(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Breviora"

f ^-1^' 



HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



DOES MOT rf!>/-f II A-rr- 



N. E. WRIGHT 

CAMBRIDGE 38. MASS 
l»»TU«N P08TA0E OUARANT.ED 






BREVIORA 



MUSEUM OF COMPARATIVE ZOOLOGY 

AT 

HARVAED COLLEGE, IN CAMBRIDGE 



Numbers 121-178 
1960-1962 



CAAIBRIDGE, MASS., T.S.A. 
1963 



Edited 
By 

Nelda E. Weight 



CONTENTS 

BREVIORA 

Museum of Comparative Zoology 

Numbers 121-178 

1960 

No. 121. Three new species of Micrathena ( Araneae. Argiopidae) 
from South America. By Arthur M. Chickering. 
11 pp. March 10. 

No. 122. Notes on certain species of Micratkcna (Araneae, Argi- 
opidae) from South America. By Arthur M. Chick- 
ering. 7 pp. March 11. 

No. 123. Alepisaurus hrevirostris, a new species of lancetfish 
from the western North Atlantic. By Robert H. 
Gibbs, Jr. 14 pp. March 14. 

No. 124. Anisian ammonoids from Malaya. By Bernhard Kum- 
mel. 8 pp., 1 pi. March 15. 

No. 125. The luminous organs of Proctoporus (Sauria, Rep- 
tilia) — a re-evaluation. By Willard D. Roth. 12 pp. 
May 27. 

No. 126. Mid-Scythian ammonites from Iwai formation, Japan. 
By Bernhard Kummel and Sumio Sakagami. 11 pp., 
3 pis. June 3. 

No. 127. Notes on the cranial anatomy of Necrolcmiir. By E. L. 
Simons and D. E. Russell. 14 pp. December 19. 

No. 128. Size of endoceroid cephalopods. By Curt Teichert and 
Bernhard Kummel. 7 pp. December 20. 

Xo. 129. Type and type locality of the Gulf Coast spiny softsheli 
turtle, Trionyx spinifer asper (Agassiz). By Robert 
G. Webb. 8 pp., 2 pis. December 21. 

No. 130. The mechanisms of carapacial and plastral hinges in 
chelonians. By R. V. Shah. 15 pp. December 22. 



No. 131. A second record of the fossil rodent Palustrinius Wood. 
By Craig C. Black. 3 pp. December 30. 

No. 132. The status of Sphaerodactylus pictus, with comments 
on the distribution of 8. sputator and S. sahanus. 
By Wayne King. 5 pp. December 30. 

1961 

No. 133. On the generic limits in the family Pilidae (Proso- 
branchia: Mollusca). By Edward H. Michelson. 10 
pp. February 27. 

No. 134. Enzymatic constitution of Alsophis saliva and its bio- 
logical implications. By George Hegeman. 8 pp. 
February 28. 

No. 135. Notes on Hispaniolan herpetology. 2. A review of the 
Anolis semilineatus group with the description of 
A7iolis cochranae, new species. By Ernest E. Wil- 
liams and A. Stanley Rand. 11 pp. April 7. 

No. 136. Notes on Hispaniolan herpetology. 3. The evolution 
and relationships of the Anolis semilineatus group. 
By Ernest E. Williams. 8 pp. April 8. 

No. 137. Notes on Hispaniolan herpetology. 4. Anolis kooprnani, 
new species, from the southwestern peninsula of 
Haiti. By A. Stanley Rand. 4 pp. April 10. 

No. 138. Pfeiffer's unfigured species of Strophocheilus (Megal- 
ohnlhnus). By T. E. Crowley and T. Pain. 8 pp., 
2 pis. June 14. 

No. 139. A new species of Sphaerodactylus from northern Haiti. 
By James D. Lazell. 5 pp. June 15. 

No. 140. A preliminary review of the Nearctic species of Siero- 
lomorpha (Hymenoptera). By Howard E. Evans. 
12 pp. June 27. 

No. 141. Three new toads from South America: Bufo manicor- 
cnsis, Bufo spinulosus altiperuvianu^ and Bufo 
quechua. By Jose M. Gallardo. 8 pp., 3 pis. June 28. 

No. 142. Australian carabid beetles VI. The tropical and some 
subtropical species of Pamborus, Mystropomus, and 
Nurus. By P. J. Darlington, Jr. 13 pp. June 30. 



No. 143. Miocene lizards from Colombia, South America. By 
Richard Estes. 11 pp. August 20. 

No. 144. A large ophiacodont pelyeosaur from the Pennsyl- 
vanian of the Pittsburgh region. By Alfred Sher- 
wood Romer. 7 pp. August 21. 

No. 145. A new species of the cetomimid genus Gyrinomimus 
from the Gulf of Mexico. By Henry B. Bigelow. 
2 pp. September 5. 

No. 146. New rodents from the early Miocene deposits of Sixty- 
Six Mountain, Wyoming. By Craig C. Black. 7 pp. 
December 14. 

No. 147. Australian carabid beetles VIII. Leiradira, especially 
the tropical species. By P. J. Darlington, Jr. 12 pp. 
December 15. 

No. 148. Australian carabid beetles IX. The tropical Noton- 
omus. By P. J. Darlington, Jr. 14 pp. December 18. 

No. 149. A preliminary study of the Silurian ceratiocaridids 
(Crustacea: Phyllocarida) of Lesmahagow, Scot- 
land. By W. D. Ian Rolfe. 9 pp. December 19. 



1962 

No. 150. The genus Bethylus in North America (Hymenoptera : 
Bethylidae). By Howard B. Evans. 12 pp. Jan- 
uary 5. 

No. 151. A new phyllocarid crustacean from the Upper Devon- 
ian of Ohio. By W. D. Ian Rolfe. 7 pp., 1 pi. 
January 12. 

No. 152. New Australian dacetine ants of the genera Meso- 
struma Brown and Codiomyrmex Wheeler (Hymen- 
optera-Formieidae). By Robert W. Taylor. 10 pp. 
January 15. 

No. 153. Anolis scriptiis Garman 1887, an earlier name for 
Anolis Icucophaeus Garman 1888. By A. Stanley 
Rand. 5 pp. February 15. 

No. 154. Notes on Hispaniolan herpetology. 5. The natural his- 
tory of three sympatric species of Anolis. By A. S. 
Rand. 15 pp. April 4. 



No. 155. Notes on Hispaniolan herpetology. 6. The giant ancles. 
By Ernest E. Williams. 15 pp. April 12. 

No. 156. The fossiliferous Triassic deposits of Ischigualasto, 
Argentina, and preliminary description of Ischigual- 
astia, a new genus of dicynodont. By Alfred Sher- 
wood Romer and C. Barry Cox. 9 pp. April 13. 

No. 157. A rhachitomous amphibian, Spathicephalus, from the 
Mississippian of Nova Scotia. By Donald Baird. 9 
pp., 1 pi. May 28. 

No. 158. A fossil gerrhosaur from the Miocene of Kenya (Rep- 
tilia: Cordylidae). By Richard Estes. 10 pp., 1 pi. 
May 29. 

No. 159. Age in a small sample of bliiefish {Pomatomus salta- 
trix (Linnaeus)). By Richard H. Backus. 4 pp. 
May 31. 

No. 160. Tvv'O new arthropod carapaces from the Burgess shale 
(Middle Cambrian) of Canada. By W. D. Ian Rolfe. 
9 pp., 1 pi. June 12. 



No. 161. 



A comparative study of the respiratory muscles in 
Chelonia. By R. V. Shah. 16 pp. July 16. 

No. 162. Australian carabid beetles X. Bemhidion. By P. J. 
Darlington, Jr. 12 pp. July 25. 

No. 163. New worm-lizards { An cylo cranium and Amphisbaena) 
from southeastern Tanganyika Territory. By Arthur 
Loveridge. 6 pp. July 26. 

No. 164. Notes on the herpetology of Hispaniola. 7. New ma- 
terial of two poorly known anoles : Anolis monticola 
Shreve and Anolis christophei Williams. By Ernest 
E. Williams. 11 pp. August 22. 

No. 165. An extinct solenodontid insectivore from Hispaniola. 
By Brjmn Patterson. 11 pp. August 22. 

No. 166. Lectotypt's of species of Ogcocephalidae selected from 
syntypes in the Museum of Comparative Zoology. 
By Margaret G. Bradbury. 4 pp. September 5. 

No. 167. BatJiyclupea schroederi, a new bathyclupeid fish from 
the western tropical Atlantic. By Myvanwy M. Dick. 
4 pp. September 5. 



No. 168. Two new species of fossil talpid insectivores. By 
Katherine M. Reed. 6 pp., 1 pi. September 7. 

No. 169. New records of inshore fishes from the Atlantic coast 
of Panama. By Ira Rubinolf and Roberta W. Rubi- 
noff. 7 pp. October 15. 

No. 170. The brain of the emu {Droniacns nouaehollandiae, 
Lath). I. Gross anatomy of the brain and pineal 
body. By Stanley Cobb and Tilly Edinger. 14 pp., 
4 pis. November 16. 

No. 171. Notes on amphisbaenids (Ampliisbaenia; Reptilia i , 
6. Redescription and range extension of Aniphishaena 
spurrelli Boulenger. By Carl Cans. 8 pp., 3 pis. 
December 14. 

No. 172. A new species of the rodent Pipcstoneomys from the 
Oligocene of Nebraska. By Raymond Alf. 7 pp. 
December 14. 

No. 173. New species of land molluslvs from the Republica 
Dominicana. By William J. Clench. 5 pp., 1 pi. 
December 24. 

No. 174. A new arctocyonid from the Paleocene of Wyoming. 
By Bryan Patterson and Paul 0. McGrew. 10 pp. 
December 24. 

No. 175. A picrodontid insectivore (?) from the Paleocene of 
Wyoming. By Paul 0. McGrew and Bryan Patter- 
son. 9 pp. December 24. 

No. 176. On the races of Kinixys hclliana Gray. By R. F. 
Laurent. 6 pp. December 27. 

No. 177. Rhipidistian classification in relation to the origin of 
the tetrapods. By Keith S. Thomson. H pp., 1 pi. 
December 27. 

No. 178. On a new species of the earthworm genus Trigaster 
Benham 1886 (Octochaetidae). By G. E. Gates. 4 
pp. December 27. 



INDEX OF AUTHORS 
BREVIORA 



T 



MUSEUM OF COMPARATIVE ZOOLOGY 
Numbers 121-178 

1960-62 

No. 

Alf, Raymond 172 

Backus, Richard H 159 

Baird, Donald 157 

BiGELOw, Henry B 145 

Black. Craig C 131, 146 

Bradbury, Margaret G 166 

Chickering. Arthur M 121, 122 

Clench, William J 173 

Cobb, Stanley 170 

Cox, C. Barry 156 

Crowley. T. E 138 

Darlington, P. J., Jk 142, 147, 148, 162 

Dick, Myvanwy M 167 

Edinger, Tilly 170 

Estes, Richard 143, 158 

Evans, Howard E 140, 150 

Gallardo, Jose M 141 

Gans, Carl 171 

Gates, G. E 178 



GiBBs, Robert II 123 

Hegeman, George 134 

King, Wayne 132 

Kummel. Bernhard 124, 126, 128 

Laurent, R. F 176 

Lazell, James D 139 

Loveridge, Arthur 163 

McGrew. Paul 174, 175 

MicHELSON, Edward H 133 

Pain, T 138 

Patterson, Bryan 165, 174, 175 

Rand, A. Stanley 135, 137, 153, 154 

Reed, Katherine M 168 

RoLFE, W. D. Tan 149, 151, 160 

RoMER, Alfred Sherwood 144, 156 

Roth, Willard D 125 

RuBiNOFF, Ira 169 

RuBiNOFF, Roberta W 169 

Russell, D. E 127 

Sakagami, Sumio 126 

Shah, R. V 130, 161 

Simons, E. L 127 

Taylor, Robert W 152 

Teichert, Curt 128 

Thomson, Keith S 177 

Webb, Robert G 129 

Williams, Ernest E 135, 136, 155, 164 



BREVIORA 



Muiseiiiim of Comparative Zoology 



Cambridge, Mass. March 10, 1960 Number 121 



THREE NEW SPECIES OF MICRATHENA 

(ARANEAE, ARGIOPIDAE) 

FROM SOUTH AMERICA 

Bv Arthur M. Chickering 

Albion College, Albion, Michigan 

During the summer of 1959 I had the privilege of examining 
the South American Mierathenae in the Museum of Comparative 
Zoology at Harvard College. Among the specimens in this col- 
lection I found representatives of what I am compelled to regard 
as new species. I am describing these in this brief paper under 
the names : Micrathena hamata sp. nov. ; M. lata sp. nov. ; and 
31. shealsi sp. nov. I have also added a few notes together with 
figures illustrating features of the epigynum of M. fissispina 
(C. L. Koch), 1836, with the hope that they will aid somewhat 
in a more precise identification of this species. The types of the 
new species are deposited in the Museum of Comparative Zo- 
ologv. 

It is with pleasure and a deep sense of gratitude that I again 
acknowledge my indebtedness to members of the staff of the Mu- 
seum of Comparative Zoology at Harvard College for their con- 
tinued encouragement over a period of more than twenty-five 
years. Persons now active on the staff of the museum and chiefly 
responsible for this encouragement may be specifically named as 
follows: Dr. A. S. Romer, Director; Dr. P. J. Darlington, Jr., 
Curator of Insects; Dr. Herbert W. Levi, Associate Curator of 
Arachnology; Miss Nelda E. Wright, Editor of Publications. 
Several other members of the museum staff have also greatly 
aided me by providing privileges of the laboratories, collections, 
and library. 



BREVIORA 



No. 121 



MiCRATHENA FISSISPTNA (C. L. Koch), 1836 

(Figures 1-3) 

Acrosoma fissispina C. L. Koeh, 1836 
Plectnna fissispina Walckenaer, 1841 
M. fissispina Simon, 1895 
M. fissispina Reimoser, 1917 
M. fissispina Roewer, 1942 
M. fissispina Bonnet, 1957 

There are three mature females in the Nathan Banks Collection 
in the Museum of Comparative Zoology. These are simply la- 
belled as having come from Brazil with no dates of collection 




External Anatomy of Micrathena 
Figures 1-3. M. fissispina (C. L. Koch). 

Figs. 1-3. Epigynum: from below; profile, right side; posterior surface, 
respectively. 



assigned. The abdominal spination is a fairly reliable feature 
for identification, but the epigynum is also important. Since this 
organ appears somewhat different than represented in Reimoser's 
(1917) figures I am offering three more drawings with the hope 
that they will help others to a more exact identification of this 
species. The male remains unknown. 



1960 



NEW SPECIES OF MICRATHENA 



3 



MiCRATHENA HAMATA sp. nov. 

(Figures 4-7) 

The two niature males treated in tliis part of this paper were 
found filed in the Banks Collection in the Museum of Compara- 
tive Zoology with females considered to belong to M. petersi 
(Tacz.). The females are probably immature specimens of M. 
sc.rspinosa (Hahn). The males appear to be new to science and. 
consequently, one has been selected as the holotype and is here- 
with described in accord with my usual procedure. 





External Anatomy of Micrathena 
Figures 4-7. M. hamata sp. nov. 

Fig. 4. Body of male, dorsal view. 

Fig. 5. Left palpal tarsus and tibia. 

Fig. 6. Palpal tarsal hook ; nearly retrolateral view. 

Fig. 7. Ibid., a different view. 



