus. Comp. Zool., vol. 69, p. 85, figs. 
46, 72. 9 syntypes from Barro Colorado Island, Panama Canal Zone, 
in the Museum of Comparative Zoology, examined. NEW SYN- 
ONYMY^ 

Adhaea index Chamberlin and Ivie, 1944, Bull. Univ. Utah, bid. ser., vol. 8, 
no. 5, p. 36, figs. 87, 105. 5 type from Brier Creek, 7 mi. north of Syl- 
vania, Georgia, in the American Musemn of Natural History. 

Archer (1946, Pap. Alabama Mus. Nat. Hist., no. 22, p. 20) 
reports this species as having been collected from A^arious habi- 
tats : sifting leaves from woods and on walls next to roofs in out- 
houses. 

Distrihutioit. Southeastern Ignited States to South America. 

Records. Florida: Collier Co.: Everglades (A. M. Nadler). 
Hardee Co.: Oua (W. J. Gertsch). Highlands Co.: nr. Sebring 
(W. J. Gertsch) ; Lake Placid (A. M. Nadler). Indian River Co. : 
Sebastian (G. Nelson). Lee Co.: Ft. Myers (W. B. Barrows). 
Orange Co.: Orlando (M. Nirenberg). Seminole Co.: Geneva 
(W. J. Gertsch). Volusia Co.: Deland (Lutz). Alabama: 
(Archer, 1946). Baldwin Co.: Silverhill (G. Nelson). Missis- 
sippi: Wilkinson Co.: Centreville (A. F. Archer). Texas: Cam- 
eron Co.: Big Tree Vine Assoc. (L. L Davis) ; Ilarlingen (L. 1. 
Davis) ; Edinburg (S. Mulaik) ; nr. Rio Hondo (L. I. Davis) : 
Brownsville (S. Mulaik). Mexico. Tamaidipas: Tampico (L. 1. 
Davis) ; nr. Nueva Morelos (A. M. Davis) ; Reynosa (S. Mulaik) : 
San Fernando (L. I. Davis). San Luis Pofosi: Tamazunchalc 
(F. Bonet). Colima: Las Humedades, Armeria (F. Bonet). 
Puehla : Huauchinango (H. M. Wagner). Veracruz: Tecolutla 



1959 



SPIDER GENUS COLEOSOMA 




be 
•S 

<D 

M 



Si 



o 



OS 









o 

O 

o 

O 

o 






6 BREVIORA No. 110 

(H. M. Wagner) ; Jalapa (J. C. and D. L. Pallisterj ; Orizaba 
(F. Bonet) ; Tuxpan (H. M. Wagner). Oaxaca: Playa Ilati. Rid 
Touto (H. Wagner). Chiapas: Caealiuatau (C. and M. Good- 
night). Guatemala: Moca (C. and P. Yanrie). Nicaragua: Musa- 
was, Waspuc River (B. Malkin). Costa Rica: Turrialba (F. 
Sehrader). Panama: Boquete (A. M. Chickering). Canal Zone: 
Barro Colorado Isl. (very common); Summit; (Jatnn; Experi- 
mental Gardens; Forest Reserve; Pedro Miguel (all A. M. Chick- 
ering). Bahama Isl.: North Bimini (A. M. Nadler) ; New 
Providence Isl. (E. B. Hayden). Cuba: nr. Havana. Venezuela: 
"in orchids from Venezuela"' (G. Becker). Colombia: [no local- 
ity]. Ecuador: [no locality]. Brazil: Rio de Janeiro (H. Sick) 
determination doubtful — lacks posterior extension on abdomen. 

("OLEOSOMA FLOKiDANUM Banks 
Figures 12-17; Map 2 

Coli'o.soiiui floriildua Banks, 1900, Canadian Eut., vol. 32, p. 98. 6 syntypes 
from Puuta Goida, Florida, in the Museum of Comparative Zoology, 
examined. Biyaut, 1944, Psyt-he, vol. 51, p. 54, figs. 3, 6, 9 , 6 . 

Theridion intevruptum Banks, 1908, ihid., vol. 40, p. 205, fig. 9. 9 type 
from Miami, Florida, in the Museum of Comparative Zoology, exam- 
ined. 

Li/ht/pJiaiites uophorns Petrunkeviteh, 1930, Trans. Connecticut Acad. Sci., 
vol. 30, p. 170, figs. 8, 9. 9 type from near Aguas Bueuas, Puerto Rico, 
probably lost. NEW SYNONYMY. 

Tlicridinn dclthile Petrunkeviteh, 1930, ibid., vol. 30, p. 206, figs. 53-5G. 
9 type from near Rio Piedras, Puerto Rico, probably lost. NEW SYN- 
ONYMY. 

I Tlierldion epiensis Berland, 1938, Ann. Soc. Eut. France, vol. 107, p. Kill, 
figs. 107-109, 9. 9 type from Epi, New Hebrides, in the Museum 
Nationale d'Histoire Naturelle, Paris. NEW SYNONYMY. 

Theridion nipanac Berland, 1942 Occas. Papers Bernice P. Bishop Mus., 
vol. 17, no. 1, p. 15, fig. 6, 9, £. o holotype from Rapa, Polynesia, 
probably in the Bernice P. Bishop Museum. NEW SYNONYMY. 

Tlicridion albovithil idh Caporiacco, 1955, Acta Biol. Yenezuelica, vol. ], 
p. 334, fig. 25. 9 type from Barcelona, Anzoategui, Venezuela, in the 
Caracas Museum, X'enezuela, examined. NEW SYNONYMY. 

'I'hcvidion ahipuld ^larples, 1955, .lour. T>inn. Soc. London, vol. 42, p. 483, 
1)1. 58, figs. 9, 13, 19, 9, 6. 9 type from Aleipata, Samoa, jirobably 
in the Otago Museum, Dunedin. NEW SYNONYMY. 
The type of Theridion albovittatum was examined ; it is all 

slii-i\-('l]('(l up Mild (lifhcuh til place. (';ii)()i'i;i('c(>"s drjiwiugs and 



1959 



SPIDER GENUS COLEOSOMA 




re 
W 



o 



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o 

o 
'o 



CO 

s 



8 BREVIORA No. 110 

proportions leave no doubt on the synonymy. The internal geni- 
talia are typical. 

There are probably numerous synonyms of this widespread 
common species in the literature ; however the poor descriptions 
of many species of "Thcridion" make it difficult to synonymize 
the names without examining the types. 

Hah its. This species has been found under stem of dead coco- 
nut leaf and under a rock (Petrunkevitch, 1930). Its habit of 
clinging to vegetation facilitates its world-wide distribution with 
cargo. Archer (1946, Pap. Alabama Mus. Nat. Hist., no. 22, p. 19) 
reports sifting it from leaf litter. 

Distribution. Cosmotropical (Map 2), probably distributed by 
man. It may be native in America. 

Records. Massachusetts: On bananas in Cambridge. New Jer- 
sey: Ramsey (W. J. Gertsch) ; probably an accidental introduc- 
tion. Florida: Alachua Co.: Gainesville (AV. J. Gertsch). Char- 
lotte Co.: Punta Gorda (S. Rounds). Dade Co.: Kendall (A. M. 
Nadler) ; Perrine (A. M. Nadler). Monroe Co. : Key West (A. M. 
Nadler) ; Stock Isl. El Salvador: San Salvador (J. B. Boursot.) 
Costa Rica: [no locality]. Panama Carial Zone: La Boca; Pedro 
Miguel; Corozal (all A. M. Chickering). Bahama Isl : North Bim- 
ini (A. M. Nadler) ; South Bimini (C. and P. Vaurie). Ciiha: 
Havana (Baker) ; Soledad (L. G. Worley) ; Ote, Chirivico (A. F. 
Archer). Jamaica: St. Ann's Bay; Kingston; Discovery Bay; 
Whitehouse; Holland Bay (all A. M. Nadler). Haiti: Trou Cai- 
man (P. J. Darlington) ; Grand Riviere ( AV. M. Mann) ; Diquini 
(W. M. Mann); Ennery (P. J. Darlington). Dominican Rep.: 
San Jose de las Matas (P. J. Darlington) : Puerto Plata (P. J. 
Darlington) ; nr. Ciudad Trujillo (A. M. Nadler) ; Boca Chiea 
(A. M. Nadler). Puerto Rico: El Yunque : Mayaguez; Cidra, 
Treasure Isl. ; Rio Piedras (all A. M. Nadler). Martinique: Fond 
la Haye (A. M. Nadler). Trinidad: Piarco (A. M. Nadler); 
Navy Base (R. Ingle). Venezuela: [no locality]. Colombia: 
[no locality]. Peru: Tingo Maria, liTO m (\V. AVeyrauch). Gala- 
pagos Isl.: Floreana. Togo: Ho, taken in Wisconsin in package 
of ])ressed plants (G. Naomasi). India: Taken in Honolulu, 
Hawaii in imported plants, ••on cargo from Calcutta," (T. F. 
Chong). New Hebrides: [no locality] (-1. S. Haeger. W. R. Emis). 



i"Hgs. 1-5. Coleosonui normale Bryant. 1. Female from side. 2. Female 
genitalia, dorsal view. 3. Epigynuni. 4. Male abdomen from side. 5. Left 
palpus. 

Figs. G-11. C. ucutiventer (Keyserliug). (i. Female from side. 7. Female 
genitalia, dorsal view. 8. Epigynuni. 9. Male abdomen from side. 10. Male, 
dorsal view. 11. Palpus. 

Figs. 12-17. C. floridanutn Banks. 12. Female abdomen, dorsal view, 
lo. Female genitalia, dorsal view. 14. Epigynuni. 1.5. Male abdomen from 
side. 16. Palpus expanded (C, conductor; E, embolus; M, median apophy- 
sis; R, radi.x). 17. Palpus. 

Fig. 18. C. bland iiui O. P. Cambridge, palpus. 



BREVIORA 

MniseiuitM of Comparative Zoology 



Cambridge, Mass. June 17, 1959 Number 111 



ON THE CAUDAL NEUROSECKETORY SYSTEM OF THE 
TELEOST PISH, FVNDIJLVS HETEROCLITUS L. 

By Uno Holmgren ^ 

Zoologifiil Institute, Uppsala, Sweden 
Biological Laboratories, Harvard University 

INTRODT'CTION 

It Avas early observed that a bulb-like structure was associated 
with the terminal portion of the spinal cord of certain fishes 
(AVeber 1S27, Rauber 1877, Arsaky 1813, Ziehen 1903, and 
P'avaro 1925). This outgrowth of the spinal cord of different 
systematical g-roups of teleosts was described by Favaro (1925), 
who studied a large number of species. Because of the histologi- 
cal and morphological resemblance of this organ to the neuro- 
hypophysis, he named it "Ipofisi caudale." 

Secretory cells in the terminal portion of the spinal cord were 
described first by Dahlgren (1914), and later by Speidel (1919, 
1922). Tlie latter also gave a histological description of some 
"irregular glandular cells," sometimes called " Dahlgren 's 
cells," in the terminal portion of the spinal cord of certain 
elasmobranchs and teleosts. Recently Enami (1955 a) investi- 
gated the caudal portion of the spinal cord of the eel, Anguilla 
japonica, with respect to the secretory activity of " Dahlgren 's 
cells" and found that neurosecretory cells were present in the 
terminal portion of the spinal cord. They extended caudally 
from the level of the last sixth or seventh vertebra. These neuro- 
secretory cells, according to Enami, resembled the secretory cells 
of the hypothalamus, and had axons which served as neurosecre- 



1- FflldW (if the t 'oiiniioinvcaltli l'''ini(1. 



2 i;i;eviora No. Ill 

tory patlnvays. The neuTosecretory niaterial was roleased at 
nerve endings, which terminate in a ventral outgroAvth of the 
s})inal cord. The outgrowth served as a storage depot for secre- 
toi-y material. This sti'ucture, which was named "Ipofisi can- 
dale" by P'avaro (1925), was called "Neurohypophysis spinalis'" 
or ■' rrohyjiophysis" by p]nami (1!)55 a). 

These observations were extended by Enami and Imai (1955. 
1956 a and b), with descriptions of the candal neurosecretory 
system of a luimber of freshwater and marine teleosts. Recently 
Sano (1958 a and b) described the " Neurophysis " and the neuro- 
secretory system of the spinal cord of Tinea ruJgaris and a num- 
ber of other teleosts. It has been suggested that the probable 
function of the caudal neurosecretory system would be osmoregu- 
lation (Enami. ]\Iiyashita and Imai 1956, and Holmgren 1!)5S b». 

Terminolofjij. Different names have been suggested for the 
above-mentioned outgrowth of the spinal cord, which servers as 
a storage organ for the neurosecretory material. Because the 
organ may be located either dorsal, ventral, or lateral to the sjiinal 
cord (Favaro 1925, Sano 1958 b), and since the term "neuro- 
hypophysis" has already been used for the "pars nervosa" of 
the pituitary gland, the name "neurohypophysis spinalis" is not 
adequate. In order to stress the fact that the organ is an ont 
growth of the terminal jiortion of the spinal cord the name 
Kroph I/sis spinalis'^ has been ado])ted for this study. This name 
stresses the position of the organ relative to the body, regai-dless 
of its varying position in relation to the spinal cord. 

Material and nufhods. Twenty specimens of the teleost tish. 
Fundulus heteroclitiis L., standard length 4-() cm., caught during 
different times of the year, were used for the anatomical descrip- 
tion. TAvelve end)ryos were also used in an attem])t to determine 
the mode of development of the urophysis. For the anatondcal 
and histological descriptions, the terminal portion of the spinal 
cord, including tli(> adjoining vertebral column, was fixed in 
Bouin's solution to which 1 per cent of CaCl2 had Ix-cn adde;! : 
Zenker's, Carnoy's and Orth's fixing fluids were also used. Tlic 
tails Avere decalcified in 7 ]ier cent IIXOm and aftci'wai'ds treate;! 
in 10 i)er cent Xa^SO-i, and the sections were cnt a1 4-S mici-ons. 
sagittally and transversely, using the usual pai-affin technique. 

1 SusKestc'l liy l»r. (J. I'riillnTL'. Zii(ilo'_'-ical Iiistitnlr. SIcirUlKilin. 



1059 CAUDAL NEUROSECRETORV SYRTEIM 3 

Tlic i'ollowiiifi- staiiiin.u' t('('liiii(|ii('s wei-e used: (ioinori 's liaema- 
toxylin-phloxin method, Ilcideiilmiii's azaii, llalnii's aldehyde 
tuehsin following performie acid oxidation (Ilolmoren 1958 a), 
Mallory's connective tissue stain, Bodian's ]m)taro'ol method, 
Masson's trichrome stain according- to Gomori (1950), periodic 
acid-Schiff (PAS), and alcian blue. 

DE8CKIPT10N 

Location of the urophysis spinalis. The nrophysis in Fundulus 
is observed as a round body, easily visible under the dissecting 
microscope. It lies at the end of the spinal cord, ventral to the 
terminal portion, above the articulation between the last vertebra 
and the hj-pural bone. This location of the urophj^sis (Figs. 1-4) 
is similar to that described for a great number of teleosts by 
Enami and Imai (1956 a and b), and Sano (1958 a and b). 
These authors also found that the nrophysis is generally situated 
ventral to the spinal cord. 

Morphology and histology. According to descriptions by Fa- 
varo (1925), tlie nrophysis consists mainly of modified glial 
tissue. The blood vessels present in the organ enter from the 
connective tissue mantle which surrounds the spinal outgrowth. 
Rauber ( 1877 ) , on the other hand, believed that the nrophysis 
consisted mainly of connective tissue. Enami (1955 a) found 
both connective tissue and modified glial tissue in the organ, and 
stressed the similarity of the nrophysis to the neurohypophysis 
of vertebrates and the sinus gland of Crustacea. 

The nrophysis spinalis of Funduhis heteroclitus is a very con- 
spicuous structure measuring about 0.2 x 0.3 mm., which ap- 
proximately corresponds to the size of the pituitary gland. It is 
covered by a heavily pigmented leptomeninx. The histological 
elements distinguished in the nrophysis are mainly of three kinds. 
In the dorsal region, nerve fibers are found which emanate from 
the neurosecretory cells, situated in the terminal portion of the 
spinal cord (Fig. 1). Orange G positive Herring body-like secre- 
tory droplets are present at the end of these axons (Fig. 3). 
These 'secretory droplets' are probably dilated nerve endings. 
The colloidal masses within the dilated nerve endings stain red 
in Gomori 's chrome haematoxylin-phloxin, thus showing affinity 



4 BREVIORA No. Ill 

for the phloxin component. The staining' reaction with azan is 
blnislired. The nerve tracts are nsnally not clistribntecl all over 
the organ as observed in some other species but merely confined 
to the dorsal region of the urophysis. 

The nrophysis is very vascular with blood vessels distributed 
throughout the organ (Figs. 4, 6). These vessels are of sinusoid 
type with thin walls, supported by argyrophilic connective tissue 
as shown with Bodian's protargol method. In the dorsal region 
of the urophysis the blood vessels frequently seem to surround 




OTTI^ Neurosecretory cells 

,1 mm. ' 



Fig. 1. Sclieniatical picture of the caudal neurosecretory system of the 
teleost. Funduhis licferoclifus L. 

intercellular sjiaces, which are most conspicuous in the area 
where the secretory tracts terminate. Besides the nerve endings, 
the urophysis consists of (n-f/ipapliilic connective tisfiuc and neu- 
roglia. The gli.d cells contain very little cyto])lasm, sometimes 
consisting only of small nuclei and surrounding membranes. The 
differentiation lietween the different types of tissue elements is 
especially well observed after using Heidenhain's stain. The 
distribution of the blood vessels indicates that the secretory ma- 
terial is taken up l)y the blood. 

Transverse sections show that the uro])hysis is surrounded liy 
a sella turcica-like connective tissue capsule (Fig. 4). This ex- 
tension of the leptomeninx is sometimes very heavily invested 
with melanophoi-es. In other teleosts, the urophysis may be at- 
tached to the spinal cord by means of a stalk, which sometimes 
is very conspifuoiis (Eiiami ;md Inuii 1956 a and 1), ?>ano 1958 b). 



1959 CAUDAL NEUROSECRETORY SYSTEM 5 

The nr()i)hy.sis of FiokIhIus lieteroclitus is very closely attached 
to the spinal cord by means of a stalk with a broad base. Most 
of the org-an, however, is separated from the spinal cord. Cross- 
sections of the urophysis (Fig. 4) show a well defined space be- 
tween the dorsal part of the organ and the ependymal parts of 
the spinal cord. Reissner's fiber is still present in this thin ter- 
minal portion of the spinal cord. 

In tlie region where the spinal curd axons i)ass into the nro- 
physis the secretory droplets are especially frequent (Fig. 3). 
The secretory material within the dilated nerve endings is here 
homogeneous and appears colloidal. The staining reaction also 
seems to have changed in a 'basophilic' direction (cf. below). 
Some of the secretory droplets were slightly aldehyde fuchsin- 
positive in the dorsal region of the urophysis, but the neurosecre- 
tory cells stained negatively. 

The neurosecretory cells. The neurosecretory cells are present 
in the terminal portion of the spinal cord, appearing first at the 
level of the sixth vertebra from the caudal end and most concen- 
trated above the third vertebra and caudally (Figs. 1, 5). The 
secretory cells, which contain Xissl substance, are obviously spe- 
cialized nerve cells of multipolar, bipolar, and also unipolar type. 
The neurosecretory cells stain negatively with both Gomori's 
chrome haematoxylin and Halmi's aldehyde fuchsin. Instead, 
they take up the pliloxin component of Gomori's stain and also 
show great affinity for the acid fuchsin of ]\Iallory's stain, thus 
confirming the findings of Enami (1955 a). 

The neurosecretory cells are also stained with the azocarmine 
of Heidenhain's stain thus .showing a marked 'acidophilic' reac- 
tion which may be due to secretory- material in the cytoplasm 
(cf. below). The spinal neurosecretory cells were observed to 
contain scattered small PAS-positive inclusions, but the secretory 
droplets within the urophysis stained negatively. A positive 
reaction would have denoted the presence in the secretion of 
considerable amounts of polysaccharides, mucopolysaccharides, 
glucoproteins or giycolipoids (Lillie ]954). The negative reac- 
tion to PAS agrees with the findings of Sano (1958b). 

The neurosecretory cells have various sizes and shapes. Those 
which are situated closer to the urophysis, are generally smaller. 
The secretory cells are generally distributed all over the spinal 



6 BREVIORA No. m 

cord which in tho tenninal portion does not show the nsual sep- 
aration into white and gray material. The secretory cells were 
found to contain a larjie nucleus with an ii-regular shape, as 
already indicated in the early descriptions by Speidel (1919, 
1922). Tlie size of the neurosecretory cells in FiduIuIks, how- 
ever, was considerably less than the ori<i'inally described spinal 
cells in skates and the eel (Speidel 1919, Enami 1955a). The 
secretory niatei-ial usually contained very fine granules, and was 
generally distributed all through the cytoplasm but sometimes 
was merely confined to the periphery of the cells. The irreg- 
ular nucleus contains one or more nucleoli and fine granular 
chromatin. 

In the sections stained with lleideidiaiu's azan, it was easy to 
distinguish tlie neurosecretory cells, with their 'acidophilic' re- 
action, from the strongly 'basophilic' motor cells. Because of the 
irregular shape of the nuclei, sometimes two or several nuclear 
areas were visible when a cell was cut in a certain plane. In the 
cytoplasm of the secretory cells the 'acidophilic' material was 
observed in various amounts, indicating a cycle in the formation 
of the secretion. The 'acidophilic' reactioii was interpreted as a 
measure of the secretory activity. Nerve cells in general and 
some neurosecretory ones from the hypothalamic system may 
show a 'basophilic' reaction (Seharrer and Scharrer 1954), per- 
haps due to a continuous synthesis of proteins, hormones and 
high amounts of ribonucleic acid. Observations on cells in dif- 
ferent stages of secretion have indicated that the cell in the be- 
ginning of the secretory phase does not show a considerable 
'acidophilic' reaction. As secretion begins the luu-lei of the cells 
begin to enlarge and 'acidophilic' granulated material arises in 
the area around the nucleus. Later, the secretory material is 
often found in the periphery of the cells or distributed all 
through the cytoplasm. It has been suggested that the heavy 
'acidophilic' reaction particularly in the periphery of the neuro- 
secretory cells is due to accumulation in these areas of 'acido- 
philic' material and Nissl-substance (>Sano 1958 b). The ob- 
servations in this study suppoi't the suggestion by Sano (1958 b) 
that the nucleus takes an active part in the formation of the 
secretory material. No selective stain is so far known for the 
secretoi'v material. 



19.')fl CAinAI, XKIROSECRETOKY SVSTKM 7 

The secretury material of tin- lUTve cells is traiisi)orle(l in the 
axons whieh serve as netirosecretory pathways and is stored in 
dilated nerve endiniis jiiid released in the dorsal region of the 
iirophysis where the axons terminate. Observations were made 
on the staining reaction of the secretory droplets to acid alcian 
bine stain. The observed negative reaction wonld mean that the 
secretion does not contain disnltide groups. These observations 
agree with those by Sano (1958 b), who found that the secretion 
in the caudal neurosecretory system of Tinea vulgaris reacted 
negatively for the astra blue and alcian blue stains. 

In the limited material examined (20 species), obtained dur- 
ing fall and wdnter, no pronounced seasonal differences were 
observed in secretory activity. 

Development of the urophysis spinalis. The caudal neuro- 
secretory cells and the urophysis spinalis were not developed 
in the stages immediately after hatching, length 15-20 mm. (Fig. 
7). The hypothalamus-pituitary system was apparently actively 
secreting at this stage. These observations thus confirm the find- 
ings by Favaro (1925) and Sano (1958 b) that the urophysis 
spinalis and the caudal neurosecretory" system develop late dur- 
ing the ontogeny. Sano (1958 b) noted that the urophysis spinalis 
was still not developed in Salmo fario at a size of 2.5-3 cm. In 
Anguilla of 6 cm. length, the caudal neurosecretory system had 
begun to function. 



-e " 



DISCUSSION 

Many investigators have pointed out the similarity in struc- 
ture between the posterior pituitary and the urophysis spinalis, 
and this analogy has also been stressed in the names suggested, 
e.g. "Ipofisi eaudale" (Favaro 1925), ''Neurohypophysis spin- 
alis" (Enami 1956), and "Neurophysis spinalis caudalis" (Sano 
1958 a, and Sano and Hartmann 1958). This similarity in struc- 
ture and organization of the urophyseal system compared to the 
hypothalamic system was emphasized in this study by the fact 
that the secretory material in the urophysis was stored in dilated 
nerve endings. 

As alread}' mentioned al)ove, there is no selective stain for the 
secretory material in the caudal neurosecretory system, although 
the dilated nerve endings show attinitv for the orange G com- 



8 BREVIORA No. Ill 

poiient of various stains. There is. however, reason to believe that 
the 'acidophilic' reaction of the cells is due to the presence of 
secretion. The terms 'acidophilic' and 'basophilic' as used in 
this study are relative, depending on fixation, stainino- and other 
treatment. The two terms have been used to indicate that the 
number of eitlier acid carboxyl or basic amino groups of the pro- 
tein molecule may be predominant (Romeis 1948). These terms 
may be useful in this case for descriptive purposes until a more 
selective stain for tlie caudal neurosecretory system has been 
developed. 

There are reasons to believe that the neurosecretory cells in the 
teleost s])iiial eoi'd, although they diffei- in size from the cells 
originally described by Dahlgren (1!)14) and Speidel (1919), 
are identical and homologous with those in the skates, in spite of 
the fact that Speidel (1922) did not find the corresponding cells 
in the spinal cord of Fundnli<s. The neurosecretory cells of 
Fundidtis do not differ greatly from unmodified nerve cells, ex- 
cept for their staining reactions (cf. above). They are far less 
conspicuous and appear less "glandular" than the secretory cells 
of the skates and the eel and can therefore be readily overlooked. 

Sano (1958 b) found that the neurosecretory material inside 
the axons could not l)e a pathological product of the cells or of 
the myelin sheath but must be considered as a true neurosecre- 
tion, as earlier indicated by Enami (1955 a, 1956). This study 
can further support these opinions by showing the cons])icuous. 
Herring body-like, secretory droplets which are formed at the 
nerve endings in the dorsal region of the urophysis spinalis 
(Fig. 3). These secretory droplets are, according to this study, 
the dilated nerve endings, which serve to store the secretory 
material. The secretory material is thus generally not stored in 
the urophyseal tissues (cf. Enami 1955 a). 

The origin of the secretory granules inside the neurosecretory 
cells has been extensively dealt with by Scharrer (1934), Palay 
(1943), Hild (1950), Scharrer and Scharrer (1954), Enami 
(1955 b), and others. According to these descriptions, the foi-- 
mation of the secretion is not fully understood. It has been ob- 
served that granules also are present in the inicleus of certain 
cells of the nucleus tubcralis lateralis of teleosts (Scharrer 1934, 
Stahl and Seite 1955. Enami 1955 b. Ortiuann 1956. Stahl 1957). 



1 !).")!) CAUDAL NEUROSECRETORY SYSTEM 9 

Acc()r(liii<i- to Stalil and Seite (1955), the iiitrHinu'lcar granules 
l)i-()])al)ly cannot be eonneeted with neurosecretion. Enanii (1955 
h), on the other hand, considered that tlie 'acidophilic' <iTaniiles 
iu the nucleus entered the cytoplasm through the nuclear mem- 
brane into the area around the nucleus or were emitted into 
neurites. The direct role of the nucleus in the secretory process 
has not yet been clearly demonstrated. As far as the caudal 
neurosecretory system is concerned, 8ano (1958 b) believes that 
tlu' nucleus vciy likely takes an important part in the formation 
of the secretion. The opinion of Scharrer and Scharrer (1954) 
and Enami (1955) that the secretory material may be formed out 
of the Nissl substance or at the expense of the latter, could not 
be confirmed in this study. As mentioned above, it was observed 
that the secretory material seemed to originate from the area 
around the nucleus, which was interpreted to mean that the 
nucleus plays an important role in the formation of the secretion. 

It was previously noted that the secretion in the neurosecretory 
cells reacted negatively with Gomori's chrome haeniatoxylin and 
Ilalmi's aldehyde fuchsin. The secretory droplets (colloid 
masses) in the dorsal region of the urophysis (Fig. 3), on the 
other hand, showed some affinity for the aldehyde fuchsin. This 
difference in staining reaction with aldehyde fuchsin in the secre- 
tory cells as compared to the urophyseal colloid may indicate that 
physical or chemical changes have occurred in the secretory ma- 
terial in association with its storage in the nerve endings. A 
stronger reaction to Gomori's stain is also generally observed in 
the hypothalamus-pituitary system for the Herring bodies com- 
pared with tlie neurosecretory cells or the tracts. In the hypo- 
thalamus-]) ituitary system, the presence of a stainable carrier 
substance or prosthetic group, has been postulated (Schiebler 
1951). If biochemical changes do occur at the release of the 
neurosecretory material at the nerve endings, it would affect the 
response to certain dyes. 

The negative reaction of the caudal neurosecretory cells and 
the secretory material in the urophysis to the periodic-acid -Schiff 
stain is in contrast with the positive reaction in the hypothalamus- 
pituitary system. Schiel)ler (1951) found that the neurosecretory 
cells of the hypothalamus of the teleost Eso.r were strongly peri- 
odic-Schiff positive. 



10 BREVIORA No. Ill 

The studies by Eiiami (lI)5o a) and Saiio (1958 a and b) have 
demonstrated the presence in fishes of a caudal neurosecretory 
system whieli morpholoo'ically is orp-anized in nnich the same 
manner as the hypothalamus-])ituitary system in vertebrates and 
the intercerebralis-corpus cardiaeum system in insects, all show- 
ing- characteristic synthesizing, transporting-, and release-storage 
elements. Such a neurosecretory system is, according to this 
study, present and actively secreting in the teleost fish, Fundulus 
heteroclitus L. 

ACKNOWLEDGEMENTS 

Dr. Gunnar Pridlierg, Zoological Institute, Stockholm, has read 
the manuscript and oft'ered valuable suggestions. The help of 
Dr. A. B. Dawson, Biological Laboratories, Harvard University, 
in preparing the manuscript is gratefully acknowledged. 