4 BREVIORA No. 121 

Male holotype. Total length 4.68 mm. Carapace 1.92 mm. lonj;, 
1.105 mm. wide opposite second coxae where it is widest; .4r)5 
mm. tall ; nearly level from behind PME to beginning of posterior 
declivity. Median thoracic fovea a shallow longitudinal depres- 
sion. Dorso-lateral foveae lacking. 

Eyes. Eight in two rows as usual in the genus; lateral ocular 
tubercles weakly developed; median ocular tubercles bearing 
ME quite pronounced. Viewed from above, both rows strongly 
recurved; viewed from in front, anterior row nearly straight, 
posterior row moderately procurved; central ocular quadrangle 
wider behind than in front in ratio of 22 : 17, about as long as 
wide l)ehind. Ratio of eyes AME : ALE : PME : PLE = 6 : 6.5 
: 8 : 5.5 (long diameter used when there are ditferences). AME 
separated from one another by four-thirds of their diameter, 
from ALE by nearly five-halves of their diameter. PME sepa- 
rated from one another by nearly three-halves of their diameter, 
from PLE by about three-halves of their diameter. Laterals 
separated from one another by two-thirds the diameter of AME. 
Clypeus strongly receding; height equal to about twice the di- 
ameter of AME. 

Chelicerae, Maxillae, and Lip. Apparently quite normal for 
males in the genus. Fragility of specimen precludes examination 
of fang groove for marginal teeth. 

Sternum. Very rugulose ; with a marked transverse groove 
between third and fourth coxae ; fourth coxae nearly touching. 

Legs. 4123. Width of first patella at "knee" .il913 mm., 
tibial index of first leg 11. Width of fourth patella at "knee" 
.12996 mm., tibial index of fourth leg 13. 

(All measurements in millimeters) 





Femora 


Patellae 


Tibiae 


Metatarsi 


Tarsi 


Totals 


1. 


.990 


.374 


.748 


.594 


.418 


3.124 


2. 


1.012 


.396 


.660 


.5.50 


.396 


3.014 


3. 


.704 


.264 


.396 


.330 


.352 


2.046 


4. 


1.078 


.330 


.706 


.660 


1.450 


3.224 


Palp 


.400 


.220 


.330 




2.682 


1.632 



1 Estimated because of loss of both fourth tarsi. 

2 Including tarsal hook. 



1960 NEW SPECIES OF MICRATIIENA 5 

There is no ventral hook on first coxa nor any corresponding 
proximal, prolateral femoral ridge and groove on the second 
feninr. There also seems to be a complete lack of modified spines 
on the legs. 

Palp. Essential features shown in Figures 5-7. The tarsal hook 
and tibia appear to be quite distinctive. 

Abdomen. General shape as shown in Figure -i. It seems high- 
ly probable that the female of the species wull be found to have 
a series of prominent paired spines, remains of which seem to 
be present in the male. 

Color in alcohol. Carapace a nearly uniform rich reddish 
brown. Sternum somewhat lighter. Legs and mouth parts with 
various shades of brown. Abdomen: Dorsum yellowish along 
each dorso-lateral margin; broadly and irregularly brownish in 
the middle with a central lighter irregular stripe and some indi- 
cation of reduced black spots; venter irregularly brownish, yel- 
lowish and nearly black. 

Type locality. Holotype male and one paratype male in the 
Nathan Banks Collection from Para, Brazil. No date of collection 
is given. The female is unlaiown. 

MiCRATHENA LATA Sp. nOV. 

(Figures 8-12) 

The specimen described below was filed in the Reimoser Col- 
lection in the Museum of Comparative Zoology at Harvard Col- 
lege as M. digitata (C. L. Koch). This is obviously an error and. 
since I can find no record of it in the literature I am compelled 
to regard it as new and am describing it as such. 

Female holotype. Total length from anterior margin of base 
of ehelieerae to posterior border of abdomen 7.15 mm. Width of 
abdomen at base of anterior spines 2.925 mm.; width between 
tips of larger posterior spines 13.325 mm. Carapace smooth; 
with median thoracic fovea a well defined pit; not markedly 
raised behind median fovea ; with no dorso-lateral foveae. 

Eyes. Eight in two rows as usual in the genus. Viewed from 
above, both rows moderately recurved; viewed from in front, 
anterior row nearly straight, posterior row gently procurved. 
Central ocular quadrangle wider behind than in front in ratio 
of about 3 : 2 ; wider behind than long in ratio of 9 : 7. Ratio 



BREVIORA 



No. 121 



of eyes AME : ALE : PME : PLE = 8 : 8 : 10 : 8.5. AME 
separated from one another by five-fourths of their diameter, 
from ALE by six times their diameter. PME separated from 
one another by three-halves of their diameter, from PLE by 
nearly six times their diameter. Laterals separated from one 
another by three-eig-hths of the diameter of ALE. Height of 
elypeus equal to about seven-fourths of the diameter of AME. 

Chelicerae. Normal to genus ; unable to observe teeth along 
fang groove because of fragility of holotype. 




External Anatomy of Micrathena 
Figures 8-10, M. lata sp. nov. 

Fig. 8. Outline of body of female holotype; dorsal view. 
Figs. 9, 10. Epigynum; profile, right side and posterior surface, re- 
spectively. 



Lip and Maxillae. Normal to genus; details considered unim- 
portant in the description. 

Sternum. Extended ; with deep lateral notches and with six 
exaggerated marginal tubercles; with posterior end extended 
between fourth coxae as a prominent tubercle nearly one-third 
as long as entire sternum. 

Legs and Spines. Only two legs remain entire, hence details 
not recorded. In general quite normal to genus. True spines 



1960 



NEW SPECIES OF MICRATHENA 



rare ; the usual numerous setigerous tubercles moderately well 
developed. 

Abdomen. Extraordinarily broadened posteriorly ; with a pair 
of nearly straight anterior marginal spines extended nearly to 
PME ; with a pair of very small lateral marginal spines ; tbo 
posterior end is broadly bifurcated and each fork is subdivided 
into two spines thus making a total of eight. 

Epigynum. General features shown in Figures 9, 10, and 12. 




External Anatomy of ilicraihena 
Figures 11 and 12, M. Juta sp. uov. 

Fig. 11. Posterior end of female holotypL'. 
Fig. 12. Epigynum, from below. 



Color in alcohol. Light reddish brown with dark streaks and 
irregular spots. 

Tyi^e locality. Holotype female from Theresapolis, Brazil, with 
no date of collection given. There are no paratypes and the male 
is unknown. 



BREVIORA 



No. 121 



MiCRATHENA SHEALSI Sp. nOV. 

(Figures 13-17) 

Female holotype. Total length from AME to tip of posterior 
spines 9.23 mm. Carapace 2.79 mm. long ; 2.08 mm. wide opposite 
second coxae where it is widest; about .98 nnn. tall behind well 




External Anatomy of Micrathena 
Figures 13-17, M. sheaJsi sp. nov. 

Fig. 13. Outline of female holotj'pe, dorsal view. 

Fig. 14. Eight lateral side of abdunien to show spines on bifurcation. 
Figs. 15-17. Epigynum : from lielow, riglit lateral side, and from posterior 
view, respectively. 



defined median fovea where it is strongly gibbous; with three 
pairs of dorso-lateral foveae as represented in Figure 13. 

Eyes. Eight in two rows as usual ; viewed from above, both 
rows moderately recurved ; viewed from in front, both rows 



Ii)(i0 NEW SPECIES OF MICRATHENA 9 

fjently procurved, measured by centers. Central ocular quad- 
rangle wider behind than in front in ratio of about 8 : 7, only 
slio'htly wider behind than long. AME separated from one 
another by their diameter, from ALE by nearly 3.5 times their 
diameter. PME separated from one another by their diameter, 
from PLE by three times their diameter. Ratio of eyes AME : 
ALE : PME" : PLE = 10 : 10 : 12 : 7 (long diameters used when 
differences exist) . Laterals separated from one another by nearly 
the radius of iVLE. Height of elypeus equal to nearly 1.2 times 
the diameter of AME. 

Chelirerae. In general, normal to the genus; promargin of 
fang groove with three teeth, the middle one bidental ; retro- 
margin with three large teeth. 

Maxillae. Apparently completely typical of the genus in all 
observed features. 

Lip. Wider than long in ratio of about 4:3; transversely 
grooved in basal third. Sternal suture procurved. 

Sternum. Elongated scutiform; moderately convex; only with 
well developed antero-lateral tubercles: not continued between 
fourth coxae ; sparsely supplied with moderately long stiff 
bristles. 

Legs. 4123. Width of first patella at ''knee" .325 mm., tibial 
index of first leg 11. Width of fourth patella at "knee" .303 
nun., tibial index of fourth leg 11. 

(All measurements in millinieters) 





Femora 


Patellae 


Tibiae 


Metatarsi 


Tarsi 


Totals 


1. 


2.860 


.990 


2.100 


1.950 


.910 


8.810 


o 


2.600 


.975 


1.820 


1.755 


.845 


7.995 


3. 


1.625 


.650 


.980 


1.040 


.715 


5.010 


4. 


3.250 


.780 


2.015 


2.080 


.910 


9.035 



Spines on legs apparentlj^ unnoteworthy ; many are lost and scars 
are difficult to locate with certainty. 

Abdomen. General features essentially as shown in Figures 
13 and 14. Moderately flattened. With no anterior marginal 
spines ; with a pair of lateral dorsal spines of moderate length ; 
bifurcated posteriorly with each fork subdivided into two spines 
with the larger below. 



10 BREVIORA No. 121 

Epigynum. Unlike that seen in any other species; anterior 
border of modified portion a granulated rim ; features essentially 
as shown in Figures 15-17. 

Color in alcohol. Legs and mouth parts a dull reddish brown, 
lighter beneath. Sternum light brownish. Carapace with a 
brownish central stripe and a broad dark brown stripe on each 
side (both represented by stippling in Figure 13) ; remainder of 
central region yellowish. Abdomen : irregularly yellowish white 
dorsally with nearly black margins ; venter light yellowish from 
genital furrow to base and lateral sides of cone surrounding 
spinnerets ; the cone with a nearly black circular ring ; remainder 
of venter irregularly dark brown or black with yellowish spots 
and streaks. Probably some loss of coloration from long preser- 
vation. 

Type locality. The holotype was simply labelled : Argentine, 
Sunchal, with no date of collection given. Apparently Cockerell 
was the collector. The species is named in honor of Dr. J. G. 
Sheals, Department of Zoologj^ British Museum (Natural His- 
tory) . 

BIBLIOGRAPHY 

Bonnet, Pierre. 

1957. Bibliographia Araneorum. Tome II, 3nie partie. T.ps Artisans 
de I'Imprimerie Douladoure, Toulouse, France. 

Cambridge, 0. P. and F. P. Cambridge. 

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

Kfyserling, Graf Eugen von. 

1892. Die Spinnen Ameiikas. Epeiridae. Verlag von Bauer & Raspe, 
Niirnberg, Germany. 

Koch, C. L. 

1836. Die Arachniden. Niirnberg, vol. Ill, pp. 1-119. 

Reimoser, Eduard, 

1917. Die Spinnengattung Micrathena. Verh. Zool. Bot. Ges. Wien, 
vol. 67, pp. 73-160, pis. 1-9, figs. 1-31. 



I960 NEW SPECIES OF MICRATHENA 11 

ROEWER, C. Fr. 

1942. Katalog der Araneae. Vol. 1. Kommissions Verlaj,' von 
"Natura." Bremen. 

Simon, Eugene. 

1892-1903. Histoire Naturelle dcs Araignees. Deuxieme Edition. 2 
Vols. Librarie Eneyclopedique de Roret, Paris. 

Walckenaer, Ch. a. 

1841. Histoire Naturelle des Insectes. Apteres. Paris, vol. II. 



/ 



BREVIORA 



Mmseiiiinti of Comparsitive Zoology 



Cambridge, Mass. March 11, 1960 Number 122 

NOTES ON CERTAIN SPECIES OP MICRATHENA 
(ARANEAE, ARGIOPIDAE) FROM SOUTH AMERICA 

By Arthur M. Chickering 

Albion College, Albion, Michigan 

During a period of work in the Museum National d'Histoire 
Naturelle in Paris during the summer of 1958 Dr. Herbert W. 
Levi, Associate Curator of Arachnology in the Museum of Com- 
parative Zoology at Harvard College, examined types of nine 
species of Micrathena Sundevall 1833, all originally described 
from South America by the great arachnologist, Eugene Simon. 
All of these species are poorly known and most of them have not 
appeared in collections since the originals were studied by their 
author. All were briefly described in 1896 ; four were mentioned 
in 1895 and accompanied by five simple figures. During the 
examination of the types mentioned Dr. Levi made free-hand 
drawings of the dorsal surface of the abdomens to show general 
form and spination. He also made careful drawings of the exter- 
nal genitalia when the latter were available ; these were made 
with the use of a reticule with squares. All of the drawings made 
by Dr. Levi were turned over to the author to use as he saw fit 
in connection with his study of the genus. The figures appearing 
in this paper were made directly from Dr. Levi's original pencil 
drawings. The outline figures of abdomens are freehand copies 
with enlargement; the drawings of genitalia were made with 
tracing paper directly from Dr. Levi's originals. It has seemed 
worth while to present these data, thus obtained, with the hope 
that they will be of some help to others who may continue the 
study of this most interesting genus. 



BREVIORA 



No. 122 



MiCRATHENA AcicuLATA Simon, 1897 
(Fiorure 1) 

M. acicvlata Petrunkevitch, 1911 
M. aciculata Roewer, 1942 
M. aciculata Bonnet, 1957 

The type is an immature female from Venezuela. Apparently 
the species has not been reported in collections since the original 
was taken. The general form of the abdomen with its spination, 




External Anatomy of Micrathena 

Figure 1, M. aciculata, abdomen, dorsal view. 

Figures 2, 3. M. gaujoni, abdomen, dorsal view and epigynum, respectively. 

Figures 4, 5. M. hamifera, abdomen, dorsal view and epigynum, respectively. 



seen in dorsal view, is shown in Figure 1. The stippled area is 
black and the posterolateral corners are white. The male is 
unknown. 

Micrathena gaujoni Simon, 1897 
(Figures 2, 3) 

M. gaujoni Petrunkevitch, 1911 
M. gaujoni Reimoser, 1917 



1960 CERTAIN SPECIES OF MICRATHENA 3 

.1/. gaujoni Roewer, 1942 
M. gaujoni Bonnet, 1957 

Simon stated that the type was 8.7 mm. long and similai* to 
J/, fissispina (C. Koch), but this hardly seems correct. There 
are four ])airs of spines, tli*' second pair the longest (Fig. 2). 
The epigynum has a pair of depressions directed posteriorly be- 
neath an overhanging rim (Fig. 3). The type is from Ecuador 
and the male is unknown. 

MiCRATHENA HAMIFERA SilllOll, 1897 

( Figures 4, 5 ) 

The length of the holotype was given as 9 mm. The general 
form of the abdomen with its spination is shown in Figure 4. 
The dorsal surface is like white enamel in general appearance. 
Figure 5 shows the form of the epigynum from "slightly be- 
hind"; just posterior to the curved boundary there is a large 
' ' sclerotized knob. ' ' The female is known only from Peru and the 
male is still unknown. 

^MiCRATiTEXA niBELLis Simoii. 1895 
( Figure 6 ) 

.1/. imbeUis Simon, 1897 

M. imhclUs Petrunkeviteh, 1911 

M. imhellis Eeimoser, 1917 

J/, imhellis Roewer, 1942 

M. imhellis Bonnet, 1957 

Simon (1895) included a tigure showing the right side of the 
abdomen with no spines. Reimoser (1917) just mentioned the 
species and did not include it in his further treatment of the 
genus. The general appearance of the dorsal surface of the 
abdomen is shown in Figure 6; the stippled areas in the figure 
are black in the type. Dr. Levi has determined that the type is 
an immature female from Venezuela. The male is unknown. 

MiCRATHENA PERLATA Simon, 1895 
(Figures 7, 8) 

M. perlata Simon, 1897 

M. perlata Petrunkeviteh, 1911 

M. perlata Reimoser, 1917 



BREVIORA 



No. 122 



M. perlata Eoewer, 1942 
.1/. perlata Bonnet, 1957 

The length of tlie female type is given as 6 mm. Dr. Levi has 
found that it also is immature. The general appearance of the 
dorsal surface of the abdomen is shown in Figure 7. The im- 
mature female is accompanied by a male which may or may not 







External Anatomy of Micrathena 

Figure 6. M. imbellis, abdomen, dorsal view. 

Figures 7, 8. M. perlata, abdomen, dorsal view and male palp, respectively. 
Figures 9, 10. .1/. pubescons, abdomen, dorsal view and male palp, respec- 
tively. 



be properly paired with it. Figure 8 shows certain features of 
the palpal tarsus. The specimens are simply labelled "ximazon." 
Simon (1895) stated that the type came from: "Brasilia: S. 
Paulo de Olwenea (de Mathan)." 



1960 CERTAIN SPECIES OF MICRATHENA 5 

MiCRATHENA PUBESCENS Simoii, 1895 
(Figures 9, 10) 

M. i)ubescens Siniou, 1897 

M. pubt'A-cciis Petrunkevitch, 1911 

M. pubescens Reimoser, 1917 

M. pubescens Roewer, 1942 

M. pubescens Bonnet, 1957 

The female type is immature. The abdomen is hairy and sug- 
gests a close relationship with M. furcula (0. P. Cambridge) 
from Central America. There are no true spines but the posterior 
end of the abdomen is somewhat bifurcate. Simon (1895) fur- 
nished figures to show the right side of the abdomen and the 
bifurcate posterior end. The immature female is accompanied 
by a mature male the palpal tarsus of which is shown in one 
position in Figure 10. Caution must always be exercised in 
matching the sexes in this genus and this male should be very 
carefully studied and compared with the growing number of 
different kinds of known males. The specimens are from Matto 
Grosso, Brazil. 

MiCRATHENA PUPA Simon, 1897 
(Figures 11, 12) 

M. pupa Petrunkevitch, 1911 
M. pupa Reimoser, 1917 
M. pupa Roewer, 1942 
il. pupa Bonnet, 1957 

The length of the type is given as 8 mm. The form of the abdo- 
men and its spination are shown in Figure 11 and some of the 
features of the epigynum in Figure 12. The type female is from 
Ecuador. Simon apparently had a male associated with the 
female but Dr. Levi did not find it in the collection. Presumably 
it is lost and the original description does not give what we now 
consider to be the important diagnostic male features. 

MiCRATHENA TOVARENSIS SiuiOU, 1897 

(Figures 13, 14) 

.1/. tucarensisFetrunkevitch, 1911 
M. tovarensis Reimoser, 1917 
M. tovarensis Roewer, 1942 
il. tovarensis Bonnet, 1957 



6 



BREVIORA 



No. V2-2 



The length of the female type is given as 7.8 mm. The general 
appearance of the abdomen and its spines, as seen in dorsal view, 
are shown in Figure 13. The appearance of the epigynum as 
shown in Dr. Levi 's drawing is given in Figure 14. The type is 
from \'enezuela. The male is unknown. 




External Anatomy of Micrathena 

Figures 11, 12. Al. pupa, abdomen, dorsal view and epigynum, respectively. 
Figures 13, 14. If. tovarensis, abdomen, dorsal view and epigj^num, respec- 
tively. 

Figure 15. M. xantliopyga. abdomen, dorsal view. 

Micrathena xaxthopyga Simon, 1895 
(Figure 15) 

M. xanthopyga Simon, 1897 

M. xanthopyga Petrunkevitch, 1911 



1960 CERTAIN SPECIES OF MICRATHENA 7 

M. xanthopyga Reimoser, 1917 
M. xanthopyga Roewer, 1942 
M. xanthopyga Bonnet, 1957 

The type is an immature female whose general appearance in 
dorsal view is shown in Figure 15. Simon (1895) published a 
figure of the type viewed from the right side. This figure shows 
four pairs of spines instead of three, as shown in Dr. Levi's 
drawing. The type is from Venezuela. The male is unknown. 

BIBLIOGRAPHY 

Bonnet, Pierre. 

1957. Bibliographia Araneorum. Tome II (3nie partie:G-M). Les 
Artisans de L 'Imprimerie Douladoure, Toulouse. 

Petrunkevitch, Alexander. 

1911. A Synonymic Index-Catalogue of Spiders of North, Central, and 
South America, etc. Bull. Amer. Mus. Nat. Hist., vol. 29, pp. 
1-809. 

Reimoser, Eduard. 

1917. Die Spinnengattung Micrathena. Yerh. Zool. Bot. Ges. Wien, 
vol. 67, pp. 73-160, pis. 1-9, figs. 1-31. 

ROEWE}R, C. Fr. 

1942. Katalog der Araneae. Yol. 1, Bremen. 

Simon, Eugene. 

1895. Histoire Naturelle des Araignees. Yol. 1, Pts. 3-4. 
1897. Descriptions d'Espeees de I'Ordre des Araneae. Annales Societe 
Entomologique de France, vol. 65, pp. 465-510. 



BREVIORA 

MTuseiaim of Comparative Zoology 



Ca^ibridge, Mass. March 14, 1960 Number 123 



ALEPISAURIS BREYIROSTRIS, A NEW SPECIES OF 

LANCETFISH 
FROM THE WESTERN NORTH ATLANTIC 

By Robert H. Gibbs, Jr. 

Department of Biology, Boston University 

Exploratory fishing for tunas in the western North Atlantic 
by the Fish and Wildlife Service vessel DELAWARE has re- 
vealed the presence of considerable numbers of lancetfishes, oenus 
Alepisaurus. Through the courtesy of Mr. James L. Squire, I 
have been privileged to accompany most of these cruises. On the 
first two, the presence of Alepisaurus was recorded and stomach 
contents sampled, but only two specimens were saved. During 
the third cruise, it became apparent that two morphological types 
were represented, and from then on an attempt was made to 
measure and make counts on all possible specimens, and to pre- 
serve a large sample. 

At first the possibility was entertained that the differences 
might be due to sexual dimorphism. Gonads were therefore 
examined on all preserved specimens and on a large number of 
fresh ones. The appearance was almost exactly the same in all 
specimens, indicating that sexual dimorphism was probably not 
a factor. This has been borne out by histological studies of the 
gonads, which lead me to the rather surprising conclusion that 
both morphological types, which I am now certain represent valid 
species, are hermaphroditic. 

A study of nearly all type specimens and of all original de- 
scriptions leads to the conclusion that all previously described 
Atlantic species are conspecific with A. ferox Lowe. The second 
form is described here. 



BREVIORA 



No. 123 



Alepisaurus brevirostris, sp. nov. 

(Figures 1-2) 

Holotype. U. S. National Museum 186197, 682 mm. in standard 
length when fresh, 684 mm. preserved ; taken on longiine with 
20-fathom buoy lines bv the M/Y DELAWARE at 38° 49' N, 
64° 02' W, on September 13, 1957. 



/,A//^/////^///A. 





Figure 1. Left lateral view of Alepisaurus hrevirostris (top), and A. ferox 
( bottom ). 



Paratypes have been distributed to the Museum of Compara- 
tive Zoology, Academy of Natural Sciences of Philadelphia, 
Cornell University, University of Miami Marine Laboratory, 
Tulane University, Scripps Institution of Oceanography, Stan- 
ford Natural Historv Museum, Museum of Zoology L'niversity 



1*J60 ALEPISAURUS BKEVIROSTKIS 3 

of Michigan, California Academy of Sciences, Britisli ^Museum 
(Natural History), Museum National d'Histoire Naturelle Paris, 
and Museu Municipal do Funchal. 

Diagnosis. A dark-hued species of Alcpisaunis with a grad- 
ually arcuate dorsal fin profile, the dorsal origin Avell forward 
of the rearmost margin of the operculum, a short head (6.5 or 
more in standard length), and a short snout (2.5 or more in 
head). 

Description. Dorsal fin high, originating over the middle of 
the opercle, its longest ray (about number 25-30) about three 
times the greatest depth of the body ; its rays flexible, easily bent 
and broken, not branched, joints difficult to observe, though 
present. Leading dorsal ray thickened, its anterior edge finely 
serrated. Anal fin highest at the second and third rays, the rays 
branched distally. Pectoral fin pointed, its middle rays longest ; 
first ray thickened, serrated anteriorly, unbranched, the rest 
branched and obviously jointed. Pelvic fin also pointed, the 
middle rays longest, the first ray thickened, serrated anteriorly, 
and unbranched, the remaining rays jointed and branched. 
Caudal fin strongly forked, with eight procurrent rays in each 
lobe, ten upper principal rays and nine lowers ; uppermost prin- 
cipal rays considerably elongated in some specimens. 

Snout inclined downward more sharply than the rear of the 
head, its length more than 2.5 times in head length. Nostrils a 
little less than halfway from tip of snout to anterior edge of 
orbit, the anterior opening round, the posterior crescent-shaped. 
Upper and lower jaws subequal. Tapper jaw with a row of many 
small thin teeth on the premaxilla, one or two large fangs on 
anterior palatine, about three smaller fangs on the rear of the 
same bone, followed by about 7-10 low, triangular teeth. Lower 
jaw with an anterior large fang, followed by about 10 small 
caniniform teeth, one to three large fangs, and about 10-15 low, 
triangular teeth. No teeth on poorly developed tongue. Two 
patches of pharyngobranchial teeth. Gill arches with 3-6 upper, 
0-1 middle, 17-23 lower groups of low, spinous rakers, totaling 
23-28 groups. Branchiostegals mostly 7. Eye about 5 in head 
length, with vertical adipose eyelids anteriorly and posteriorly. 
Preopercle smooth. Opercle large, sculptured with lines radiat- 
ing from its antero-dorsal corner ; subopercle also with striae 



4 BREVIORA No. 123 

radiating' from its anterior point ; interopercle apparently absent. 
Interorbital space bonnded by prominent parietal ridg'cs, tlie 
area forming a flat to slightly concave surface, gradually widen- 
ing posteriorly. 

Body elong-ate, its greatest depth, at level of pectoral fins, 
about 12 in standard length. A low lateral keel occupying most 
of the rear half of each side. Lateral-line pores opening along 
the keel and continuing forward l)eyond it ; lateral line on head 
forming prominent supraorbital, suborbital, and preoperculo- 
mandibular systems; supratemporal apparently lacking. Anus 
posterior to pelvic insertion by less than half the length of the 
pelvic fins. 

Fin-ray counts and morpliometric data are given in Tables 1 
and 2. ' 

Coloi-afioii. Body iridescent brownish-black dorsally, becom- 
ing gradually lighter laterally. Region al)Ove lateral line liberal- 
ly sprinkled with both large and small melanophores, many of 
the former ocellated. Below the lateral line, and particularly on 
the belly, many small melanophores present. General coloration 
decidedly dark in comparison with A. ferox. Lateral keel black. 
Dorsal fin membrane iridescent black, often with a horizontal 
row of white spots a short distance above base. Other fins, 
including adipose dorsal, black. Head dark above, becoming 
lighter ventrally. Abdominal cavity marked externally l)y alter- 
nating light and dark bands, the light ones representing strips 
of muscle, between which the dark peritoneal lining shows 
through. 

Visceral Anatomu. Peritoneal lining black. Liver relatively 
small, covering only the most anterior portions of the stomach 
and intestine. Stomach black, highly distensible, forming a long, 
blind sac. Intestine arising at the anterior end of the stomach, 
continuing straight, without bends, to the anus, divided at al)out 
one-third of its length into anterior thick-walled and posterior 
thin-walled portions. Kidneys occupying the entire length of the 
body cavity, lying retroperitoneally along the ventral side of the 
vertebral column. Ureters enter a thin-walled urinary bladder 
which extends about from the level of the pelvic insertion to the 
anus. Gonads consisting of a prominent pair of elongate, con- 
tinuous ovaries, which occupy the posterior third of the body 



1!)()0 ALEPISAFRT^S BREVIROSTRIS 5 

cavity above the intestine, and a pair of thin testes, ahnost in- 
visible, lying in the dorsal groove formed by the two ovaries. The 
dncts of the ovaries, and presumably also those of the testes, join 
the urinary bladder and open by a urogenital pore immediately 
behind the anus. Swinibladder absent. 

Related Species. Other than A. hrevirostris, the only recog- 
nized Atlantic species of the genus Alepisaurus is A. ferox Lowe. 
The two species are distinguishable by many characters. The 
most trenchant ones are shown in Table 3, and may be visualized 
in Figures 1 and 2. In addition to these, many less-perfect ones 
are demonstrable. Characters associated with head length show 
significant differences (snout to dorsal origin, snout to pectoral 
insertion, see Table 2). The pectorals average about one-fifth 
of the standard length in A. hrevirostris and are slightly shorter 
in A. ferox. The eye is relatively larger in A. hrevirostris, doubt- 
less correlated with the shorter snout. Meristic characters show 
consistent modal differences : dorsal rays most commonly 42-45 
in A. hrevirostris, 39-42 in A. ferox; anal raj^s usually 14-15 vs. 
15-17 ; pelvic rays 13-14 vs. 14-15. 

Alepisaurus is common in the Pacific, but at present it is not 
possible to ascertain the species. I have examined seven Pacific 
specimens in the U. S. National IMnseum and find them extremely 
similar to, if not conspeeific with A. ferox (see Table 1). The 
only disconcerting element was the low number of dorsal rays 
in four specimens, which indicates at least some degree of dif- 
ferentiation. I have seen no examples of a form resembling A. 
hrevirostris from the Pacific. 

Young Specimens. Five specimens, 85.0 to 190.5 mm. standard 
length, present a confusing array of characters which defy posi- 
tive identification. Tables 1 and 2 show for these specimens many 
characters within or beyond the ranges displayed by adults of 
both A. hrevirostris and A. ferox. There is obviously a great 
change between 200 and 500 mm. in the relative proportions, the 
caudal end in particular increasing proportionally greatly in this 
time. Presumably the shape of the head also changes, as these 
five are all extreme, even to A. hrevirostris, in having short heads 
and snouts. I am inclined to call them all A. ferox, but with 
considerable doubt. 



BREVIORA 



No. 123 







be 



p 



™ 



=t-l 











K O 

^ o 



be 



OD 
O 



c 
bJD 



F^' 



1960 ALEPISAURUS BREVIROSTRIS 7 

3Iensural Discrepancies. Counts and a few selected measure- 
ments were made at sea on most of the specimens which came 
aboard the M/V DELAWARE. A later check has shown that 
the measurements cannot be used in conjunction with others 
taken on preserved specimens. Among the 17 preserved speci- 
mens of each species used in the descriptions here, were 11 A. 
ferox and 16 A. hrevirostris which were also measured at sea 
when fresh. In these, the measurements of standard length and 
head length were consistently less in preserved specimens. In 
A. ferox, the standard length of one specimen was 0.7 per cent 
greater, the others 1.0-16.2 per cent less, averaging 6.6 per cent 
less. The head lengths were 3.4-6.6 per cent less, averaging 5.1 
per cent. The corresponding figures for A. hrevirostris w'ere : 
standard length 0.3 and 1.1 per cent greater in two, others 2.0- 
13.1 per cent less, averaging 4.5 per cent less ; head length 0.5-2.8 
per cent greater in three, 0.5-9.2 per cent less in 13, averaging 
2.7 per cent less. 