SUMMARY 

An examination of the terminal portion of the spinal cord of 
the teleost Fioxluhis heteroclitus L. has confirmed the presence 
of a caudal neurosecretory system. The neurosecretory cells are 
distributed from and posterior to the sixth vertebra and are more 
concentrated above the third vertebra from the caudal end of the 
spine. Histologically, these neurosecretory cells are modified 
nerve cells of multi- ])i-. and unipolar type. They show affinity 
to 'acidophilic' stains such as acid fuchsin, phloxin, and azocar- 
inine. In contrast with the neurosecretory cells of the hyjiothala- 
nnis, these s[)inal cells are Gomori-iiegative and show no reaction 
for the alcian blue stain, the latter staining reaction indicating 
the absence in the secretion of sulfonate groups. The secretory 
matefial, wliich also shows negative periodic-acid-Schiff reaction, 
is transported from the neurosecretory cells in axons serving as 
secretory ]iathways, and the neurosecretion is stored in dilated 
nerve endings wliich terminate in a conspicuous organ, the U)'o- 
plnjsis spinalis (neurohypophysis si)iualis caudalis, neurophysis, 
ur()hyi)oi)hysis), which is a ventral outgrowtli of the spinal cord. 
At the iici-ve endings. Herring l)()dy-like secretory droplets ai-e 
formed. No seasonal changes in the secretory activity of the 
caudal ncMirosecretory system were observed. Tlie uro]ihysis 
shows histologically the same structure as the neurohypophysis 



1959 CAUDAL NEUROSECRETORY SYSTEM 11 

of the i)ituitary ^'laiid, containing nerve eiulinj^s, modified <rlial 
tissue and blood vessels. 

The nrophysis and the caudal neurosecretory system must de- 
velop rather late durin<; the ontop:eny since it is not present in 
end)ryos immediately after hatehin*^, at which time the hypo- 
thalamie-|)ituitary system is actively secreting;. 

REFERENCES 

.\ksakv, a. 

1813. Uc pisum fciobro ct nu'dulln spinali. Disseitatio inauguralis. 
Halae, p. 6. 

Hahlgren, U. 

1914. The electric motor nerve centers in the skates (Ra.iidae). Science, 
40:802-863. 

EXAMI, M. 

1955a. Studies in neurosecretion. II. Caudal neurosecretory system in 
the eel {AnguiUa juponica). Gunma Journ. Med. Sci., 4(1): 
23-36. 

1955b. Nuclear secretion in the cells of the preoptic nucleus in the eel 
{AnguiUa japonica). Endocrin. Jap., 2:33-40. 

19o(). Studies in neurosecretion. VIII. Changes in the caudal neuio- 
secretory system in the loach (Misgurus aiiguillicaudatiu'i) in 
response to osmotic stimuli. Proc. Jap. Acad., 32(3 ) :197-200. 

Ena-mi, M. and K. Imai. 

1955. Studies in neurosecretion. V. Caudal neurosecretory system in 
several freshwater teleosts. Endocrinologica Jap., 2(2):105-116. 

195(5n. Studies in neurosecretion. VI. Xeurohypophysis-like organize 
tion near the caudal extremity of the spinal cord in several 
estuarine species of teleosts. Proc. Jap. Acad., 32:197-200. 

1956b. Studies in neurosecretion. VII. Further observations on the 
caudal neurosecretory system and the neurohypophysis spinalis 
(urohypophysis) in marine teleosts. Proc. Jap. Acad., 32(8): 
633-38. 

En'ami, M., S. Miyashita and K. Imai. 

1956. Studies in neurosecretion. IX. Possibility of occurrence of a 
sodium-regulating hormone in the caudal neurosecretory system 
in teleosts. Endocrinologies Jap., 3(4) :280-90. 

Favaro, G. 

1925. Contributi alio studio morphologico dell' ipofisi caudale dei 
tcleostei. Mem. K. Accad. Naz. Lincei, 6(l):29-72. 



12 BREVIORA No. Ill 

(iOAIOKI, G. 

1950. A rapid one-step tiiclironie stain. Amer. Jonr. Clin. Path., 20: 
661-664. 

HiLi), W. 

1950. Zur Frage der Neurosecretion ini Zwischenhirns der Sehleie 
(Tinea vulgaris) iind ihrer Beziehung- zur Xeurohypophyse. 
Zeitschr. Zellforsth., 35:33-40. 

Halmi, S. N. 

1952. Differentiation of two types of basophils in the adenohypophysis 
of the rat and the mouse. Stain. Tech., 27(l):61-64. 

Holmgren, U. 

19o8a. Secretory material in the pineal body as shown by aldehyde- 

fuchsin following performic acid oxidation. Stain. Tech., 33(3) : 

147-149. 
19581). On the caudal neurosecretory system of tlie teleost fish, Fun- 

liitiiLs hffcrnriitii.s. (Abstract.) Anat. Rec. 132:454. 

Lll,LIE, li. U. 

1954. Histopathologic technic and practical histoclicmistry. Blakiston 
Co., New Yoik, Toionto. 

Ortmax.v, R. 

1956. Ilistochemische Untersucluingen an den Kerneinschlusskorperu 
der Ganglienzellen ini Nucleus praeopticus dcs Karpfen (Cypri- 
ntis carpio L.). Zeitschr. Anat. Entw. Ges., 119:485-499. 

I'ALAV, S. L. 

1943. Neurosecretion. A'. The origin of neurosecretory granules from 
the nuclei of nerve cells in fishes. Jour. Comp. Neurol., 79: 
247-275. 

IvArBKR, A. 

1877. Die letztcu spinalen Nerven und (ianglien. Morph. Jahrl)., 3: 
603-624. 

Ko>tEis, B. 

1948. Mikroskopische Teehnik. Munich 15th ed. 

Saxo, Y. 

1958a. ITeber die Xeuiophysis (sog. Kaii(lalliy|io])liyse, "UrohyjJO- 

physe") lies Teleostieis, Tinea nih/ari.s. Zeitschi-. Zellforsch., 

47:481-497. 
19.")8li. W'eitere Unteisuchungen iil)er den Feinliau der Neuroi>liysis 

spinalis caudalis. Zeitsdir. Zellforsch., 48:236-260. 



If).')!) CAUDAL NEUROSECRETORV SYSTEM 1 :> 

Sano, V. ;uid F. Haktmann. 

1!)58. Zur voigleiehondcii Ilistolojiic \()ii Neuropil^ sis spiimlis cini- 
il;ilis iiiul Xciiroh.vpdpli.vsis. Zcitsclir. Zcllt'orscli., 48:r);!S-r)47. 

SCHARKKK. K. 

1!>;>4. Uehor den r'oitoilisjunf>t'n des ZeUkciiis .-m sckictorisclicn 
Voifjiin^en in Ni'ivcn/.cUcii. P^niikf. Zcitsclir. l':itli., 47:14IMm. 

SrnAKRER, E. and B. Scharrkr. 

1954. Xi'nrosecTction. In Ilnndli. inikr. Anat. Mciisch., 4(r) ) : !»r):M050. 

SCHIEBLKR, T. H. 

19;')!. Zur Histoflu'niie dos nonrosecrotoriscdicn hypotlinlainiscdi-neuro- 
hypophysaren Systems. Acta Anat., 13:233-tir)r). 

Speidel, C. C. 

1919. Gland-cells of internal secictiou in the spinal cord of skates. 

Carnegie Inst. Washington, Pul)l. 1181. 
W^'l'l. Further comparative studies in other fishes of cells homologous 

to the large irregular glandular cells in the spinal cord of the 

skates. Jour. Comp. Neur., 34:303-317. 

STAilL, A. 

1957. Kecherches sur les elaliorations cellulaires et la neuiosecretion 
dans I'eneephale des poissons teleosteens. Acta Anat. (Basel), 

31(suppl.) :28. 

Wtahl, A. and R. Seite. 

1055. Sur la presence d'une elaboration figuree d'origine nucleaire 
dans les cellules neurosecretoires du noyau preoptique chez les 
poissons teleosteens. C.R. Soc. Biol. (Paris), 149:382. 

WEiiEi;, 10. II. 

1827. Knoten uud uupaaieu Faden iiiit dem sich das Iliickenuiaik liei 
einigen Fisclien endigt, namentlich l>eim Ci/priiiu.s cdrpid. Mcch- 
el's Arch. Anat. u. Physiol., 2:31(i-317. 

Ziehen, Th. 

1903. rVutralnervensystem, 1 Aid. Jena. Gustav Fischer. 



■n i:i' 



PLATE I 

Fig. L'. Sagittal section of the terminal portion of the spinal cord, showing 
the nrophysis spinalis, ventral to the spinal cord. The area inside the rec- 
tangle is shown in Figure '.). Fixation Benin's solution. Halnii's aldehyde 
fuchsin stain. 8 microns, 100 x. 

Fig. 3. The area, shown in the rectangle in Figure -. The dilated nerve 
endings of the neurosecretory cells, terminating in the dorsal parts of the 
urophysis. These dilated nerve endings are similar to the Herring bodies of 
the neurohypophysis of the pituitary gland. Fixation Bouin's solution. 
Halmi's aldehyde fuchsin stain. 8 microns, 500 x. 

Fig. 4. Transverse section of the urophysis spinalis and the ependymal 
terminal portion of the spinal cord. The leptomeninx covered with melano- 
phores surround the organ. Note that the urophysis is heavily vascular. 
Fixation Bouin's solution. Mallory's triple stain. 8 microns, 500 x. 

Fig. 5. 'Acidophilic' neurosecretory cells from the terminal portion of 
the spinal cord. Fixation Bouin's solution. Gomori's chrome haematoxylin- 
phloxin method. 8 microns, 1200 x. 












fc' 



<gOi;^ '"tf^.. %'ii^*^S^^°W^S^ 













PLATE 1 




^mi^ 



'-13 




.■f" .uj-lf. 



V .*y ,:~f-:'S^'^'-^'^ 



V 













?\% v-^ 




, 4 * 



,f 



- ^ 




PLATE II 

Fig. G. Median .seftiou tluougli tlio urophysis, showing- the bk)od vessels. 
Fixation Bouin's solution. Boilian's piotargol method. S niic-rons, ."00 x. 

Fig. 7. Sagittal section of the terminal portion of the spinal cord in a 
newly hatched Fundulu.'i embryo. Xote that no urophysis nor any secretory 
cells are yet developed. Fixation Bouin's solution. T5odian's protargol 
method. 500 x. 



BREVIORA 

MmseiLioi of Coimpsirative Zoology 



CAMBKiiKiE, Mass. Octobek 15, 1959 Number 112 



A MorXTED SKELETON OF THE GIANT 
PLESIOSAUR KR0N0SAVRU8 

By Alfred Sherwood Romer 

and 

Arnold D. Lewis 

In 1924 Longman described a plesiosaur mandibular fragment 
of large size from the Lower Cretaceous of Queensland as the 
type of a new genus and species, Kronosaurus queenslandiciis. 
Later (Longman 1932), some further fragmentary material, 
mainly the proximal ends of two limb bones, was obtained from 
the type locality near Ilughenden, and a letter from Dr. Long- 
man to Mr. W. E. Schevill in November 1932 mentions the dis- 
covery there of eight caudal vertebrae which appear not to have 
been described. The limb bone material, added to the evidence 
of the jaw fragment, indicated a plesiosaur of very large size. 
The remains suggested relationship to PeJoneustes, and Longman 
(1932) published a hypothetical restoration of Kronosaurus 
based mainly on the work of Andrews (1913, fig. 26) on 
Peloneustes philarchiis. 

In 1931-32, Mr. William E. Schevill, a member of an expedition 
sent to Australia by this museum, explored the Queensland Lower 
Cretaceous deposits in the Richmond region, west of Ilughenden. 
In Grampian Valley, 30 miles to the north of Richmond, he 
found a rostrum of a young individual of Kronosaurus (MCZ 
No. 1284). His attention was called, by a local resident, to the 
Army Downs region, five miles farther north. Here was found the 
skeleton (MCZ No. 1285) which is described below. 

The specimen had been entombed, in an articulated state, 
dorsal surface up, in a limestone matrix. In recent times, how- 
ever, the skeleton had been subject to erosion, so that it consisted 



2 BRE^^ORA No. 112 

essentially of a series of limestone nodules, freed from the under- 
lying strata and nearly completely l)uried in the soil. Skull, 
neck, trunk and part of the tail were contained in a linear series 
of 15 nodules of varied size. Of these the first had beiMi displaced 
and overturned; the others, however, appear to have undergone 
little or no displacement. Erosion had destroyed much of the 
outer parts of the nodules, so that, for example, most of the 
superficial bones of the skull had been destroyed, part or all of 
many of the neural spines had vanished, and the girdles, ribs and 
abdominal armor are incomplete. Erosion had, further, destroyed 
some of the contacts between successive blocks, but because of 
their seemingly undisturbed position, interpolation of materials 
once filling the gaps can be made with considerable confidence. 
No trace of the pectoral limbs was preserved. The pelvic limbs 
were present in normal articulation, extending out on either side 
from the large block containing the pelvic region, but even the 
femora were badly weathered and the more distal regions of the 
"flippers" were very poorly preserved. 

The blocks, weighing in all nearly four tons, were excavated 
and shipped to Cambridge by Mr. Schevill. Two years later the 
skidl was developed by Dr. T. E. White, and a description pub- 
lished by him (1935). Because of the very considerable outlay 
of time, energy — and money — necessary for the lal)ori()Us task 
of preparation, the postcranial material lay untouched in storage 
for tw^o decades. Three years ago, however, Mr. Godfrey Cabot, 
who is deeply interested in the subject of marine reptiles, fur- 
nished us with funds sufficient to pay the costs of preparing and 
mounting the skeleton. The work was directed by the junior 
author, with the assistance of Messrs. James A. Jensen and David 
Fuller. 

As noted above, erosion had destroj'cd a fail' fraction of this 
on-je complete and articulated skeleton, so that approximately a 
third of the specimen as exhibited is plaster restoration. There 
are, nevertheless, few points in which any great deviation from 
verity can have occurred in mounting the skeleton. As noted 
below, the exact number of cervical plus trunk vertebrae is un- 
certain, as is the lengtli of the short tail. Ti-unk rib lengths are 
for the most part uncertain, l)ut the curvature of the ribs as 
pre.served furnished a rather safe guide to trunk proportions. 



195f) KRONOSAURUS 3 

The front padcllcs nvc la('kiii<,', hut can he restored with consid- 
erahh^ eontidence by eomparison with preserved parts of the 
pelvic a{)pendafies. Tpon completion of the mount, the only 
structural feature which we would wish to modify, given the 
()j)portunity. Mould he to mo\'e the pelvic girdle upward and 
backward by perhaps 20 cm. 

Lon<iinan, we have noted, compared Kronosanriis, on tlie basis 
of tlie fragmentary remains available to Jiim, to Peloneustes of 
the Oxford Clay. Our skeleton confirms his belief in the main. 
As will be seen from the description below, we are dealing with a 
pliosaur of somewhat similar nature, although of much larger 
size — our animal lias about four times the linear dimensions of 
Andrews' restored skeleton with a total length, as restored, of 
1280 em. — about 42 feet. The head, however, is relatively larger 
in Kronosaurus, making up slightly over one-fourth of the total 
length, whereas in Peloneustes the skull is but one-fifth the length 
of the animal. Further, the trunk is relatively long in the Aus- 
tralian giant, and the neck extremely short ; although the number 
of presacral vertebrae in Kronosauriis and Peloneustes is ap- 
proximately the same, the beginning of the dorsal rib series and 
the position of the shoulder appears to have been aliout 10 verte- 
brae farther forward in our specimen. Kronosaurus thus appears 
to be the most stouth' built of plesiosaurs. 

As a short-necked pliosaur, it is of course exceeded in length 
by some of the long-necked elasmosauroid types. In bulk, how- 
ever, it seems certainly to be larger than any described plesiosaur, 
although Dr. L. B. Tarlo {in litteris) tells us that material of a 
Kimmeridge Clay pliosaur indicates an animal of comparable 
size. 

As far as preserved, the Kronosaurus postcranial skeleton does 
not depart structurally in any marked manner from the general 
pliosaurid pattern, and hence need not be figured or described 
in detail. The notes given here can be followed by reference to 
the excellent descriptions and figures given by Andrews (1913) 
for Oxford Clay pliosaurs. 

Vertebral column. The series of nodules containing the skele- 
ton yielded part or all of 46 vertebrae between the atlas-axis and 
the proximal part of the tail. The sequence of the preserved 
vertebrae can be determined in every case from the sequence of 



4 BREVIORA No. 112 

the blocks eontaiiiiug them. But whih^ various short series are 
in articulated position within a given block, and certain of tlie 
blocks were in contact, there are obvious gaps in the series, and 
we have restored 11 vertebrae in tilling out the column to the 
proximal part of the tail. For the most part the number of miss- 
ing vertebra? in these gaps can be determined with considerable 
confidence, through the relation of adjacent vertebrae to otlier 
structures, or the length of the eroded interval between blocks. 
It is possible that our restored vertebral count may vary l)y one 
or two segments from the true condition. 

Anteriorly, vertebrae 1-5 were found in {Position behind the 
skull. Following a short gap, 4 further vertebrae were present 
in another block. The first of this succeeding series is nearly com- 
plete; the third has a complete rib (on the right side). From 
the evidence of rib lengths, position of rib facets, centrum dimen- 
sions, and the length of the gap between blocks, interpolation of 

8 vertebrae between the first and second series appears reason- 
able, and we shall consider the second series to include vertebrae 

9 to 12. An isolated neural spine and arch appears from its 
proportions and the nature of the rib facet to have occupied a 
short gap following vertebra 12. The next block contained verte- 
lirae 14-16. Lt was necessary to break up, in the field, the large 
luass containing the pectoral girdle region; This caused two 
further gaps, the first of which can be accurately determined to 
involve one segment ; the second, following a series of 5 vertebrae 
— presumably 18-22 — seems certainly to have been occu])i(Ml by 
8 vertebrae. There follows a series of 4 vertebrae, our numbers 
26-29. Behind this is a block with 2 vertebrae sejiarated by gaps 
both fore and aft. The first gap is of a length appropriate for 2 
vertebrae ; the second gap is short, but because of the imperfect 
nature of the vertebra behind it, it is uncertain whether a seg- 
ment should be intercalated here, as we have done in the mount. 
Back of this point, an unbroken series of vertebrae, which we 
have restored as numbers 35-43, can be traced to a point close to 
tlie puboischiadic suture on the underlying pelvic girdle. Here 1 
vertebra is definitely missing, followed by vertebrae 45-48, found 
above the ischium, and 49-50 close behind them. Three further 
adjacent blocks contain 4 vertebrae and parts of the centra of 8 
others, bringing the total as restored to this point to 57 vertebrae. 



1!>5JI KRONOSAURUS 5 

b'foiu the rapid (Icci-casc in si/c seen within llic [)()st-])('lvif. scries, 
it is obvious that the tail was siiort ; we have restored it to jiive 
a total luuuhei- of 7!' vertehi-ac As will lie seen IVoni the descrip- 
tion of the rib attaehuuMits u'iveii below, there appear to be 12 
true et'rvieal vertebrae, 2 "pectorals," probably 80 doi-sals, giv- 
in^^ a total oP approximately 44 presacrals, followed by .'{ or 4 
saerals. 

Centra. Measurements (in nnn. ) of some of the best preserved 
centra are as follows: 



Centrum 


Li nglli 


WIdtli 


Hciylit 


4 


102 


181 


193 


9 


98 


180 


210 


15 


114 


182 


233 


20 


128 


202 


225 


27 


145 


213 


217 


35 


143 


205 


208 


42 


136 


182 


201 


50 


118 


180 


198 


54 


108 


157 


172 



As can be seen, the length and width of the centra reach a maxi- 
mum in the mid -dorsal region, with somewhat decreasing- meas- 
urements toward the sacrum, and a sharp drop in size — partic- 
ularly in length — in the proximal part of the tail. The height 
of the centrum is nnich more uniform, with a fairly marked 
increase along the neck to the anterior dorsals, and a very gradual 
i-eduction back of this point to the proximal caudals. In conse- 
(juence of the diiferences in width-height relations, the propor- 
tions of the vertebrae, as seen in end view, ditil'er considerably 
from one region of the column to another. Mid-dorsals and pos- 
terior dorsals are nearly, circular in outline ; anterior dorsals are 
considerably taller than broad, giving an oval contour, and both 
the cervicals and the posterior vertebrae, from the posterior 
dorsals back, are moderately taller than broad. The cervicals 
and anterior dorsals are about twice as high as long ; however, 
the length of the centrum increases posteriorly along the dorsal 
series, while the height decreases, so that towards the sacral 
region the height is but half again the length ; in the tail, with 



6 BREVIORA No. 112 

reduction taking place more rapidly in length than in height, 
there is some increase posteriorly in relative height. 

Typical dorsal centra have the simple outlines of a short spool, 
with a smoothly curved lateral surface between raised anterior 
and posterior rims. Anteriorly and posteriorly, however, the 
presence of rib attachments modifies the lateral surfaces, as noted 
in more detail in the next section. In the cervicals the rib 
facets take the form of a circular "crater," with raised rims, 
occupying nearly the whole length of the lateral surface of the 
centrum. In the caudal region the low rib facet is of smaller 
dimensions. Above, it is joined by a vertical ridge, half way be- 
tween anterior and posterior margins, which ran upward to the 
neuro-central suture. Below the rib facet, the preserved caudals. 
except the first two, are excavated laterally below a curved longi- 
tudinal ridge which is convex ventrally. 

No chevrons were found with the few caudals i)resent ; how- 
ever, presumed caudal 4 is bevelled ventrally on either side for 
chevron attachment. 

Neural arches and spines. The axis neural spine is of large 
size. Many of the cervical spines behind this point are preserved. 
They are slender at the base, well tilted back and, in at least the 
posterior cervicals, broadened distally with a pronounced median 
groove, presumably for the attachment of ligaments. The an- 
terior margin has a well-developed median keel for most of its 
height. Many of the dorsal spines are incomplete or absent ; how- 
ever, their general nature is apparent. They are broader at the 
base, and less inclined backward than the cervicals ; the anterior 
keel is less pronounced and the distal grooving is absent. Three 
proximal caudals have partially preserved spines which slant 
strongly toward the rear. In the cervical region there are well 
developed zygapophyses. In the anterior part of the dorsal re- 
gion zygapophyses are present, but reduced in size. Many of the 
neural arches of the more posterior vertebrae are imperfectly 
preserved or are absent, but it is significant that no trace of 
zygapophyses has been seen on any vertebra po.sterior to No. 27 
(including several seemingly complete ])()sterior dorsal and 
proximal caudal arches). Apparently connections between neural 
arches were made here, in piscine fashion, by ligaments alone. 



] 959 KBONOSAURUS 7 

Rib articidatiun.s. On the atlas-axis, a small conical tuherclc is 
present, low down on either side at about the level of the suture 
between atlas intereentruni and centrum; presumably this repre- 
sents a small fused lib. With the axis centrum there begins a 
series of cervical rib facets, oval in shape, with raised borders. 
No clear distinction could be seen between presumed dorsal and 
ventral areas of attachment. In the most anterior vertebrae the 
facets are situated low down on the centrum, with their lower 
rims about 2.5 cm. above the bottom margin of the centrum. The 
anterior ones are short ovals in outline, with the longer axis 
vertical, and measuring about 7 by 8 cm. Anteroposteriorly these 
facets cover most of the lateral surface of the centrum. On the 
9th and 10th vertebrae the facets have moved upward to some 
extent, so that their lower margins are about 3.5 cm. above the 
level of the bottom of the centra. Here the surface is more 
markedly oval in shape, with diameters of about 5.5 and 10 cm. 
At vertebra 12, the upward tendency is more pronounced; the 
dorsal margin of the facet is at the line of suture between arch 
and centrum, and the surface measures 6 by 10.8 cm. As noted 
above, centrum 13 is missing. However, vertebra 14 is of the 
transitional type which has been termed ''pectoral" (this verte- 
bra appears to have lain above the scapula). The articular rib 
facet measures 5.5 by 11 cm.; its upper third lies definitely on 
the neural arch. Further, there is the beginning of a "transverse 
process"; the lower part of the articular area extends outward 
about 3 cm. beyond the level of the surface of the centrum, and 
the upper, neural arch portion is similarly produced. 

Presumably the ' ' pectoral ' ' shift began with vertebra 13 ; by 
vertebra 15, there is a developed transverse process. This extends 
directly laterally to a distance of 19 cm. ; the upper margin of 
the articular surface, which measures 6.5 by 13 cm., is at the level 
of the zygapophysial facets. The centrum forms a considerable 
part of the root of the process, but whether it reaches the facet 
surface is uncertain. On vertebra 16 the centrum enters broadly 
into the formation of the root of the transverse process, but fails 
to reach its distal end. On vertebra 18 the centrum buttresses a 
small area of the base of the process, with a ventral keel extend- 
ing out from this area along the underside of the process. By 
vertebra 19 this keel has disappeared, the lower surface of the 



8 BREVIORA No. 112 

process is smoothly rounded, and the process springs from a 
point well above the arch pedicel. Here the dorsal surface of 
the process slants somewhat upward distally, extending out about 
20.5 cm. from the midline. The processes in this region have 
become circular in section, measuring about 5 cm. in diameter at 
half-length, but somewhat expanded distally to an oval articular 
facet measuring about 7 by 11 cm. From this point back there is 
little change to be seen over much of the dorsal region in such 
processes as are well preserved. By vertebra 36, for example, the 
process, which here extends 19.5 cm. from the midline, has a 
diameter at midlength of about 6.5 cm., but expands distally to 
a nearly circular facet with a 7.5 cm. diameter. There is a slight 
posteroventral ridge development on the underside of the process, 
which is here directed somewhat posteriorly but does not rise 
appreciably toward its distal end. Similar processes are present 
to vertebra 43 ; but on this last vertebra the ventral ridge has 
increased in depth, so that the base of the process extends doAvu 
to a point just above the suture between arch and centrum. 

Vertebra 44 is missing. The following vertebra lacks the distal 
end of the process but its well preserved basal portion is nanoAv 
but tall, measuring 10.2 cm. in height, and although the line of 
suture is not clear, the lower two-fifths of the process definitely 
arises from the centrum. We are dealing with a sacral, and it is 
probable that vertebra 44 was a sacral as well. Vertebra 46 is 
also a sacral; the rib process is still deeper, measuring 11.8 cm. 
in height distally, and with a width not exceeding half tlie height. 
Approximately half the articular surface is furnished by the 
centrum. The process is nuich shorter than that of the dorsal 
vertebrae, extending only 13 cm. from tlie mid-line; further, its 
upper margin .slants slightly downward and outward. The trans- 
verse process of 47 is smaller than that of 46, although similarly 
built, extending outward only 12 cm. and with a distal height of 
8 cm. Possibly this was a further sacral. There is, however, no 
marked break between sacral and caudal elements. 

Caudal rib facets can be followed for 7 vertebrae fui-ther. On 
vertebra 48 the lower margin of tlie base of the process has 
retreated upward a centimeter or more from the position found 
in 47 ; from this region backward, however, the ventral margin 
of the base moves ventralh- so that bv vertebi-a 55 it lies below 



1959 KRONOSAURUS 9 

the luidclk' of llu' lutcrul surlHet' ol' tlie eentruni. Bvt'U luovv 
nuirked is a downward movement of the upper margin of the 
rib facet. Beyond vertt'hra 50 the neural arch takes no part in 
rib support, the depth of the transverse process diminishes, and 
from vertebra 53 back its attaclnnent is an essentially circular 
I'acet centrally placed on the lateral surface of the centrum. There 
also takes place a gradual reduction in length of the transverse 
process, so tliat by vertebra 54 it is little more than a circular 
area projecting but slightly from the general contoui-s of the 
centrum. 

Ribs. On the first five vertebrae the ribs are fused to the 
centra and, except for the loss of most of rib 5 on the right side, 
were nearly completely preserved. The first rib, as noted, is 
represented by a short spine fused, without evidence of suture, 
to the vertebra ; on ribs 2-5 the line of suture is evident, although 
rib and centra were coossified. These anterior cervical ribs are 
short and stout. They extend outward and somewhat backward, 
flattened dorsoventrally. The distal ends are somewhat expanded, 
the expansion being directed backwardly from the line of the 
shaft. Rib 9 on the right side is nearly complete, with an esti- 
mated total length of 140 nun. The head is large, measuring 99 
by 55 mm., with the greater diameter vertical ; the rib extends at 
first directly outward, but then turns sharply backward, with a 
tAvist such that the dorsal margin of the head turns to the back 
margin of the shaft. Distally, the rib thins to an oval, with diam- 
eters of 57 and 30 mm. at 13 cm. from the head, the major axis 
running from posterodorsal to anteroventral margins. Rib 1 1 
is similarly built. 

The more posterior cervical and "pectoral" ribs are poorly 
preserved. One distal rib segment Avliich appears to have lain in 
the pectoral region shows a much flattened shape, 86 by 35 mm. 
in section, with a terminus which is little expanded. Presumably 
this rib bore part of the serratus musculature, which one may 
expect to have been present — even if feebly developed — in 
plesiosaurs. Of the dorsal ribs, considerable material was pre- 
served, but most had been disarticulated and badly broken. 
Those of vertebrae 22 and 23 alone are articulated and nearly 
complete. The heads are large, that of rib 23 having diameters 
of 102 by 88 mm. The total length of rib 23 was approximately 



10 



BREVIORA 



Xo. 112 



98 cm. The rib extends straight out from the transverse process 
for about 18 cm. ; in the next 20 cm. there is a marked downward 
curvature to the extent of about 60°, followed by a further more 
gentle curvature to reach a total of 90° from the line of the trans- 
verse process and head. The shaft is somewhat flattened, with 
diameters, in section, of 58 and 47 mm. Distally the ril) ex])ands. 
terminating in an unfinished surface, for the ventral rib segment, 
which measures 80 by 65 mm. 

Many of the fragments of more posterior ribs suggest similar 
proportions, except that the preserved heads of some of these 
ribs are more nearly circular in section. The sacral ribs are poorly 
preserved, and caudal ribs are absent in the material. 




Fig. 1. Shoulder girdle of the mounted specimen in dorsal view. J latched 
areas restored. X 1/30. 



Girdles. White (1940, fig. 11a) has given a sketch of the 
shoiddcr girdle (Fig. 1). No remains of dermal elements were 
recovered. Tlie I'ight scapula, similar in shape to that of 
Peloneustcs, is nearly complete, except for the dorsal blade; the 
left scapula was not found. Of the coracoids, much of the an- 
terior half was [)resent on both sides. Farther back, three i)or- 
tions of the nu'dial margin of the ritrht coracoid. shown in the 



1959 



KRONOSAURUS 



^^ 



tigure, were in place. Fruiu tliein the outlines of the coraeoitl can 
be fairly aeenrately restored, except for the degree of develop- 
ment at the posteroexternal angle. As restored, the coracoid is 
extremely long (160 cm.). This figure is 270 per cent of the 
width of the bone, measured from the outer margin of the glenoid 
process to the midline (.19 em.). In Prloncnsfcs pliilai-chus and 
Situolcstes, as figured by Andrews (191."j, figs. 8, 21), the coi'acoid 
length is about twice the width. The Kronosaurus coracoid is 
thus even longer proportionately than in typical pliosaurs. 




Fig. 2. Pubis and ischium of the mounted specimen in dorsal view. 
Hatched areas restored. X 1/30. 