Synonymy. Previous descriptions of Alepisaurus leave much 
to be desired. Eight nominal species have been described. I be- 
lieve all these descriptions refer to a single, perhaps polytypic, 
species. These names are discussed below. 

Alepisaurus ferox Lowe, 1833. The original description was 
made from two specimens from Madeira (Lowe, 1835a). In all 
important characteristics the written description is nearest A. 
ferox as recognized in the present study, but the accompanying 
drawing shows an arcuate dorsal fin profile, an absolute character 
of A. hrevirostris. The matter was further complicated by a 
description and drawing of a third specimen from Madeira 
(Lowe, 1835b) which shows a dorsal fin profile characteristic of 
A. ferox. 

N. B. Marshall has kindly examined two types in the British 
Museum. His observations leave no doubt that both are A. ferox 
as I understand it. In the specimen labeled "TYPES!" (no 
registered number) the standard length is 1125 mm., head length 
(to tip of lower jaw, as upper is damaged) 191 mm. (17%), 
dorsal rays 39 or 40, anal 17, pectorals 14. In the specimen 
labeled "SYNTYPE" (number 1852.9.13.98) the standard 
length is 1225 mm., head length 206 mm. (17%) ; snout 90 mm. 
(44% of head) ; dorsal 41, anal 17, pectorals 15. In both, the 



8 BREVIORA No. 123 

dorsal origin is over or sliglitly Ix'liiud the posterior edge of the 
operculum and no large, oeellated melanophores are present; 
both are larger than any A. hrevirostris I have seen. The name 
Alepisauriis ferox is thus reasonably established for the long- 
snouted species. 

Alcpisaitnis aznreus Valenciennes, 1849. No type is extant, 
but the following parts of the original description strongly sug- 
gest A. ferox: ". . . la dorsale est d'egale hauteur jusque vers 
le trentieme rayon ..." (Cuvier and Valenciennes, 1849 : 531) ; 
dorsal rays 38, anal 16. The pectoral count of 10 is presumably 
in error. The length of 5 feet 3 inches is larger than any known 
A. hrevirostris. The description was based on a specimen from 
the Canary Islands. 

Alepisaurus richardsonii Bleeker, 1855. Based on the descrip- 
tion by Sir -John Richardson (1844) of a head from Van Diemens 
Land. The drawing of the head shows a snout most like A. ferox. 

Alepisaurus alfiveJis Poey, 1861. It is difficult to be certain of 
this description. The anterior rays are all said to be the same 
height, the posterior ones decreasing rapidly ; dorsal rays 40, anal 
17, pectorals 16 ; all most like A. ferox. The pelvic count of 13 is 
presumably in error. Based on a Cuban specimen. 

Alepidosaurus horealis Gill, 1863. Based on a head, dorsal, 
caudal, and pelvic tins from a Pacific specimen. I have examined 
this specimen and find it close to A. ferox except for a dorsal 
count of 35. The pelvic count of 13 is apparently an error. The 
stated snout/head ratio of 2/5 definitely precludes A. hrevi- 
rostris. 

Alepidosaurus serra Gill, 1863. Described from a head, caudal, 
and pelvic fins of a Pacific specimen. I can detect no significant 
differences between Gill's descriptions of this species and of 
A. horealis. He places much emphasis on opercular sculpturing, 
which seems to be quite variable and not a good specific charac- 
ter. Again the pelvic count of 13 is presumed to be an error. The 
distance from eye to snout, stated as 2/5 of head length, excludes 
A. hrevirostris. 

Alepidosaurus poeiji (iill, 1863. Described on the basis of 
drawings (which I have not seen) of the second specimen from 
Cuba mentioned by Poey (1861) in his description of A. altivelis. 



1960 ALEPISAL'KUS BREVIKOSTRIS 9 

Tlic specinuMi was described as liaviii<i tlie fii-st dorsal rays be- 
coming lon<i'er, the fourtli very long, rays 6-24 high and e(iual. 
This suggests A. ferox, as does the dorsal ray count of 41. Poey's 
presumably erroneous pelvic count of 13 is restated. 

AhiJidosaiirus aesciilapius Bean, 1883. I have examined the 
type and find it similar to A. ferox. The dorsal rays could not 
be counted accurately, but Beau (1883) gave 39; anal ra3^s 16, 
both A. ferox characters. The snout length (41 ^o of head length) 
rules out A. hrevirostris with absolute certainty. Described from 
a Pacific specimen. 

Disirihution of Alepisaitrus in the Atlantic. Among the speci- 
mens examined and definitely identified, the most southerlj- was 
one A. ferox from off southern Puerto Rico, now in the Museum 
of the University of Miami Marine Laboratory. Several of the 
same species from the Gulf of Mexico are in the U. S. National 
Museum, and specimens from the Gulf of Maine (La Have Bank 
the most northerly) were seen at the Museum of Comparative 
Zoology. The remaining positively identified specimens of A. 
ferox and all those of A. hrevirostris are from the region of the 
Gulf Stream {scnsu strict o), almost all collected by the M/V 
DELAWARE. Some authors have documented their records w^ell 
enough so that Ahpisannis ferox can be said with certainty to 
be found in the following additional regions: Madeira (Lowe, 
1835a; Maul 1946); Canary Islands (Cuvier and Valenciennes, 
1849) ; Cuba (Poey, 1861)'; coast of North Carolina (Brimley, 
1938 ; also a specimen in the U. S. National Museum) ; Greenland, 
Iceland, Faroes (Jensen, 1948). 

In the long-lining activities of the M/V DELAWARE, both 
species were taken on the same set on several occasions. Surface 
temperatures at stations at which A. ferox were taken ranged 
from 69-83°F, for A. hrevirostris 58-84°F. The buoy lines, at- 
tached to the end of each section of long-line gear, have been 
10-20 fathoms long, mainly the latter ; the specimens have thus 
been taken quite near the surface. It is noteworthy that, while 
A. ferox and A. hrevirostris have been taken in good numbers in 
late summer and fall, they have been extremely scarce in June. 
The only spring specimen still available, though many were 
taken, turned out to be A. hrevirostris. Only A. ferox was taken 
in winter. 



10 



BREVIORA 



No. 123 


























h^' ^M'i^-^"  



•X ~ 



► Vst 







'!%^^. 



Figure 3. Gonads of Alepisaunis fero.r, typical of both species. Top: 
cross-section through entire structure, testes the small, lobular structures in 
center. Bottom: enlargement sliowing two lolmles of testis above and 
darkly-stained eggs )jelow. 

Reproductive Condition. When the possihility was being co.n- 
sidered that the two species of Alepisaurus might be dimorphic 
sexes, gonads were immediatel}' examined on all available speci- 
mens, and all others possible were thereafter inspected at sea and 



1J)60 ALEPISAURUS BREVIROSTRIS 11 

many were preserved. All a])peare(l to be immature females. No 
specimens even approaching' ripeness have yet been seen. 

Results of histological examination have shown the sui-prising 
fact that both male and female gonadal structures are present in 
both species (Fig. 3). The testes lie dorsal to the much-larger 
ovaries and in the groove between them. The ducts of both 
organs appear to unite before reaching the urogenital opening. 
When the testes are examined under oil immersion, meiotic ac- 
tivity is visible in peripheral cells of the crypts in spite of poor 
fixation. The eggs of the ovaries are fairly well-developed. The 
conclusion can hardly be escaped, therefore, that both species of 
Alcpisaunis are hermaphroditic. 

ACKNOWLEDGMENTS 

I am particularly indebted to James L. Squire, chief of North 
Atlantic Fishery Exploration and Gear Research of the U. S. 
Bureau of Commercial Fisheries and to field personnel and 
members of the crew of the M/V DELAWARE for many cour- 
tesies rendered in connection with the exploratory cruises of the 
DELAWARE. Specimens at the T^. S. National Museum, Mu- 
seum of Comparative Zoology, and the Museum of the University 
of Miami Marine Laboratory were examined through the kind- 
ness of Giles W. Mead, Myvanwy ^l. Dick, and C. Richard 
Robins. I have profited from correspondence or discussions with 
Norman J. Wilimovsky, Giles W. Mead, and G. E. ^Nlaul. My 
sincere appreciation goes to M. Blanc of the Museum d'llistoire 
Naturelle in Paris for his efforts in assuring the lack of types 
there and to N. B. Marshall for information on the types of Lowe 
in the British Museum. Gail G. Pasley of Woods Hole Oceano- 
graphic Institution made the full-length drawings. 

LITERATURE CITED 

Bean, Tarleton H. 

1883. Description of a new species of Alepidosaunts (A. ac.scitlapius) 
from Alaska. Proc. U. S. Nat. Mus., 5: 661-663. 

Bleeker, Pieter 

1855. Over eenige vissclicii van Van Diemcnsland. Uitg. K. Akad. 
Wetensch. Amsterdam, 2: 31 pp. 



12 BREVIORA Xo. 123 

Brimley, C. S. 

1988. The huicet fisli, Ah'ijisauruK fero.r, on the North Cjirolinn co.-ist. 
J. Elisha Mitchell St-i. Soc, 54: 2-i;v246. 

CUVIER, (iEORGES ailcl ACHILLE VALENCIENNES. 

1849. Histoire naturelle des poissons. Vol. 22, pp. i-xx, 1-532. 

GiLi-, Theodore 

1863. Descriptions of new species of Alepidosaurida. Proc. Acad. Nat. 
Sci. Philadelphia, 14 (1862) : 127-132. 

Jensen, Ad. S. 

1948. Contributions to the ichthyofauna of Greenland. Spolia Zoologica 
Mus. Hauniensis, 9(20) : 114-116. 

Lowe, E. T. 

1835a. Description of a new genus of acanthopterygian tishes. Trans. 

Zool. Soc. London, 1: 123-128. 
1835b. Additional observations on Aleinsaurus fero.r. Trans. Zool. Soc. 

London, 1: 395-400. 

Maul, G. E. 

1946. Mouogratia dos peixes do Museu Municipal do Funchal. Bol. 
Mus. Mun. Funchal, 2(2) : 1-61. 

PoEY, Felipe 

1861. Meniorias sol>re la historia natural de la isla do Cuba. Havana. 
Vol. 2: 1-442. 

KicHARDSON, Sir John 

1844. Ichthyology of the voyage of the HMS Erebus and Terror. Lon- 
don. Pp. 1-139. 

Table 1. Pin-ray and gill-raker eonnts of Alepisaurus 

Dorsal Kays Pelvic Eays 

34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 8 9 10 
A. hrcvirontris 313 5 46111 1193 

A.ferox 2 3 10 Id 10 7 4 2 1 2 43 7 

Pacific (USNM) 2 2 11 3 3 

Small 1 13 1 

Anal Kays Pectoral Kays Gill liakers 

13 14 15 16 17 18 12 13 14 15 16 23 24 25 26 27 28 29 

A. brcvirostris 19 12 1 1 13 10 2 3 4 113 2 

A. ferox 1 14 21 15 1 1 5 30 16 14 4 5 2 1 
Pacific (USNM) 6 1 4 3 
Small 3 2 2 2 1 



1960 



ALEPISAT'RIS BREVIROSTRIS 



13 



Table 2. Proportional dimensions of Alcpisaurus expressed as 
per cent of standard lenoth. Range, with mean in parentheses. 
Based on seventeen adnlt specimens of each species and five small 

specimens. 



Standard li'iiptli, iiiiii. 

Per (Tilt of 

standard Iciigtli 
Snout to anal origin 
Snout to polvie insertion 
Snout to pectoral insert 
Snout to dorsal origin 
Head length 
Greatest depth 
Caudal peduncle depth 
Pectoral length 
Pelvic length 
Anal liase 
Anal height 
Per cent of 

liead length 
Snout to tleshy orbit 
Snout to lioiiy oi-\nt 
Fleshy orbit length 
Bony orbit length 
Least interorbital width 
Snout to anterior nostril 
Xumlier of gill raker 

groups (total) 



brevirostris ferox small 

551-894(670.6) 431-1088(777.5) 85.0-190..1 (144.2) 



77-83(79.5) 76- 
42-50(46.3) 43- 
13-16(14.7) 17- 
9.4-13(11.4) 16- 
12-16(14.5) 16- 
7.2-10.4(8.5) 8.0 
1.9-3.2(2.4) 2.3 
14-20(16.3) 17- 
7.2-12.4(9.5) 7.5 
8.5-11.4(9.8) 9.2 
5.9-7.9(6.9) 7.4 



88(80.1) 
53(47.5) 
23(19.4) 
22(17.9) 
23(18.6) 
-12.5(9.5) 
-4.0(2.8) 
24(20.5) 
-10.4(9.0) 
-13.8(10.7) 
10.3(8.6) 



31-37(34.5) 
26-31(28.4) 
19-23(20.9) 
23-33(27.8) 
15-20(17.5) 
14-16(14.9) 
23-28(25.4) 



41-46(43.3) 
35-42(38.1) 
13-20(17.8) 
17-28(23.4) 
14-18(16.2j 
21-24(22.4) 
23-29(25.8) 



80-81(80.0) 

53-58(55.3) 

22-25(24.0) 

21-25(23.2) 

23-30(25.6) 

13-16(14.4) 

2.7-4.0(3.4) 

15-19(17.3) 

7.4-8.5(7.9) 

9.5-13(11.1) 

6.5-9.1(7.8) 



36-41(38.4) 

27-37(31.4) 

25-28(26.2) 

31-36(33.4) 

15-17(16) 

14-18(17.2) 

25-27(26) 



14 



BREVIORA 



No. 123 



Table 3. Principal differential characters of A. hrevirostris and 

A. ferox 



A. hrevirostris 
Coloration dark, more and 
larger melanophores, many of 
them ocellated (Fig-. 2) 
Dorsal fin gradually arcuate, 
without free anterior rays 
(Fig. 1) 



Irregular horizontal row of 
white spots often present on 
dorsal membrane 
Dorsal origin well in advance 
of rear margin of the opercu- 
lum 

Head 6.5 or more in standard 
length 

Snout 2.5 or more in head 
length 



A. ferox 
Coloration light, fewer, most- 
ly small melanophores, few or 
none ocellated (Fig. 2) 
Dorsal fin Avith several ante- 
rior rays elongated, free from 
membrane; rest of fin about 
equal in height until sudden 
posterior drop, or slightly 
liigher before drop (Fig. 1) 
No white spots on dorsal mem- 
brane 

Dorsal origin about level with 
the rear margin of the opercu- 
lum 

Head less than 6.5 in standard 
length 

Snout less than 2.5 in head 
length 



BREVIORA 

Mmseiiiii of Comparative Zoology 



Cambridge, Mass. Makch lo, i960 Xu.mi'.kk 124 

ANI8IAX A:\LM()N()1DS FROM MALAYA 
By Bernhard Kummel 

Possiliforous marine strata are sparsely represented or known 
from Malaya. Even the Triassic system which is one of the better 
known systems to yield fossils is represented by extremely small 
faunas consisting- mainly of pelecypods, and only indeterminate 
ammonoids have been reported by Ne\^i:on (1923, 1925). The 
first discovery of determinable ammonoids was reported by 
Savafje (1950) from mudstones near Knala Li]iis, Pahano-. The 
initial collections were submitted to L. F. Spath of the British 
Museum (Natural History) who made the following report (in 
Savage, 1950) -. "Quite a number of common Middle Triassic 
(Anisian) genera can be recognized in the collection, including 
Paracerofifes (dominant), ^Sturia, Piijchifes, and Acrochordi- 
ccras. so that the age of the assemblage is the trinodosus zone. 
Specific identifications would be more difficult but are unneces- 
sary; Paraceratifcs trinodosus (Mojsisovics) and such close allies 
as the Himalayan Ceraiiics fhnilleri (Oppel) and P. winterhot- 
tomi (Salter) are probably all represented." 