The pelvic girdle (Fig. 2) is exceptional!}' long. The left pubis 
is represented only by a small piece at the posterior end and two 
symphysial fragments, but the right pubis is nearly complete. 
The element measures 128 cm. in length, with a greatest width 
posteriorly of 74 cm. This is, again, an elongate bone, with the 
width 173 per cent of the length, contrasted with 122 per cent. 
118 per cent and 122 per cent in Andrews' figures of rdoneustcs. 
Simolestes and Pliosaicnis (Andrews 1913, figs. 24, 10, 7). Ex- 
cept for part of the external and posterior margins, the ischia 
are nearly complete. The greatest length is 103 cm., the Avidth to 
the outer margin of the acetabulum 52 cm. The pi-oportions of 



12 BREVIORA Xo. 112 

the ischium (in eoiitrast to the pubis) are not far from tliose of 
certain other pliosaurs, for the greatest length is 198 per cent 
of the width, as compared with figures of 189, 186, and 163 foi' 
the three pliosaurs mentioned above. In consequence, the bone is 
somewhat small in proportion to the pubis, for it is but 80 per 
cent of the length of the pubis in our skeleton whereas it is 118 
per cent, 112 per cent, and 103 per cent in the three other 
pliosaurs described by Andrews. There is a well-defined but 
small puboischiaclic vacuity. As in plesiosaurs generally, the 
pubes and ischia are considerably thickened medially lielow the 
acetabulum, indicating the presence of a very stout diamond - 
shaped symphysial cartilage. Of the ilium, the basal portion, 
witli a breadth of 17.7 cm. and a greatest height of 38 cm., is 
preserved. The contours of this portion indicate that the iliac 
blade was considerably expanded. 

Linhbs. As noted, no remains of the pectoral paddles were 
preserved. Both pelvic limbs had been in position, but were in 
great measure eroded. The femora were present, but the middle 
of the shaft was missing on both. However, the gap in the left 
femur was short; as the bone lay, this was less than 10 cm., and 
the bone has been restored to this extent, with a total length 
of 106 cm. The contacts between left femur and tibia and fibula 
are lost, but most of the tibia and the anterior part of the fibula 
are present. Articulated with them are the two elements some- 
times termed intermedium and fibulare and, beyond, two large 
distal tarsals and metatarsals III and IV. A fragment fi'oiii the 
right limb includes the more posterior of the two distal tarsals 
seen on the left, and attached to it distally, metatarsals III and 
IV in addition to a fragment of metatarsal \'. 

Described i)lesiosaurs in general have the forelimb and meso- 
podial elements arranged (to borrow a phrase from mammalian 
osteology) in an '"alternating" fashion, so that (as in the carpus 
of advanced ungulate mammals) the elements in each row alter- 
nate to some degree with those proximal and distal to them, and 
facets between successive elements may be tilted at an angle from 
a transverse line of the bone. Here, on the contrary, there is. 
in the elements preserved, a one-to-one relation of bones of the 
three proximal rows, and the lines of articulation bctwc n suc- 
cessive segments are directly transverse. A similar construct'on 



1!)5!) KRONOSAURTTS 13 

is present iu a picsiosaur from the Iviiumeridge Clay ut" Strethain 
now heinp; described l)y Dr. Tarlo. 

Ill both jiaddles a number of disarticulated and varial)ly 
weathered phalan<i'es, measurin*]^ on the order of 10 to 12 cm. in 
lenoth, were present. The size of the limb elements [)resent sup- 
ii:est a paddle, beyond the femur, of about twice the lengtli of 
that of Peloneustcs pliilarchtis, i.e., about 120 cm. to 140 cm., and 
a total length of the limb of about 230 cm. 

Fijioids. Beneath the cervical vertebrae was found a symmet- 
rical pair of bones with expanded, diagonally-facing- "heads"; 
neither is complete distally ; the one with the longer shaft meas- 
ures 262 mm. as preserved. These appear to be a pair of basihyoid 
elements ; they differ markedly, however, from the slender ele- 
ments generally reported in plesiosaurs. 

GastraJia. A portion of the ventral armor was preserved, but 
the materials are too imperfect to warrant detailed description. 
There is evidence indicating that at least a part of the gastralia 
were of complex form, in contrast to the primitive rod-like shape. 



14 BREVTORA No. 112 

BIBLIOGRAPHY 

Andrews, C. W. 

1913. A descriptive catalogue of the marine reptiles of the Oxford 
Clay. Based on tlie Leeds collection in the British Museum 
(Natural History), London, Part II. London, xxiv + 206 pp. 

Longman, H. A. 

1924. Some Queensland fossil vertebrates. Mem. Queensland Mus., 8: 

16-28. 
1930. Kro7iosaurus queenslandicus. A gigantic Cretaceous pliosaur. 

Mem. Queensland Mus., 10: 1-7. 
l!i32. Restoration of Kronosaurits queenslandicus. Mem. Queensland 

Mus., 10: 98. 

White, T. E. 

1935. On the skull of Kronosaurus queenslandicus Longman. Oceas. 

Pap. Boston Soc. Nat. Hist., 8: 219-228. 
1940. Ilolotype of PJesiosaurus longirostris Blake and classification 

of the plesiosaurs. Jour. Paleont., 14: 451-467. 



1950 



KRONOSAURUS 



15 




BREVIORA 

Museiuiim of Comparative Zoology 

('AMUUiDdi:, Mass. Xovkmukk 2, 1959 NrMHKK 1 1;{ 

A NEW PllVLLOMEDUSA FROM BOLIVIA 
(SALIENT I A, HYLIDAE) 

By Benjamin Sitreve 



In 1954 Carl Gans and Padre F. S. Pereira, C.M.F. collected 
amphibians and reptiles in the state of Mato Grosso in Brazil 
and the Department of Santa Cniz in Bolivia. W. C. A. Boker- 
man of the Departaniento do Zoologia, Sao Paulo. Brasil, did 
some preliminary work on identifying' the amphibia, while I was 
called upon to complete this task. While doing so, I discovered 
the novelty described here. Dr. Gans, who is planning to publish 
a complete report on his collection, has graciously presented 
about half of the am})hibians to the Museum of Comparative Zool- 
ogy, including the type of the new form, the remaindei- going 
to the Carnegie Museum of Pittsburgh, Pennsylvania. 

P II YLLO MEDUSA PAILONA sp. UOV. 

Type. Museum of Comparative Zoology 29677, an adult male, 
from El Pailon, 5 kilometers from the eastern shore of the Rio 
Grande, altitude 350 meters. Department of Santa Cruz, Bolivia, 
collected by Carl Gans and P. S. Pereira March 5, 1954. 

Pamtffpes. M.C.Z. 29(578-7!) and Carnegie Museum 36278-80, 
all adult males, with the same data as the type. 

Diagnosis. Apparently most closely allied to Pkyllomedusa 
sauvagii Boulenger but differing in apparently smaller size, 
longer hind limbs, immaculate venter and other aspects of colora- 
tion. 

Description. Tongue slightly nicked behind; vomerine* teeth in 
two slightly oblique groups between the choanae; snout longer 



2 BREVIOKA Xo. 118 

than the diameter of the eye, obliquely tnineated from behind 
forwards (male only?, shape of snout may be different in fe- 
male) ; loreal region slightly oblique: interorbital space broader 
than the upper eyelid; tympanum (longest diameter) two-thirds 
the diameter of the eye (two-thirds or slightly less in paratypes) ; 
fingers free, first very slightly shorter than second, fourth some- 
what shorter than third ; toes free, first longer than second ; disks 
of fingers and toes much smaller than tympanum ; inner meta- 
tarsal tubercle fairly prominent ; the tibio-tarsal articulation of 
the adj^ressed hind limb reaches to the tympanum ; upjx'i' surfaces 
minutely granulate ; parotoids large, short, a larger portion ex- 
tending about as far as axilla, a rather obscure extension extend- 
ing farther back (not evident in some ]iaratypes) ; belly and 
lower surface of thigh granulate, other parts of lower surface 
less granulate; a subgular vocal sac aiul a brown nuptial pad 
on inner side of thund). 

Coloration in alcohoL Above, grey blue (quite light in some 
of the paratypes) ; inner two fingers and outer three toes yellow- 
ish suffused with brown, disks of other fingers and toes brownish ; 
eye more or less narrowly bordered with white suffused with 
orange, decidedly more extensive and continuous on upper eye- 
lid than lower (possibly bright orange in life). Below, yellowish ; 
a white stripe on tarsus and forearm (that on the former curving 
ai'ound the tibio-tarsal articulation), separating dorsal and 
ventral pattern, continuing on to the adjoining digit ; a similar 
stripe, but short, above anus; a wash of more or less jnirplish 
brown under limbs particularly in the vicinity of the stripe, the 
same i-olor below anal stripe ; also a Avhite stripe on under side 
of upper arm curving around insertion anteriorly. Some of the 
paratypes show obscure white spotting under the thighs. 

Head and hody lengih. Type 67nnn. Paraty])es 60-69mm. 

Remarks. A transforming individual (M.C.Z. 29680), api)ai-- 
ently rcfcr-able to this species, was taken from the stomach of a 
liatchling Chironius pyrrhopogon (Wied). This toad had the tail 
still present but the limbs appeared fully developed. Owing 
to some possible doubt about the identification, this individual is 
not included as a parat.ype. The datfi is the same as that of the 
type and pa i-a types. 



!!),')!) m:\v riivi.LDMKDi'SA 



3 



The raii^e o\' sail ra(/il is yivcii l)y the most I'ccciit reviser of tlic 
genus, A. Fuiiklioiisei- (1957, p. 54), as Argentina, southern 
Brasil, and southern Bolivia. While this roughly includes the 
type locality of the species d(^sci"il)ed here, there apjiears to be 
no evidence that the two forms are sym]:)atric. The nearest 
locality. El ("armen, at which Tarl (ians collected saitrofjii is 
about 488 kilometers to the east of El Pailon. L. ]\Iiilh'r and \V. 
Ilellmich (19.S6, ]). 107) record saiiragii, mentioning some varia- 
tion in color, from "Curriche von San Ramon" north of San Jose 
de Chiquitos. "Curriche von San Ramon" is not specifically in- 
dicated on Midler and Ilelhnich's map {loc. cit., p. 115), though 
a short sjnir from the main itinerary extending north and west 
from San dose ch' Chiquitos may include or terminate at "Cur- 
riche." In any event, this locality is much closer to El Pailon 
than is El Carmen, San Jose de Chiquitos itself being about 218 
km. to the east. In the absence of evidence of sym])atry, the 
judgment that sauvagii and pailoiia are specifically distinct is 
l)ased on the several dififereuces cited in the diagnosis. 

P. pailona seems also to be related to /'. holiviana from which 
it appears to differ in having smaller discs on the fingers and 
toes, differently shai)ed parotoid glands, and in different colora- 
tion, consisting chiefly in the absence of white spotting or vermi- 
culation on the sides, and a nearlv immaculate ventral coloration. 

According to Funkhouser {loc. cit., p. 57), the range of holivi- 
ana is the eastern Andean foothills of Ecuador and Bolivia, thus 
apparently occurring at higher altitudes than that (350m.) at 
which pailona occurs. Again there appears to be no sympatry. 
and specific distinction is inferred from the extent of morphologi- 
cal and color differences. 

LITEEATUBE CITED 

FrxKHorsEK, Axxe 

19;")7. A review of the neotropical tree-frogs of the genus P}iyUo))udiit>a. 
Occ. Papers Nat. Hist. Mus. Stanford University, Xo. 5, pp. 1-90, 
figs. 1-40, maps 1-4. 

MuLi.ER, L. and W. Hellmich 

1936. Wissensc-haftliehe Ergebnisse der deutschen Gran Chaco Expedi 
dition. Part 1, Amphibia, Chelonia, Loricata. Stuttgart, pp. 
I-XVr, 1-120, pis. 1-8, text figs. 1-35, 2 maps. 



r' 








I'liilllniiii 'lit.sa pailona, paratype, MCZ l29()79, vcntial view to sluiw iiimmcii- 
latc venter. Carl Gans, photographer. 



'~^t^\J 



BREVIORA 

Miiseiuiiii of Comparative Zoology 



Cambridge, Mass. November 23, 1959 Number 114 



ANOMALOPHIS BOLCENSIS (MASSALONGO), A 

NEW GENUS OP FOSSIL SNAKE FROM THE 

ITALIAN EOCENE 

By Walter Auffenberg 

Department of Biology and Florida State Museum, 
University of Florida 

The genus Archaeophis as defined by Massalongo (1849) was 
based on two fossil specimens of presumed snakes originally in 
the collection of DI Canossa from the well known Middle Eocene 
teleost and plant beds of the Monte Bolea limestone near Verona, 
Italy. 

One specimen was designated as the type of Archaeophis 
proavus. It was smaller than the second specimen and much 
more complete, including almost the entire skeleton as well as 
an impression of the skin. The larger specimen lacked a head, 
nor was there any indication of an impression of the skin. This 
specimen w^as made the type of his second new species, Archae- 
ophis holcensis. Both were considered to be primitive snakes. 
Massalongo thought that they were merely quite distantly related 
to the only fossil snakes known at that time {Palaeophis and 
Falaeryx). They were not compared with any Recent genera. 
On the basis of the round form of the vertebrae as well as the 
general shape of the tail, Massalongo believed that Archaeophis 
was a terrestrial genus. 

In the following years the specimens became separated. The 
type of A. proavus was purchased by the Berlin Museum and the 
type of A. holcensis was deposited in the Museum of Compara- 
tive Zoology, having been purchased by Louis Agassiz. 



2 BREVIOBA No. 114 

A recent study of the two species formerly attributed to 
Archaeophis shows that A. bolcensis differs radically from A. 
proavus, and necessitates erecting a new family and genus based 
on the characters exhibited by the type and only specimen of 
bolcensis. 

In 1904, Janensch briefly described the skull, teeth, vertebrae 
and ribs of Archaeophis proavus. but in somewiiat greater detail 
than had been previously done by Massalongo. It was also in 
this paper that Janensch first suggested that the genus may have 
been a highly specialized marine form. This idea was based on 
his interpretation of the shape of the tail and body which he 
thought were very compressed in life. 

In 1906 he redescribed, in much more detail, the type of 
Archaeophis proavus. Various parts of the skeleton and particu- 
larly the skull liad been further prepared, and all elements were 
examined with the aid of a binocular microscope. The descrip- 
tion of the skull is far better than any previously given. Un- 
fortunately, the skull capsule is badly crushed, so that the indi- 
vidual bones are very difficult, if not impossible to trace. The 
bones of the jaw apparatus are all fairly well preserved. Certain 
parts are missing, Init in some cases the shape of the missing 
elements can be determined from stains or impressions appearing 
on the limestone slab on which the specimen is preserved. The 
head is unusually pointed for a snake, a fact of w^hich Janensch 
was quite cognizant. The teeth are most non-snakelike in that 
in cross-sectional view they are definitely 5-sided. Janensch 
interpreted certain elements partially hidden by matrix as quad- 
rates and squamosals. McDowell and Bogert (1954:67) suggest 
that the "quadrate'" agrees fairly well with the ))ranchial bones 
of an eel, but from ^vhat can l)e seen of them they are not con- 
siderably different from the same elements in Recent snakes. 
However, the ma.ior parts of the squamosals ( ?) are hidden by 
matrix, so that their exact shape, or method and place of articu- 
lation with the skull cannot be determined. 

The vertebrae and ribs of Archaeophis proavus are very im- 
portant, since only these elements can be compared with cor- 
i-esponding structures in Archaeophis bolcensis. Janensch points 
out that the exact form of each of these elements is very difficult 
to ascertain. This is due to several factors. The bone making up 



1959 ANOMALOPHIS BOLCENSIS 3 

the ueural areli of each element is very thin, and thus most of 
the elements are badly broken. When the limestone block in 
which the specimen was imbedded was split some of the bone 
making up the individual vertebrae was aceidently removed. 
Furthermore, the outer layer of bone has been lost in most of the 
vertebrae along the column, so that only the deeper part remains. 
This means that only the more general structure of the vertebrae 
can be determined in most parts of the column. Reasonably 
complete elements occur in only a few areas. For the most part 
these are regions which were originally completely covered by 
matrix until prepared at the Berlin Museum. Unfortunately, 
full preparation was not possible due to the fragile nature of 
the elements. No single vertebra could be profitably removed 
intact, so that anterior and posterior views of separate elements 
are not available. In addition, because of the overlapping nature 
of these vertebrae, the exact shape of either the neural arch, 
cotyle or condyle is not determinable. Janensch prepared 
a reconstruction of the side view of one of the middle thoracic 
elements based on several vertebrae in this area. His ventral 
view of a vertebra from the same part of the column is taken 
from only one prepared element (Fig. IC) . One of the vertebrae 
was split when the block was separated into two slabs, providing 
a fair cross section ( Fig. II). 

The exact number of vertebrae is difficult to ascertain with any 
degree of certainty due to broken or missing segments. Massa- 
longo had estimated this number as 507. Janensch believes his 
estimate of 565 is much closer. One hundred and eleven of these 
are caudal members. The entire length of the specimen is about 
95.5 cm., of which the tail makes up about 10.5 cm. 

The individual vertebrae show considerable variation in size 
and proportion along the length of the column. The more an- 
terior members are small, and higher than long. The middle 
vertebrae are largest. The posterior vertebrae are smaller, and 
longer than high. All of the segments appear to be procoelous. 
The condyle is strongly oval, with the main axis in the horizontal 
plane. From the side the condyle is flattened at its posterior end, 
not rounded as it is in all fossil and modern snakes. However, 
this difference may be due to breakage. The cross section of a 
thoracic element illustrates the unusual thinness of the neural 



BREVIORA 



No. lU 






o 

o 



E 





H 





Fifj. 1. Certain stnietural features of the type of Archeophis proavns 
-Massalongo {fide Jancnsch, 1906). A, Skull, from below. B, Reconstruction 
of tlif jaw apparatus. C, Vertebra 78 seen from below. D, Vertebra 46, seen 
r' 10111 the side. E, Cross sections of a single rib. F, Vertebral centrum and 
liypapophysis. G, Vertebral centrum and haemapopliysis. H, Cross section 
of tooth. I, Cross section of vertebra. J, Rib from the middle of the body. 



195P ANOMALOPHIS BOLCENSIS 5 

arch, even eonsicleriiig- that the outer lamellae were aeeidently 
removed in splitting the rock. There is no well developed neural 
spine, but rather a small keel that begins near the anterior edge 
of the neural arch and continues posteriorly to near the posterior 
edge of the element. The neural canal is roughly pentagonal in 
shape when viewed in cross section. The centrum is seen to be 
subtriangular in cross-sectional view. This condition is not seen 
in any fossil or Recent snake, where the centra are alw^ays oval 
to round. The zygosphene is very small and its articular surfaces 
are little developed. The articular facets of the post- and 
prezygapophyses are horizontal, not vertical as McDowell and 
Bogert (1954) claim. This is obvious not only from Janensch's 
illustration, but is clearlj^ stated in the text as well. The para- 
diapophysial articulations are single and rather small, located 
anterior to the middle of the vertebrae, and placed low on the 
centrum. Janensch states that several individual vertebral ele- 
ments indicate that these small structures are not the result of 
erosion, but that they may have been topped by a cartilaginous 
surface of considerable extent in life. Whether or not they 
represent an entire, or partial, synapophysis or are even homolo- 
gous to part of one is unknown. Accessory processes are ap- 
parently absent throughout the column, as they are in several 
families of fossil and Recent snakes. Well developed, laterally 
compressed hypapophyses are developed along the entire pre- 
sacral portion of the vertebral column. From the side they vary 
from sigmoid-sbaped structures in the anterior part of the 
column, to much low^er, broad-based triangular structures pos- 
teriorly. A midventral keel runs anteriorly from the antei'ior 
edge of the base of the hypapophysis to near the anterior edge 
of the centrum. The posterior portion of the base of the hypa- 
pophysis does not come in contact with the base of the condyle. 
Paired haemapophyses are found on all of the caudal elements. 
They are long, spine-like structures, placed far back on the 
centrum and directed almost straight downwards. The ribs are 
([uite long, bent posteriorly and considerably compressed lat- 
erally. There are no costal processes on any of the ribs. 

These are the most important structural features of the verte- 
brae and the ribs, as described by Janensch. Many of the char- 
acters are not found in other snakes, such as the triangular 



6 BREVIORA No. 114 

centrum in cross section, the very small articular processes, the 
poorly developed zygantrum and zygosphene, and the strongly 
oval condyle. The ribs are considerably longer, less curved, and 
more compressed than in any other fossil or Recent snakes 
Imown. For a complete description of these structures as well as 
those of the skull and skin the reader is reTerred to Janensch 
(1906). 

On the basis of his study of the tyj)e of Archacophis proavus, 
Janensch concluded that: (1) the type of A. proavus is a highly 
specialized marine snake; (2) the type of A. holcensis belongs 
to the same genus, perhaps even to the same species as the type 
of A. proavus; and (3) Archacophis represents a new family of 
snakes, the Archaeophidae, distinguished mainly on the shape 
of the teeth. 

McDowell and Bogert (1954: 66-67) point out several 
remarkable features of the vertebral structure of Archacophis. 
These are: (1) the smooth and unsculptured vertebrae; (2) the 
fact that the prezygapophj^sis is reduced to a small spine; (3) 
the plane of the articular surface of the postz.ygapophysis is 
vertical, not horizontal; and (4) there is no neural spine. To 
these authors the combination of characters seemed to suggest 
that Arcliacopliis was not a snake. They point out that some of 
the features of the genus are found in certain eels, and other 
characters are similar to those found in some snakes. They state 
that, "Looking at Janensch 's figures, we are led to suspect that 
Archacophis might not even be reptilian, for there is much to 
suggest (though not enough to prove) affinities with the teleost 
fishes, particularl}^ some eel of the OphicJitJius-Mke group." They 
go on to suggest that Archacophis might best be placed as a 
vertebrate of unknown affinities. Some of the characters they list 
as remarkable (if Archacophis is a snake) are certainly not very 
important, or very remarkable, such as the degree of smoothness 
and sculpturing of the vertebrae. As pointed out above, some of 
the outer lamellae of bone have been lost. Only general slia])e 
of the structures is really determinable. Much detail has been 
lost, and absence cannot be used as any sort of criterion, except 
as regards degree of mineralization or breakage. Furthermore, 
details of configuration of Recent snake vertebrae vary greatly 
from genus to genus. The absence of ;i well developed neural 



1959 ANOMALOPIIIS BOLCENSIS 7 

spine is characteristic of several groups of Recent and fossil 
snakes. As pointed out previously, McDowell and Bogert mis- 
interpreted Janensch's figure of the side view of the middle 
thoracic vertebra as well as his text discussion, since they stated 
that the articular surface of the postzj-gapophysis is vertical. 
Both the figure and the text clearly indicate that this surface is 
horizontal, as it is in all snakes. However, this in itself proves 
nothing, since a zygapophysial articular surface when present in 
fishes is sometimes horizontal. 

McDowell and Bogert, as well as Janensch, assumed that the 
type of Archacophis bolccnsis was structurally very similar to 
the type of A. proavus. Massalongo stated that this was true, but 
his analj'sis of the characteristics of both species was rather 
superficial. Janensch examined proavus verj^ closely, but failed 
to examine the type of holcensis. McDowell and Bogert relied 
only on Janensch's description of proavus for the generic char- 
acters. The present study of the type of A. holcensis has shown 
that this species is quite unlike proavus in many of its structural 
details. The two species are very different in a number of im- 
portant features. I wish to thank Drs. E. E. Williams and A. S. 
Romer for permission to examine the type of Archacophis holcen- 
sis, and especially for allowing me to prepare it further. 

The specimen in the Museum of Comparative Zoology is repre- 
sented b}' sections of the anterior, middle and posterior parts of 
the vertebral column and their associated ribs. These were origi- 
nally located on three limestone slabs (MCZ 1001, 1002 and 
1003). One of the slabs was intentionally sawn in half and one 
l)ecame broken sometime in the past. The specimen is consid- 
erably larger than the type of A. proavus. There is no trace of 
the skull. A dark stain on the limestone matrix in several places 
may represent portions of the flesh, or skin, but no details of 
scalation, if it exists, can be made out. The middle and posterior 
parts of the specimen are resting on their dorsal sides, so that 
only the ventral surfaces of the vertebrae are visible. The an- 
terior section rests on its ventral side. Unfortunately^ through- 
out most of the column the vertebrae are crushed, or badly 
broken, on the exposed surface. However, most obvious at once 
is the fact that, broken as they may be, there are no hypa- 
pophyses on any of the middle or po.sterior vertebrae. On this 



8 BREVIORA No. 114 

basis alone it is obvious that two genera are involved. As in 
A. proavus the ribs are all long, curved backwards only slightly, 
and lack costal processes. 

To aid in uncovering certain details of structure of both the 
vertebrae and ribs, acetic acid had been applied to several areas 
along the vertebral column, in the past. This was a most unsatis- 
factory undertaking, since the loss of the encasing matrix caused 
ends of ribs and the dorsal or ventral surfaces of some of the 
vertebrae to disintegrate. Fortunately, dry preparation was still 
possible in several areas. Two vertebrae were completely freed 
from the matrix so that anterior and posterior views could be 
drawn and studied. The surfaces otherwise hidden were exam- 
ined and the structures compared with those in fossil and Recent 
snakes. The isolated vertebrae are considerably cracked. Some 
processes had been broken off in the past, and a few slightly 
damaged in extricating the delicate elements. It was thus neces- 
sary to reconstruct, very slightly, the missing parts of the isolated 
middle thoracic element. The reconstructed portions were drawn 
from other vertebrae, in the immediate area, in which these 
particular processes were still complete. The quite reasonable 
complete reconstruction of at least the middle dorsal member 
allows a thorough comparison with A. proavus and with all fossil 
and Recent snakes. Most important, it proves beyond doubt that 
Aychaeophis holcensis is a snake, regardless of the taxonomic 
position of A. proavus. 

Unlike A. proavus, the vertebrae of A. holcensis possess well 
developed zygantral and z.ygosphenal articular surfaces. There 
is a well developed neural spine. The pre- and postzygapophysial 
articular surfaces are also well developed. The vertebrae are 
much more robust than are those of A. proavus. The paradiapo- 
physial articulating surface is better developed than in A. 
proavus. The elements are well ossified throughout the column, 
and tlie outer laminae of bone are still present. 

To facilitate future comparisons of Arrhacophis holcensis with 
other fossil or Recent snakes, a description of the partly recon- 
structed middle thoracic vertebra is given below. 

Description of vertebra : — Centrum long, the sides converging 
posteriorly, a truncated triangle from below. ]irovided with a 



1959 ANOMALOPHIS BOLCENSIS 9 

well developed, slightly oval cotyle anteriorly, and a well de- 
veloped condyle posteriorly, which is only slightly separated 
from the basic portion of the centrum by a small constricted area. 
A haemal keel is present, gladiate to spatulate in shape, and 
slightly flattened ventrally. It extends from near the lip of the 
cotyle to just anterior to the condyle. The ventral surface of the 
eentrum is slightly- flattened on either side of the haemal keel. 
There is a faint subcentral ridge extending posteriorly from just 
behind the synapophysis, or the paradiapophysial articular facets 
on either side, to near the base of the condyle. The paradiapo- 
physial articular facets are oval in shape, and located low on the 
centrum, well in front of the middle of the vertebra. From the 
side, the ventral surface of the centrum is reasonably straight. 
not concave, or convex, as it is in many snakes. The buttresses of 
the prezygapophyses are well developed. In anterior view they 
are seen to possess a sharp anterior edge. The buttress is closer 
to the median line ventrally than dorsally. Between the buttress 
and the projected lip of the cotyle there is a small scooped out 
area. Whether or not this small depression possessed a nutritive 
foramen is unknown. There is no extended accessory process 
below the prezygapophysial facet. The facet is slightly liigher 
anteriorly and outw^ardly than posteriorly and medially. From 
above, the articulating surface is ovoid to sub-triangular. The 
articular surfaces of the postzygapophyses are developed on the 
laterally expanded posterior portion of the neural arch, as they 
are in all snakes. The facets are oval, the main axis being antero- 
posteriorly. The neural arch is fairly long and not greatly 
emarginate from above. The interzygapophysial ridges are fairly 
prominent, especially posteriorly. The neural spine is long at its 
base, extending from just above the base of the zygosphene to 
the anteriorl}' indented posterior edge of the neural arch. It is 
slightly thickened and truncated dorsally. The posterior edge is 
slightly lower than the anterior one. There are no overhanging 
edges at the top of the spine either anteriorly or posteriorly. 
From above, the dorsal edge of the spine is relatively thin, not 
overly thickened into a peg-like or oval-shaped structure, as in 
some primitive snakes. The zygosphene is well developed, robust, 
but small for the length of the centrum when compared to the 
Boidae. From above, the anterior edge of the zygosphene is 



10 



BREVTORA 



No. 114 




1959 ANOMALOPHIS BOLCENSTS 11 

almost straig-lit, perhaps with a small median notch. From the 
front, the zygosphene is somewhat thickened dorsoventrally, the 
dorsal edge slightly convex, presumably with a small notch 
medially. The zygosphenal articular facets are oval, the anterior 
edge being at an angle of about 45 degrees when viewed from 
the front. The neural canal is fairly small, roughly triangular 
in shape. The medial edges of the prezygapophysial articular 
facets are even with the floor of the neural canal. It is impossible 
to determine the position and size of the zygantral foramina, if 
they are present at all. Whether or not a medial anapophysis 
occurs on the floor of the neural canal cannot be determined, 
since the canal is filled with matrix, which, if reinoved, would 
weaken the isolated fossil considerably. 

The partially reconstructed middle thoracic vertebra is illus- 
trated in Figure 2. Table 1 gives all of the pertinent measure- 
ments possible on the vertebra. 

TABLE I 

APPROXIMATE MEASUREMENTS (IN MM.) OF A SINGLE 

THORACIC VERTEBRA FROM THE TYPE OF 

AFCHAEOPHIS BOLCENSIS 

Centrum length 11.0 

Centrum width at its narrowest part 8.0 

Pre-prezygapophysial width i 22.0 

Post-prezygapophysial length ^ IG.O 

Cotyle width i.o 

Cotyle heiglit 4.0 

Length of neural spine along dorsal edge -4.0 

Height of neural spine along anterior edge 3.;") 

Zygantrum width at its widest part 5.5 

A single vertebra was also removed from the anterior part 
of the column of the type of A. holcensis. Unfortunately, the 
anterior elements are even more crushed than the middle and 
posterior ones. Unlike the middle thoracic region, the anterior 

1 The pre-iirezysapoph.vsial width is moasiired from the outer edge of one 
prezygapophysial facet to the outer edge of the opposite prezygapophysial facet. 

2 The post-prezygapophysial length is measured from the posterior edge of the 
postzygapophysial facet to the anterior edge of the prezygapophysial facet. 