The rarity of Triassic ammonoids in Southeast Asia, lying as 
it does at the eastern end of Tethys between the richly fossil- 
iferous Triassic horizons of the Himalayas and the island of 
Timor, warrants a more substantial record of these faunas than 
Spath Avas able to give. Through the kindness of Dr. M. K. 
Howarth of the British Museum (Natural History) all of the best 
preserved material from Kuala Lipis, Pahang, was loaned to the 
writer. Close examination of this small collection (29 specimens) 
showed that Spath 's conclusions as to genera present and age 
assignment are correct in spite of the rather poor preservation. 



2 BREVIORA No. 124 

The fauna contains the following species : 

Paraceratites trinoclosus (Mojsisovics) 
Sturia sanftovinii Mojsisovics 
Acrochordiceras sp. indet. 
Ptychitcs sp. indet. 

Data on the geographic and geologic occurrence of this fauna 
can best be quoted from Savage (1950, p. 76) : "One of the newly 
recorded areas is some 10.5 miles south-south-Ave.st of Kuala Lipis 
on one of the branches of the Sungei (= River) Tua, where it 
flows through Budu Estate (approximately lat. N. 4°02'30", 
long. E. 102°00'15"; Malayan Survey Department Topographi- 
cal Sheet No. 2 0/13) . The rocks are mudstones, rarely laminated 
sufficiently to be called shales. They are fairly homogeneous, 
slate grey, almost black and carbonaceous ; but some slightly 
sandier strata (muddy siltstones) weather to a pale buif or 
brown. The beds show minor flexures but in the main strike 
40°-220° and dip to the north-west at angles of about 40°. They 
are strongly jointed along several directions, the two main joint 
systems being vertical and striking 60° -240° and 155° -335°. The 
beds are fossiliferous over a distance of at least 50 yds. in a 
horizontal direction normal to the strike, equivalent to a strati- 
graphical thickness of about 100 ft." No data are available with 
the specimens as to their precise position in the fossiliferous 
horizon. 

The species recognized in this small fauna are common forms 
widely distributed in Tethys and the general cireum-Pacific 
region. Paraceratites trinoclosus and Sturia sansovi)ui were 
originally established on Alpine specimens. The specimens as- 
signed to PtycJiitcs and Acrochordiceras are too poorly preserved 
to enable specific identification but there is no doubt as to the 
generic assignment. 

The two previous records of Triassie annnonoids from Malaya 
are not as satisfactory. A small fragment of an ammonoid was 
recorded by Newton (1923, p. 302, pi. 9, fig. 29); it is quite 
indeterminable. The specimen came from argillaceous sandstone 
at Mount Faber, Singapore, and was reported to have a "de- 
pressed whorl with indications of straight ribs and furroAvs con- 
necting with some well-separated knob-like tubercles situated 
within a short distance of the inner margin." Newton likewise 



!!)()() AXISIAN AAi:.ION()lI)S FROM MALAYA 3 

(lid not believe the speeiiiieii w;is determinable but thoiiji'lit that 
it i'es(Miiblo(l the genus B(il(ih>iiil( s. There does not appear to be 
any .justification for this sugo'estion. The second record of am- 
monoid i-emains from ^lalaya is also in a paper by Xewton 
(1925) on a small I'l^per Triassic fanna from the Province of 
Kedali. The fii-st of the two specimens available to Xewton was 
cited as Aiiuiioinfis sp. indet. "A" and he sug-gested tliat it may 
be referred to a form of tlie An-estidae. Tt seems more likely that 
it is a Jui'drltcs or ])ossibly an A H(it<))iiif( s but the specimen lacks 
the eharacteristic consti'ictions of these genera (Spath, lOol, p. 
10(5. footnote). The second specimen was listed by Xewton as 
Aninwuifcs sp. indet. "B" and suggested a resemblance to 
BaJafoiiih s. Spatli (1!'.")1. ]). 15) thought that this specimen 
might be Ihoinoceras )uisf uri in in (INIojsisovics) . 

in the adjoining regions of southeast Asia only Indochina has 
yieUled a large and varied fauna of Triassic anunonoids wliich, 
however, are generally not well preserved. The literature on the 
stratigraphy and faunas of this area is very large and need not 
be reviewed here. A brief summary can be found in Saurin 
(1956). The only other really new discovery of Triassic am- 
monoids in southeast Asia has been nmde in Thailand where 
Anisian and Karnian fannas are now known. The geology of this 
Triassic region has been described by Pitakpaivan (1955) who 
includes a preliminary list of the species present, identified by 
Kummel. Full description of this fanna will be published shortly. 

In the following description of the species from Kuala Lipis 
the extensive synonymies for Paraccratiics trinodosus and Sturia 
saiisovinii have been omitted; essentially complete synonymies 
can be found in Diener (1915a) and Kutass}' (1933). 

SYSTEMATIC DESCRIPTIONS 
Family CERATITIDAE Mojsisovics, 1879 

Genus ParacERATITES Hyatt, 1900 

Paraceratites trinodosus (Mojsisovics) 

Plate 1, figures 3-6 

The collection contains no less than twenty crushed and in- 
complete specimens that can be assigned to the well known 



4 BREVIORA No. 124 

Paraceratites ti'inodosus (Mojsisovics). Allowing for the general 
faulty preservation, these specimens agree well in most details 
witli tlie type of this species and with other specimens assigned 
to it. In his preliminary examination of the fauna, Spath (in 
Savage, 1950, p. 76) considered that in addition to Paraceratites 
trinodosus the fauna also contained P. tJivilleri (Oppel) and 
P. winter!) ottomi (Salter). These are very closely allied forms 
occurring with P. trinodosns in the Himalayas. However, con- 
sidering the poor preservation of the specimens, it seems that a 
more conservative approach is desirable and I am recognizing 
only the better known and more widely distributed P. trinodosus. 

This species is particularly widespread in the Alps, Balkans 
and the Middle East. It is likewise recorded from the Himalayas 
and Nevada. Identical or closely related species are also known 
from Japan. This species gives its name to the upper Anisian 
trinodosns zone. 

Repository. BMNII — C 55672, C 55673, C .15674, C 55675 
(figured specimens) ; C 55653, C 55654, C 55655, C 55657, 
C 55658, C 55661, C 55662, C 55663, C 55666, C 55667, C 55668, 
C 55670, C 55671, C 55676, C 55678. 

Family PTYCHITIDAE Mojsisovics, 1882 

Genus PtYCHITITES Mojsisovics, 1875 

Ptychites sp. indet. 

Plate 1, figure 7 

The collection contains three crushed and fragmentary speci- 
mens that without (juestion belong in Ptijchites but identification 
at the specific level is not possible nor advisable. The most 
complete specimen is actually only the impression of one side of 
a conch. The illustration on Plate 1, figure 7 is of a latex cast 
of this impression. It shows the funnel-shaped umbilicus, broad- 
ly arched lateral areas, and the radial ribs — all features which 
are very characteristic of the genus PfijcJiites. 

Ptychites is known from Middle Triassic strata througliout the 
world. It likewise includes a very large number of species based 
largely on differences in shape of the conch, character of ribs, 
degree of involution, and details of the suture. In the Himalayan 



1960 ANISIAN AMMONOIDS FROM MALAYA 5 

Musc'hclkalk, PiycJiitcs, of tlie jiTuup of F. nKjifcnts (Oppelj to 
which these Malayan specimens most likely belong, is one of the 
most abundant forms present, hciiiu represented by seven species 
(Diener, 1895, 1907). In southeast Asia the record of Ptychites 
is very fragmentary and represented mostly by indeterminate 
species. Even the rich Middle Triassic faunas of Timor appear 
to have only one species, P. aniarassicus Welter (1915; Arthaber, 
1928). The genus is, however, also present in Thailand and Indo- 
china. It is also known to be present in Japan and New Zealand. 
Repository: BMNH — C 55659 (figured specimen) ; C 55664, 
C 55656. 

Genus Sturia Mojsisovics, 1882 
Sturia sansovinii Mojsisovics 
Plate 1, figure 2 

The most easily recognizable species in the collection is this 
strigate form which is widely distributed throughout Tethys. 
The specimen consists only of the impression of slightly more 
than one third volution of one side of a w^horl ; no suture or 
whorl section is preserved. In spite of this fragmentary preser- 
vation the ornamentation is so characteristic that there is no 
reason to doubt its identity with this species. The ornamentation 
consists of broad, flattened strigations separated by broader, 
rounded grooves which bear a fine spiral line down the center. 
The pattern of ornamentation on the Malayan specimen is identi- 
cal to the fine specimen from the Shalshal Clififs in the Himalayas, 
figured and described by Diener (1895, pp. 61-62, pi. 15, figs. 
la, b). In his description of the Himalayan specimen Diener w^as 
quite emphatic as to the identity of his form with the type from 
the Alpine Middle Triassic. In this conclusion he had confirma- 
tion from Mojsisovics who also examined the Himalaj'an speci- 
men. 

Sturia sansovinii is known from Anisian and Ladinian strata 
at many localities in the Mediterranean region. Bibliographic 
citations to these can be found in Diener (1915a) and Kutassy 
(1933). The distribution in the region of eastern Tethys and 
in the circum-Pacific region is not so well known and is of special 



6 BREVIORA No. 124 

interest here. This species is the only form of Stm-ia from the 
Himalayas proper in the so-caUed Ilimahiyaii faeies, where 
Dieiier (1895, 1907) has recorded specimens from tlie upper 
Musehelkallv at the Shalshal Clift* and at Si)iti. However, in 
Tibet in some of tlie exotic bloclvs of Alalia -Jobai- near Cliitichun 
Peak No. 1 (17,74Q ft.), Diener obtained a specimen of Sfuria 
whicli lie identified as Sturia monogolica (Diener, 1895, p. 113, 
pi. 29, tig". 4). This form is quite distinct from other species of 
Sturia in its open umbilicus and the suture, characterized by 
long', slender, pyramidal saddles. At a later date Diener (1916) 
erected the genus Fsilusiuria with S. mougolica as the type 
species. 

Indeterminate species have been recorded from Middle Triassic 
horizons in upper Thailand (Kummel, in Pitakpaivan, 1955). 
These particular forms are small, poorly preserved, and crushed 
specimens whose relationship to N. sausoriiiii is impossible to 
determine. Among the inimerous jMiddle Triassic faunas de- 
scribed from Indochina, Sfuria has as yet not been recorded. 
Welter (1915) records S. cf. sansovinii from a Ladinian horizon 
on Timor based on a frag-mentary specimen. 

Sturia japonica Diener (1915b, pp. 18-20, pi. 6, figs. 1-2) is 
based on a highly distorted specimen from Middle Triassic forma- 
tions at Inai, Japan. It is (luite similar to *S^. sansovinii differing 
in minor features of the suture and character of the strigations. 

Sturia sansovinii is thus found widely distributed throughout 
the Tethyan geosyncline where it occurs in strata of Anisian and 
Ladinian age. 

Repository: BMNH — C 55669 (figured specimen) . 

Family ACROCHORDICERATIDAE Arthaber, 1911 
Genus AcEOCHORDICEEAS Hyatt, 1877 
AcROCHORDicERAS sp. iudet. 
Plate 1, figure 1 

A single, large, crushed and elongated specimen can be as- 
signed to the genus Acrochordiceras but its poor and incomplete 
preservation prevents determination of its specific relationship. 
The whorl sides bear strong radial to slightly curved sharj) ribs. 



1900 AXISIAX AM MONOIDS FROM MALAYA 7 

Some ul' tlu' ribs bcyiii at tlu' uinbilie-ai .slioulder.s where the}' 
increase in height forming somewhat of a tubercle beyond which 
tlu'v bifurcate. Other ribs lack tlie umbilical protuberances and 
are slightly less prominent. The poor preservation prevents de- 
tei-mining the pattern of alternation of these two types of ribs. 
Tlie conch was no doubt slightly evolute but the shape of the 
wliorl section is not possible to determine nor is any part of the 
suture preserved. 

Close comparison of this Malayan specimen with the known 
species of Acrochordiccras is not very satisfactory but one fea- 
ture is notable — that is, the rather sharp ribs on the Malayan 
form. The extent to which these sharp ribs may be due to the 
deformation of the specimen is hard to determine, however. In 
most species of Acrochordiceras the ribs tend to be rounded and 
in some cases broadly rounded. Even though specific comparisons 
are not possible there is no question of the generic assignment of 
this form. 

The genus Acrochordiceras is widely distributed in the Tethy- 
an belt from the Alps to Timor and is likewise known from a 
number of localities in the circum-Pacific region. 

Reposifo)>i. R]\IXII — C 55660 (figured specimen). 

KEFERENCES 

.\kthaber, G. V. 

1928. Ammonoidea Leio.stiaca aus der obereu Trias von Timor. 2. 
Nederl. Timor Expedite 1916 oiidcr leiding von Dr. IT. G. Jonker. 
Uitgegeveu door Dr. H. A. Bromver. IV. Jaarb. Mijnw. Nederl. 
Ind., vol. LV, no. 2. pp. 1-174, pis. 1-20 (1926). 

DiENER, Carl 

1895. Himalayan Fossils. The Cephalopoda of the Muschelkalk. India 

Geol. Survey, Palaeont. Indiea, ser. 1.5, pt. 2, pp. 1-118, pis. 1-31. 
1907. Fauna of the Ilimalaj'an Muschelkalk. India Geol. Survey, 

Palaeont. Indiea, ser. 15, pt. 5, pii. l-tlO, pis. 1-17. 
1915a. Fossilium Catalogus. I. Pt. 8. Cephalopoda triadiea. 369 pp. 

Berlin. 
19151). Japanisehe Triasfaunen. Denksehr. Akad. Wiss. Wien., vol. 92, 

pp. 1-30, pis. 1-7. 
1916. Einige Bemerkungen zur Nomenklatur der Triaseephalopoden. 

Centrabl. Min. Geol. Paltiont., pp. 97-105. 



8 BREVIORA No. 124 

KUTASSY, A. 

1933. Fossilium Catalogus. I. Animiilin. Pt. 56, Cephalopoda triadica 
II., pp. 371-832, Berlin. 

Newton, R. B. 

1923. On marine Triassic shells from Singapore. Ann. Mag. Nat. Hist., 

ser. 9, vol. 12, pp. 300-321, pi. 9. 
1925. On marine Triassic fossils from the Malayan Piovince.s of Kedah 

and Perak. Geol. Mag., vol. 62, pp. 76-85, pi. 3. 

PiTAKPAIVAN, KaSET 

1955. Occurrences of Triassic Formation at Mae Moh. Royal Dept. 
Mines, Bangkok, Rept. Investigations No. 1, pp. 1-11, pis. 2-4, 
map. 

Saubin, E. 

1956. Lexique Stratigraphique International, vol. 3, Asie, Fasc. 6a. 
Indochine, pp. 1-141. 

Savage, H. E. F. 

1950. Triassic fossils from near Kuala Lipis, Pahang (Malaya). 
Colonial Geol. and Min. Resources, vol. 1, no. 1, pp. 76-77. 

Spath, L. F. 