12 



BREVIORA 



No. 114 



portion of the specimen is resting in a natural position, so that 
the ventral surface of each vertebra is imbedded in the matrix 
and thus is somewhat protected. The dorsal and lateral surfaces 
are almost completely crushed. By careful preparation of one 





B 




Fig. 3. Reconstruction of assumed cross section of the mid-bodj' region 
in Anomalophis bolcensis (A), compared with that of a Recent Constrictor 
constrictor (B).The lower drawing (C) illustrates the general shape and 
position of the single hypapnphysis present in the more anterior vertel)rae 
of the tjTpe of Anomalophis bolcensis. 



in.'iO ANOMALOPHIS BOLCENSIS 13 

of tlu'so elouieiits it was jjossible to remove it from tlie articulated 
|)o.sition. The ventral surface of the centrum is provided with a 
well developed hypapophysis. It is only slij2:htly compressed 
laterally. From the side it forms a very gentle sigmoid-shaped 
structure, directed more ventrally than posteriorly. Anteriorh', 
it is continued as a Ioav midventral keel to the lip of the cotyle. 
The spine is located ])osteriorly on the centrum, the posterioi- 
edge beginning just anterior to the slight constriction near the 
base of the condyle (Pig. 3). 

A well developed ridge extends from the posterior edge of 
each paradiapophysis to near the base of the condyle. These 
ridges converge only slightly posteriorly. The ventral surface 
of the centrum is quite flattened between the two ventrolateral 
subcentral ridges. The lateral borders of the centrum are almost 
parallel, but converge slightly posteriorly. No nutritive foramina 
are visible. Very little of the structure of the lateral or dorsal 
surfaces of the element are determinable. The prezygapophysial 
articular surfaces are quite evident. They are horizontal. From 
above, they are oval, with the long axis nearly parallel to the 
axis of the centrum, but diverging anteriorly. 

The ribs vary considerably along the column in regards to their 
length, curvature, and degree of compression. Anteriorly, these 
elements are about 80 mm. long, only slightly curved, and very 
little compressed. They are quite attenuate. The middle thoracic 
members are somewhat more robust and more curved, particu- 
larh^ proximally. The entire length of a single rib in this area 
is about 90 mm. These members are proportionately more com- 
pressed proximally than are more anterior or more posterior 
ribs. Near the proximal end they are somewhat subtriangular 
in cross section. The more posterior ribs are about 110 mm. 
long, very filiform, little curved and hardly compressed at all 
along any part of their length. Views of the individual ribs and 
cross sections of them are given in Figure 4. The cross sections 
were obtained by taking out small broken sections of ribs in the 
various areas and then replacing them after examination. The 
proximal rib ends vary only slightly throughout the column from 
almost round to decidedly oval. The middle thoracic members 
are provided with a small indentation near the middle, pre- 
sumably marking a very weakly developed costal process in these 



14 



BREVIORA 



No. 114 



elements. More anterior and posterior ribs lack any evidence of 
such a structure. 

The shape of the rib, i.e., its length and degree of curvature, 
indicates that Archacophis holcensis possessed a body which was 
much more laterally compressed than any of the living snakes 






:■ dnnD 



c 




Fig. 4. Lateral and cross sectional views of the ribs from several regions 
along the body of the type of Amomalophis holcensis. Left, anterior rib ; 
middle, thoracic rib; right, posterior thoracic rib. 

(Fig. 3). Its body shape is probably approached most closely 
hy the posterior portions of some of the living hydrophids. Jan- 
ensch's conclusion that this compression indicates that Archaco- 
phis represents a highly modified marine form seems quite rea- 
sonable. 



1959 ANOMALOPIIIS BOLCENSIS 15 

The type of Archacopliis bolccnsis has been .shown to differ 
in a number of structural features from that of Archaeophis 
proavus. The most important differences are: (1) the presence 
of a much better developed zygantrum, (2) the absence of hypa- 
pophyses throughout the middle and posterior portions of the 
vertebral column, (3) a condyle which is mucli more rounded, 
and (4) a well developed neural spine. Archaeophis holcensis is 
unquestionably a snake. The type is to be referred to a genus 
other than that which includes the type of A. proavus. It has 
been suggested that the latter may be a fish (McDowell and 
Bogert, 1954). This idea was based on several characters already 
mentioned above. However, as these authors have pointed out, 
certain features of A. proavus are quite snakelike. The phyletic 
position of A. proavus is thus uncertain. Furthermore, this 
species is generically distinct from the type of A. holcensis. The 
type of Archaeophis has been fixed as proavus (Kuhn, 1939). 
The species holcensis is thus left without a generic name, for 
which I propose the following : 

ANOMALOPHIS new genus 

Diagnosis. A genus of Eocene snake, differing from all Recent 
and most fossil genera in lacking a costal process of the ribs. In 
the absence of this structure Anomalophis is approached by the 
Paleophidae.i It differs from members of this family in lacking 
well developed pterapophyses on the posterior part of the neural 
arch. 

Genotype. Anomalophis holcensis. 

Anomalophis bolcensis (Massalongo) 

Diagnosis. Same as for the genus. 
Holotype. MCZ 1001, 1002 and 1003. 

Type Locality and Horizon. Near Verona, Italy; Monte Bolca 
limestone. Middle Eocene. 

1 Romer has recently (1956:563) provisionally placed the Paleophidae in the 
Lacertilia. The family was formerly considered to include only Palcophifs and 
Ptn-ospheiius, though' he added SimoUnphix and Pachiiophis. The two latter 
senera may indeed be lizards, since their vertebrae are quite varanoid in general 
appearance. These two jrenera are placed in the Sinioliophinae (Pachyophinae). 
However, Paleophis and Ptrrosphcnus (Palaeophinae) are most certainly snakes, 
thouffh perhaps somewhat aberrant. I propose that Paleophis and Ptoosphenus 
be returned to the Suborder Ophidia (Serpentes), and that they constitute a 
distinct family distinsrnisbed lar.sely on the basis of well developed pterapophyses 
and double hypapophyscs in some of the vertebrae. 



16 BREVIORA No. 114 

The combination of the differences, pointed out above, in both 
the vertebrae and ribs su<i'gests that Avoinolophis should be placed 
in a separate family of snakes, the Anomalophidae, containing 
only one genus at the present time, Anomalophis. The single 
specimen of the type species of the genus lacks a skull, but cer- 
tain vertebral and rib characters are quite distinctive. The new 
familial characteristics are as follows : A single hypapophysis 
occurs in each of the more anterior vertebrae ; in the middle and 
posterior vertebrae the hypapophysis is reduced to a low keel ; 
centrum long; neural spine with a long base and not overly thick- 
ened ; neural arch normal, without aberrant processes ; no well 
developed acecssory processes; ribs long, filiform, no costal proc- 
esses, slightly compressed, with little curvature. 

Cranial and pelvic girdle material of all of the earlier fossil 
snakes is sorely needed. Until this becomes available it seems best 
to defer a complete comparison of all of the later Mesozoic and 
early Cenozoic boid-like snakes (of which Anoni'alophis seems to 
be a member) until a later date. 

LITERATURE CITED. 
.Tanensch, W. 

1904. Eine fossile Schlange aus dem Eoean des Monte Bolca. Zeitschr. 
deutsch Geol. Gesell., vol. 56, pp. 54-56. 

1906. liber Ardhaeophis proavits Mass., eine Schlange aus dem Eoean 
des Monte Bolca. Beitrage zur Palaontologie Osterreich-Un- 
garns, vol. 19, pp. 1-33. 

KUHN, O. 

1939. Fossilium Catalogus, Pars 86, Ophidia, p. 13. 

McDowell, S. and C. M. Bogebt 

1954. The systematic position of Lanthanotus and the affinities of the 
Anguinomorph lizards. Bull. Amer. Mus. Nat. Hist., vol. 105, 
art. 1, pp. 1-82. 

Massalongo, a. 

1849. Specimen photographicum animalium quorundam plantarumque 
fossilium agri Veronensis. Verona, pp. 1-55. 

ROMEE, A. S. 

1956. The osteology of the reptiles. Chicago: University of Chicago 
Press, xxi + 722 pp. 



y •^■^^-\^/\^ 



BREVIORA 



seminri of Comparative Zoology 



Cambridge, Mass. Nove.mber 25, i;)59 Number 115 



THE LEMON-COLORED PLEXAURIDS FROM THE 
WEST INDIES AND BRAZIL 

By 

Elisabeth Deiciimann 

Museuni of Comparative Zoology 

and F. M. Bayer 

United States National Museum 

In Kiikenthal's monograph of 1924 are listed two yellow plex- 
aurids, referred to different genera and each definitely falling 
outside the normal limits of the genus in which it is placed. 
These two forms are Plexaura flavida (Lamarck) — which, inci- 
dentally, also appears in a dark purple phase — and a species 
which, hitherto, most commonly has been listed as Eunicea 
humilis Milne Edwards and Haime. The latter, thanks to the 
junior author's delving into old, forgotten literature, has now 
been given the older name sulphurea Donovan. Lamarck's 
species appears to be common in the West Indies, from the 
northern shores of South America to the eastern coasts of Flori- 
da, but apparently it does not reach Bermuda (see Verrill, 1907, 
p. 261), while Donovan's species is one of the most characteristic 
elements of the Brazilian reef fauna, ranging from 7°S to 20° S 
latitude with a few, rather dubious, records from Curacao, 
Aruba, and San Bartholomew Island, in the West Indies. 

As the Museum of Comparative Zoology possesses a large 
amount of material of both forms, and since a number of con- 
fusing statements concerning them have crept into the literature, 
there seems good reason for revising these two species and allo- 
cating them to their proper place, and at the same time reviewing 
their history. 



2 BREVIOKA No. 115 

Verrill established in 1907, page 309, a new genus Plexauroi)- 
sis, to accommodate what he considered a new species, P. hicolor, 
taken from shalloAv water in Bermuda. The species was not 
found again until 1951, when it was realized that this small form 
merely represented the immature stage of the tall Pse^idoplex- 
aura crassa (Ellis and Solander), with a few flat rods still re- 
tained in the tentacle bases, and Avith a greater number of clubs 
and spindles with spines on the external side than are usually 
present in the older colonies. Hence Plexauropsis is a straight 
synonym of Wright and Studer's Pseudoplexaura 1889. 

Verrill 's genus and species was, however, included in Kiiken- 
thal's monograph of 1924 (p. 118), with a somewhat distorted 
description : The presence of the few tentacle rods was omitted, 
and too great emphasis was placed on the one-sided spinulation 
of the external spindles — a feature which occurs more or less 
pronounced in a number of plexaurids, but often has not been 
specially noticed. Neither Kiikenthal nor his students had, of 
course, access to Verrill 's material. 

As a result of Kukenthal's misleading account, Stiasny in- 
cluded a number of species in Plexauropsis, in his paj^ers from 
1935 to 1951. 

His tricolor, from Bermuda (1935 a, p. 241 ; 1935 b, p. 69, pi. 3, 
fig. 13 and text fig. R), is identical with Plexaura homomalla 
(Esper), redescribed by Verrill in 1907. 

His hicolor, based on some old material labelled humHis (from 
Curagao and "No locality," 1935b, p. 73), was later transferred 
back to humilis (1935c, pp. 107-115, text figs. 1-3), with a de- 
tailed analysis of all earlier records appended. 

His Jmmilis (1935b, pp. 74-78, pi. 3, fig. 14, pi. 7, fig. 32, text 
fig. T), was later changed to Eunicea hicksoni (1935c, p. 115), 
but it represents merely the widespread form Plexaura flexuosa 
Lamouroux. Most of this material had been identified by Gordon 
(1925, p. 17) as mutica Duchassaing and Michelotti. 

In addition, Stiasny had included the following feather-shaped 
forms in the genus Plexauropsis: 

P. flavida n. sp. (1935a, p. 242: 1935b, p. 71, pi. 1, fig. 5, text 
fig. S) from Curasao. 

P. sp. (1935b, p. 72) from "Antillen." 

P. puertorealis n. sp. (1942, p. 101, pi. 1, figs. 5-8) from 
Venezuela, taken with numerous colonies of his new '' flavida." 



1959 LEMON-COLORED PLEXAURIDS 3 

All these species are, however, identical with Lamarck's 
variable and \\4despread Gorgonia flavida. The latter is briefly 
mentioned by Stiasny (1935b, p. 57), but he merely states that 
the spicules agree with Kiikenthal's description and, as the latter, 
he places it in the genus Plexaiira. 

The result of this survey is that of all the species referred by 
Stiasny to the now defunct genus Plexauropsis, only two remain, 
namely, Milne Edwards' humilis, now sulphur ea of Donovan, 
and Lamarck's flavida, and for these two forms the proper 
generic name must be found. 

In 1931, Aurivillius established a new genus, Muriceopsis, 
to which humilis (sulphurea) belongs — in fact it ends up by 
being the type species — and this genus has now had its diagnosis 
emended so it also accommodates Lamarck's flavida. 

Whether any member of the genus occurs outside the tropical 
Western Atlantic remains to be seen. Possibly some little known 
West African species may have to be included in the genus. 
Furthermore, the Museum of Comparative Zoologj^ possesses a 
branch which hardly can be distinguished from the typical 
flavida from the West Indies, but which bears the label Mauritius 
(N. Pike). While the possibility of mislabelling is not excluded, 
it must, nevertheless, be remembered that in the case of the 
crustaceans several AVest Indian forms have been taken in 
Mauritius and Madagascar, so this disjointed distribution is 
not unique. The branch from Mauritius bears some similarity to 
Lamouroux' figure of oUvacca from the Indian Ocean and may 
possibly be referred to that species. 

Muriceopsis Aurivillius 1931 (emended) 

Muriceopsis Aurivillius, 1931, p. 115. 

Diagnosis. Colonies either low, bushy, with crooked branch- 
lets or tall, feather-shaped with crowded, straight pinnulae. 
Poh'ps small, either completely retractile so the pores are flush 
with the surface, or they form low mounds or crescent-shaped 
lower lips. Spicules arranged in an inner layer of slender, 
pointed spindles with distant warts, yellow^ or purple in color, 
a middle layer of slender or stout spindles and an external layer 
of clubs or spindles with large, mostly flattened, spines on the 



4 BREVIORA No. 115 

external side. Polyps with a delicate operculum of flattened, 
Avarted rods, bent or straight. Color lemon yellow, buff or purple. 

Type species. Gorgonia sulphurea Donovan 1825. 

Remarks. Milne Edwards and Ilaime's name, Eunicea humilis 
1857, has been most commonly used and that name takes in 
Muriceopsis tuherculata Aurivillius 1931, nee Gorgonia tuhercv- 
lata Esper 1791. 

Aurivillius selected, quite properly, what he considered to be 
p]sper's tuherculata as the t.ype, on the basis of some material 
from the "West Indies which had been identified by KoUiker. 
However, Esper 's species came from the Mediterranean and his 
figure (1791, pi. 37) agrees completely with a number of fans 
received from the Mediterranean in 1957 (in U.S.N.M.). These 
have been referred to Eioiicclla lata Kiikeuthal, but the clubs, 
0.07 mm. long, are rather similar to those found in Eiinicella 
verrucosa (Pallas) — from the same lot — hence lata possibly 
should be withdrawn as a synonym of Pallas' species. On the 
other hand, some of Esper 's material — possibly that which Avas 
examined by Kolliker — may well have been a West African 
species, which is known to range from Congo to Cape Blanco, 
and possibly farther north. This species appears in several of 
Stiasny's papers, first as Plexauropsis mauritaniensis, but finally 
withdrawn by himself as a synonym of Mnriccides chuvi Kiiken- 
thal (see Stiasny 1951, p. 17). From the excellent figures which 
Stiasny has given, one has the impression that he is dealing 
with a form which externally is quite similar to Esper 's tuhercu- 
lata. As for the spicules, Stiasny indicates that some are deep 
yellow, often with purple center, and that purple spicules, in- 
cluding some red ones, occur scattered but do not form a con- 
tinuous inner layer. We do not know whether Kolliker had any 
material of such a species, but he referred two colonies from St. 
Bartholomew Island, West Indies, to Esper 's tuherculata and 
gracilis n. sp. Both colonies, in the Zoological Museum in Stock- 
holm, were re-examined by Aurivillius, who considered gracilis 
a variety of tuherculata. 

In this connection the question is of little importance since 
Aurivillius' "tuherculata" is based on specimens of what com- 
monly has been called Eunicea hiimilis Milne Edwards and 
Ilaime, now sulphurea Donovan. 



1959 LEMON-COLORED PLEXAURIDS 5 

The three other species which Aurivillius lists as valid (in his 
key) are all synonyms of the type species (Muricea bicolor 
Wright and Studer 1889, Muricea acropora Verrill 1912, and 
Eunicea humilis Milne Edwards and Haime 1857). 

As dubious species, he lists three. Of these Muricea granulosa 
Verrill (1870, p. 373, text fig. 4) came from Sherbro Island, West 
Africa ; it has clumsy spindles and clubs, which are stated to be 
all yellow or white, with tendency to form flat spines with bi- 
lobed or tri-lobed ends; the largest spindles measure 0.69 mm., 
while the clubs measure 0.49 mm. From Verrill 's description 
it looks as if the colony is fan-shaped and it may possibly repre- 
sent a colony of Muriceides chuni Kiikenthal in which no purple 
or red spicules were observed. 

The second, Muricea vatricosa (Valenciennes) — not Esper, 
as Aurivillius writes — from Bissago Archipelago, West Africa, 
has coarser spicules; the spindles are up to 0.84 mm. long, the 
clubs 0.66 mm., and the slender spindles 0.72 mm. long. (Verrill's 
measurements [1870, p. 374] are based on material received from 
Kolliker, who lists the species [1865, p. 136]). Nothing is said 
about its outer form and its position is quite uncertain ; it is not 
even mentioned by Kiikenthal in 1924. 

The third, Muricea humosa (Esper) (1794, p. 36, pi. 6, figure 
erroneously labelled "placomus"!) is stated to have come "most 
likely" from Curacao. Its crumbling, earth-like tissue and its 
general shape suggest PJexaura JwmomaUa (Pallas), a form not 
uncommon along the northern shores of South America. The 
spicules of hnmoso are — according to Kolliker — small, 0.44- 
0.67 mm. long, which fact agrees reasonably well with what one 
finds in homomaUa, where the clubs and spindles have a tendency 
to fomn long, flat, narrow spines on the external side. 

Key to the species accepted in the genus Muriceopsis Aurivillius 

1931 

Low, 10-20 cm. high, soiubby bushes, with crooked branchlets, strongly 
waited ; polyps completely retractile, with minute pores, scattered over the 
surface. Inner layer slender, pointed rods with distant warts, about 0.35 
mm. long; middle layer with hea\-y waited spindles, mostly 0.60 mm. long; 
uuter layer clubs and spindles 0.25-0.35 mm. long with laciniated leaves on 
the external side. All spicules j^ellow or those in the inner layer purple. 



6 BREVIORA No. 115 

Shallow water, coasts of Brazil, possibly in some localities farther north. 

(St. Bartholomew, Venezuela.) 

Muriceopsis sulphur ea (Donovan) 
Tall, up to 40 cm. high, feather-shaped colonies, often secondary feathers 
arising in the same plane; pinnulae 3-6 cm. long, crowded, not all in one 
plane; polj-ps with minute pores, rarely forming low warts or half moons. 
Inner layer slender, pointed rods averaging 0.27 mm. in length; middle 
layer mostly slender rods with distant warts, often slightly bent, mostly 
0.3 mm. long, rarely up to 0.5 mm. long; outer layer clubs and spindles with 
tall spines on the external side, up to 0.40 mm. long. Spicules in inner layer 
purple; in middle and outer layer yellow, or partly or all purple. Shallow 

water, West Indies, possibly also Mauritius. 

Muriceopsis flavida (Lamarck) 

Muriceopsis sulpiiurea (Donovan) 
Plates 1, 3 

Gorgonia sulphtirra Donovan, 182."), p. 126, and a colored plate. 

Eunicca humilis Milne Edwards and Haime, 1857, pp. 149-150, pi. B 2, fig. 1 ; 

Verrill, 1866-1871, p. 360, pi. 4, figs. 4, 4a-b; Kunze, 1916, p. 5.10 

(not examined). 
Muricea humilis and var. humilis, var. mutans and var. macra Verrill, 1912, 

pp. 377-379, pi. 29, figs. 1-la, pi. 32, figs. 4-5, pi. 35, fig. 2, text fig. 1. 
Muriceopsis humilis Aurivillius, 1931, p. 115 (not examined). 
Muricea aeropora Verrill, 1912, p. 379, pi. 32, fig. 3, pi. 35, figs. 1, la. 
Muriceopsis aeropora Aurivillius, 1931, p. 115 (not examined). 
Xee Eunicea mutica Duchassaing and Michelotti, 18G0, p. 28, pi. 3, fig. 10; 

Gordon (partim) 1925, p. 17 (=Plexaura flexuosa Lamouroux 1816). 
Nee E. humilis Stiasny, 1935b, p. 74, pi. 3, fig. 14, pi. 7, fig. 32, text fig. T 

(=Plexaura fi.exuosa Lamouroux 1816). 
Eunicea Htrinn Valenciennes, 1855, p. 13. 
Veo Gorgonia citrina Esper, 1794, p. 129, pi. 38 [—Pterogorgia citrina 

(Esper)]. 
Muricea bicolor Wright and Studer, 1889, p. 134, pi. 23, fig. 11, pi. 25, fig. 8; 

Verrill, 1912, p. 380 (not examined). 
Muriceopsis bicolor Aurivillius, 1931, p. 115 (in key). 
Plexauropsis bicolor, Stiasny, 1935b, p. 73. 
Xec Plexauropsis bicolor Verrill, 1907, p. 265, pi. 33B, figs. 2b-c, pi. 35A, 

figs. 1-2; text fig. 149 [=Pseudoplexaura crassa (Ellis and Solander") 

1786]. 
Muricea tuberculata Kolliker, 1865, p. 136. 
Xec Gwgonia tub ercidata Esper, 1794, p. 137, pi. 37. \—Eumcella s]i. 

(parlim) and ? Muriceides chuni Kiikenthal, 1919]. 
Muriceopsis tiihcrcnlata Aurivillius, 1931, p. 115. 



1959 liEMON-COLORED I'LKXAL'RIDS 7 

Diaynosis. Low, scrubby coluuit's, 10-20 cm. liigli, with crooked 
branchlets, 3-5 nun. in diameter; surface knobby and the small 
pores mostly upwardly directed, toward the tip of the branch- 
lets; sometimes a short, hook-like lower lip is formed. Branching 
dichotomously or loosely ])innately, wdtli branchlets alternate or 
opposite, occasionally all developed from one side. Color bright 
yellow, whitish or pale buff. 

Si)icules in inner l-dyer slender, pointed rods with distant 
warts, usually about 0.35 mm. long; in middle layer a varying 
number of moderately thick spindles, about 0.6 mm. long watli 
comi)osite warts of fairly uniform distribution and size ; a few 
may be 0.7 mm. long. In some colonies this middle layer is 
weakly developed. Outer layer with large asymmetrical clubs 
and spindles with large irregular spines or laciniated leaves on 
the external side while the inner side has low, composite warts. 
The poh'ps have a weakly developed operculum of flat rods of 
which the largest are curved, with warts along the edge ; length 
varying from 0.10-0.12 mm. 

The spicules are either all 3'ellow or those in the inner layer 
— and occasionally a few in the middle and outer layer — are 
purple. 

Type. Possibly in some collection in England. 

Type locality. "Brazilian Sea." Collected in numbers by 
8enor Eibello. 

Distribution. Taken under various names in Bahia: hu)nili.s, 
bicolor, acropora. According to Verrill, common in Porto Seguro 
and i-anging at least from Parahiba, N. of Pernambuco, to 
Guarapary in the South, that is, from about 7°S to 21°S latitude. 

In addition, the species has been listed from San Bartholomew, 
W.I., hy Kolliker and Aurivillius (same material), and from 
Curasao (old material in Holland) by Stiasny. Both these non- 
Brazilian localities may possibly be wrong. At least the species 
has not been reported from north of Pernambuco in any recent 
collection. 

Specimens examined. One fragment with purple inner spicules 
and no locality, probably from Kolliker (labelled no. 51 — not a 
M.C.Z. number). Twelve lots from Brazil: Murraio, Armagao, 
Victoria, Guarapary, Porto Seguro, Bahia and ''Brazil." 



8 BREVIORA No. 115 

Remarks. Donovan's material is brilliantly sulphur-colored 
and, as the plate indicates, falls into two varieties: an elongate 
and a more scrubby one. It is not known whether the type of 
humilis lacked the purple inner spicules, but this was apparent!}' 
assumed by Wright and Studer, who realized that their hicolor 
otherwise w^as almost identical with humilis (sulphurea). Verrill 
described, in 1912, the three varieties of Muricea humilis as 
having purple spicules in the inner layer, while his own species, 
M. acropora, has only yellow or white spicules. 

The material in the Museum of Comparative Zoology lacks in 
all cases the purple spicules, except the old fragment, without 
locality, "no. 51" — -in all likelihood received from either 
Kolliker or the Paris Museum. The material shows great varia- 
tion in the size of the colonies as well as in the number of branch- 
lets and their thickness. Here and there are found colonies with 
very slender branchlets, and in these the middle layer of thick 
spindles is poorly developed, as in Muricea gracilis of Kolliker 
(^ forma gracilis of Aurivillius) — ^ perhaps the result of unfavor- 
able ecological conditions. 

MuRiCEOPsis FLAViDA (Lamarck) 
Plate 4 

Keratophyton Seba, 1734-1765, III, p. 198, pi. 107, fig. 8. 

Crorgonia flavida Lamarck, 1815, p. 158; 1816, p. 318; 1836, p. 496; 

Lamouroux, 1816, p. 402; Dana, 1848, p. 664. 
Plexaura flavida Valenciennes, 1855, p. 12; Milne Edwards and Haime, 

1857, p. 153; Kolliker, 1865, p. 138, pi. 13, fig. 6 (cross-section of axis) ; 

Verrill, 1907, p. 138, pi. 36A, fig. 4 (table explanation to 35A), text 

fig. 148; Kiikenthal, 1917, p. 335; 1924, p. 117; Moser, 1921, p. 114; 

Stiasny, 193.jb, p. 57 ; 1951, p. 54. 
Gorgonia spicifera Dana, 1848, p. 117. 
Nee Gorgonia spicifera Lamouroux, 1821, p. 36, pi. 70, figs. 1-2. (=M%iricea 

spicifera) . 
Plexauropsis flavida Stiasny, 1935a, p. 242; 1935b, p. 71, pi. 1, fig. 5, 

text fig. S; 1942, p. 106, pi. 1, figs. 6-7; 1951, pp. 51, 56. 
Plexauropsis pii,ertorealis Stiasny, 1942, p. 107, pi. 1, figs. 7-8, text fig. 15. 
Plexauropsis sp. Stiasny, 1935b, p. 72. 
? Plexaura olivacea Lamouroux, 1816, p. 431, pi. 16. 

Diagnosis. Colonies tall, up to 40 em., feather-shaped, often 
with secondary liranehes developing into new feathers, placed in 



1 !);')}) l.E.MON-COLOKEI) I'LlLXAUKinS 9 

the same plane as the original one. Pinnulae croAvded, round, 
straight, 3-6 mm. long and about 3 nun. in diameter. Polyps 
numerous on all sides oi" the pinnulae usually completely re- 
tracted so only a fine ])ore is visible, as a pin-prick, rarely form- 
ing low warts; openings usually at right angles to the branchlet. 
Color of colonies varying from bright lemon-colored to buff or 
light to dark purple. 

Spicules in inner layer slender, purple, spindles with distant 
warts, length about 0.27 mm. Middle layer with numerous, 
slender, spindles with similar armature of warts and often slight- 
ly curved or bent, length about 0.35 mm., rarely up to 0.50 mm. 
in length, and extremely rarely becoming stout. External layer 
clubs and spindles, with long spines on the external side and low, 
composite warts on the inner side ; length up to 0.40 mm. Tenta- 
cles with delicate operculum of flattened rods, often curved and 
with marginal warts, length up to 0.20 mm., mostly around 0.13 
mm. Spicules in middle and external layer yellow or purple or 
mixed; those in the operculum always colorless. 

T\jpe. Possibly in the Natural History Museum in Paris. 

Tijpe locality. "Antilles"; Seba's Keratophyton came from 
Nova Hispania (=Mexico). 

Distrihution. Seems to be not uncommon in the West Indian 
region, ranging from the coast of Venezuela, Mexico, Dominica, 
Tobago, Puerto Rico, Florida to the Bahamas. Some old records 
from Bermuda mentioned by Verrill are rejected, as the species 
has not been taken in that well-explored region during the last 
50 years. Verrill 's figures are based on spicules from a specimen 
from Dominica. 

From Mauritius, the Museum of Comparative Zoology pos- 
sesses a fragment, buff in color, with slightly more delicate 
spicules (Plate 5, figs. 25-34). It may possibly represent 
Lamouroux' olivacea from "Indes orientalis" (Plate 2). Wright 
and Studer have with some doubt placed the latter as a synonym 
of their Euplexaura pinnata, 1889 (p. 144, pi. 33, fig. 5), from 
Japan, 8 and 50 fathoms depth. From the figures given of the 
spicules, I doubt that their species is identical with the one from 
Mauritius. There is of course the possibility that the M.C.Z. 
specimen has had a locality label misplaced and merely repre- 
sents the West Indies form. The spicules are figured here and 



10 BREVIORA No. 115 

it is hoped that future workers may be able to procure more 
material — from Mauritius or localities in the vicinity — and 
thereby settle the question. 

Specimens examined. A total of 20 colonies or fragments from 

11 different lots: Gulf of Mexico, Guadeloupe, Cuba, "Antilles" 
Tobago, Puerto Rico, Bahamas, Key West, Florida, Indian River, 
Florida, "Florida" and "W.I.", besides the branch from 
Mauritius. 

Remarks. The species varies in color as well as in spicules, but 
it does not seem possible to separate the material into different 
groups. As far as color is concerned, those from the Caribbean 
region have all been dull brown or buff in color, as are the colo- 
nies of Plexaura flexuosa Lamouroux and Antillogorgia acerosa 
(Pallas), while in other parts of the West Indies both purple and 
yellow or buff colonies are found. 

LIST OF REFERENCES 

AURIVILLIUS, M. 

1931. The gorgonarians from Dr. Sixten Bock's expedition to Japan 
and Bonin Islands 1914. Kgl. Svenska Vet. Akad. Haiidl., ser. 3, 
vol. 9, no. 4, pp. 1-337, pis. 1-6, text figs. 1-65. 

Dana, J. B. 

1848-1849. Zoophytes; United States Exploration Expedition. Phila- 
delphia, pp. 1-740, pis. 1-61. 

Donovan, E. 

1825. (Gurgonia sulplmrea, description and colored plate.) Naturalists 
Repository, London, vol. 4, p. 126 and plate. 

DuCHASSAiNG, P. and G. Michelotti 

1860. Memoire sur les Coraillaires des Antilles. Acad Sei. Torino, 

vol. 19, pp. 1-89, pis. 1-10. 
1864. Supplement, nid., vol. 23, pp. 97-206, pis. 1-11. 