1951. Catalogue of the fossil Cephalopoda in the British Museum 
(Natural History). Part 5, the Ammonoidea of the Trias (D). 
London, pp. 1-228. 

Welter, O. A. 

1915. Die Annnoniten und Nautiliden der ladinischen und anisischen 
Trias von Timor. Paliiont. von Timor, vol. V, pp. 71-136, pis. 
83-95 



Explanation of PLATE 

The specimens illustrated on this plate are from mudstones of Anisian 
age from near Kuala Lipis, Pahang, Malaya. They are deposited in the 
British Museum (Natural History), London. 

Figure 1. Acrochordiceras sp. indet. BMNH — C 55660. X 0.5. 

Figure 2. Stiiria sansovinii Mojsisovics. BMNH — C 55669. X 0.5. 

Figure 3-6. Paraceratites trinodosus (Mojsisovics) BMNH — C 55672 — 
C 55675. X 1. 

Figure 7. PtydJiitcs sp. indet. BMNH — C 55659. X 1. 




\ 



t:'g. 










/^ 



3«i 



BREVIORA 

MuseiLiinii of Comparative Zoology 



Cambridge, Mass. May 27, 1960 Number 125 



THE LUMINOUS ORGANS OF PBOCTOPORUS 
(SAURIA, REPTILIA) —A RE-EVALUATION 

By Willard D. Roth 

Department of Anatomy, Harvard Medical School, Boston, Mass. 

and 

Carl Gans 

Department of Biology, The University of Buffalo, Buffalo, X. Y., 

and Carnegie Museum, Pittsburgh, Pa. 

INTRODUCTION 

The lierpetological literature contains two reports describing 
the first luminous organs in a terrestrial vertebrate. The two 
papers discuss identical specimens of the Trinidad lizard Procto- 
porus shrevei Parker. Sanderson (1939, and observations cited 
by Parker, same date) claimed that light was emitted by black 
bordered ocelli on the sides of a male, and Parker (1939) sup- 
ported this on the basis of his histological examination of the 
preserved animal. 

No new observations have been published since that time, 
but a number of workers have commented upon the original 
observations. Thus Pope (1955, p. 306) remarked that "other 
teiids have spots somewhat like those of P. shrevei, so it is highly 
probable that they, too, can light up." In contrast to this, 
Harvey (1952, p. 494) in his monograph on bioluminescence 
stated that he believed "all reports of luminescence in higher 
vertebrates to be false or spurious due to reflection of light or 
infection by luminous bacteria." 

The divergence of opinion on this interesting point prompted 
a re-examination of this question. The present paper reports a 
few additional field observations, and includes as well a detailed 
examination of the histological structure of the ocelli. Since 
Proctoporus shrevei is very rare, this re-examination had to be 
carried out on two related and superficially similar forms. 



2 BREVIORA No. 125 

OBSERVATIONS ON LIVING SPECIMENS 

Parker (1939, p. 659) mentioned that Sanderson's tield notes 
contained onh- a brief reference to color pattern involving "five, 
black spots each containing- a small, vivid Avliite, sometimes 
Inminons bead." Parker fnrther stated that Sanderson in con- 
versation informed him "that the animal was kept alive in 
captivity and could be stimulated to emit light from the lateral 
spots : the light was of a pale greenish hue, similar to that pro- 
duced by the hands and figures of a luminous watch. Excitement 
produced by flashing an intermittent beam of light on the lizard 
was found to be a very effective stimulus to light production." 

A more specific first-hand report was given by Sanderson 
(1939, pp. 41-43) in his popular, considerably amplified and 
slightly different, account. When initially observed the lizard 
"turned its liead away from me and both its sides lit up for a 
few seconds like the ])ortholes of a ship." "After one brilliant 
display on the night of its arrival in camp it refused to shine 
with full brightness though the beadlike spots remained plainly 
discernible in a darkened box when the rest of the animal was 
invisible." A "loud Avhistle, sudden winds, and flashes of light 
greatly agitated our lizard, causing it to switch on its "port- 
holes". . . . The light was much brighter the first time it was 
switched on after the animal had been quiescent for a period, and 
more especially after it (the lizard) had previously been sub- 
jected to intense illumination." 

We have been able to obtain four sets of further observations 
on live specimens of the genus Proctoporiis. 

Julian S. Kenny (V. C. Quesnel, personal communication) 
some time ago repeated Sanderson's experiments on the original 
species (P. shrevei) with entirely negative results. Kenny's field 
notes also indicate that the lizards are diurnal and inhabit rela- 
tively open spaces on El Tucuche, Trinidad. 

Dr. Janis Roze (in lift.) states that a specimen of Proctoporiis 
achlyens Uzzell (M.C.Z. 53128. later used for histological exami- 
nation reported herein) did not glow when placed in a darkened 
room after capture. lie adds that exposure to ultraviolet light 
did cause the spots to shine faintly. The test was carried out in 
an incompletely darkened room, and the results seem to be 
open to some question. 

Harold Heatwole and Owen J. Sexton (personal communica- 
tion) performed a nuinber of experiments at a field station in 
Venezuela. They tested one adult male of P. Jucfuosuf; (Peters) 



1 !»()() THE LUMINOUS ORGANS OF PROCTOPORUS 3 

and two adult males of /'. achlyens for a period of one month. 
The spots of the first species were yellow and those of the second 
were red in life, both series of spots bleaching to white after 
formalin preservation. The specimens were repeatedly moved 
from light to dark environments and were observed at night. 
The animals were disturbed. No luminescent effect was ever 
noted, intraviolet illumination was not attempted. 

The most extensive series of observations on live animals was 
made in Ecuador by James A. Peters {in litt.). Specimens of 
Prionodactijlus vertehralis (O'Shaughnessy) n.\\(\.Neiisticnrus ec- 
plcopiis Cope were observed while free in the field, during the 
collecting process, and for several weeks in the laboratory. He 
reported that neither luminescence nor any other kind of light 
could be noted in broad sunlight, dim or artificial light, or in the 
complete absence of light. No reflection could be noted under 
various types of lighting (sun, fluorescent and incandescent), in 
quiescent, active or deliberately disturbed animals. The evidence 
is most valuable because Peters was aware of the lizards' reputa- 
tion and was deliberately testing the hypothesis of luminescence. 

SUPERFICIAL APPEARANCE AND PHYLOGENETIC 
DISTRIBUTION OF THE OCELLI 

The supposedly luminous ocelli (Fig. 1) are rather similar in 
the species of Procioporns and Ncusfictirus here discussed. They 
are ari-anged in a single row along the side of the animal ; each 
ocellus always shows a light-colored, sharply -defined, black- 
bordered, circular center. Thev mav be restricted to adult males, 
with juvenile specimens and females showing onh^ traces. There 
is usually a size decrease of the ocellar center posteriorly along 
the series. There may be a marked irregularity in the width of 
the black border. There is no correlation between the ocellar 
and the scale patterns. 

While sharply-defined ocelli are commonly Avell developed only 
in Boulenger's (]885, p. 332) teiid group II, a check of the more 
than 130 species of teiid lizards as well as forms of other families 
represented in the collection of the Museum of Comparative 
Zoology at Harvard College indicated that patterns with sharply 
contrasting light and dark rolors are extremely common. A 
complete morphological series may be demonstrated in the 
Teiidae. This series ranges from ]iatterns with alternating light 
and dai'k stri])es. thi-ough those in which the stripes alternately 
fuse and bi'eak up. to ]iattefiis with well-defined light circles on 



BREVIORA 



No. 125 



a dark haekgTOund. Such spots may be found over the entire 
body or may be restricted to the sides. The condition found in 
the males of Proctoporus and Neusticurus represents only one 
extreme development of color variation. Similar conditions may 
also be observed in certain geckonids and iguanids. Here the 
ocellar pattern may occur all along the side, with the color 
contrast emphasized around a limited number of spots. 







Fig. 1. Proctoporus achlyois Uzzell. Lateral view to show shape, size 
and anangeiuent of ocelli. The scale is graduated in mm. (M.C.Z. 5.3128 — 
C.G. photo.) 



H1ST0L0C4ICAL AND HISTOCHEMICAL EXAMINATION 

OF THE SKIN 

Methods 

Two specimens were used for histological examination. These 
were M.C.Z. 43764, Neusticurus ecpleopus ocellatus Sinitzen from 
Hacienda Pampayacu, Departamento de Huanuco, Peru, and 
M.C.Z. 53128, Proctoporus acJiIyens Uzzell from Choroni, Estado 
de Aragua, Venezuela. The museum specimens were reported to 
have been fixed in 10 per cent formalin and transferred to 
70 per cent alcohol for storage. 

Sections Avere cut from the second (and largest) ocellus of the 
left side of each specimen. A sample of faintly pigmented skin 
from the ventral surface of M.C.Z. 53128 was sectioned for 
comparison. Small blocks of tissue, including both the center 
and the black margin, in the case of the ocelli, were excised. 
These tissue blocks, including the epidermis, dermis, and a small 
aiiioiiiit of undcrlvino' .skeletal muscle were hvdrated through a 



1960 THE LUMINOUS ORGANS OF PEOCTOPORUS 5 

descending- alcohol series, post-chromated in saturated aqueous 
potassium dichromate at room temperature for three days, 
washed overniglit in running tap water, dehydrated in ethanol, 
cleared in chloroform, and embedded in tissue mat (56-58°). 
Five-micron sections were mounted individually so that various 
staining techniques could be carried out on adjacent sections. 

The techniques used were : Harris hematoxylin and eosin and 
the paraldehyde fuchsin method as modified by Halmi (Halmi, 
1950) for general morphology, Wilder 's modification of the 
Bielchowsky silver impregnation method (Romeis, 1948, p. 355) 
for nerve fibers and endings, Sudan black B with acetone con- 
trols for lipid compounds, the periodic acid-Schiff technique for 
1-2 glycol linkages, the azo dye methods for protein bound sulf- 
hydryl and disulfide groups (Barrnett and Seligman, 1952, 
1954 j, dilute methylene blue at pH's 4 to 9 and dilute light 
green at pH 's 3 to 8 for a rough approximation of pH signature 
of proteins (Singer, 1952) and buffered toluidine blue and thio- 
nine for metachromasia. 

GENERAL MORPHOLOGY OF THE SKIN AND OCELLUS 

On the basis of the appearance of nuclei, blood cells, striated 
muscle fibers, small nerves, and the cells of the epidermis, the 
fixation was judged to be good. Presumably the external loca- 
tion of the tissue with the consequent immediate exposure to the 
formalin had been advantageous. In addition, the tanning with 
dichromate seems to have been successful in preventing the 
shrinkage that normally results from paraffin embedding of 
formalin -fixed tissues. 

As revealed by hematoxylin and eosin (Figs. 2, 3), the epi- 
dermis is composed of a very thin stratified squamous epithelium 
showing a prominent basement membrane. The epithelium con- 
sists of a single-layered cuboidal stratum germinativum covered 
by only one or two layers of flattened squamous cells. The surface 
is covered by dense keratinous scales. 

The dermis can be subdivided into tw^o layers. Superficially, 
it is composed of a rather loose fibroelastic tissue which contains 
a dense accumulation of melanin pigment except in the region 
of the ocellus. The pigment appears to be contained in chroma- 
tophores, but this cannot be stated categorically for all of it. 
Often fine strands of pigment granules extend into the epidermis. 
In some instances these granules seem to be in processes between 
the epidermal cells, but in others they seem to occur within the 



6 



BREVIORA 



No. 125 



e-ytuplasni of tiie epithelial cells. Possibly they occur in both 
locations. The deep layer of the dermis is composed of typical 
dense collagenous connective tissue with a rather regular orienta- 
tion parallel to the surface of the skin. Both layers of the dermis 
contain an extensive network of elastic and reticular fibers. 




Fig-. 2. Cross-section through the center of ocellus (l)et\veen the arrows) 
and surrounding pigmented skin taken from Procfnpnriis (irhhifns Fzzcll. 
Ilematoxvlin and eosin. .jOX. 




Fig. 3. Cross-section through the center of the ocellus shown in Fig. 2. 
Xote the thin epidermis (E), the vacuolated appearance of the superficial 
layer of the dermis (S), and the dense collagenous deep layer of the dermis 
(D). Hematoxylin and eosin. 400X. 



1 ;)60 



I in: LUMINOUS organs of proctoporus 



The dermis is underlain by typical loose connective tissue con- 
tainintj- small blood vessels, fat cells, and small peripheral nerves. 

The white center of the ocellus, which is the prime object of 
this study, differs histologically from the rest of the skin only 
ill tJie fact that no melaimi pigment occurs in the superficial layer 
of the dermis or epidermis. The sections thronoli the ocellus do 
not show any obvious differences in thickness or arrangement of 
skin structures as compared with the normal pigmented areas. 

It should be noted especially that no nerve fillers or specialized 
nerve endings were recognized in the dermis of either the ocellar 
or the pigmented regions of the skin, although distinct nerve 
fibers could be seen in the subcutaneous tissue and in tlie under- 
Ijang muscle bundles. It should lie noted further that no exten- 
sive or unusual vascular ai'rangement occurs in the region of the 
ocellus. 

THE IIISTOCHEMLSTRY OF TPIE SECTION 

Ilistochemically, the epidermis shows nothing striking. Its 
surface gives a moderate reaction for sulfhj'dryl groups and an 
intense reaction for disulfide linkages, as would be expected if its 




Fig. 4. Cross-seetioii tlirough tlie same ocellus tis shown in Figures 2, 3. 
Xote the intense PAS-positive reaction of tlie sn]iei-(icjai layer of the 
dermis. I'eriodic-acid-Scliiff reaction connterstained witli hematoxylin 
•400X. 



8 IJREVIORA No. 125 

scales were keratinized. The epidermis covering the white spot is 
identical witJi that covering pigmented dermis. The deep layer 
of the dermis also seems to be identical below the white and 
pigmented skin. The reactions of both collagenous and elastic 
tissue present nothing novel. The superficial dermis of the white 
spot is of special interest, however. In hematoxylin and eosin 
preparations, it is much more lightly stained than the deep 
portion of the dermis, contains less collagen and is far more 
cellular. However, the cytoplasmic limits (and the cell boun- 
daries) of these cells cannot be made out. The impression, 
therefore, is of tenuous, presumably branched cytoplasmic proc- 
esses. 

Histochemicaliy, this superficial region of the dermis (the cell 
cytoplasm?) shows a number of characteristics. Thus it gives 
an intense positive PAS reaction which is diastase resistant 
(Fig. 4). Further, it shows a moderately strong reaction for 
sulfhydryl groups which is not greatly intensified Avith the 
disulfide test. AVith controlled pH staining it shows only a weak 
staining with light green even at low pH's, but a moderate (pll 
5) to heavy (pH 8) staining with methylene blue or thiouine. 
From these results it seems possible to conclude that the cells 
of this region are not vacuolated (as might appear from H and 
E sections), but contain a substance or substances not readily 
stained by routine methods. The results of the PAS method shoAv 
that this material contains numerous 1-2 glycol linkages, but is 
not glycogen. The results of the sulphydryl and the controlled 
pH methods indicate a moderately basophilic protein which con- 
tains appreciable cystine or cysteine or both. A complete absence 
of staining witli Sudan black B indicates that it is not a 
phospholipid, a glycolipid, or a lipoprotein. Thus these findings 
suggest the presence of a mucoprotein or mucopolysaccharide. A 
heavy accumulation of connective tissue ground substance would 
account for these observations except for the absence of meta- 
chromasia. While the fixation in this instance is not optimum 
for a critical evaluation, the ap]iearance of the sections favors 
an intra- rather than extracellular localization. 

Furthermore, upon the examination of unstained sections, the 
region shows no mai'ked granulation under either the light or 
phase-contrast inicroscope. The reflecting pigment guanine is 
supposedly insoluble in all of the reagents used for fixation and 
embedding. Guanine crystals, if present, should therefore be 
evident in unstained sections, and their absence rules out this 
pigment as the source of the white appearance. 



1960 THE LUMINOUS ORGANS OF PROCTOPORUS 9 

DISCUSSION AND CONCLUSIONS 

Basically there are no important differences between the pres- 
ent more extensive histological description and that originally 
fnrnished by Parker (li)39, p. GrtS) . He accented the differences 
l)et\veen the tissue underlying- the glistening white spot and that 
below the remainder of the dermis. The points emphasized were 
tive : (1) a reduction of the epidermis to one-half its normal 
thickness, (2) absence of the chromatophore layer, (3) presence 
of a mass of spongy mesenchymatous tissue, (4) large intra- or 
intercellular spaces in the spongy tissue, and (5) absence of nerve 
endings. Parker emphasized the poor preservation of the ma- 
terial. Neither the stains nor the method of preservation were 
specitied, but the photomicrographs suggest that only H and E 
or a similar routine staining technique was employed. 

We differ in failing to find either vacuolated spaces in the 
s])()ngy tissue or a reduction in thickness of the overlying epi- 
dermis. It seems clear that the presence of "vacuolated spaces" 
could be accounted for entirely by the fact that Parker 's material 
was poorly preserved for histological purposes and that these 
are presumably fixation or shrinkage artifacts. With regard to 
tlie variation in thickness of the epidermis. Parker's photographs 
indicate that such a reduction does not coincide with the absence 
of melanin. Our sections indicate similar and regional differences 
in the thickness of the epidermis of some of the pigmented 
scales. 

On the basis of the additional evidence reported in this paper, 
the following statements may be made : 

Sanderson's observations and Parker's comments thereon con- 
tain a number of inconsistencies. Sanderson reported that the 
light is (1) under the control of the animal and can be turned 
on and off, (2) much brighter the first time it is used after a 
period of quiescence, and (3) brightest .inst after the spot has 
been subjected to illumination. In spite of this Parker suggested 
that the organs are reflectors rather than truly luminous. A 
reflector might be capable of producing the first of the three 
effects, but the last two would be characteristic of true lumin- 
escence. 

None of the subsequent field observers has reported similar 
results. Their experiments cover more specimens and a longer 
period than does Sanderson's report. However, in all but one 
case the species involved are diff'erent. though externally very 
similar and probably closely related to the form on which the 
original report had been based. 



10 BREVIORA No. 125 

The lateral ocelli of rroctuponm luiglit represent one of four 
types of structures: independent light-generating organs (either 
innervated or under liormonal control), receptacles for light-gen- 
erating organisms, specialized reflecting structures, or simplj^ 
nonpigmented "white" spots with a hlack margin. 

The first of these possil)ilities is made unlikely by a number of 
factors. The center of the ocellus shows no gross difference from 
the surrounding skin and can be recognized only by the absence 
of melanin. The ocellus does not, in any manner, resemble pre- 
viously described luminescent organs. The cells of the white 
spot do not have an epithelioid appearance and in no way 
resemble cells previously described for luminescent organs. There 
is neither special innervation nor vascularization. This is par- 
ticularly important since Sanderson indicated that the light was 
turned on (|uite rapidly. 

The absence of any vacuoles or staining reactions character- 
istic of bacteria seems to rule out the possibility of storage of 
luminous microorganisms. 

There are certain difficulties in distinguishing between re- 
flecting structures and plain white spots. It seems certain that 
the white spots do not represent one of the more complicated 
reflecting systems since specialized epidermal cells and similar 
structures are lacking. 

It is, therefore, concluded that the white ajipearance is pro- 
duced by an inter- or intracellular substance, which lies at the 
same level of the skin as the dermal melanophores, and which 
may or may not have sjiecial reflecting properties. The further 
possibility exists that a local accumulation of connective tissue 
ground substance, or a specific intracellular mucoproteiu or 
mucopolysaccharide could be strongly reflective. 

This conclusion is in good agreement with all reports of obser- 
vations on live animals but Sanderson's. If any of the species 
of Proctoporus or related teiids are luminescent, they would seem 
to glow only under very special circumstances and by a yet un- 
described mechanism. HoAvever, the inconsistencies of the initial 
reports by Sanderson and Parker and the completely negative 
result of the investigations presented here force us to reject, for 
the present, any interpretation of these "portholes" as l)io- 
luminescent organs. 

Our findings, furthermore, suggest a quite different and inter- 
esting possibility. The location and appearance of the protein- 
containing cells in the superficial dermis suggests that they 
could be potential melanophores which have formed no pigment. 



1960 THE LUMINOUS ORGANS OF PROCTOPORUS 11 

Clearly, proof of this would involve a study of the histochemistry 
of such "prepiofmented" melanophores. Nevertheless, it seems 
possible to speculate that tliis color pattern in lizards might be 
achieved through a precise local control of the chemistry of the 
melanophore cells and hence, might prove an interesting area 
for the study of specified control of cellular differentiation. 

ACKNOWLEDGEMENTS 

It is a pleasure to acknowledge the original observations kindly 
contributed by Messrs. H. Heatwole, 0. J. Sexton, J. Kenny 
and V. C. Quesnel, and Drs. J. A. Peters and J. Roze. We are 
grateful to A. B. Dawson, D. W. Fawcett, T. S. Parsons, T. 
Uzzell and E. E. Williams for critical comments on the manu- 
script. Many of tlie histochemieal slides were prepared by Miss 
Grethe Aas, and Figures 2 to 4 are from photographs taken by 
L. Talbert. This study was completed under grant NSF G-9054 
of the National Science Foundation. 



LITEEATUEE CITED 

BaRRNETT, E. J. AXD A. M. SELIGilAX 

1952. Histoeheniit-al demonstrations of proteiu-bound sulphydiyl 

Sioii])s. Sc-ien-e, n.s., vol. 116, pp. 323-327. 
19-ji. Ilistoeliemical demonstration of the siilfhydiyl and disulfide 

groups of protein. Jour. Nation. Cancer Inst., vol. 14, pp. 

769-803. 

BOULEXGER, G. A. 

1885. Catalogue of the lizards in the British Museum (Natural His- 
tory). 2nd ed., London, vol. 2, xiii + 497 pp. 

Halmi, N. S. 

1950. Two types of basophils in the anterior pituitary of the rat and 
their respective cytophysiological significance. Endocrinology, 
vol. 50, pp. 140-142. 

Harvey, E. N. 

1952. Bioluminescence. Academic Press, New York, xvi + 649 pp. 

Parker., H. W. 

1939. Luminous organs in lizards. Jour. Linn. Soc. London, Zool., vol. 
40, pp. 658-660. 

Pope, C. H. 

1955. The reptile world. Alfred A. Knopf, New York, xxv + 325 
-f xiii pp. 



12 BREVIORA No. 125 

EOMEIS, B. 

1948. Mikroskopisehe Techiiik. Oldenbourg, Munich, xi + 695 pp. 

Sanderson, I. T. 

1939. Caribbean treasure. Viking Press, New York, 292 pp. 

Singer, M. 

1952. Factors which control the staining of tissue sections with acid 
and basic dyes. Int. Eev. Cyt., vol. 1, pp. 211-255. 



BREVIORA 

Mmiseiiim of Coimpsirsitive Zoology 

Ca:sibridok, Mass. .Itne 3, 19()() NiniBEu 126 

MJD-SC'VTIUAX AMMONITES FROM IWAI FORMATION, 

JAPAN 

By Bernhard Kummel and Sumio Sakagami 



Lower Triassic ammonoids are known from only a few locali- 
ties in Japan. The first fair-sized fauna to be recorded was that 
from the Taho formation on the island of Shikoku described by 
Yehara (1928) . The majority of ammonite species from the Taho 
formation belong in A^iasihirites and Hemiprionites; Yehara had 
erroneon.sly assigned these ammonites to species of Meekoceras, 
Kijmaiitcs, Ophiceras, and Xenodiscus. This fauna is clearly 
representative of the zone of Anasihirites multiformis, which is 
known from Timor, Kashmir, the provinces of Kiangsu and 
Hupeli in China, British Columbia, Queen Elizabeth Islands, 
and western United States. 

A most imi^ortant contribution to our knowledge of the Lower 
Triassic of Japan was made by Sumio Sakagami who in 1955, 
described a small fauna of ammonites from the Iwai formation, 
Kaizawa Valley, Hinode-mura, Nishitama-gun, Tokyo-to. Saka- 
gami had specimens from two fossiliferous beds, seventeen meters 
apart. The lower fauna was concluded to be of early Scythian 
age and the upper fauna to be mid-Scythian (Meekoceras zone) 
in age. 

Correspondence between the authors about this fauna and the 
studj^ of additional material bear out the conclusion that the 
faunas of both fossiliferous beds are of Meekoceras zone age. 
The object of this paper is to further document the species 
present and the age of the Iwai formation. 

The Iwai formation is well exposed in the Kaizawa Valley 
where Sakagami has recognized four members. In ascending 
ordci- these members are: (a) more than 40 m. of black and 
bluish sandstone, (b) 10 m. of shale, (c) 10 m. of sandstone, and 



2 BREVIORA No. 126 

finally (d) about 25 m. of black shale which contains the two 
fossiliferous units. The lower fossil bed occurs about 2 meters 
above the base of the upper member, where the fossils occur in 
lenses of black limestone. The species identified from this horizon 
are: 

Dieneroceras iwaiense (Sakagami) 

Diencroceras sp. indet. 

Owcnitcs shhnizui (Sakagami) 

Farmiannites sp. indet. 

Aspeiiites sp. indet. 

Juvenites sp. indet. 

The upper fossil bed lies about 17 meters above the lower fossil 
bed and consists of marl lenses from which Sakagami obtained 
a single specimen that has been assigned to Aspenites. 

The genera and species in both the lower and upper beds are 
forms very characteristic of the mid-Scythian MccJwccras zone. 
Faunas of this age are well known from several localities in 
California, Nevada, Utah, and Idaho (Smitli, 1932; Kummel, 
1954) ; from the Queen Elizabeth Islands of Arctic Canada 
(Tozer, 1958) ; from the Island of Timor (Welter, 1922) ; from 
Southland, New Zealand (Kummel, 1959) ; from the northern 
Caucasus Mountains, Russia (Kiparisova, 1958) ; and finally 
faunas of this age appear to be represented in the Kolyma River 
region of northeastern Siberia (Popov, 1939) and in Yugoslavia 
(Petkovic and Mihajlovic, 1935). 

Of all the forms represented in the Iwai faunas the single 
specimen of Owenites furnishes the best clue as to their age. 
Oivcnites shimizui (Sakagami) is an immature form that, how- 
ever, compares very closely with Oivenites Ti-oencni from the 
Meclfoceras beds of western United States. Owenites is also 
known from Timor, New Zealand, and the northern Caucasus 
Mountains, Russia. Dieneroceras is a longer ranging genus but 
Dieneroceras iwaiense (Sakagami) is close in its general conch 
morphology to D. eiieneri from the Meekoccras beds of the west- 
ern United States. The Iwai specimens j^laced in Dieneroceras sp. 
indet. are more involute than D. iwaiense and of very different 
appearance. These specimens lack any sign of a suture and the 
identification can only be considered as tentative. Juvenites is 
another genus of wides])read occurrence in the Meekoccras zone 
of western United States but it does range above and below this 
zone. The single specimen from the upper fossiliferous bed which 
Sakagami identified as Aspenites sp. was not available for study. 



1960 



AMMONITES FROM IWAI FORMATION, JAPAN 



The specimen is fragmentary but appears from the illustration 
to be reasonably placed in Aspenites. Another specimen has been 
uncovered from the lower fossiliferous bed that is of much 
better preservation and is without any doubt an indeterminate 
species of Aspenites. 

Previous assessments of the age of the Iwai formation rested 
largely on the identification of the most common species in the 
lower fauna — Diencroceras iwaiense (Sakagami) -as an Ophi- 
ceras. Shimizu (1932) appears to have been the first to comment 
on the Iwai ammonites, though Fujimoto (1926) discussed some 




Fig-. 1. l)iagi;iiimi;iti(.- representation of i\w suture of (a) Oiccnitcs 
shimizui (Sakagami), holotype (from Sakagami, 1955, pi. '2, tig. 2c) X 7; 
(b) Owcnites koeneni Hyatt and Smitli, from a paratype of 15 mm. in 
diameter (from Hyatt and Smith, 1905, pi. 10, tig. 10), X 8; (e) Diencro- 
ceras iioaiense (Sakagami), partial suture of holotji^e (from Sakagami, 
1955, pi. 1, fig. le), X 10. 



Pseudomonutis, earlier. Forms like Dieneroceras iwaieMse are 
extremely difficult to place stratigraphically, and it was not until 
the presence of such genera as Owenites and Aspe)titcs was estab- 
lished that both the age and generic assignment of the "Ophi- 
ceras" could be properly evaluated. 



4 BREVIORA No. 126 

SYSTEMATIC DESCRIPTIONS 

Family DIEXEROCERATIDAE Kiimmel, 1952 

Genus DiEXEROCERAS Spatb, 1934 

DiENEROCERAS iWAiENSE (Sakagaiui) 
Plate 1, fio'ures 3-5 ; Plate 2, figures 7-9 

Oplticvran iwaiense Sakaganii, 19.").j, pp. 135-136, pi. 1, figs. 1-9. 
Ophiceras sp. Sakaganii, 1955, pp. 136-137, pi. 1, figs. 10-1]. 
Dienerocera'i iwaicnsr, Kumniel. 19.~9, p. 430. 

The dominant element in the lower ammonoid bed at Iwai is a 
new species of Dieuerocrros. Sakagami (1955) had eleven speci- 
mens that he described and illustrated, and there are now four 
additional specimens in the collections of the Museum of Compar- 
ative Zoology. The conch is very evolute, each whorl embracing 
the preceding- one only slightly. The whorls are compressed with 
broadly arched flanks which converge slightly toward tbe venter. 
The ventral shoulder is subangular and distinctly marked and 
tbe venter is a low broad arch. The umbilical shoulders are 
broadly rounded. The serpenticone coiling of the conch exposes 
all of the inner whorls whicli are more rounded (and less com- 
pressed) than the outer volutions. The conch is smooth, except 
that on some of the specimens there appear to be extremely faint 
radial folds. The measurements of the better preserved speci- 
mens are as follows : 



*MCZ 5282a (Topotype) 
TUE 5255 (Paratype) 
TFE 5254 (Paratype) 
TFE 5252 (Paratype) 
TUE 5257 (Paratype) 
TUE 5251 (Holotype) 
MCZ 5282b (Topotype) 
TUE 5256 (Paratype) 
TUE 5253 (Paratype) 

The suture is faintly and only partially visible on the holotype 
and on one of the paratypes and consists of a large first lateral 
lobe, and a much smaller second lateral lobe on the umbilical 
wall. It is not possible to determine whether or not the lobes are 
denticulated. 

♦MCZ = MusiMiiu (pf ('iiiuiiar.irivc Zooliigy : TUB =: Tokyo I'uiversity of Education 



D 


II 


w 


T' 




( .Mcasurciiicin 


s in mm.) 




22.5 


7.7 


5.3 


11.0 


21.0 


15.8 




10.1 


20.7 


7.0 




10.6 


20.5 


6.5 




9.0 


18.0 


6.2 




8.7 


15.5 


4.5 


3.7 


7.5 


15.0? 