Ellis, J. and D. Solander 

1786. The natural history of many curious and uncommon zoophytes. 
London, pp. 1-208, pis. 1-83. 

ESPER, E. J. C. 

1788-1829. Die Pflanzenthiere. Niirnberg, vols. 1-3, text and plates. 
Gorgonians, 1894, part II, pp. 1-132, pis. 1-39; 1897, part I, 
pp. 152-179, pis. 40-50; 1806, Additional remarks, pp. 25-37, 
pis. 51-55. 



1959 LEMON-COLORED PLEXAURIDS 11 

Gordon, I. 

1925. Gorgonids fioiu Curacao Island. Bijd. Dierk. Amsterdam, vol. 
24, pp. 15-24, pis. 4-5. 

KoLIilKER, A. VON 

1865. leoues histiologicae oder Atlas der vergleichenden Gewebelehre. 
Leipzig, Abt. 2, Heft 1, pp. 85-182, pis. 10-19. 

KUKENTHAL, W. 

1912. Gorgonaria. Wis. Ergebnisse der Valdivia Expedition, vol. 

13, Part 2, Heft 1-2, pp. 1-946, pis. 30-80, and 318 text figs. 
1917. System luid Stammesgescliichte der Plexauridae. Zool. Anz. 

vol. 48, pp. 330-336, 340-347. 
1924. Gorgonaria. Das Tierreich, Berlin, Leipzig, 47 Lief., pp. I-XII 

+ 1-478, text figs. 1-208. 

KUNZE, G. 

1916. Die Gattung Eunicea Lamouroux. Zool. Jahrb., Suppl. 11, 
Heft 4, Part 4, pp. 505-ri51, pis. 24-26, 25 text figs. 

Lamarck, J. de 

1815. Suite des polypiers cortieiferes. Mem. Mus. Hist. Natur. Paris, 
vol. 2, pp. 157-164. 

1816. Histoire naturelle des animaux sans vertebres. Paris, 7 vols. 
1836. Histoire naturelle des animaux sans vertebres. Paris, Edition 2, 

11 vols. 

Lamouroux, J. 

1816. Histoire des polypiers coralligenes flexibles, vulgairement 

nomuies zoophytes. Caen, pp. LLXXXIV, 1-559, 19 pis. 
1821. Exposition methodique des genres de I'ordre des polypiers, etc., 

Paris, pp. I-A^II, 1-115, pis. 1-84. 

Milne Edwards, H. and J. Haime 

1857. Histoire naturelle des corallinaires. Des Alcyonaires. Paris, 
vol. 1, pp. 96-220. 

MOSER, J. 

1921. Ergebnisse einer Eevision der Gattung Plexaura Lamouroux. 
Zool. Anz., vol. 53, pp. 110-118. 

Seba, a. 

1734-65. Locupletissimi rerum naturalium thesauri accurata descriptio, 
etc., Amsterdam, vols. 1-4, 449 plates with text. 



12 BREVIORA No. 115 

Stiasny, G. 

1935a. Diagnosis neuer uud alter Arten der Gorgonarien Familie 

Pk'xauridae. Zool. Anz., vol. 109, pp. 236-245. 
1935b. Die Gorgonacea der Siboga Expeditie. Suppl. 1, Revision der 

Plexauridae. Leiden, Siboga Expedition. Monograph 13b, no. 7, 

pp. 1-106, pis. 1-7, and 27 text figs. 
1935i'. Der Plexauropsis humilis (Milne Edwards) und Eunicea hicksnrii 

nov. spec. Zool. Anz., vol. 112, pp. 107-116, 3 text figs. 
193(>a. Diagnosen einiger neuen Gorgonarien Arten von Cap Blanco 

(Westafrika). Zool. Anz., vol. 113, pp. 201-206. 
1936b. Gorgonaria von Cap Blanco (Westafrika, Mauritanien). Capita 

Zool., vol. 8(2), pp. 1-44, pis. 1-6, text figs., map. 
1939. Gorgonaria von Cap Blanco, Senegal und Rio d'ouro. Rev. Zool. 

Bel. Afrika. Tervueren, vol. 32, pp. 285-328, pis. 10-18, text. 

figs. 
1942. Aleyonaria von Venezuela (Inseln Blanquilla und Los Frailes). 

Arch. Neer. Zool., vol. 6, pp. 101-116, pis. 1-2, text figs. A-D. 
1951. Alcyonides et Gorgonides des collections du Museum National 

d'liistoire Naturellc, Part II. Mem. Mus. Hist. Nat. Paris, 

Zool., vol. 3, part 1. pp. 1-80, pis. 1-22. 

V.\LENCIENNES, M. 

1855. Extrait d'une monographie de la familie des gorgonidees de la 
classe des polypes. C.R. Acad. Sci. Paris, vol. 41, pp. 7-15. (Re- 
print in Ann. Mag. Nat. Hist., ser. 2, vol. 16, pp. 117-183.) 

Verrill, a. E. 

1866-71. Notice of the corals and echinoderms collected by C. F. Hartt 

at the Abrolhos Reefs, Province of Bahia. Trans. Connecticut 

Acad. Arts Sci., vol. 1, pp. 351-376, pi. 4. 
1870. Contribution to Zoology from the Museum of Yale College, no. 7. 

Descriptions of new corals. Ainer. Jour. Sci., vol. 49, Art. 43. 

pp. 370-375, text figs. 1-4. 
1907. The Bermuda Islands, Part 5, section 1: Coral reefs. New 

Haven, pp. 160-304, pis. 28-40, text figs. 71-181. Also in Trans. 

Connecticut Acad. Arts Sci., vol. 12, pp. 204-348, pis. 28-40. 
1912. The gorgonaiians of the Brazilian Coast. Jour. Acad. Nat. Sci., 

Philadelphia, vol. 15 (2), pp. 373-404, pis. 29-35, text figs. 

Wright, E. P. and T. Studer 

1889. Aleyonaria, reports on the scientific results of the VGy;igi> of 
H.M.S. "Challenger." Zoology. Edinburgh, vol. 31, pp. I- 
XLVIIT -f 1-134, pis. 1-43. 






^■'».- 







■*. - ■"■•••, 









;! 



^. 



^ 










.5^ 












fell V 




'•¥»«fi^.: 



Plate 1 



iluriceopsis sulphurea (Donovan), two colonies from Brazilian waters. Ke- 
production of Donovan's original plate. 




Plate 2 



Muiiceopsis olivacea (Lamouroux), a braiuli from "Ocean Indien." Ke- 
production of Lamouroux' original figure. 




Plate 3 



Muriceopsis sulphurea (Donovan), from Brazil; M.C.Z. material. 

1-4. Spindles from the inner layer. 

5-6. Spindles from the middle layer (low magnification). 

7-9. Spicules from the external layer. 

10-12. Eods from the tentacles. 

13. Short warted rod from the middle layer. 
All except 5-6 drawn with high magnification. 
Division of scales 1/10 mm. 




Plate 4 



Muriceopsis flavida (Lamarck), from West Indies; M.C.Z. material. 

14-16. Spindles from the inner layer. 

17-21. Spicules from the external layer. 

22-24. Rods from the tentacles. 
All drawn with high magnification. 
Division of scale 1/10 mm. 




Plate 5 



Muriceopsis flavida (Lamarck), from Mauritius (?) ; M.C.Z. material. 

25-29. Spicules from the inner layer. 

30-34. Spicules from the external layer. 
All spicules drawn with high magnification. 
Division of scale 1/10 mm. 



BREVIORA 



Meseiuiitii of Comparative Zoology 



Cambridge, Mass. January 6, 1960 Number 116 



INSECTIVORES OF THE MIDDLE MIOCENE SPLIT 
ROCK LOCAL FAUNA, WYOMING 

By Katiierine Milmine Reed 

During' the .years 1947, 1948 and 1951, field parties from tlie 
University of Wyoming-, Amherst College, the Royal Ontario 
Museum, and the Chicago Natural History Museum made col- 
lections of a microfauna from the Middle Miocene Split Rock 
locality near Muddy Gap, Fremont County, Wyoming. The 
present paper is the third in a series dealing with this material. 
Black (1958), and Black and Wood (1956) have published on 
sicistine and mylagaulid rodents. Studies are in progress on the 
lagomorphs by Dr. Mary R. Dawson and on the reptiles by Dr. 
Walter Auffenberg. Mr. Craig C. Black is currently working on 
the rodents and carnivores. The collection includes both teeth 
and postcranial elements; I have not recognized any remains 
definitely referable to insectivores among the latter. A mole, 
a hedgehog and a shrew are present in the fauna, the former two 
represented by numerous isolated teeth, the latter by one incom- 
plete ramus. The scarcity of shrews in so large a collection is 
surprising. 

Material for this study was made available by Amherst Col- 
lege, the University of Wyoming, the Royal Ontario Museum 
and the Chicago Natural History Museum, to all of which I am 
g-rateful. I wish to express my thanks to Professor Bryan Patter- 
son for his suggestion that this work be undertaken and to him 
and to Mr. Black and Mr. Clayton E. Ray for help and advice 
during the study. I am indebted to Dr. R. A. Stirton for the 
loan of the type of Met echinus nevadensis, to the American 
Museum of Natural History for specimens of Prosed ops, and to 
the Chicago Natural History Museum for the type of Mctcchinus 



2 BREVIORA No. 116 

)ii(irsla}idc)isis. M3' thanks are also due to the jMaminal Depart- 
ment of the Museum of Comparative Zoology for access to Recent 
talpid and erinaceid material. T am very grateful to Dr. H. B. 
Whittington for his generous assistance with the photography of 
the specimens. 

The following abbreviations are used: 
A. CM. — Andierst College Museum 
C.N.H.M. — Chicago Natural History Museum 
K.O.M. — Royal Ontario Museum 

ILC.M.P. — University of California Museum of Paleontology 
r.W. — T'niversity of Wyoming 

Family TALPIDAE 

Subfamily SCALOPINAE 

Mesoscalops^ new j^eiius 

Type species. Mesoscalops scopelotemos, new species. 

Diagnosis. In general, similar to Proscalops but differing as 
follows : protocone of upper molars more rounded and somewhat 
larger than hypocone, situated nearlj- directly internal to para- 
cone, imparting a square appearance to outline of teeth: P^ with 
lingual face faintly grooved and with wide lingual shelf; lower 
molars Avith small median cingulum just above gum line be- 
tween hypo- and protoconids; talonid of Mj wdder than tri- 
gonid. 

Mesoscalops scopelotemos new species 

Type. A. CM. No. 10461, isolated left M^. 

Ilifpoeliejm. Tvpe and A.C.M. Nos. 10483-10499, 10503-10522. 
10456-10459, 11322-11324, 11304-11414, 11326-11328, 11420- 
11426, 11431-11433; U. W. No. 1074; R.O.M. No. 2077, C.N.H.M. 
Xos. PM 2125-2129, 2136-2159. These numbers include numerous 
isolated specimens of right and left P-^-M^ and Mi-;.,. 

Horizon and locality. Middle Miocene; NWV4, Sec. 36, T29N, 
UilOW, Fremont County, Wyoming, from the vicinity of the 
Brarhycrus quarry (Schultz and Falkenbach, 1940, p. 251), 
seven miles west of Muddy Gap filling station, in a draw abont 
V, mile south of T\S. highway 287. 

1 The generic name is given in reference to the Middle Tertiary age of the 
^enus. 



1960 MIOCENE INSECTIVORES 3 

Dia<j)wsis. As for tlu> genus. For iiieasiireuKnits see Table J. 

Description. P"* consists of a blade-like paracone with a niota- 
stylar area and a lingual cusp in the form of a wide shelf. Some 
specimens show a partial division of the lingual cusp into two 
cuspules, and several also show some irregularities on the inner 
margin. One tooth has a very small accessory cuspule on the 
j)()sterior side of tlie labial cusp. There is usually a faint groove 
running down the lingual face of the tooth, and when the 
lingual cusp is divided, this groove runs between the cuspules 
(PI. 1, fig. 1). Ml is as described in the generic diagnosis above. 
Two unworn specimens show a small metastyle. An accessory 
cuspule is often found at the base of the anterior face of the 
tooth, which, when the tooth is worn, forms an irregularity in 
the outline. The size of this cuspule varies, and it is not present 
in some specimens (PI. 1, figs. 2, 3, 4). IVP exhibits the same 
general size relationships of hypocone and metacone as does 
Ml, and both this tooth and M^ show no pinching, such as is 
seen in Proscalops miocaenus, in the unworn protocone ; with 
wear, however, the pinched shape becomes noticeable. M- is 
more nearly quadrate than M^, the antero-posterior and trans- 
verse dimension being nearly equal. Both metacone and paracone 
are distinctly V-shaped in M-, whereas in M^ only the former 
has this shape. The paracone of M^ is slightly the larger of the 
two. There is a small notch between the cusps on the labial side 
of the tooth. The valley between the arms of the metacone opens 
externally nearer the mid-line of the tooth than in M^, due to 
the presence of an anteriorly -curving projection from the poster- 
ior arm of the metacone; this projection is, however, variable and 
not always present (PI. 1, fig. 5). In M^, the protocone is much 
larger than the hypocone, the latter being rudimentary and 
assuming a variety of shapes. The paracone is larger than the 
metacone, which is simply an oblique blade across the posterior 
border of the tooth. The two cusps are separated by a labial 
notch. The protocone is situated almost directly lingual to the 
paracone. M-^ is less quadrate than M^ or M-, but is, neverthe- 
less, not triangular in contour. Between the arms of the para- 
cone on the labial side there may be irregnlarities in the enamel 
in the form of swellings that are variable in size and number. 
M^ is the smallest of the upper molars (PI. 1, fig. 6). All the 



4 BREVIORA No. IIG 

upper molars have two short, slender, labial roots and one stout 
lingual root. 

In Ml the talonid is wider than the trig'onid. The protoconid 
is the hig'hest cusp and the hypoconid next in prominence. The 
paraconid is widely divergent from the metaconid. The hypo- 
conulid is small and posterior to the entoconid and at the inner 
end of the low posterior cingulum as is usual in clilambdodont 
teeth. The crest between the hypo- and entoconid shows some 
variation; it may not be continuous and may bear a swelling 
in the enamel midway between the cusps. There is also variable 
development of a small anterior cingulum, which appears in the 
majority of specimens ; at most this is a very narrow median 
shelf that slopes downward labially. All lower molars have a 
small basal cingulum between the hypoconid and the protoconid 
(PI. 2, fig. 1). The trigonid and the talonid are nearly equal in 
width. The protoconid is generally but not invariably hig-her 
than the hypoconid. Metaconid and paraconid are much less 
divergent than in ]\Ii. There is an accessory cuspule, formed by 
the anterior cingulum and the base of the paraconid. The meta- 
conid in unworn specimens shows a rudimentary metastylid 
which is not continuous with the crest from metaconid to hypo- 
conid. M2 is similar to Mi in leng-th (PI. 2, fig. 2). In Mg, the 
trigonid is wider than the talonid ; the basin of the latter opens 
anteriorly. The protoconid is the highest cusp. There is an 
anterior cingulum and accessory cuspule as in Mo. M3 is the 
smallest of the lower molars (PI. 2, fig. 3). 

Discussion. Mesoscalops appears to be related to and is prob- 
ably derived from Proscalops. A study of the Proscalops group 
of talpids, based partly on new material accumulated since 
Matthew's work (1901, 1909), will be presented in a forthcoming 
paper. 

Family ERINACEIDAE 
Subfamily ECHINOSORICINAE 

Metechinus Matthew 1929 

Metechinus marslandensis Meade 1941 

In addition to isolated teeth that are similar in every respect 
to those preserved in the type, there are numerous, certainly eon- 
specific, teeth that considerably enlarge our knowdedge of the 
dentition of this species. 



1960 MIOCENE INSE( TIVORES 5 

Txjpe. C.N.II.M. No. P 263i)9, incomplete left luaiulible wiili 
Ml andM^. 

Ilypo'digm. Type and A. CM. Nos. 1()45!)-U)4(i0, 10462- 
10482, 11325, 11315-11321, 11427-11430; R.O.M. Nos. 207S, 2071); 
U.W. Nos. 1072. 1073, 1075; CN.II.M. Nos. PM 2130-2135, 2108- 
2124. These numbers inehide numerous isolated specimens of 
P^-M-andMi-o. 

Horizon and localiti). As for Mesoscalops scopelotemos, above. 

E molded diagnosis. P^ and molars as in M. 7ievadensis, dif- 
fering as follows : slight re-entrant between liyoeone and meta- 
cone in M^ ; hypocone rather well separated from U-shaped crest 
joining metacone and paracone ; labial cusps of P'* subequal in 
size. Lower molars as determined hy Meade (1941, p. 43) : 
•'Talonid and trigonid on Mj-o of equal width . . . Hypoeonid 
of Ml less elongated antero-posteriorly and higher than in M. 
ncvadcnsis. Mo proportionately smaller and with trigonid 
slightly longer antero-posteriorly than in M. nei'adrnsis.'" 

For measurements, see Table 11. 

Description. The specimens of Mi agree closely with the type. 
Meade has stated that the hypoconulid of Mi is an "inconspicu- 
ous eminence"; in at least four of the less worn specimens there 
is a distinct protuberance in the position of the hypoconulid. 
It is best developed on the larger specimens but is easily seen 
on others as well. I was unable to find a specimen in which the 
antero-posterior dimension was as large as that given by Meade, 
namely, 4.6 mm. Indeed, I was unable to duplicate his measure- 
ments on the type. From the anterior edge of the paraconid to 
a point between the hypoeonid and the entoconid, I get a meas- 
urement of 3.8 mm. Measuring from the paraconid to the 
postero-external edge of the entoconid, the dimension is still only 
3.9 mm. Among 25 specimens, the greatest length obtained was 
4.0 nnn. There is nothing to add to Meade's description of M^. 
In this material, as in the type specimen and in M. ncvadcnsis, 
there is no trace of M|. 

In the upper dentition, P^ is three-rooted, with one lingual 
and two labial roots. The posterior labial root is the stoutest. The 
paracone and metacone are laterally compressed, equal in height 
and form a very broad V. The lingual part of the tooth is in the 
form of a shelf that mav extend slightlv antero-internal to the 



B BREVIORA No. 116 

paracoiie. This shelf bears two euspules, one situated on the 
anterior edge, the other and larger at about the mid-line of the 
tooth. This euspule development shows variation and does not 
seem to be of taxonomic importance. The shelf widens pos- 
teriorly and there are slight re-entrants where it joins the para- 
eone and metacone (PL 2, fig. 4). In AP tlie metaeone is more 
internally situated than the paraeone, with a erest running from 
it postero-externally. There is a small parastyle. The protoeone 
is situated on the lingual side of a U-shaped crest running from 
a point below the center of the metacone to the antero-internal 
side of the paraeone. There is a small protoconule that varies 
in the degree of development. The hypocone is more externally 
situated than the protoeone and is connected to the metastyle by 
a short, antero-externally directed spur (PI. 2, fig. 5). M^ is tri- 
angular in shape and consists of three main cusjis, probably 
the para-, proto- and hypocones. There is a small parastyle 
situated directly external to the paraeone and a very small 
eingulum running along the posterior edge of the tooth for about 
half its length. 

There are numerous antemolar teeth, largely premolars, but I 
am uncertain as to their position and accordingly omit deserij)- 
tion. 

Discussion. This new material, although frohi a higher hori- 
zon, supports Meade's reference of the species to the genus 
Met echinus. The upper teeth show only minor differences as 
compared with M. nevadensis; the major difference is the size, 
M. nevadensis being half again as large as the earlier form. 
Other dilferences are the slightly wider, relatively speaking, 
lingual cusp of P"*, the smaller metastyle and the degree of 
separation from the U-sliaped crest in M^ mentioned above. M- 
differs in the shift to the posterior of the exterior (stylar) cusi) 
in M. niarslandcnsis. 

Family SORICIDAE 

LmxoECUS Stirton 1930 

LiMNOECUS ? sp. 

Referred specimen. C.N.H.M. PM 2167, incomplete left ranuis 
with Ml and partial Mo, both badly worn. 



1 !>()() MIOCENE INRECTIVORES 7 

llorilon and I'xdlih/. As lor M ( soscdlops scopclod iikis, aboNC. 

Description. Tlic trigonid of IMj is narrower than the tak)ni(l. 
There is a very small anterior ein<>nhim whieli widens slijilitly to 
the exterior. A uiiiuite posterior ein<i'nluni is also present. There 
is a small swellin<i' in the talonid at the ])osition of the entoconid.- 
The renuiant of ]\I.> shows a lari>er anterior einpidnm and su^- 
ji,ests that the trif^-onid of the tooth is wider than the triyonitl of 
Ml. The trigonid and talonid valleys open antero-internally. A 
foramen is present below the anterior root of Mj. Examination 
of the specimen in ultra-violet light showed no traces of pig- 
mentation whieh is in agreem(^nt with the findings of ^Tac- 
Donald (1947). 

Measurements : in millimeters a-p trig. tal. 

Linniorcus f sp. 
L. niohrarensis 



1.3 


0.8 


0.9 




1.1 


0.50 


0.78 U.C.M.P. 


36172 


1.1 


0.75 


0.82U.C.M.P. 


36171 


1.0 


0.62 


0.69 U.C.M.P. 


31047 



L. trie us pis 

The last three measurements are from figures given by Mac- 
Donald (1947) and Stirton (1930). 

Discussion. Tlie specimen is tentatively referred to this genus 
because of the agreement in size and what can be seen of the 
structure, in particular the cingula. It is closer to L. niohrarensis 
than to L. tricuspis. It is unlikely on a size basis that the 
specimen should be referred to Mystipterus. Because of its poor 
condition, the specimen has not been figured. 

Table I 

Measurements of cheek teeth of 31csoscalops scopclotrnios, 

in millimeters 



Tofi 


tl 


1 


N. 


Range 


Mean 


S.D. 


V. 


P^ loft. 




a-p 


7 


2.65-2.1 


2.23 


.221 








trans. 




1.9-1.55 


1.8 


.119 




P^ light, 




a-p 


17 


2.7-2.05 


2.24i!i.03 


.199±.03 


8.8±1.5 






trans. 




2.1-1.75 


1.92±.02 


.079±.01 


4.1ib.7 



-Following tile tcriiiiiiology of Patterson and McGrew (1937). 



8 BREVIORA No. 1]() 

Table I (Continued) 



Tooth 


N. 


Range 


Mean 


S.D. 


V. 


.\[l left, 


a-p 


21 


3.35-2.45 


2.80±.04 


.191±.02 


6.8±1.0 




trans. 




2.9-2.25 


2.42+.03 


.144±.02 


.").9±.9 


Ml right, 


a-p 


6 


3.2-2.7 


2.96 


.197 






trans. 




2.8-2.25 


2.49 


.171 




M- left, 


a-p 


8 


2.62-2.2 


2.41 


.111 






trans. 




2.45-2.25 


2.33 


.062 




M- right, 


a-p 


10 


2.65-2.2 


2.47±.03 


.116±.02 


4.7±.l 




trans. 




2.6-2.0 


2.49±.05 


.180±.03 


7.2±1.8 


M3 left, 


a-p 


9 


2.5-2.15 


2.30 


.125 






trans. 




2.0-1.6 


1.85 


.148 




M3 right. 


a-p 


8 


2.6-2.0 


2.21 


.186 






trans. 




1.95-1.8 


1.81 


.083 




]\ri ieft, 


a-p 


27 


3.1-2.55 


2.81 ±.02 


.115±.01 


4.0±.5 




trig. 




2.0-1.5 


1.73it.02 


.136±.01 


7.3±.9 




tal. 




2.2-1.6 


2.03±.03 


.130±0.1 


6.4±.S 


Ml right, 


a-p 


27 


2.95-2.5 


2.71±.02 


.133±.01 


4.9±.6 




trig. 




2.05-1.55 


1.70±.01 


.095±.01 


5.5±.7 




tal. 




2.25-1.65 


2.07±.02 


.150±.02 


7.2±.9 


M- left. 


a-p 


12 


3.05-2.5 


2.75±.04 


.140±.02 


5.0±1.0 




trig. 




2.1-1.55 


2.01 ±.0.1 


.186±.03 


9.2±1.8 




tal. 




2.2-1.7 


1.93±.04 


.145±.02 


7.5±1.5 


M:. right, 


a-p 


13 


2.9-2.45 


2.65±.03 


.126±.02 


4.7±.9 




trig. 




2.2-1.7 


1.95±.03 


.122±.02 


6.2±1.2 




tal. 




2.2-1.8 


1.95±.04 


.152±.02 


7.7±1.5 


M;: left, 


a-p 


9 


2.4-2.0 


2.15 


.122 






trig. 




1.6-1.3 


1.45 


.091 






tal. 




1.4-1.0 


1.17 


.155 




M;{ right. 


a-p 


S) 


2.25-1.9 


2.07 


.133 






trig. 

tal. 




1.5-1.35 

1.25-.85 


1.42 
1.01 


.056 
.140 





a-p =^ antero-posterior dimension 
trans. = transverse dimension 
trig. = transverse dimension of trigonid 
tal. = transverse dimension of talonid 



1960 MIOCENE INSECTIVORES 

Table II 

Measiiroments of cheek teeth of Mctccliiiuts )iiarsl(iii(l( nsis. 

in niillinieteris 



Tooth 


N. 


Kange 


Mean 


S.D. 


V. 


P^ left. 


a-p 


9 


3.2-2.8 


3.02 


.133 






trans. 




2.5-2.05 


2.30 


.089 




P^ right. 


a-p 


8 


3.5-2.6 


3.04 


.21 






trans. 




2.5-2.15 


2.31 


.098 




Ml left. 


a-p 


1-2 


3.05-2.8 


2.9±.05 


.179±.03 


6.1±.12 




trans. 




3.55-3.1 


3.31±.04 


.143 ±.03 


4.3±.8 


Ml light, 


a-p 


10 


3.05-2.7 


2.92±.l 


.319±.07 


10.2±2.2 




trans. 




3.7-2.5 


3.29±.08 


.264±.05 


8.0±1.7 


M'- left, 


a-p 


!) 


2.05-1.55 


1.86 


.160 






trans. 




2.4-2.15 


2.19 


.125 






pa-hyc 




2.4-2.1 


2.29 


.131 




M- right, 


a-p 


7 


2.5-1.6 


1.86 


.171 






trans. 




2.5-2.15 


2.29 


.156 






pa-hyc 




2.55-2.05 


2.29 


.208 




:*Ii left. 


a-p 


19 


4.0-3.2 


3.70±.06 


.27±.04 


7.3±1.2 




trig. 




1.95-1.6 


1.80±.00 


.016±.00 


8.8±1.4 




tal. 




2.0-1.6 


1.85±.04 


.13±.02 


7.0±1.1 


Ml right, 


a-p 


4 


3.85-3.4 


3.64 


.217 






trig. 




1.9-1.8 


1.85 


.057 






tal. 




2.0-1.8 


1.88 


.24 




M2 left, 


a-p 


11 


2.7-2.25 


2.53it.04 


.134i!i.02 


5.2±1.1 




trig. 




1.55-1.3 


1.46±.02 


.076±:.U1 


5.2±1.1 




tal. 




1.5-1.05 


1.28±.04 


.164±.03 


i-2.8±-2.; 


M2 right. 


a-p 


11 


2.7-2.1 


2.45±.06 


.177±.03 


7.2±1.8 




trig. 




1.6-1.1 


1.42±.03 


.26±.02 


8.8±1.9 




tal. 




1.4-0.9 


1.19±.04 


.141±.03 


11.8±2.5 



Measurements of elieek teeth of M. nevadensis (Matthew, 1929), 

in millimeters 



Tooth 


Dimension 


P^ a-p 


4.5 


trans. 


4.2 


Ml, a-p 


3.9 


trans. 


5.05 


Mr, a-p 


2.7 



10 > BREVIORA No. IK) 

Table II (Continued) 



Tooth 


Dimension 


trans. 


3.7 


Ml, a-p 


5.3 


trans. 


2.7 


-Ml.., a-p 


3.6 


trans. 


O i) 



a-p =: antero-posterior dimension 
trans. = transverse dimension 
trig. = transverse dimension of trigonid 
tal. = transverse dimension of talonid 
pa-hyc = paracone-hypocoue dimension 

EEFEEENCES 

Black, C. C. 

1958. A new sicistine rodent from the Miocene of Wyoming. Breviora, 
Mus. Comp. Zool., no. 86, pp. 1-7. 

Black, C. C. and A. E. Wood 

1956. Variation and tooth replacement in a Miocene mylagaulid rodent. 
Jour. Paleont., vol. 30, no. 3, pp. 672-684. 

MacDonald, J. R. 

1947. A new shrew from the Niobrara River Upper Miocene of 
Nebraska. Amer. Jour. Sci., vol. 245, no. 2, pp. 123-126. 

Matthew, W. D. 

1901. Fossil mammals of the Tertiary of northeastern Colorado. Mem. 

Amer. Mus. Nat. Hist., vol. 1, part 7, pp. 355-447. 
1909. Carnivora and Insectivora of the Bridger Basin Middle Euceue, 

Mem. Amer. Mus. Nat. Hist., vol. 9, part 6, pp. 291-576. 
1929. A new and remarkable hedgehog from the later Tertiary ot' 

Nevada. Univ. Calif. Publ., Bull. Dept. (ieol. Sei., vol. 18, no. 

4, pp. 93-102. 

Meade, G. E. 

1941. A new erinaceid from the Lower Miocene. Ceol. scr., Field Miis. 
Nat. Hist., vol. 8, no. 7, pp. 43-47. 

Patterson, B. and P. O. McGrew 

1937. A new sorieid and two erinaceids from the White River Uligo- 
cene. Geol. ser. Field Mus. Nat. Hist., vol. 6, no. 18, pp. 245-272. 



1960 MIOCENE INSECTIVORES 11 

SCHULTZ, C. B. AND C. II. FaLKENBACH 

1940. Merycochoerinae, a new subfamily of oiouduiits. Hull. Anicr. 
Mus. Nat. Hist., vol. 77. pp. 213-30(5. 

Stirton, E. a. 

1930. A new tjenus of Soricidae from the Barstow .Miocene of Cali- 
fornia. Univ. Calif. Tubl., Bull. ])ei)t. Geol. Sci., vol. lit, no. S. 
pp. 217-228. 



Plate 1 

Stereoscopic photographs 

Fig. 1. Mesoscalnps scopelotemos, P^ left, A. CM. No. 10456, crown view. 

Fig. 2. Mesoscalops scopelotemos, M^ left, A. CM. Xo. 10461, crown view. 

Fig. 3. Mesoscalops scopelotemos, M^ left, A.C.M. No. 11323, crown view. 

Fig. 4. Mesoscalops scopelotemos, M^ right, CN.H.M. No. PM 2127, 
crown view. 

Fig. 5. Mesoscalops scopelotemos, M^ left, A.C.M. No. 10458, crown view. 

Fig. 6. Mesoscalops scopelotemos, M^ left, CN.H.M. No. PM 2126, crown 
view. 

In all figures, the exterior is to the top of the figuje. All xlO. 



Vx 



Plate 2 
Stereoscopic photographs 

Fig. 1. MrsoNcalops sco2}clotcmo.s, Mi loft, A.C.M. No. 11431, lingual view. 

Fig. 12. Mcsoscalnp.s scoiJclotemos, Mo right, A.C.M. No. ll.'?22, lingual 
view. 

Fig. 3. Mesoscalops scopelotcmos, My left, C.N.H.M. No. PM 2128, 
lingual view. 

Fig. 4. Metechinus marslandt'nsis, P* left, C.N.H.M. No. PM 2130, crown 
view. 

Fig. 5. Metecliimis marslandensis, M^ left, C.N.H.M. No. PM 2133, crown 
view. 

Fig. 6. Meteehinu.s marslandensis, M- right, U.W. No. 1074, crown view. 

In Figures 4, 5, (5, the exterior is to the top of the figure. All xlO. 




■M 



u 



BREVIORA 

MmseiLim of Comparative Zoology 



Cambridge, Mass. January 20, 1960 Number 117 



NOTES ON HISPANIOLAN HERPETOLOGY 

1. ANOLIS CHRISTOPHEI, NEW SPECIES, FROM THE 

CITADEL OF KING CHRISTOPHE, HAITI 

By Ernest E. Williams 

Re-examination of the three specimens on the basis of which 
Cochran (1928 and 1941) postulated the existence of Anolis 
coelestinus in the north of Haiti near Cap Haitien has revealed 
that one of them (MCZ 25485) represents an undescribed species. 
(The other two are also not A. coelestinus and will be discussed 
at another time.) 

The specimen in question is poorly preserved and was, when 
first examined, soft-bodied and much contorted. It has now been 
straightened and hardened in 90 per cent alcohol. 

A majorit}^ of the scale characters usually employed in the 
identification of anoles do indeed appear to ally MCZ 25485, a 
juvenile $ , with A. coelestinus. 

It has smooth ventrals, somewhat squarish, juxtaposed, ar- 
ranged in transverse rows, larger than the dorsal and flank 
scales which are relatively coarse and nearly homogeneous. The 
scales of two middorsal rows are very slightly larger than the 
adjacent scales. There are multiple keels on the scales of limbs 
and digits. The number of loreal rows and of rows between the 
supraorbital semicircles, as well as the number of rows between 
interparietal (pineal) scale and the supraorbital semicircles fall 
within the known range of A. coelestinus. As in the latter, the 
scales on the snout are small and numerous. 

However, there are several features of the specimen that are 
difficult to reconcile with its identification as coelestinus. 

1. Its locality: The Citadel of King Christophe near Cap 
Haitien is very remote from any other mainland record of 



2 BREVIORA No. 117 

coelestinus, a species characteristic of the Cul-de-sac plain and 
Tiburon and Barahona peninsulas. 

2. Its color: MCZ 25485 has an elaborate pattern well de- 
scribed by Miss Cochran (1941, p. 179). A number of specimens 
of undoubted coelestinus of similar or smaller size are available 
which show no trace of such a complex pattern. 

3. The digital clilations: In the specimen these are conspicu- 
ously narrower than in typical specimens of A. coelestinus. Tbo 
lamellae under the fourth toe are also fewer: about 19 under the 
second and third phalanges, (as compared with 27 in coelestinus). 
about 32 under the whole toe (as compared with 47 in coelesti- 
nus). 

The last feature is crucial. The locality might be valid, and 
only appear anomalous because of insufficient collecting in the 
intervening area. The color also might be explained by the 
chance preservation of one phase in the repertoire of color 
changes possible in the species. However, the narrowness of the 
digital dilations suggested that it might not even be a member 
of the arboreal chlorocyanus group, that it might instead l)e a 
ground anole or spend a large part of its time on the ground. 

The specimen was therefore subjected to very careful exami- 
nation and direct comparison with chlorocyanus and coelestinus 
and other Hispaniolan forms. It proved to differ from all of them 
in a combination of significant features and seems not to be iden- 
tical with any known species of anole. A specimen from the 
unidentified Hispaniolan collection of the American Museum of 
Natural History from the same locality proves to belong to the 
same species, which because of its occurrence in the vicinity of 
the famous Citadel built at the command of Henri Christophe is 
named : 

Anolis christophei new species 

Type. MCZ 25485, 9 

Type locality. "At or near the Citadel of King Christophe, 
Cap Haitien, Haiti." 

Collector. W. S. Eyerdam, 4. x. 1927. 

Paratype. AMNH 49736, same locality, collected by \V. G. 
Hassler 16. vi. 1935, 9 

Diagnosis. An Anolis resembling Anolis coelestinus in many 
of its scale characters, but differing in the following: narrower 



1960 



ANOLIS fllRISTOl'lIEI 



digital dilations and fewer lamellae under fourth toe ; the longer 
legs; the presence of a distinct frontal depression; a pattern of 
canthal scales with the anterior four or five abruptly smaller 
than the posterior two or three; a long rather than a wide 
mental ; larger sublabials ; in the continuation of strongly en- 
larged scales of the suboeular series behind the eye and in a 
complex color pattern of bands and lines. 

Description. Head: Head scales mostly smooth, a few feebly 
or bluntly unicarinate ; about 10 scales across snout at level of 
second and third canthals. A moderate frontal depression, the 
scales in the depression relatively small and numerous. 





Fig. 1. Mental and neighboring scales. Left, Anolis christophei n. sp. ; 
light, Anolis coelestinus. 

Supraorbital semicircles separated from one another by one 
scale and from the supraocular disks by 1-2 rows of granules. 
Supraocular disks consisting of about 9 to 11 keeled scales which 
are separated from the elongate supraciliaries by at least three 
rows of granules. Canthus low, canthal scales seven, the anterior 
four or five abruptly smaller than the posterior two or three. 
Loreal rows 6-7. Temporal scales granular, as small as the scales 
of the lower flanks, not bounded dorsally by any well-marked 
supratemporal line of larger scales. Supratemporal scales granu- 
lar, grading into larger scales surrounding the interparietal. 
Interparietal scale smaller than ear opening, separated from 
supraorbital semicircles by 5-6 slightly enlarged scales, in con- 
tact posteriorly with granules like those of the dorsum. 

Posterior frontal small, about one-third the size of the anterior 
supraorbital. Two scales as large or larger than the posterior 
frontal between the latter and the canthals. 



4 BREVIORA No. 117 

Suboculars separated from supralabials by a single row of 
scales. One scale intervening between subociilar series and 
canthal row. Six to seven supralabials to the center of the eye. 

Mentals longer than wide, 2-4 scales inserted between the tips 
posteriorly. One sublabial on each side in contact with infra- 
labials. Central throat scales granular, rounded. 

Trunk: Middorsal scales granular, not keeled, hardly larger 
than the flank scales into which they grade very gradually. 
Ventrals in transverse rows, larger than dorsals, squarish, 
smooth, juxtaposed. 

Limbs and digits: Hand and foot scales multicarinate, about 
19 lamellae under phalanges 2 and 3 of fourth toe, about 32 
under whole toe. Scales of anterior upper arm smaller than 
ventrals, unicarinate. Lower arm scales about as large as ven- 
trals, tricarinate. Anterior thigh scales and lower leg scales 
about as large as ventrals, unicarinate. 

Tail: Tail subcircular in section; verticils distinct, surmounted 
by 6 keeled scales, very slightly larger than the lateral caudal 
scales ; ventrally four keeled still larger scales per verticil. 

Size: Type 9 44 mm. Paratype 9 39 mm. 

Color: The paratype was preserved too long in formaldehyde 
and is a uniform dingy brown. The pattern of the type was well 
described by Doris Cochran (1941), whose observations may be 
quoted in full : ' ' Above mottled with dark gray and sepia ; a 
light dorsal line with diamond-shaped dark brown patches 
approaching it from the side at intervals and set off by light 
posterior margins ; a dark scalloped band on the nape of the 
neck with a verj- definite dark posterior margin which is further 
accented by a pale tan area directly following it; a butterfly- 
shaped dark spot across the occipital region ; two dark bands 
across the supraorbital region ; top and sides of .snout marbled 
with dark and light ; a wide light gray stripe leaving posterior 
border of ear and continuing about halfway to shoulder, at which 
point it abruptly ends ; a light stripe issuing from beneath this 
ending continues above the shoulders and fades out gradually 
behind the axilla; limbs marbled with tan and brown; tail dark 
gray above, rather uniform; under surfaces of arms, legs and 
tail pale yellowish white, throat with pale bro\\ni reticulations 
over it ; belly liglit blue green, highly iridescent ; heavy brown 



11)60 ANOLIS CllRISTOPIlEl 5 

marbliiijrs all over lower labials and sides of chin; center of 
throat with two {)ale brown stripes marking oft* each side of where 
the gular fan will develop. ' ' 

Relationships: I have provided tables of the condition in llis- 
paniolan species of the most conniionly used scale characters of 
anoUnes in order to provide a conspectus of the genus and its 
close relatives in this island. I have added in the final column 
of the table one or two special or unique diagnostic characters 
for each species or species group. It should be evident from this 
table that the resemblances of A. cliristophei to coelestinus are 
not greater than its resemblances in some respects to other forms. 
The complex pattern of the type is somewhat like that of juvenile 
cyhotcs but there is again no sufficient warrant for inferring 
special relationship, since christophei differs sharply from 
cyhotcs in its squarish rather than cycloid ventrals, slenderer 
head, and in having the middorsals not enlarged. The elongate 
mentals of christophei are quite distinctive as compared with 
coelestinus but similar to those of distichus, which also has the 
ventrals squarish and arranged in transverse rows, but the new 
species clearly differs from distichus in the longer head and in 
the absence of the two parallel rows of scales on the front, in 
the number of scales between parietal and semicircles, etc. 

From the other Hispanolian species with ventrals arranged in 
transverse rows the new species differs in much the way it 
differs from coelestinus, or in other features as well. Further 
speculation on its relationships seems unwarranted with only 
two female specimens at hand and no knowledge of the species 
in life. 

Acknowledgments: I am grateful to Mr. Charles M. Bogert 
for the privilege of studying the unidentified Hispaniolan collec- 
tions in the American Museum of Natural History. Miss Margaret 
Estey made the sketches for Figure 1. Mr. A. Stanley Rand 
made a number of measurements and counts that have only 
partly been used here. 



BREVIORA 



No. 117 



REFERENCES 

Cochran, Doris M. 

1928. The herpctolosical collections made in Haiti and its ad.joininu 

islands by Walter J. Eyerdam. Proc. Biol. Soc. Washington, 41: 

53-60. 
19-11. The Herpetology of Hispaniola. Bull. U. S. Nat. Mus., no. 177: 

1-389. 













labial ji 




S V length 


lamellae 






to center 




adult $ 


4th toe 


canthals 


loreals 


of eye 


Anolis 












ricoidii 


137 


33 


3+ 


4-8 


9 


seniilineatus 


38 


20 


4 


4-5 


5-6 


olssoni 


40 


20 


4 


3-5 


6 


distichus 


51 


20 


4 


4 


5-6 


eybotes 


67 


20 


3 


4-10 


7 


armouri 


46 


18 


3-4 


4-6 


6-7 


shrevi 


58 


17 


3 


6 


6-7 


nionticola 


40 


17 


5 


7-9 


7 


dailingtoni 


42 


14 





6 


6 


chloracyanus 


71 


29 


4 


3 


7-8 


coelestinus 


64 


28 


6 


5 


8-9 


cluistophei 


(44)-^ 


19 


7=4(+3) 


6-7 


6-7 


hcndeisoni 


46 


21 


6 


6-7 


6 


baliarucoensis 


46 


19 


7 


7 


6-7 


Xiphoeercus 












dailingtoni 


7;j 


23 


4-5 


2-3 


7 


("haniaelinorops 












wetniort'l 


116 


ea 11 


o 


4 


9 



No adult S availaljlc. 



Table I: Comparison of certain characters in Ilispaniolan anolines. IJata 
mostly from Cochran (1941). Where a single number is given, this is to be 
considered the mode around which variation is to be expected. Extreme 
variants are usually not listed. Lamellae under fourth toe are counted under 
phalanges II and III only. 



196U 



ANOLIS CIIRISTOPHEI 



DO 






^ 






f^ 












a 






tC 


cr: 




a 


a> 




o 


'S 




in 


o 




,2 


qj tW 




"3 


O-TS 






o _a; 


y — V 






'j: 


« 


C "oj 






o S 


"3 


5d 


f^ M 


■+^ 


+ 




3 


W 


i fcfi 


j-» 






II 








•-^ 


- a 


!3 


^ 


~^ <o 


SJ 



c - 
S 



c3 1-1 to  



~ > S :^ 
°^ i * 'w 



m 



o 






K o 






2 o 



o 



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ca 



o 



:i = S 



>-, 






o 3 






•* 



0) 




o 




o 


93 


s 


tc 


03 


o 






0) 




Hi 


■*-> 






4- 


o 






SJ 


'x 


I) 




0^ 


^ 



bXJT 



cr 

CO 



'^ 



m 

o 



tn 

--^ O 

bb_fct 

"S i 
1 + 



cr 



in 



o 






bo 



t> 


^ 


> 


> 


<o 


(D 


Ti 


T^ 



cr 









M 

a 






a 3*' r- « 






OH S 

^ -2 

&T3 



M 



03 
<U 

I— t 

O tJ 

2 -^ 



5 != 









m 
o 

O 



r^ 


c 


a> 


0) 


« 


-l-a 






O 


'o 




c3 


« 







"" 


o 


SZ 1 1 




^ II 


03 


o ^ 


J5 


o >. 


o 


fi "^ 


X 


c 




K 






rt 


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a; 



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CS 

& 

CO 



c3 






x/ 



BREVIORA 

Muiseinini of Coimparative Zoology 



Cambridge, Mass. February 24, 1960 Number 118 

A SURVEY OP TIIP] LEPTODACTYLID FROGS, 
GENUS EVP80PHV8, IN CHILE 

By .Iose M. Cei* 

INTRODUCTION 

The interesting leptodactylid genus Eupsophus Fitzinger, 1848, 
consists of three allopatric Neotropical species groups Avidely 
disjunct in distribution. One, the peruanus-wetisteini group, is 
found in the plateau of central Peru. The second is composed of 
several species from southeastern Brazil (TJioropa of some au- 
thors). A third stock, the subject of the present report, now oc- 
curs only in Chile, although fossil evidence (Schaeffer, 1949) 
suggests that during late and middle Tertiary species of the group 
ranged much farther to the east. The Chilean forms have been 
variously referred to the genera Borhorocoetes, Cacotus or Cy- 
stignathus previous to their current allocation to Enpsnpkns 
(Parker, 1932). 

The following nominal forms of Eupsophus have been recog- 
nized from Chile: hihroni (Bell) ; calcaratus (Glinther) ; coppin- 
geri (Glinther) ; grayi (Bell) ; hidalgi (Espada) ; kriegi (Midler) : 
niaculatus (Glinther) ; masareyi (Roux) ; nodosus (Dumeril and 
Bibron) ; roscns (Dumeril and Bibron) ; taeniatus (Girard) ; 
verrucosus (Pliilippi). In addition, a number of names were pro- 
posed by Philippi (1902) but his descriptions are inadequate for 
identification and, since his types are no longer in existence, I 
follow Vellard (1947) and Schmidt (1954) in disregarding these 
names. The majority of the species in the genus are known only 
from their original descriptions, life history information is scanty 
or wanting, all are of doubtful status, and their affinities are 

*Insfituto de Biologia. Universidad Nacional de Cuyo, Mendoza, Argentina. 



2 BREVIORA No. 118 

unknown (Vellarcl, 1947; Schmidt, 1954). A detailed revision of 
Eupsophus is thus badly needed. 

MATERIAL AND ACKNOWLEDGMENTS 

During my extensive stay in Chile dnrinp; 1956-1957 I concen- 
trated on a preliminary review of the systematics of the Chilean 
amphibian fauna and particularly emphasized field observation 
and life history data in my work. On the basis of experience with 
living Eupsophus, together with a study of preserved materials 
from world collections, it is now possible to critically survey the 
Chilean portion of the genus. Accordingly, the Chilean forms 
may be placed into three species groups and only five forms may 
be accepted as valid. These conclusions are based upon a com- 
parison of all type specimens with material collected at the type 
localities and throughout the range of the genus in Chile. Meas- 
urements are based upon a total of 115 sexually mature examples : 

3 E. coppingeri (west Patagonian coast) ; 53 E. grayi (Valdivia, 
Cliiloe, Concepcion) ; 11 E. nodosus (Valdivia, Chiloe, Zapallar) ; 
10 E. roscKs (Valdivia, Chiloe) ; 38 E. faniiafus (Valdivia. 
Chiloe, Mall^eo, Puerto Blest, east slope of the Cordilleras). Ad- 
ditional material, especiallj^ of taeniatus, nodosus, and grayi, was 
examined but not measured. All measurements were taken with 
calipers with an accuracy of 0.5 mm. Characteristics of the pec- 
toral girdle, prevomerine teeth and external morphology Avere also 
examined. 

Eupsophus is poorly represented in herpetological collections 
but through the courtesy of Mr. C. M. Bogert, Dr. R. F. Inger, 
and Dr. E. E. AVilliams, I have been able to examine and studj" 
the samples of this genus in the American Museum of Natural 
History, the Chicago Natural History Museum, and the Museum 
of Comparative Zoology at Harvard LTniversity, respectively. 
Specimens in the Instituto de Biologia, Universidad Nacional de 
Cuyo, Mendoza, Argentina, and Instituto Miguel, Lillo, Univers- 
idad Nacional de Tucuman, Argentina, as well as in Centro In- 
vestigaeiones Zoologicas Universidad de Chile were also studied. 
Comparative descriptions and photographs of all existing pri- 
mary types w^ere provided through the efforts of Miss A. C. C. 
Grandison (British Museum, London), Dr. J. Guibe (Museum 
National d'Histoire Naturelle, Paris), and Dr. E. Forcart 
(Naturhistorische Museum, Basel). I wish to acknowledge this 
valuable assistance. 



1960 riTILEAN EUPSOPIIUS 3 

Other important information relative to the current pr()l)leni 
was obtained from the follo^Ying museums and institutions: 
Academy of Natural Sciences, Philadelphia ; Carnef^ie Museum, 
Pittsburpjh ; Musoo Bulloek, Ansi'ol. Cliile; Museo de Concepeion, 
Chile; Museo Naeional B. Rivadavia, Buenos Aires, Museo Na- 
tional Plistoria Natural, Santiago de Cliile; Museu Nacional, Rio 
de Janeiro; Museum d'llistoire Naturelle, Geneve; Museum of 
Zoology, University of Michigan : Naturhistoriska Museum, 
Stockholm ; Peabody Museum, Yale University ; Senckenberg 
Museum, Frankfort a.M. ; United States National Museum ; 
Zoologisehe Museum, Berlin; Zoologische Museum, Hamburg; 
Zoologische Sammlung des Bayerisehen Staates, Munich. Finally, 
Dr. Jay Savage, Department of Biology, University of Southern 
California has read over the completed manuscript and offered 
numerous helpful suggestions. 

SYSTEMATIC NOTES 

Species groups. Morphological and biological data support the 
concept of three phyletic lines in Chilean Eupsophus: (1) cop- 
pingeri-nodosns, (2) grayi-roseus, (3) faeniatus. 

Figure 1 illustrates the anatomical features of the pectoral 
girdle and prevomerine teeth in the five valid species. Similar- 
ities in the structure of omosternum, xiphisternum, and in the 
prevomerine teeth are evident between E. coppingeri (Giinther. 
1881) and E. noclosus (Dumeril and Bibron, 1841). E. taen- 
iatus (Girard, 1854) differs markedly from these forms in the 
unnotched xiphisternum. E. grayi (Bell, 1843) and E. roseiis 
(Dumeril and Bibron, 1841) belong to a quite different stock on 
the basis of their greatly developed omosternum, the peculiar 
rounded form of the carinate and calcified xiphisternum (most 
accentuated in grayi), and more posterior placement of the 
vomerine teeth. Affinities between coppingeri and noclosus are 
also indicated by other morphologic features (see key). 

Synonymous species. Eupsophus kriegi, E. maculatus and E. 
verrucosus are synonyms of E. noclosus. E. hihroni, E. calcaratus 
and E. masarcyi are synonyms of E. grayi. The type of E. hidalgi 
(Jiminez de la Espada, 1875) is probably lost and the position of 
this very poorly described species (only the t.vpe is known) ap- 
pears uncertain. It may be synonymous with E. faeniaius on the 
basis of morphological characters, but no final decision is possible. 



BREVIORA 



No. 118 




6 mm 



5 mm 



Fig. 1. Pectoral girdles and prevomerine teeth in Chilean Eupsophus. 
A, E. coppingeri, Puerto Eden, Patagonia; B, E. nodosus, Valdivia; C, E. 
tacniatus, Angol; D, E. roseus, Valdivia; E, E. grayi, Concepci6n. 



!!)()() CHILEAN EUPSOPIIUS 5 

Identity of the pectoral girdle and prevomerine teeth and an 
analysis of many characters, both in the holotypes of K. maciiJa- 
fus (British Museum 60-9-23-4-1947-2-19-99) and E. nodosua 
(Paris Museum, Kill) and in my samples, fully support the pro- 
posed synonymy of E. maculatus (Giinther, 1808) with E. nodosus. 
Some local differences due to geographic variation may have in- 
fluenced Giinther in describing maculatus. Eupsophus kriegi 
(Miiller, 1926) was described from male specimens, and macu- 
laius and nodosus were at that time known only from females, 
as pointed out by Philippi (1902). A remarkable sex-dimorphism 
in nodosus apparently has resulted in the continued recognition 




If^ ^^ 




Fig. 2. Secondary sexual features of male Eupsophus. A, E. nodosus; 
B, E. grayi; C, E. taeniatiis. 

of kriegi. Only the two type specimens of Miiller 's species were 
known (Valdivia, Munich 141/1925; Quebrada de Penalolen, 
Santiago de Chile, Munich 35/1931) ; both were destroyed during 
World War II. The excellent description by Miiller and my obser- 
vations on living specimens from the central zone of Chile, are 
sufficient to identify E. kriegi as the adult male of E. nodosus, 
characterized by tremendous development of secondary sex 
characters (Fig. 2). 

A direct examination of the neotype (Instituto Miguel Lillo 
00162) of Vellard, and a preliminary analysis of geographic 
variation in nodosus make it possible to recognize the redescribed 



6 BREVIORA No. 118 

E. verrucosus (Philippi, 1902) as a juvenile specimen of nodosus 
from the Valdivian forest population. 

Synonymy of Eupsophus hihroni (Bell, 1843) with E. grayi is 
easily verified by study of the original descriptions of Bell and 
a re-examination of the holotypes (British Museum 45-5-25-43- 
1947-2-19-26, and 45-5-25-25-1947-2-17-83, respectively). In my 
opinion E. calcaratus (Giinther, 1881) is based upon a juvenile 
gj'ayi from the Chiloe Islands. All external characters correspond 
in both forms, and the peculiar characters of calcaratus (V- 
shaped glandular dorsal line, tibio-tarsal inner tubercle) occur 
frequently as individual variants, in insular (Chiloe) and con- 
tinental populations (Malleeo) of E. grayi. Besides the holotype, a 
specimen collected by Cunningham (British Museum 68-9-22-8- 
1947-2-20-1), only two young individuals from Ancud (Chiloe) 
are listed in the British Museum as calcaratus (personal com- 
munication by A. C. C. Grandison). The holotype of E. masarcyi 
(Roux, 1910) (Basel 2786) matches extreme individual variants 
in my Valdivian sample of grayi. Specimens such as Museum of 
Comparative Zoology 13742, from Valdivia, agree quite well with 
the description by Roux (1910) and also with the type re- 
examined through the courtesy of Dr. Forcart. 

Geographic variation. Geographic variation is an important 
feature needing further study in populations of these Chilean 
batrachians. Statistically significant differences are indicated 
in cephalic shape between the samples from the continent and 
Chiloe Island (males and females), both in E. grayi and E. 
taeniatus. The head of insular specimens is noticeably more 
elongated. A statistical analysis of Chilean E^ipsophus will be 
presented later in a more detailed paper. 

Color. Color patterns exhibit some interesting trends in all 
Chilean Eupsophus (Fig. 3). Eupsophus nodosus (Fig. 3A) pre- 
sents characteristic enlarged dorsal spots, dark bands on the legs, 
and a triangular brownish or greenish spot. Eupsophus coppin- 
geri (Fig. 3B) shows a reduced dorsal pigmentation ; the enlarged 
spots and bands are not indicated, but the interocular greenish 
spot is present and brilliant in life. The manner of the evolution 
of typical color pattern of nodosus is suggested in E. grayi (Fig. 
3 D and E). In some specimens a whitish dorsal line is evident 
(Fig. 3E). A line also appears in 40 })er cent of the observed 



15)60 



CIIILICAN KlJI'SOI'JllS 










k; 






o 






O 



c 



K5 






a, 

as 






o 

a 



bJD 






8 



BREVIORA 



No. 118 



20 




• E. GRAYI  E. ROSEUS 

Fig. -i. Distribution of Cliileau Enpsopltzis. Grai/i groui). 



1 !»()() CHILEAN EUPSOPTIUS 9 

specimens of E. roscus (Fig. HF), a species whicli agrees in 
coloration with some examples of grayi but with the spotting re- 
duced or indistinct. The dorsal surface is generally' ochraceous, 
greenish or gray in Eupsophus, but in some specimens (the 
masareyi form of grayi) a brilliant reddish color has been re- 
ported. Individual variation ])robably is involved. Eupsoi:)hus 
tacniatus (Fig. 3C) is the mo.st distinctive Chilean form in color 
pattern. The interocular spot and brownish dark bands on the 
legs are occasionally indicated and two peculiar lateral dorsal 
brownish stripes are also present. Inguinal dark spots are typical 
of tacniatus and sometimes occur in grayi on Chiloe Island. 

Distribution. Apparently' Eupsophus nodosus and E. taeniatus 
are primitive, well adapted and formerly widespread species. 
Similarities in shoulder girdle and prevomerine teeth indicate 
the relationship between the two forms. Eupsophus copping eri 
appears to be a specialized form derived from nodosus and is sym- 
patric with the southern populations of EupsopJius nodosus over 
a wide area. The distribution of Eupsophus nodosus and E. tac- 
niatus is significant. Present relict populations of both species are 
found in the Valparaiso coastal forest and in the valleys of 
Santiago and ])robably reveal a late Tertiary invasion, under 
other climatic and ecological conditions (Capurro, 1952). 
Eupsophus nodosus and E. tacniatus also are found, with E. 
grayi, on the eastern slopes of the Cordilleras (Bariloche, Argen- 
tina), and Scliaeffer (1949) suggests a late Tertiary occurrence 
of Eupsophus in Patagonia. 

Eupsophus grayi and E. roscus are sympatric in the Valdivian 
rain forest, but the interspecific and physiological limits of both 
forms are not yet known. Similarities in the pectoral girdle, pre- 
vomerine teeth and color patterns of grayi and roscus have been 
pointed out above (Figs. 1 and 3). Evolution of their color 
patterns from a primitive nodosus-Uke pattern is probable (see 
Fig. 3). Other significant biological features of Chilean 
Eupsophus are the male secondary sex characters. In the grayi 
and tacniatus groups the dark (^ueratinous pads (first and second 
fingers) are similar, but (juite different from the spiny compli- 
cated pads of nodosus, in which circular spiny areas are also 
present. 



10 



RREVIORA 



No. 118 




72 70 ' 6^ 



C_";e. TAEN1ATUS#E. NODOSUsB^- C0PPIN6ERI 

Fig. 5. Distribution of Chilean Eupsophus. Nodosi^ and taeniatus 
groups. 



lf)60 CHILEAN EUPSOPIII'S 11 

Life historij. 'Vhv life liistory and development of Chilean 
Eiipsophus are poorly known, but Eupsophus tacniatus shows 
similarity (see Cei and Ca{)urro, 1958) to the Brazilian species 
of Thoropa which was called Borhorococtcs by Boulencrer (1891) 
and Noble (1931) but is referred to Eupsophus in the recent 
paper by Cochran (1955). Eii:gs of Eupsophus tacniatus are pi*r- 
mented but they are without pigment in the grayi and nodosus 
groups. 

SUMMARY 

Chilean species of Eupsophus are reviewed. Only three funda- 
mental groups are recognized : grayi-roseus, nodosus-coppi^igeri , 
taeniatits. Sj^nonymy of E. kriegi, E. maculatns and E. ver- 
rucosus with E. nodosus, and of E. hihroni, E. calcaratus and E. 
masarcyi with E. grayi is demonstrated. 

Morphological affinities in pectoral girdle, prevomerine teeth 
and color patterns in Chilean species are analyzed. Secondary 
sex characters of grayi, nodosus and taeniatus are compared and 
illustrated. The present distributions of nodosus and taeniatus 
express their probable position as ancient and formerly wide- 
spread forms. Biological affinities between E. taeniatus and Bra- 
zilian Thoropa are mentioned. 

Eupsophus coppingeri and E. roscus are specialized and rare 
forms, sympatric with grayi and nodosus in the southern Chilean 
rain forest belt. 

A Key to the Cliilean Species of Eupsophus 

la. Prevomerine teeth between anterior edges of ehoanae; (•arti]ag:inous 
omosternum very reduced, much shorter tlian epicoracoid cartilages; 
xiphistcrnum not calcified, notched or irregular at posterior margin, 
longer than epicoracoid cartilages -' 

lb. Prevomerine teeth between or behind posterior edges of ehoanae; 
cartilaginous omosternum dilated, only slightly shorter than epicoracoid 
cartilages; xiphisternum partially calcified, rounded at posterior mar- 
gin, shorter than epicoracoid cartilages 4 

2a. Xiphisternum irregularly rounded posteriorly; tympanum evident; 
adults not exceeding 46 mm. in standard length; slender habitus; skin 
smooth; snout prolonged; gray or greenish, with two lateral dark 
stripes; belly yellowish tacniatus 



]2 BREVIORA No. 118 

2b. Xiphisternum deeply notched posteriorly; tympanum not evident, cov- 
ered by granular skin; adults more than 46 mm. in standard length; 
stout habitus 3 

3a. Skin tuljercular; snout obtuse; a glandular tympanic ridge; head not 
markedly depressed; gray or greenish with dark irregular dorsal spots; 
belly whitish nodosum 

3b. Skin smooth; snout truncate; no glandular tympanic ridge; lica<l 
markedly depressed ; olive-brown, with obscure brov\aiish markings ; 
belly salmon-pink coppingeri 

i;\. Prevomerine teeth far liehind choanae; snout prolonged; skin smooth, 
no glandular ridges; bro^vnish or greenish with dark dorsal blotches; 
lielly grayish-white; adults no more than 50 mm. in standard length . . 

grai/i 

-ill. Prevomerine teeth between posterior margins of choanae; snout 
rounded; skin smooth but with a few glandular ridges; brownisli with 
obscure dark markings; belly whitish; adults more than 50 mm. in 
standard length roseus 

LITERATURE CITED 

Bell, Thomas 

1843. In DarAvin, The zoology of the voyage of H.M.S. Beagle, 1832-3(5. 
Pt. 5, Reptiles, 51 pp., 20 pis. 

r.oi-LENGER, George Albert 

ISyi. Notes on American batrachians. Ann. Mag. Nat. Hist., (6) 8: 
453-457. 

Capurro, Luis F. 

1952. Eupsoplins lacniatus Girard anfibio del bosque relicto de Quia 
tero. Invest. Zool. Chilenas, 1 (8) : 3-6. 

Cei, Jose Miguel and Luis F. Capurbo 

1958. Biologia y desarrollo de Eupsophus taeniattts Girard. Invest. 
Zool. Chilenas, 4: 159-182, pis. 1-4. 

Cochran, Doris Mable 

1955. Frogs of southeastern Brazil. Bull. United States Nat. Mas., 
206, xvi + 409 pp., 34 pis., 28 text-figs. 

DuMERiL, Andre Marie Constant and Gabriel Bibron 
1841. Erpetologie generale. Vol. 8, 784 pp. 



1J)6() CHILEAN EUPSOPIIUS \'.j 

(liRARD, Charles 

18."4. Ahstraft of a report to I^icut. Jnuies ^^. Gilliss, T'''.S.N., upon tlio 
reptiles collected during the U.S.N. Astronomical Expedition 
to Chile. Proc. Acad. Nat. Sci. Philadelphia, 7:226-227. 

GtTs-THER, Albert Carl Ludwig Gotthilf 

1868. First accoimt of species of tailless batrachians added to the 

collection of the British Museum. Proc. Zool. Soc. London, 

1868: 478-490. 
1881. Account of the zoological collections made during the survey 

of H.M.S. Alert in the straits of Magellan and on the coast of 

Patagonia. III. Reptiles, batrachians and fishes. Proc. Zoo). 

Soc. London, 1881: 18-22. 

Jimi'nez de la Espada, Marcos 

1875. Vertebrados del viaje al Pacifico de 1862 a 1865 por una 
comision de naturalistas. Batracios. Madrid. 208 pji., 6 pis. 

MULLER, LORENZ 

1926. Neue Reptilien und Batrachien der Zoologisehen Samnilung des 
Bayrischen Staates. Zool. Anz., 65 (7/8) : 195-197. 

XOBLE, GlADWTN KiNGSLET 

1931. The biology of the Amphibia. McGraw-Hill, New York, 577 pp., 
174 text-figs. 

Parke^i, Hampton Wildman 

1932. The systematic status of some frogs in the Vienna Mu^ieum. 
Ann. Mag. Nat. Hist.; (10) 10: 341-344. 

Philippi, Rudolph Amandus 

1902. Suplemento a los Batraquios chilenos descritos en la Histori;i 
Fisica i Politica de Chile de don C. Gay. Santiago, xi + 1"! 
pp. 

Roux, Jean 

1910. Eiue neue Cystignathidenart aus Chile. Zool. Anz., 86 (4/5) : 
111-112. 

Schaeffer, Bobb 

1949. Anurans from the early Tertiary of Patagonia. Bull. Amer. 
Mus. Nat. Hist., 93 (2) : 41-68. 

Schmidt, Karl Patterson 

1954. Reports on the Lund University Chile Expedition 1948-1949, 13. 
Amphibia Salientia. Lunds Univ. Arsskrift., (N. F. 2) 49 (19) : 
3-6. 

Vellard, Jean 

1947. Dos batracios intersantes de la region de Bariloche. Acta Zool. 
Lilloana, 4: 145-154. 



BREVIORA 

Meseiiaim of Compsirsitive Zoology 



('AMUKiixiK. Mass. March S. HKiO Xi:.mi',i;k llH 

APv( TIC AKCHIBEXTIIAL AND ABYSSAL MOBLl SKS 
FROM DKlFTlXi; STATION ALPHA 

By Arthur 1L Clarke, Jr. 

National ^Iiuseuni of Caiuula 
Ottawa, Ontario, Canada 

INTKODUCTION 

Duriii<i' the summer of 19,38 a floating Arctic ice station (Sta- 
tion Alpha) was manned b^^ personnel from the Camhritlge Air 
Force Research Center and the Lamont Geological Observatory. 
While the station drifted slowly northeasterly in the high Arctic 
region about 800 miles north of Point Barrow, Alaska and 800 
miles from the North Pole, eight representative bottom samples 
were taken from deep archibenthal and abyssal de})ths. 

All of these samples contained molhisks (see Tables 1 and 2), 
a total of seventeen species. L^nfortunately, the sami)les were 
not placed in preservative and it is now impossible to ascertain 
which, if any, were taken alive. Notwithstanding this limitation, 
the collections are significant in that, (1) they come from a region 
that is very poorly known biologically; (2) they represent major 
extensions in the known geographic and bathymetric ranges of 
many of the species; (3) they contain three new species; and 
(4) they indicate the presence of large scale sediment transfer 
processes within the area studied. 

There is only one other report on archibenthal and abyssal 
mollusks from this part of the Arctic Ocean. This is by 0. A. 
Scarlato in a report by K. A. Brodskii and M. M. Nikitin of the 
hydrobiological work done at the Russian Scientific-Research 
Drifting Station of 1950-1951. Fourteen species of mollusks are 
listed from six stations scattered between points lying approxi- 
mately 350 miles N.W. and 550 miles north of Point Barrow, 



2 BREVIORA No. 119 

Alaska. For completeness, the station data and the mollusks 
obtained are listed beloAV (see Tables 3 and 4). 

Table 1. U. S. Station Alpha Bottom Samph^ AVhieh Yielded 

Mollusks 1 

Sample Depth Latitude Longitiule liistriniieiit Quantity of 

No. (fills.) ° N ° W Bottom used iiiateiial taken 

83° 59' 151° 44' brown ooze 10 gal. can 3 gal. 

84° 09' 150° 23' mud, and 1 M. trawl 1 gal. 

liel)l>les 

84° KV 149° 11' pebliles 1 M. trawl 10 cc. 

84° 23' 148° 51' It. brown Ekinan grab li/o qts. 

sediment 

84° 28' 148° 28' mud & rocks 1 M. trawl 1 gal. 

84° 34' 146° 24' l)rown mud 1 M. trawl 1 qt. 

84° 30' 145° 00' brown clayey 1 M. trawl 1 gal. 

mud 
10 1000 85° 01' 138° 00' rocks & silt 1 M. trawl 1 pt.Vt 

rocks 

Table 2. Mollusks in Bottom Samples Taken from U. S. Drifting 

Station Alpha - 



o 


1370 


3 


1364 


4 


907 


5 


971 


6 


924- 




934 


7 


1208 


8 


1257 



Sample No. 


4 


6 


5 10 7 8 3 2 


Depth (fathoms) 


907 


924- 
934 


971 1000 1208 1257 1364 1370 



Oolus hunkinsi, n. sp. — 

Siphonodentaliuin lobatum Sby. — 
Nucula zophos, u. sp. 1 

Portlandia intermedia Sars — 

P. lenticula Mollcr ~ 

Malletia abyssopolaris, n. sp. — 

Bathyarca frielei Friele 2 

Hyalopecteii frigidus Jensen — 
C^y^clopecten (Delectopecten) 

greenlandicus Sowerby — 

Astarte borealis Schumacher — 

A. montagui Dillwyn — 
Serripes groenlandicus 

Bruguiere — 

Macoma sp. — 

Iliatelia arctica Linne — 

Lyonsiella alaskana Dall — 

Poromya sp. (fiagment) — — —  1 —  — — — 

Cuspidaria sp. (fragments) — 1 — — 1 — — 

TOTALS 3 201 4 66 137 18 38 10 

1 Out' live brittle star and 4 live holothurians occiirrcd in samples 7 and 10 
respectively. No otlier living aninuUs were specifically niited by the eolleeter. 

- .Samples are aiTan(,'e(l in order of increasin;: depth. (Quantities refer to the 
number of specinK'ns ({jastropods and scaphopoils) or valves (bivalves) obtained. 



1 


— 


1 


4 


— 


— 


— 


7 


— 


— 


, — 


— 


— 


— 


73 


— 


11 


— 


— 





— 


3 


— 


— 


14 


— 


11 


— 


1 


— 


—  


— 


— 


— 


— 


30 


— 


12 


31 


4 


.) 





50 


4 


34 


34 


6 


8 


— 


25 


— 


3 


36 


o 


— 


— 


5 

1 

O 


— 


3 


13 


() 


10 


4 


^ 


— 


— 


— 


1 


— 


1 
1 


— 


1 


3 


^ 


3 


— 









1 











1960 



ARCTIC ARCniBENTHAL AND ABYSSAL MOLLUSKS 



Table 3. Russian Bottom Samples Which Yielded Mollusks -^ 



SAMPLE 
No. 



DEPTH 

(fms.) 



LATITUDE 

°N 



LONGITUDE 
°W 



R 1 


400 


76' 25' 


167° 14' 


R 2 


1400 


76° 44' 


170° 15' 


R 4 


550 


78° 22' 


167° 10' 


R 6 


1000 


78° 52' 


165° 00' 


R 7 


650 


78° 54' 


162° 00' 


R 11 


1800 


80° 45' 


161° 00' 



Table 4. Mollusks Reported Prom Russian Bottom Samples ^ 



Sample No. 


R ] 


R4 


R 7 


R 6 


R 2 


R 11 


Depth (fathoms) 


400 


550 


650 


1000 


1400 


1800 


Lora violacea Mighels 


— 


— 


2 


— 


— 


— 


Lora sp. 


— 


2 


— 


— 


— 


— 


Sip ho sp. 


— 


— 


sev. 


— 


— 


— 


Cylichna alba (Browu) 


— 


— 


o 


— 


— 


— 


Siphonodentalium lobatum Sby. 


sev. 


sev. 


many 


sev. 


1 


— 


Nucula tumidula Malm 


— 


— 


1 


— 


— 


— 


Nueula sp. 


—  


sev. 


sev. 


1 


— 


— 


Portlandia sp. 


— 


1 


1 


— • 


— 


sev. 


Area pectunculoides Scacchi 


— 


— 


— 


1 


15 


 — • 


Area f rielei Jeffries 


— 


— 


sev. 


— 


— 


— 


Dacrydimii vitreum (MoUer) 


— 


—  


— • 


sev. 


— 


— 


Propeamussium f rigidus (Jensen) 1 


— 


— 


— 


o 

i^ 


— 


Cuspidaria glacialis (G. Sars) 


frag. 


— 


2 


— 


— 


— 


Cuspidaria sp. 


— • 


— 


— 


— 


1 


— 



SYSTEMATICS 
Family NEPTUNEIDAE 

CoLus HUNKiNSi, new species 

Plate I, figure 9 

Shell fusiform, approximately one inch long, white, rather 
thin, and with revolving carinae. Whorls 6I/2, convex, rather 
strongly shouldered, and with sutures deeply impressed. Sculp- 
ture consisting of about 18 well defined carinae on the body whorl 
(only the upper 5 or 6 of which are prominent), about 6 on the 



3 Locations and depths are from a map showing the stations. The values are 
tlierefore only approximate. 

4 The abbreviations "sev." and "frag." are for "several" and "fragments," 
respectively. 

5 Taken alive. All other mollusks represented by empty shells only. 



BREVIORA 



No. 119 



penultimate whorl, and 3 on the upper whorls. Fine, incremental 
lines are also present below the protoconch. Spire produced at 
an angle of about 42°. Parietal lip with a thin callus. Palatal 
lip thin or slightly thickened and convex. Columella broadly 
sigmoid. Aperture ovate, medium sized. Siphonal canal short 
and open. Umbilicus absent. Periostracum very thin, light 
brown. Nuclear whorls about 3%, with no distinct demarcation 
between the nuclear whorls and the postnuclear whorls. First 
whorl small (0.8 mm. in diameter), planospiral, and unsculp- 
tured; second and later whorls turreted and rapidly increasing 
in size. On the second whorl the upper carina and numerous fine 
longitudinal riblets appear. On the third whorl the second and 
third carinae appear and the longitudinal riblets become stronger 
and evenly spaced, equaling the carinae in height and thickness. 
On the fourth whorl the carinae become stronger, the riblets 
gradually give way to fine, incremental lines, and the whorl 
assumes adult sculpture. Operculum not seen. 



Height 



Width 



Length 


Number 


of 


of 


\.perture 


Whorls 



Holotype (sample 6) 


24.4 mm. 


12.0 mm. 


12.0 mm. 


61/2 


Paratype (sample 10) 


20.06 


12.2 


11.2 


— 


Paratype (sample 7) 


10.0 


6.0 


5.8 


4y2 


Paratype (sample 7) 


7.3 


5.0 


4.6 


4 



Types. The holotype, a dead specimen from Station Alpha 
sample 6 (924-934 fathoms, 84° 28' N, 148° 28' W), is No. 222066 
in the Museum of Comparative Zoology. Paratypes from samples 
7 and 10 are at the National Museum of Canada, the Museum of 
Comparative Zoology, and Lamont Geological Observatory. 

Remarks. C. hunkinsi has the aspect of a typical archibenthal 
or abyssal neptuneid. It is similar to C. parvus (Verrill and 
Smith 1882) from off Martha's Vineyard in 312 to 506 fathoms, 
but that species is smaller (14 mm. in length, 7 whorls), much 
less strongly shouldered, and longitudinal riblets are not present 
on the nuclear whorls or at least they are not mentioned in the 
description. Another similar species is C. krampi (Thorson 1951) 
from 1028 fathoms between Disko Island, Greenland, and Baf- 



6 Top of spire broken, only 2^4 whorls remaining. 



1960 ARCTIC ARCIIIBENTHAL AND ABYSSAL MOLLUSKS 5 

finlaiid, but that is sijjfnificantly larger (36 to 39 mm., 5 to 6 
whorls), the constriction between the aperture and sii)honal 
canal is much less prominent, and the whorls are much less 
strongly shouldered. 

The species is named for Mr. Kenneth Hunkins of the Lamont 
Geological Observatory who collected the Station Alpha material 
treated in this report. 

Spccimc7is examined. This species is known only from six dead 
specimens collected from Station Alpha in samples 6, 7, and 10, 
taken in 924 to 1208 fathoms about 800 miles north of Point 
Barrow, Alaska. The "Sipho sp." reported from Russian sample 
R 7 (650 fathoms, 470 miles N.N.W. of Point Barrow) may be 
C hunkinsi, but this is uncertain. 

Family SIPHONODENTALIIDAE 

SiPHONODENTALIUM LOBATUM SoWCrby 1860 

The seven dead specimens from Station Alpha sample 6 (924- 
934 fms.) are all typical, measuring up to 13 mm. in length. 
8. lohaium occurs throughout the Arctic Ocean from shallow 
water to 2000 meters (approximately 1100 fathoms) on soft silty 
and silty-sandy bottoms with a bottom temperature of —1.8 to 
+4° C (Brodskii and Nikitin 1955). It is also recorded from the 
North Atlantic in 60 to 1813 fathoms (La Rocque 1953). 

Family NUCULIDAE 
NucuLA ZOPHOS, new species 

Plate I, figures 15-18 

Shell subtriangular, compressed, rather large for the genus 
(over 1/2 inch long), finely sculptured, with medium sized, well 
developed taxodont hinge teeth and a brownish or blackish 
periostracum. Outline triangular, with prominent, elevated 
beaks located near the posterior third of the shell and with a 
flattened base which is usually nearly straight centrally. Surface 
sculptured with numerous, rather fine, closely spaced concentric 
ridges and grooves, and fine, closely spaced radial lines which 
cause the surface to appear cancellate when viewed under the 
microscope. Exterior stained with brown and black, but portions 
of what appears to be a thin brown periostracum remain. Inner 



6 BREVIORA No. 119 

surface uacreous and exliibitiug- the radial lines which end at 
the strongly crenulated shell margin. Anterior and posterior 
muscle scars and simple pallial line present but not prominent. 
Hinge plate Y-shaped, with an anterior and a posterior row of 
somewhat compressed, columnar, taxodont teeth. The two rows 
are separated by a prominent, narrow, anteriorly directed 
chrondrophore. Taxodont teeth somewhat heavier anteriorly, 
and numbering about 15 to 17 in the anterior row and 7 to 10 
in the posterior. Umbones deeply excavated. 



Length Height " Width (1 valve) 



Holotype (sample 6) 


10.0 mm. 


7.6 mm. 


2.5 mm. 


Paratype (sample 6) 


14.G 


9.9 


2.8 


Paratype (sample 6) 


14.5 


10.0 


3.0 


Paratype (sample 10) 


9.7 


7.3 


2.2 



Types. The holotype, a left valve from Station Alpha sample 6 
(924-934 fathoms, 84° 28' N, 148° 28' W), is number 222067 in 
the Museum of Comparative Zoology. Paratypes from samples 
4, 6, and 10 are at the National Museum of Canada, the Museum 
of Comparative Zoology, and Lamont Geological Observatory. 

Remarks. Because of its peculiar elongate shape, flattened 
base, sub-central umbones, and reticular sculpturing, N. zophos 
does not closely resemble any other species. In sculpturing it is 
somewhat similar to iV. nucleus Linne, but nucleus is smaller, 
more rounded, with the beaks far forward, and much different in 
general appearance. N. iphige^iia Dall 1908 from 259 fathoms in 
the Bay of Panama is probably morphologically closer to zophos 
than is any other Nucnla, but that is much larger and more 
ponderous, the base is more rounded, and the radial sculpturing 
is different and much more conspicuous. 

Specimens examined. The only known specimens are the 85 
single valves dredged from Station Alpha, about 800 miles north 
of Point Barrow, Alaska, in 907 to 1000 fathoms. It is possible 
that the "Nucula sp." collected from the Russian Drifting Sta- 
tion at localities R4, R6, and R7 (550 to 1000 fms.) also represent 
this species, but those specimens have not been examined by the 
writer. 



1960 



AKCTIC ARCHIBENTllAL AND ABYSSAL MOLLUSKS 



Family MALLETIIDAE 
Malletia abyssopolaris, new species 

Plate I, figures 19-22 

Shell siibovate, inflated, of medium size (length about i/^ inch) 
and thickness, weakly sculptured and with small, well developed 
taxodont hinge teeth and a brown (?) periostracum. Outline 
varying from nearly circular to ovate-rhomboid and with or 
without a postbasal swelling. Beaks rather prominent and in- 
flated. Surface sculptured with narrow lines and rays. Ligament 
groove narrow and elongate, running from below the umbo to 
near the middle of the posterior row of teeth. Exterior stained 
brown or blackish, but portions of what appears to be a thin 
brown periostracum remain. Inner surface iridescent and show- 
ing the external sculpturing. Anterior muscle scar impressed, 
posterior scar and pallial line less prominent but usually clearly 
visible. Pallial sinus absent. Hinge plate concave anteriorly and 
convex posteriorly. Hinge teeth taxodont, V-shaped when viewed 
apically, continuous under the umbones, larger and higher an- 
teriorly, smaller posteriorly, and numbering about 32 to 40 in 
adult specimens, of which 10 to 12 are anterior and 22 to 28 are 
posterior. Umbones excavated. 



Length 



Height 



Width (1 valve) 



Holotype (sample 6) 12.6 mm. 

Paratype (sample 6) 14.0 

Paratj-pe (sample 6) 12.9 

Paratype (sample 7) 11.0 



11.0 mni. 
10.0 

9.6 

8.0 



3.6 mm. 
4.3 
3.7 
3.5 



Types. The holotype, a left yalve from Station Alpha bottom 
sample 6 (924-934 fathoms, 84° 28' N, 148° 28' W), is No. 222068 
in the Museum of Comparative Zoology. Paratypes from samples 
2, 3, 6, 7, 8, and 10 are at the National Museum of Canada, the 
Museum of Comparative Zoology and Lamont Geological Ob- 
servatory. 

Remarks. M. abyssopolaris resembles M. dbyssorum Verrill 
and Bush 1898 taken off Chesapeake Bay in 2620 fathoms, but 
that species is only 5 mm. long, the teeth are not continuous 
under the beaks, and it is without radial lines. M. dunkeri Smith 



8 BREVIORA No. 119 

1885 from off Japan in 1875 fathoms is also somewhat similar, 
but it too is only 5 mm. long, it is without radial lines, and the 
beaks are proportionately much smaller. 

This species is apparently more closely related to Malletia 
than to other existing genera, notwithstanding the absence of the 
pallial sinus, which according to most authors is present in all 
species of Malletia. This is a case similar to that of Tindaria 
erehus Clarke, 1959, in which the pallial sinus is present, al- 
though it is supposedly absent in species of Tindaria. For reasons 
given there, this single character is not considered sufficient 
to disrupt an otherwise acceptable generic placement, and rather 
than place 31. ahyssopolaris in a new genus or subgenus, it is 
left in Malletia. 

Specimens examined. Known only from 88 single valves col- 
lected from Station Alpha between 924 and 1370 fathoms about 
800 miles north of Point Barrow, Alaska. 

Family NUCULANIDAE 

PORTLANDIA INTERMEDIA SarS 186.") 

Plate I, figures 6-8 

Single valves of this species occurred in samples 3, 6, and 7 
in depths ranging from 924 to 1364 fathoms. It is panarctic in 
distribution and has been recorded alive from depths ranging 
from 4 to 5 fathoms in the Siberian Ice Sea to 630 fathoms near 
the Shetland Islands. Dead shells have been recorded in the 
North Atlantic from depths as great as 1273 fathoms (Ockel- 
mann, 1958). It usually occurs on clay or mud bottoms, some- 
times with admixtures of sand and gravel. 

PORTLANUIA LENTICULA Mollcr 1842 

Plate I, figure 4 

A single valve in sample 6 (924-934 fathoms) seems to belong 
to this species. It is another panarctic species and has been 
recorded from zero fathoms in eastern Greenland to about 76-" 
fathoms north of the Shetland Islands. It is reported as usually 
occurring on clay or mud which may or may not contain addi- 
tional sand or gravel, as in Portlandin int<'r)u< ditt. 



1960 ARCTIC ARCIIIBENTIIAL AND ABYSSAL MOLLUSKS 9 

Family ARCIDAE 
Batiiyarca frielei ''Jeffreys" Friele 

Plate I, figures 10-14 

Ana frit'Ui (Jeffreys m.s.) Friele 1877, Nyt Magazin for Naturvidenska- 
berne, 23:2. Type locality: Norwegian Sea, Norwegian 1876 Expedi- 
tion stations 40 (1180 fnis.), 51 (1130 fms.), and 53 (1500 fms.) ; 
also Porcupine Expedition. 

? Area imitata Smith 1885, Challenger Keports, Lamellibranchiata, p. 321 + 
text figs. Type locality: Challenger station 244, Mid-North Pacific 
in 2900 fathoms. 

The specimens collected from Station Alpha are variable in 
shape, and since the extremes are all connected by intergrades it 
is impossible to separate them objectively. It is probable that 
the Russian report of both Area pectunculoides Scacchi and Area 
frielei is the result of such an attempt. The type figures of 
pectunculoides in Scacchi 1835 (pi. 1, figs. 12 a, 12 b) are poor 
but certainly do not represent this species. We are left with the 
choice of using frielei "Jeffreys" Friele 1877 or imitata Smith 
1885, both of which represent forms which are very similar to 
each other and to the present species, but not identical to it. 
Smith's figures of the two forms of imitata resemble the most 
frecpient variations seen in the present species, but since frielei 
is more frequently used and is the earlier name, it is used here. 
The problem should be studied further. 

Dead specimens of this species were found in seven of the eight 
samples collected from Station Alpha, in depths ranging from 
907 to 1364 fathoms. The Russian report of a living specimen 
of A. pectunculoides at 1400 fathoms probably represents this 
species. It is recorded from the North Atlantic and adjacent 
Arctic Ocean in depths ranging from 10 to 12 fathoms (north- 
east Greenland) down to 1540 fathoms (off western Norway), 
and as living only in regions having a temperature of —1° to 
+1° C. It is also reported from the Barents and Kara seas on 
brown, soft silt at depths of approximately 80 to 110 fathoms 
and deeper (Brodskii and Nikitin). According to Ockelmann 
(1958), on the coast of eastern Greenland it is much more com- 
mon below 130 fathoms than in shallower deptlis. 



10 BREVIORA No. 119 

Family PECTINIDAE 
Hyalopecten frigidus Jensen 

Plate I, figures 1-3 

Pecten fragilis Jeffreys 1876, Annals and Magazine of Natural Ilist<»ry, 
(4) 18, 424. Type locality: Valorous Expedition stations 9, 12, and l(i, 
between Ireland and Greenland, 1450 to 1785 fathoms and NorweKiini 
Expedition of 1876, 1000 to 1500 fathoms, Greenland and Norwegian 
Seas. Includes three species: P. undatus Verrill 1884, P. greenlandicun 
Sowerby 1845, and P. frigidus Jensen 1912. Figured by Jeffreys (Pro- 
ceedings of the Zoological Society of London for 1879, pi. 45, figs. 1, 1). 
Not Peoten fragilis Montagu 1808. 

Pecten frigidus Jensen 1912, The Danish lugolf Expedition, 2, part 5 
(Lamellibranchiata, part 1), p. 33, pi. 1, figs. 7 a-f. New name for 
P. fragilis Jeffreys, in part. 

Jensen 1912 has clarified the previously confused taxonomic 
status of this species by pointing out the following facts: (1) 
Jeffreys' original description of fragilis was based on three 
species: P. undatus Verrill (=P. biscayensis Loeard 1898), frag- 
ments of which were collected by the Valorous Expedition be- 
tween Ireland and Greenland ; P. greenlandicus Sowerby, from 
the same source, believed by Jeffreys to be the lower valve of 
fragilis; and a third species from the Norwegian Sea went to 
Jeffreys by Friele, and later named frigidus by Jensen. (2) 
Jeffreys' two figures of fragilis are of tliis third species but show 
a right valve in the right-hand figure and the mirror image of 
the right valve in the left-hand figure. 

The right-hand figure (Jeffreys 1879, loc. cit.) is hereby se- 
lected as the type figure of fragilis Jeffreys 187G. It therefore 
becomes automatically also the type figaire of frigidus Jensen 
1912. Pending possible location of the specimen, with locality 
data, from which the figure was drawn, the type locality cannot 
be restricted within the region explored by the Norwegian Nortli 
Atlantic Expedition of 1876 to 1878, viz., the Norwegian and 
Greenland seas. 

Seventy-six specimens and fragments from Station Ali)lia 
samples 6, 7, 8, and 10 (924-1257 fathoms) appear to belong to 
this species. It has been reported alive from the Norwegian and 
Greenland seas in 579 to 1539 fathoms in localities where bottom 
temperature is between and — 1.1° C. 



1960 ARCTIC ARCIIIBENTHAL AND ABYSSAL MOLLUSKS 11 

Cyclopecten (Delectopecten) greenlandicus Sowerby 1845 

Forty-one single valves and fragments of this fragile species 
occurred in Alpha Station samples 2, 3, 6, 7, 8, and 10 (924 to 
1370 fathoms). Tlie species is recorded from the subarctic North 
Atlantic Ocean and from localities generally distributed over the 
entire Arctic coasts of North America, Europe, and Asia. It 
occurs alive from 2 fathoms in east Greenland (Ockelmann, 
1958) to 1100 fathoms between Greenland and Jan Mayen (Hiigg 
1905), usually on clay containing stones, gravel, or shells. 

Family ASTARTIDAE 
AsTARTE BOREALis Schumachcr 1817 

Only one small, damaged valve was collected from Station 
Alpha of what is assumed to be this species. This occurred in 
sample 6 (924-934 fathoms). It is a panarctic, circumpolar 
species, recorded also from the North Atlantic. Living specimens 
are known from fathoms in east Finmark to 254 fathoms north 
of Spitzbergen. Dead shells have been recorded from 1482 
fathoms in the North Atlantic. 



AsTARTE MONTAGUi Dillwyn 1817 

Plate I, figure 5 

Jensen (1912) and Ockelmann (1958) have done much to 
clarify the status of this variable species. Three valves from 
Station Alpha samples 3 and 6 (924 to 1364 fathoms) appear 
to belong to it. A. montagui occurs in the North Atlantic and 
throughout the Arctic in depths ranging from the low^ tide line 
(western Baltic Sea) to 244 fathoms (western Greenland). 
Usually found on sand or clay, the species also occurs on rocky 
or muddy bottoms (Ockelmann, 1958). 

Family CARDIIDAE 
Serripes groenlandicus Bruguiere 1789 

Three fragments from station 7 (1208 fathoms) belong to this 
species. aS^. groenlandicus is panarctic, extending south in the 
Pacific to Hokkaido, Japan, and Puget Sound, Washington, and 



12 BREVIORA No. 119 

south in the Atlantic to Cape Cod, Massachusetts, and northern 
Norway. 

In depth it ranges alive from just below the low tide line 
(Iceland) to 166 fathoms (western Greenland), and single valves 
have been found as deep as 1340 fathoms in the North Atlantic. 
Although it is most abundant on sand or mud bottoms, it occurs 
on other substrates also. 

Family TELLINIDAE 
Macoma sp. 

This genus is represented by a fragment collected in Station 
Alpha sample 6 (924 to 934 fathoms). It is a panarctic and 
circumboreal group and occurs alive from low tide to depths of 
only a few fathoms. 

Family SAXICAVIDAB 
HiATELLA ARCTicA Liune 1767 

Five valves from Station Alpha samples 3, 6, and 10 (924 to 
1364 fathoms) show the double row of spines on the i)osterior 
slope Avhieh is characteristic of H. arctica (see Abbott 1954, fig. 
92, after Lebour, 1938). The species has an apparently enormous 
geographic range, being recorded as panarctic and south to 
western Panama, the West Indies, the Mediterranean, and even 
from the Cape of Clood Hope. Probably several sibling species 
are involved however. In the Arctic it occurs principally at the 
low tide line or in shallow water, although a doubtful record for 
live specimens in 1200 fathoms off Ireland has been reported 
(Ockelmann, 1958). Single valves are reported from the North 
Atlantic in about 1300 fathoms. A common habitat is among the 
holdfasts of kelp although the species is also known to bore into 
soft rock. 

Family VERTICORDIIDAE 
Lyonsiella (Laevicordia) alaskana Dall 1895 

One valve from sample 7 (1208 fathoms) apparently belongs 
to this species. It was described from a living specimen found in 
1569 fathoms in tbe Gulf of Alaska south of Sitka, Alaska, in 
green ooze and later recorded also from off Catalina Island, 
California, in 600 fathoms. 



li)60 



AKCTIC ARCIIIBENTHAL AND ABYSSAL MOLLUSKS 



13 



Family POROMYIDAE 

POROMYA Sp. 

A single unidentifiable fragment of Poromya occurred in 
sample 10 (1000 fathoms). Poromya is a cosmopolitan genus of 
archibenthal and abvssal bivalves. 



Family CUSPIDARIIDAE 

CUSPIDARIA sp. 

An unidentifiable fragment of Cuspidaria occurred in samples 
6 (924 to 934 fathoms) and 7 (1208 fathoms). Like Poromya, 
Cuspidaria is also a cosmopolitan archibenthal and abyssal genus. 

CONCLUSIONS 

Samples 3, 6, 7, and 10 (see Tables 2 and 5) contain shells 
which clearly were derived from shallow water. All of the 
samples contain species that might have come from shallow water 
and it is therefore probable that transport from shalloAV water 
has occurred at most, or possibly all, of the stations on which this 
study was based. 

Table 5. 

Species which could be living 
where collected or could have 
been transported from 
shallow water 



Species probably 
living where collected 



Species probably 
transported from 
shallow water 



Colus hunkinsi 
Nucula zophos 
Malletia abyssopolaris 
Ilyalopeeten f rigidus 
Lyonsiella alaskana 



Portlandia intermedia 
Portlandia lenticula 
Astarte borealis 
Astarte montagui 
Serripes groenlandicus 
Macoma sp. 
Hiatella arctica 



Siphonodentalium lobatum 
Batliyarea frielei 
(Vclopecten greenlandicus 
Poromya sp. 
Cuspidaria sp. 



Supporting evidence for the transport theory lies in the fol- 
lowing observations. (1) Many of the specimens, especially those 
which are considered to have been transported and those in the 
questionable category, are predominantly fragmentary and give 



14 BREVIORA No. 119 

evidence of having been subjected to much more friction and 
mechanical wear than is observed in specimens from stable en- 
vironments. (2) Most of the species which are thought not to 
have been transported, because of agreement between observed 
depth and recorded depth range, including some which are very 
fragile (e.g. Hyalopecten frigidus), are noticeably less worn and 
fragmentary. (3) The character of the sediments in the samples 
from which the mollusks were removed shows them to be unsorted 
(see Table I) and composed of several kinds of rock. 

Friele and Grieg (1901) and others report similar mixtures of 
shallow and deep water species from archibenthal and abyssal 
depths off northern Norway, south of Spitzbergen, and between 
Jan Mayen and Iceland. Large stones were also observed scat- 
tered over the bottom in the Greenland Sea and elsewhere. They 
also ])oint out that ice masses are frequently seen to carry large 
quantities of mud and other sediment derived from shallow 
water areas, and postulate that it is the melting of such sedi- 
ment-laden ice masses far from land that causes shallow water 
mollusks, stones, etc. to be deposited in these regions. 

Similar mechanisms must have operated in the high Arctic 
and probably account for the presence of shallow water species 
in the archibenthal and abyssal samples dredged from Station 
Alpha. Most of the specimens were covered with a thin coating 
of magnesium oxide when collected, indicating that they had been 
exposed at the surface of the sediment for a protracted period. 
This observation, together with (1) data derived from gravity 
cores indicating a slow rate of sedimentation and (2) observa- 
tions made at Station Alpha that the present Arctic Ocean ice 
cover contains little or no rock fragments or sand, has led Donn 
ct al. (1959) to the conclusion that the rafting must have oc- 
curred during an interA'al of open water which predated the 
Recent Epoch. 

In 1954 the author examined mollusk shells and other inverte- 
brates taken along with stones and dirt from one of the floating 
ice islands (Arctic Ice Island T-3, see Crary, 1958 and Fletcher, 
1953). Fifty-seven valves of Astarte crcnata Gray and three of 
Batlniarca frielei Friele were present in the sediment as well as 
approximately 150 serpulid worm tubes and four colonies of 
an unidentified ectoproct bryozoan. The mollusks are common, 
panarctic, sub-tidal to archibenthal species. Clearly, some ice 



1960 ARCTIC ARCIIIBENTHAL AND ABYSSAL MOLLUSKS 15 

transport of sediment, shells, etc. from shallow water is still 
going on and the process has not been confined to pre-Recent 
epochs. 

BIBLIOGRAPHY 

Abbott, R. T. 

19.^4. Aiiieriean Seashells. D. Van Nostrand Company, Inc., New York, 
i-xiv + 1-541 pp., 40 pis. 

Brodskii, K. a. and M. M. Nikitin 

1955. nydrobiological Work (English translation), Materialy nabliu- 
denii nauehno-issledovatel skoi dreifuiushchei stantsii 1950/51 
gada, ed. M. M. Somov, 1 zd. 'Morskoi Transport'. Vol. 1, 
Sect. 4. Translated by American Meteorological Society. ASTIA 
document No. AD 117135. 

Clarke, A. H., Jr. 

1959. New Abyssal Mollusks from Off Bermuda Collected by the 
Lamont Geological Observatory. Proc. Malacological Soc. Lon- 
don, 33(5) : 231-238, pi. 1, fig. 1. 

Crary, a. p. 

1958. Arctic Ice Island and Ice Shelf Studies. Arctic, 11 (1) : 3-42. 

Dall, W. H. 

1895. Report on Mollusca and Brachiopoda Dredged in Deep Water, 
Chiefly Near the Hawaiian Islands, with Illustrations of Hither- 
to Unfigured Species from Northwest America. Proc. U. S. Nat. 
Mus., 17:675-733, pis. 23-32. 

DoNN, W. L., ]\L EwiNG, and R. J. Menzies 

1959. Cliaracteristics of the Late Quaternary Arctic Ocean. Inter- 
national Oceanographic Congress, Preprints, pp. 19-20. 

Fletcheir, J. O. 

1953. Three Months on an Arctic Ice Island. The National Geographic 
Magazine, 103(4) : 489-504. 

Frielb, H. 

1877. Preliminary Report on Mollusca from the Norwegian North 
Atlantic Expedition in 1876. Nyt Magazin for Naturvidenska- 
beme, 23(3):1-10. 

FsiELE, H. and I. A. Geieg 

1901. Zoologi, Mollusca 3, Den Norske Nordhavs-Expedition 1876- 
1878, 28: 10-131, map. 



16 BREVIORA No. 119 

Ha'gg, R. 

1905. Mollusca and Brachiopoda gesammelt von dcr schwedischen 
zoologisehen Polarexpedition nach Spitzbergen, dem nordost- 
lichen Gronland und Jan Mayen i. J. 1900. Arkiv for Zoologi, 
2(13):1-136, pi. 

Henderson, J. B. 

1920. A Monograph of the East American Scaphopod Mollusks. Bull. 
U. S. Nat. Mus., Ill: i-vi + 1-177, 20 pis. 

Jensen, A. S. 

1912. Lamellibranchiata, Part 1. The Danish Ingolf -Expedition, 2(5) : 
1-121, 4 pis. 

Knudsen, J. 

1949. Scaphopoda. The Zoology of Iceland, 4(62) : 1-7. 

La Rocque, a. 

1953. Catalogue of the Recent Mollusca of Canada. Bull, Nat. Mus. 
Canada, 129: i-ix + 1-406. 

Leche, W. 

1883. Arktiska Hafsmollusker. 1. Lamellibranchiata. Vega Expedi- 
tionens. Velenshapliga lakttagelser, 3; 433-529, 3 pis. 

Ockelmann, W. K. 

1958. Marine Lamellibranchiata. The Zoology of East Greenland. 
Meddelelser om Grcmlaoid, 122(4) : 1-256 + 3 pis. 

Oldroyd, I. S. 

1924. Marine Shells of Puget Sound and Vicinity. Publications Puget 
Sound Biological Station, Univ. of Washington, 4:1-272. 

Posselt, H. J. and A. S. Jensen 

1898. Crr0nlands Brachipoder og Bkrddyr. Conspectus Faunae Groen- 
landicae. Meddelelser om Gr0nland, 23: i-xix -f 1-298, map. 

Sars, G. O. 

1878. I. Mollusca Regiones Arcticae Norvegiae. Bidrag til Kundaska- 
ben om Norges Arktiske Fauna. Christiania, Norway, i-xiii + 
1-467, 52 pis. 

SCACCHI, A. 

1834-1835. Notizie Intorno AUe Conchiglie ed a' Zoofiti Fossili. Annali 
Civili (Sicilie) (Nov. -f Dec. 1834) 6:75-84. Ibid., 7:5-18, 2 pis. 
(Jan.- April 1835). 



1!)(J0 ARCTIC ARCIIIRENTIIAL AND ABYSSAL MOLLUSKS 17 

Smith, E. A. 

1885. Report on tlio Laiiu'Ililiranoliia.ta. Challenger Reports, Zoology, 
13:1-341, 25 pis. 

Thorson, G. 

1941. Marine Gastropoda Prosobranchiata. The Zoologj- of Iceland, 
4(60): 1-150. 

1944. ^Marine Gastropoda Prosobranchiata. The Zoology of East Green- 
land. Meddelelser om Grmiland, 121 (13) :1-181. 

1951. The Godthaab Expedition 1928. Meddelelser om Gr0nland, 
81(2):1-118. 



Plate 

Arctic Archibenthal and Abyssal MoUusks collected from Station Aljjha 

1 - 3 Eyalopecten frigidus Jensen, sample 6 ; 

4 Portlandia lenticula M0ller, sample 6 ; 

5 Astarte montagui Dillwyn, sample 6; 

6 - 8 Portlandia intermfidia Sars, sample 3; 
9 Colus hunhinsi Clarke, holotype, sample 6 ; 

10-14 Bathyarca frielei ' ' Jeffreys ' ' Friele, station 6 ; 
15 - 18 Nucula zophos Clarke, holotj-pe (16) and paratypes, all sample 6 ; 
19 - 22 Mallctia abyssopolaris Clarke, holotype (22) and paratypes from 
sample 6 (19, 20, 22) and sample 7 (21). 

Figures 1 - 8 and 10 - 14 are 2.4 X, figure 9 is 3.2 X, and figures 15 - 22 
are 2.1 X. 




t' 



..^.^^^^ ^v-//' 




18 



J"' 



21 




Plate 



BREVIORA 



Miiseiiiim of Coonparative Zoology 



Cambridge. Mass. March 9, IDGO Number 120 



TAVO SPECMES OF TORTOISES IN 
NOKTIIEKN SOUTH AMERICA 

By Ernest E. Williams 

In 1825 Thomas Bell beautifully figured and clearly dis- 
tinguished two species of land tortoise from northern South 
America. His plates (later reproduced in Sowerby and Lear) 
give an excellent idea of most of the characters by which they 
are even now to be distinguished. Vet 130 odd years later the 
two forms are still customarily synonymized, though occasional 
(not fully documented) suggestions that they may be distinct 
have been made (e.g., Luederwalt, 1926). 

I was first led into the study of this problem when, during my 
work on a fossil Cuban tortoise, I was struck by the variability 
t)f the position of the gular-humeral sulcus in relation to the 
entoplastron in American Museum specimens of "Tesfudo dcn- 
ticnlata." At that time (1950, p. 14) I published a mention of 
this remarkable (as I then thought) intraspecific variability. 
However, dnring my visits to Eurojiean museums during tenure 
of a Guggenheim Fellowship in 1952-53, T became aware that 
two forms, each defined by several correlated characters, were 
being confused under the names "dcnticulata" or "tahnlata," 
used synonymously, but T was not then sure whether subspecies 
or species were involved. 1 have since examined all northern 
South American tortoises in the Museum of Comparative 
Zoology (MCZ), the American Museum of Natural History 
(AMNH), the Chicago Natural History Museum (CNHM), the 
Philadelphia Academy of Natural Sciences (PANS), and the 
United States National Museum (USNM), as well as those of 
the Departamento dc Zoologia (DZ), Sao Paulo, Brasil. It has 
become evident that the two forms are in several places sympatric 



BREVIORA 



No. 120 



or nearly so and that they are best interpreted as two sj)ecies 
— the same two species distinguished by Bell in 1825. 

The species are not dil^cnlt to se])arate and are not in any 
gennine sense sil)lin<is, despite some tendency to overlappinp' 
variability. Variability tending to produce overlaps in individ- 
ual characters is characteristic of all closely related species of 
turtles — that is, of all forms whicii have not at some time been 
separated generically. In tliis case as in others of this sort, 
recognition of species is never to he made on any single supposed 
key character but on the balance of chai'acters in the charactei" 
complex. If determination of species is made on total characters, 
no individual should be at all doubtful or difficult to place. 

I present below in parallel colunnis the differences wdiich seem 
to me useful in diagnosing the two species. 



denticulatii 

Adult (lursal .slirll liruwn, nearly 
uniform oi- with vaguely bounded 
lighter areolae -- Juveniles nearlj' 
uniform yellow brown 

Prefrontal scales elongate 

Frontal scale usually- broken up 

Juveniles with a finely denticulate 
margin 

Concentric grooving on carapace 
shields weak or absent 

Posterior angle of gular scutes well 
forward of the entoplastron 

Dorsal surface of each gular scute 
usually divided {=:4. dorsal gulars) 

Humeral iiiecliau suture usually 
longer than femoral median suture 

inguinal narruwl\ in contaci with 
fenujial on ventral plane of plas- 
tron, i.e. inconspicuous in ventral 
view 



carhiiiiiiiid 

Adult dorsal shell Mack, usuall\ 
with small shaiply lioundcd yellow 
areolae — juveniles like adults Ida k 
and yellow 

Prefrontal scales short 

Frontal scali' usually entire 

Juveniles with a nearly sniootli 
maigin 

Concentric grooving on carapace 
shields usually strong 

Posterior angle of gular scutes en- 
croaching on the entoplastron 

Dorsal surface of each gular scute 
usually undivided (^- dorsal gu- 
lars) 

Femoral median suturt' usually 
longer than humeral median suture 

Inguinal hidadly in contaci with 
femoral on ventral |dane of plas- 
tron, i.e., consi)icuous in vi'iitral 
view 



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4 BREVIORA No. 120 

Each of these characters shows variation and therefore re- 
quires comment : 

Color. The color of the carapace is very strikingly distinct in 
most specimens of the two species, and, so far as I know, is in- 
variably diagnostic in juveniles. However, a few of the older 
denticulafa may show a strongly contrasting pattern of orange 
and dark brown not unlike the black and yellow of carhonaria; 
the colors, however, are duller and the light areolae wider and 
less sharply bounded. 

Prefronial and frontal scales. These differences will hold in 
most specimens but not in all. Some tendency to a break-up of 
the frontal occurs in a few carhonaria; while the converse 
tendency to a nearly entire frontal is present in some denticu- 
lata. 

Dcnticidate margin in juveniles. This is a very consistent 
character in spite of the fact that, examined under magnifica- 
tion, hatchlings of carhonaria show faint traces of the denticula- 
tions so characteristic of its sibling. 

Concentric grooving of carapace shields. A character often but 
not invariably useful. 

Gular scutes encroaching on entoplastron or not. So far as 
it has been possible to check, a consistent character. 

Dorsal surfaces of gidar scutes subdivided or not. Only sta- 
tistically useful. 

Humeral/ femoral ratio. I^sually a very good character, but a 
few carhonaria have the femoral and humeral subequal. 

Inguinal-femoral relationsliip. An excellent diagnostic char- 
acter if attention is paid to the precise relationship. In both 
carhonaria and denticulata the inguinal scute is rather triangu- 
lar, not narrowed anteroposteriorly as in the two other species 
of neotropical tortoises, chilensis and elephantopus. In denticu- 
lata, however, the scute is smaller than in carhonaria and not at 
all conspicuous in direct ventral view. There may be in denticu- 
lata a relatively broad contact with the femoral but this is on 
the edge of the plastron at the inguinal, not as in carhonaria on 
the main ventral plane of the plastron. To see the inguinal- 
femoral contact clearly in denticulata. it is necessary to turn the 
animal to the side to examine the plastral edge. In carhonaria 
the broad contact of inguinal and femoral is best seen in straight 
ventral view. 



1!)G0 



SOrni AMERICAN TORTOISES 



5 



Two characters not tabulated may be significant : 

1. Size. Specimens of dcnticulata from eastern Peru (Bassler 
collection, American Museum) are the lar*jest South American 
tortoises I have seen. AMNll 58084 has a carapace length of 
G73 nun. and AMNIi 58085 is but little smaller (637 mm.). No 
carhonaria approach this size, thoug-h both species of northern 
South America are larger than cliilensls, the third species in 
southern South America. 

2. Carapace shape. Both species tend to have parallel-sided 
shells. However, carhonaria appear on the average to have nar- 
rower shells, often in fact indented midlaterally to give a dumb- 
bell shape in dorsal view. Denticulaia may have more rounded 
contours. There is, however, much variability. 

Many of the South American tortoises in eollections have no 
or very poor data. It is therefore difficult to get a precise idea of 
the geographic relationships of the two species. I list below in 
parallel columns the most precise localities that 1 have been able 
to obtain for specimens examined by me. 



denttculata 
Colombia 



carhonaria 
Colombia 



Dept. Caquetd: 
S. of Florencia 
Morelia 



Dept. Meta: 
Villavicenc'io 



Venezuela 



Dept. Ailantico : 
Cienega de Guajaro 
Puerto Bello 

Dept. Antioguia: 
Golfo de Uraba 



Dept. Cxindinamarca , 
W. of Honda 

Dept. Magdalena: 
Rio Frio 
Totumal 



Venezuela 



Barinas State . 
Barinas 



BREVIORA 



No. 120 



(l< III iciildid (ooiit.) 



Monagas State: 

Juanipa Eiver near Caripito 
Territorio do Amazonas : 

foothills Mt. Dueda 



Trinidad 
British Guiana 

Deinerara 

Essequibo Eiver 
Kamakusa 
Kartabo 
Oho Mtii. 

Surinam 

' ' Paramaribo ' ' 

French Guiana 



Peru 



' ' Cayenne ' 

Dept. Junin: 
Atalaya 



Dept. Loreto : 
Alto Bio Pisqui 
Iquitos 
Pucallpa 
Rio Ucayali 
Eio Xapo 
Yarinacocha 

Brasil 

Amn~on(is State: 

Lago Alc'ixo (Thayer Exped.) 
Lago .la una 11 (Thayer Exped.) 
Manaus 
Rio Xeo-ro 



carhonaria (eoiit.) 

Caraboba State : 
Maracay 



no state: 

Orinoco Region 
Rio Apiire 
Los Testigos 



British Guiana 

Essequibo Eiver ur. Onara 
(70 miles from Georgetown) 

Kamakusa 

head of Rupononi Eiver 



Surinam 



Cottica 



Brasil 

Amazonas State: 
Manaus 

\illa I'.clla (Tliayer Kxped.) 



I})(i0 



SOUTH AMERICAN TORTOISES 



(hiiticiihitii (oont.) 

Eio Jiirua 
Tefe 

Para Statr: 

Belem (= "Para") 
Fordlandia 

Mavanhdo State : 
Chatao, Rio Gunipi 

Golds State: 
Anapolis 



iiirhoiitirUi (cont.) 



Para Stat( : 

Boloni (="rara") 

Fordlandia 

Sao Mateus 

Maranlido State : 
Barra do Coida, Rio Mearim 

Golds State: 

Bana do Rio Sao Domingo 
Cana Brava 



Esphritu Santo State: 
Rio Doce 



state unknown 

Fia (W. James — Tliayer Exped.) 
Xeberos (Brit. Mus. — purch. 
Higgins) 

Bolivia 



Dcpt. Pando : 

Baracca, Rio Madidi 

JJcpt. Santa Cruz: 
Fortin Caiiada Larga 



Mato Grosso State : 
Maracuju 
Miranda 

Nioac 



P,io Branca Prov.: 

between Frechal and Limao 
on Rio Surumu 

Dlstrlto Federal : 

Recreio de Bandeirantes, 
8. of Rio de Janeiro 



Bolivia 



Dept. Chacn : 



Dept. Santa Cms: 

San Jose de Chiquitos 

Paraguay 

Asuncion 
Rio Paraguay 
Ybabopo 



8 BREVIORA No. 120 

It is easily seen that the localities, more restricted than state 
or department, from which both species are currently recorded, 
are few. Even in these cases it is doubtful whether the two 
species have been taken together in any literal sense : it may well 
be that these place names are merely central points for large 
collecting areas. 

However, the localities for the two forms are so interwoven 
and the forms themselves so distinct that any interpretation 
except that of species distinction appears difficult. In any event, 
the biological situation is an extremely interesting one, and it 
would be wortli while to have precise information on their 
habitats and habits. No detailed information of this sort is at 
present available. Dr. P. E. Vanzolini comments on the Bra- 
silian localities as follows : 

"Some are in deep forest, some in the cerrado. I have collected 
the beast in the cerrado area (Barra do Corda), but it was in 
small wooded spots or gallery forests. Our eastern Goias speci- 
mens are in the same case — in the Blaser collection are several 
typical forest forms. I flew over the area on purpose and found 
out that there are quite extensive wooded areas. 

Localities in loidouhteclly forested areas: All in Amazonas and 
Para, Chatao, Rio Doce, Recreio dos Bandeirantes. 

Localities where I know there is for sior at least gallery forest : 
Every single other ! 

Incidentally, in Maranhao (Barra do Corda) several of my 
specimens were caught in dens (terribly damp and muddy) l)ut 
not by myself. The habit is known by all there.'''' 

It is interesting that in eastern Peru only denticidata is 
known. This w^as reported already by Sclater in 1871 (p. 744) 
from observations by E. Bartlett. The situation in Amazonian 
Colombia appears similar. By contrast, only carbo)taria is known 
from northern Colombia, Paraguay and southern Brasil (Rio 
de Janeiro). 

I have thus far used the names d( iitlcidala and rayh())taria 
without justifying this procedure. A few coiiimciits on the 
nomenclature are required. 

Tcstudo denticidata, was described by Linnaeus in tlic 12tli 
edition of the Sjf sterna Natura (1766) on the basis of a sjieci- 
men in tlie Museum de Geer. The brief description woubl not 



1960 sorrii .\.mi:ki(AN tortoises 9 

suffice foi- idcnt itical ion if Scliocjjff liad not piiblislKMl in 1792 
a fifi'iirc of a spccimt'ii wliirli may well lie the one citcil hy 
Linnaeus. Aceoi'diuu' to Selioepff (p. 140), tlie Museum de (ieer 
had had not one l)ut two speeinieus. one of which, at the lime 
of Sc'lioepfl"s wi'itiu>i\ was h)catt'd in Stockholm and the othei' 
in llpsala. The latter specimen is that figured !)>• Schoei)ff ( in 
color, in some editions). Selioepff compares this specimen care- 
fully with Tjinnaeus' description and considers it to be TJnnaeus' 
type; Andersson (1900), re-examining the specimen, lias agreed 
with Sehoepff. 

Schoeptf's figure (plate 28), in spite of a certain ei-udity, is 
unquestionably the species here called denticulato. The denticu- 
lations that gave the species its name are clearly shown and the 
colored editions show the characteristic yellow brown of juveniles 
of this species. Thus there can be no doubt about the name of 
this form. 

The next name proposed was Testudo fabiilnta Walbaum 
1782. No figure exists, but the description of the color of the 
shell as "castaneo et sulphureo'' in the Latin text (p. 122), or 
"castanien brauner und hellgelber Parbe" in the (4erman text 
(p. 75), sufficiently identifies this as the brown form and thus 
a synonym of dcnticulata Linnaeus. 

Testudo tessellata Schneider 1792 is the next name available. 
It is, however, a composite, based partly on an older description 
which has always been cited in the synonymy of carhoiiana and 
partly on a description and figure which as clearly ajiply to 
doiticulata. Schneider's detailed description is taken from the 
figure and it is best to assign his proposed name to this concept 
and thus to the synonymy of denticnlata. 

Four names w^ere proposed by Spix (1824), all illustrated hy 
adequate figures. Three — hercuJcs, scidpta, and vauado — are 
clearly difit'ering sizes and minor color varieties of the brown 
tortoise of northern South America. The fourth — carhonaria 
— is clearly the black and yellow species of this jiaper. 

The adjoining plates of sculpta and carhomiria show, in addi- 
tion to the difference in coloration (here at its most extreme 
because an essentially unicolor young doificidafa is ])ictured), 
the characteristic dift'erences in humeral-femoral ratio and in the 
relation of inguinal to femoral. Carhonaria Spix is unmistakably 



10 BREVIORA No. 120 

tlio name to Itc niijilicd to tlie second species in nortlicni Sonth 
America. 

T. hoiei Wag-ler 1833 is a later name and cannot disturb the 
nomenclature here adopted. Wag'ler's plate clearly identifies 
his s])eci('s as carhoiuiria. Once again the hlacU and yellow of 
I he dorsal shell is a\cI1 shown and on the plastron the diagnostic 
linnici'al IVmofal ratio and inLiuinal jiattern. 

Other Neotropical Species 

The tortoises of South America and the Galapagos form a 
natural group for Avhicli the subgenus Chdonoidis of the genus 
Gcoclidoiic^ may be employed. There are four living species: 

G(()ch<l()))f dcnticuhita Linnaeus, G. carhonaria Spix, G. chil- 
(iisis (iray. G. cUphantopus Harlan. 

The following- key will aid in the se])aration of these sjiecies : 

1. Dorsnl colcir iniifonii Ijrown or Itliick, or brown with va^iu'ly l)Ouiided 
orange areolae. Humeral median suture longer than femoial - 
Dorsal color black and yellow, the areolae bright yellow sharply de- 
lineated. Femoral median suture longer than humeral. 

Ct. rarbonaria Spix. 

Northern South America south to Paraguay and 

Rio de Janeiro but not Amazonian Peru. 

2. Inguinal narrow antero])osteriorly with a conspicuous contact with 
the femoral. Shell rounded elliptical in adults, not denticulate in 

hatchlings 3 

Inguinal smaller, nioic tiiaugular, not conspic)iousl.\- in cinilacl with 
femoral in straight ventral view. Shell elongate elli|itical in adults, 
denticulate in hatchlings. (i. (Iciiiiculaia Linnaeus. 

Northern South America except northern ('i)loniliia, 
south to Bolivia but not Paraguay, and to Es])iritu 
Santo but not Rio de Janeiro. 

3. Size snuill (to 220 mm.), (iular region biliil, shell margin subsi'iiate. 

G. cliilensi.s (ii-ay. 

Paraguay, S. Brasil and N. Aigentiiin. 
Size giant (to more than 1 m.). Gular region liuncate, slidl margin 
cntiir. (1. chplinnlopus llailan. 

( ialapagos Islands. 

1 l<\)r (hf use of acochclone ralhcr than Tcslinio sit LovitIiI-c mihI Williams, 
I'.ir.T. \>]>. 1^1 1 lilO. 



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12 BREVIORA No. 120 

Acknowledgments. T am indebted for assistance and many 
courtesies during- my visits to their institutions to Miss Alice 
Grandison and Mr. J. C. Battersby of the British Museum 
(Natural History), Dr. Robert Mertens of the Senckenberg 
Museum, Dr. Walter Hellmich of the Zoologische Staatssamm- 
lung des Bayerischen Staates ; Mr. C. M. Bogert and Theresa A. 
Curtin of the American Museum of Natural History, Dr. Robert 
Inger and Mr. Hymen Marx of the Chicago Natural History 
Museum, Dr. J. Bohlke of the Philadelphia Academy of Natural 
Sciences, and Dr. Doris Cochran of the United States National 
Museum. Dr. Paulo Vanzolini of the Departamento de Zoologia, 
Sao Paulo, Brasil, has permitted me to examine all the tortoises 
in the collection of that institution and has assisted me in other 
ways. Miss Margaret Estey has made dra\ving:s for the present 
paper. Dr. Samuel B. McDowell, Jr., has provided the carapace 
measurements of the two largest G. denticidata at the American 
Museum of Natural History. 

REFERENCES CITED 

Andersson, L. G. 

1900. Catalogue of Linnean type-specimens of Linnaeus 's Eeptilia in 
the Eoyal Museum of Stockholm. Bihang K. Sven. Vet.-Akad. 
Handl., vol. 26, pp. 1-29. 

Bell, Thomas 

1825. A monograph of the Testudinata. London. 
(Pages and plates not numbered.) 

Linnaeus, Carolus 

1766. Systema Naturae, ed. 12, vol. 1 "Holmiae." 532 pp. 

LOVERIDGE, A. AND E. E. WILLIAMS 

1957. Revision of the Africans tortoises and turtles of the suborder 
Cryptodira. Bull. Mus. Comp. Zool., vol. 115, pp. 160-557. 

LUEDERWALT, H. 

1926. Os chelonios brasileiros com a lista dos especies do Museu 
Paulista. Rev. Mus. Paulista, vol. 14, pp. 405-468. 

Schneider, G. 

1792. Beschreibung und Abbildung einer neuen Art von WasserschUd- 
krote nelist Bestimmungen einiger Insher wenig beknnnten 
fremden Arten. Rclirift. Ges. naturf. Freunde Berlin, vol. 10, 
pp. 259-284. 



li'tJO SOrill AMKKICAX TORTOISES 13 

SCHOEPPF, J. D. 

1792. Naturgc'scliirhte der ISc-hiliikroteii. ErlaiifU'ii, IGO i)i). 

SCLATER, P. L. 

1871. Notes oil rare or little known animals now or lately living? in 
the Society's Gardens. Part TIT. Reptiles. Proc. Zool. Soe. 
London, 1871, pp. 743-749. 

SOWERBY, J. DEC. AND EdWARD LeAR 

1872. Tortoises, Terrapins and Turtles. London, pis. 1-lx. 

Spix, J. B. DE 

1824. Aiiimalia nova sive species novae Testudinum et Ranarum. 
"Monachii. " 53 pp. 

Wagler, J. 

1833. Descriptiones et Icones Amphibiorum. Part 2. Plates xiii-xiv. 

Walbaum, J. J. 

1782. Cheloiiosraphia oiler Beschreibuiig einiger Schildkroten iiach 
naturlichen Urbildern. Lubeek and Leipzig, 132 pp. 

Williams, Ernest E. 

1950. Testitdo cubeiisis and the evolution of Western Hemisphere tor- 
toises. Bull. Anier. Mus. Nat. Hist., vol. 95, pp. 1-36. 




I'Jatt' 1. 'I'dp: III, -1(1 aiiil aiitcrioi- r;uai)aee of (!< nclKlonc den I iciilaUi 
juv. Xdic (Iriiticiilal idii of ('(Ig'os of first marginals, elongate prefrontal 
sliiclils ami fidiital sliicid liidkrii up into smaller scales. IJotlom: Head 
and anterior earai)ace of ( iiochrhniv curbunaria juv. Note absenoe of 
dentieulation of edges of first marginals, the short prefrontals and the 
large frontal. F. White phot. 





Plate 2. Dorsal and vcntial views of Geochelonc dmticiihtid Jin . !•'. White 
phot. 





Plate 3. Dorsal and ventral views of Geochelone carhonay'in jnv. F. White 
i)]iot. 



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