4.5 




7.7 


15.0 


5.0 




7.2 


9.4 


3.0 




4.5 



l!)(i() AMMONITES P^ROM IWAI FORMATION, JAPAN 5 

Rcniorks. Many of the mid- and late Scythian ammonoids, that 
have on various occasions been assi<irned to Ophiccras, are now 
commonly placed in Dicncroceras whicli is interpreted as a per- 
sisting generalized stock out of the early Scythian ophiceratids 
(Spath, 1934, p. 12-i; Kummel, 1952," p. 849, 852). In the 
Meekoceras fauna of western United States, Dicncroceras is rep- 
resented by D. clicncri (Hyatt and Smith), D. knccJiti (Hyatt 
and Smith), D. suhquodratunt (Smith), and some as yet nnde- 
scribed species. The specimens assigned by Smith (1932, p. 50, 
pi. 54, figs. 1-17; plate 56, tigs. 13-18) to Ophiccras saluntala 
Diener are also a species of Dicncroceras. As now interpreted 
the species assigned to Dicncroceras show a wide range in 
morphological features, especially marked in the cross-section of 
the whorls. 

Dicncroceras iwaioise is morphologically most similar to the 
genotype, D. clicncri. This is especially noticeable in the angular 
ventral shoulders and low arched venter. The illustrations of the 
holotype and paratype of D. dicneri originally described l)y 
Hyatt and Smith (1905, pi. 8, tigs. 16, 17, 19, 20) are inaccurate 
drawings; these types are re-illustrated here on Plate 3, figures 
1-4. The Japanese species, however, is more compressed and the 
whorls converge toward the ventral region more than in D. 
dieneri. The genotype bears faint strigations which are most 
often not preserved, as already noted by Smith (1932, p. 49). 
None of the Japanese specimens show any trace of strigations. 

Occurrence. Lower fossiliferous bed of upper memlier of Iwai 
formation, Kaizawa Valley, Iwai, Ilinode-mura. Xishitama-gun, 
Tokyo-to, Japan. 

Reposifory. MCZ 5282a,b,c, (PI. 1, figs. 3-5) ; TUE 5254, 
paratype (PI. 2, fig. 7) ; TTTE 5251, holotype (PI. 2, figs. 8, 9). 

DiENEROCERAS Sp. iudct. 

Plate 1. figure 1 : Plate 2. figure 10. 

risltiiaitts .sp. Sakaganii, 1955, p. 137, pi. 1, fig. 12. 

In the original collection described by Sakagami (1955), he 
had a single, small, incomplete specimen of which only one side 
of a half of a volution was preserved (PI. 2, fig. 10 of this 
report). Four additional, though fragmentary, specimens are 
now available. The l)est specimens are illustrated on Plate 1, 
figure 1. The conch is small, compressed, and involute ; the whorl 
flanks are broadlv arched merging witli a well rounded umbilical 



6 BREVIORA No. 126 

shonld(M- (111 the one side, but the ventral shoulders are sub- 
angular. The venter is broadly arched. The whorl sides liear 
weak, slightly sinuous, narrow folds which are most i)ronounced 
on the dorsal half of the whorl side. Thifortunately, no suture is 
preserved on any of the specimens. 

Ronarl-s. The incompleteness of the specimens and the absence 
of any sutures makes identification of forms like this extremely 
uncertain. The specimens could very possibly be juveniles of 
larger forms. Under these circumstances the assignment to 
Diencroceras can only be considered as tentative but the most 
reasonable conclusion for the moment. 

Occurrence. Lower fossiliferous bed of upper member of Iwai 
formation, Kaizawa Valley, Iwai, Hinode-mura, Xishitama-gun, 
Tokyo-to, Japan. 

Rcpositortj. MCZ 5283 (PI. 1, fig. 1); MCZ 5286 (unfigured 
specimens) ; TUE 5260 (PL 2, fig. 10). 

Family PROPTYCHITIDAE Waagen. 1895 

Subfamily OAVENITINAE Spath, 1984 
Genus OwENITES Hyatt and Smith, 1905 

OwENiTEs SHiMizui (Sakagami) 
Plate 2, figures 5, 6 

l\ni'litiH .sliiiiii~ui Sakayanii, 1955, p]). 13S-]39, \>\. -, li;.;;;. -;[ c. 
f/'/rr/n'/r.s- shi}ni~ui (Rakagaiiii ), Kuniniel, 1959, p. 43(1. 

The holotype and only specimen of this si)ecies is a small, 
juvenile specimen that can with confidence be assigned to 
Oivenites. The specimen measures 21.0 mm. in diameter, 11.3 mm. 
for the height of the last whorl, 8.0 mm. for the width of the 
last whorl, and the umbilicus is 1.5 mm. in diameter. The conch 
is involute with broadly arched whorl sides that converge, form- 
ing a sharp acute venter. The only ornamentation consists of 
radial growth lines. 

The suture (Fig. la) is ceratitic and typical of that found in 
species of Owenitcs. It consists of a narrow, denticulated ventral 
lobe, a large denticulated first lateral lobe, a smaller second 
lateral lobe and a series of small auxiliary lobes. This suture 
is almost identical in its basic plan to that of a specimen of 15 mm. 
in diameter of Owenifes koeneiii Hyatt and Smith (1905, pi. 10, 
fig. 10) reproduced here on Figure lb. 

Rcnwrks. Mature specimens of Owcnites show marked excen- 
truinbilication on the outer whorls producing a deep funnel- 
shaped umbilicus. Tlie immature volutions (roughly up to 25-30 



19(iO AMMONITES FROM IWAI FORMATION, JAPAN 7 

mm.) form a tightly involute coiieh and the small nmbilicns 
shows no tendency toward excentrumbilieation. Owenitcs shitni- 
zui is almost identical in conch shape and proportions to speci- 
mens of comparable size of 0. koeneni of the Meekoceras zone of 
western United States. A paratype of 0. koeneni originally illus- 
trated by Hyatt and Smith (1905, pi. 10, figs. 7-9) by a poor 
drawing is illustrated here on Plate 3, figures 5-7. This speci- 
men measures 15 mm. in diameter and is the specimen from 
which the suture of Text-figure lb was obtained. The suture is 
very similar in these two species at about the same diameter, 
ditfering only in minor details. The resemblance to Kingites in 
conch shape and suture is more apparent than real. 

Owenitcs is one of the best mid-Scythian zonal markers in the 
Circum-Pacific region. In western United States (California, 
Nevada, Utah, and Idaho) the genus is very common in the zone 
of Meekoceras gracilitatus. The genus was first established for 
specimens from the Meekoceras limestone in the Inyo range, Cali- 
fornia (Hyatt and Smith, 1905, p. 82). Smith (1932) recognized 
a number of additional species of Owcnifes in western United 
States but most of these appear to be merely intraspecific variants 
of 0. koeneni. 

The Timor Owenites cgrediens Welter (1922) has a narrow, 
rounded keel-like venter formed by the shell, but the internal 
cast has a sharp venter. Likewise the Timor species is generally 
more inflated, producing a broader and deeper umbilical funnel. 
The suture also differs slighth" in the shape of the lobes and the 
auxiliary series. Eecently, a specimen of Owenites cf. koeneni 
Hyatt and Smith, has been described from beds of Pre-Etalian 
age in western Southland, New Zealand ( Kummel, 1959) . Outside 
of the Circum-Pacific region. Owetiites has been recorded only 
from the northern Caucasus Mountains. 

Occurrence. Lower fossiliferous bed of ui)per iiK'inlR'r of Iwai 
formation, Kaizawa Valley, Iwai, Hinode-mura, Nishitama-gun, 
Tokyo-to, Japan. 

Repositorii. TUE 5262. holotype (PI. 2. figs. 5, 6). 

Family PARAXAXXITIDAE Spath, 1930 

Subfamily PAKAXANXITINAE Spath, 1930 

Genus PaRAXANNITES Hyatt and Smith, 1905 

Paranannites sp. indet. 
Plate 2, figures 1, 2 

I'vopt yclntvs ;iff. i-osi iilrd ii Irl Spntli. S;ik;if;-;iiiii, ]9.').1, ]iii. l.'iT-liiS, pi. H, 
figs, la, 1). 



8 BREVIORA No. 126 

Paranannitcs i\i. iudct., Kuniuu'l, ]!».'!), ji. 4'M). 

This form i.s represented by a single specimen of only moderate 
l)reservation. The eonch is involute, compressed, with flattened, 
l)araHel Avliurl sides and a broadly rounded venter. It measures 
33.1 mm. in diameter, 15.3 mm. for the heig'ht of the last whorl, 
9.3 mm. for tlie width of the last whorl, and 6.5 mm. for the 
diameter of the umbilicus. Unfortunately no suture is preserved. 

Lower Triassic ammonoids of this conch morphology are dif- 
ficult to identify, and without the suture generally impossible 
to recognize. The fact that the associated fauna includes species 
of Owenitis and Juvenites precludes the probability that this 
specimen could represent a species of Propfychitcs, which is gen- 
erally an earlier Sc^^thian form. The associated genera indicate 
a mid-Scythian age for the fauna and, of the ammonites of this 
age, Paranajuiitcs comes the closest in its conch morphology to 
this specimen from Iwai, Japan. Paranannitcs aspcncnsis from 
the Meckoceras zone of western United States has a conch of the 
same degree of involution and rounded venter which, however, 
is more inflated — ^the whorl width being just slightly less than the 
whorl height. Paranannitcs prrteiuiis Smith (1932, p. 99, pi. 31, 
figs. 13-15) has a laterally compressed conch with flattened sides 
like the Iwai specimen (PI. 3, figs. 9, 10). This species of Smith 
is believed to be a synonym of P. aspene7iesis. The tentative 
placement of tliis specimen in Paranannites is, of course, based 
on the assum])tion that it is a mature specimen. If, however, it is 
a juvenile form it is most likely not a Paranannitcs, and then 
could possibly be the inner whorls of a Flcmingites or Arctoceras, 
or other such larger ammonoids of mid-Scythian age. 

Occurrence. Lower fossiliferous bed of upper member of Iwai 
formation, Kaizawa Valley, Twai, Ilinode-mura, Nishitama-gun, 
Tokyo-to, Japan. 

Repository. TUE 5261 (PI. 2, figs. 1, 2). 

Genus JuVEXITp:s Smith, 1927 

Juvenites sp. indet. 
Plate 1, figure 2 

The collection contains a single specimen in which only one 
side of a half volution is preserved. The specimen measures ap- 
proximately 16.4 mm. in cliameter, 7.3 mm. for the height of the 
last whorl, and th(> umbilicus is 5.0 mm. in diameter. The conch 
is involute Avith l)roa(l, de]iressed Avhorls and a broadly rounded 



19(i0 AMMONITES FROM IWAI P^ORMATION, JAPAN 9 

vonter tliat <>rade.s imperceptibly onto the lateral areas. The 
eoiieli bears very eoiispiciioiis forward projecting constrictions. 
There appear to be six such constrictions on the half volution. 
A^o indication of a suture is preserved. 

Remarks. The lack of a suture and the incompleteness of the 
specimen necessarily make the present identification tentative. 
Even so, in consideration of the association with Owenitcs and 
Aspcnitcs which are clearly mid-Scythian in age, the assignment 
of this specimen to Juvenites appears reasonable. The constric- 
tions on the Iwai specimen are similar in depth and distinctness 
to those in Juvenites septentrionalis Smith (1032, pi. 31, figs. 
31-32) but in the latter species the constrictions are radial. 
Strongly projected constrictions somewhat similar to those on 
the Iwai specimen are present on Juvenites thcrmarum (Smith, 
1932, pi. 21, figs. 11-12, 16-17, 19-20). 

Occurrence. Low^er fossiliferous bed of upper member of Iwai 
formation, Kaizawa Valley, Iwai, Ilinode-mura, Nishitama-gun, 
Tokyo-to, Japan. 

lifprmtoyjj. MCZ 5284 (PL 1, fig. 2). 



Family IIEDENSTROEMIIDAE Waagen, 1895 

Subfamily ASPENITINAE Spath, 1934 

Genus ASPENITES Hyatt and Smith, 1905 

AsPENiTES sp. indet. 
Plate 2, figures 3, 4 ; Plate 3, figure 8 

A.siK nil( K .s[). S;ik;if;'aiiii, 1935, \>. lo9, pi. '2, ligs. 3a, 1). 

One of the authors (Kummel) has not had the opportunity 
to examine the single representative in the collection originally 
described by Sakagami (1955, p. 139) who states that it agrees 
with Aspenites of the western United States but the suture is not 
preserved. As well as one can tell from the illustration, this 
identification appears to be reasonably correct. Another specimen 
has since been uncovered from the lower ammonite bed that 
appears without doul)t to he a juvenile representative of Aspe- 
nites. This specimen (MCZ 5285) measures only 8.0 mm. in 
diameter and is a completely involute conch, greatly compressed, 
with broad arched flanks which converge to a narrow, keeled 
venter (PI. 3, fig. 8). No suture is preserved. The specimen 
is almost identical with specimens of comparable size of Aspenites 
acutus Hyatt and Smith (Smith, 1932, pi. 30, figs. 6-7, 9, 11-12). 

Aspenites is fairly abundant in tlie Mrekoerrns beds of Avestern 
United States and in Timor. 



10 BBEVIORA No. 126 

Occurrence. Botli iipi)er and lower fossiliferous beds of upper 
member of Iwai formation, Kaizawa Valley Twai. liinode-mura, 
Nishitama-s'un, Tokyo-to, Japan. 

Ecpositonf. tup: 5268 (PI. 2, %s. :], 4); MCZ 5285 (PI. 3, 
fig. 8). 

KEFERENCEW 

FUJIMOTO, 11. 

1926. A new Locality of " Pseudomonotis." Jour. Geol. Soc. Japan, 
vol. .S.3, no. 390, p. 113. (In Japanese.) 

Hyatt, A. and .1. P. Smith 

1905. The Triassie Cephalopod (ienei'a of America. U. S. Geol. Survey, 
Prof. Paper 40, pp. 1-394, pis. 1-85. 

KU'AKISOVA, L. 

1958. Geologic Structure of the USSR, vol. 1, Stratigraphy. Moscow, 
588 pp. (In N. A. Belyaevsky, et. al.) 

KUMMEL, BERNHARD 

1952. A Classification of the Triassie Ammonoids. Jour. Paleont., vol. 
26, pp. 847-853. 

1954. Triassie Stratigraphy of Southeastern Idaho and Adjacent 
Areas. U. S. Geol. Survey, Piof. Paper 254-H, pp. 165-194, pis. 
34-40. 

1959. Lower Triassie Annnonoids from Western Southland, New Zea- 
land. New Zealand Journal Geol. Geog., vol. 2, no. 3, pp. 429-447, 
figs. 1-7. 

Pktkovic, K. V. and P. Mihajlovic 

1935. La faune des cephalopodes trouvee dans le Trias Inferieur en 
Montenegro (Yougoslavie) ses caracteristiques et son importance. 
Ann. geol. Penins. Balkan., vol. 12, pp. 253-269. 

Popov, G. 

1939. New Species of Ammonoids from the Triassie of the Okhotsk- 
Kolyma Land. Arctic Institute, Leningrad, Problems of the 
Arctic, no. 12, pp. 72-82. 

SAKAGAill, S. 

1955. Ijower Triassie Ammonites from Iwai, Oguno-Mura, Nishitama- 
gun, Kwanto Massif, Japan. Science Report, Tokyo Kyoiku 
Daigaku, sec. C, no. 30, pp. 131-140, pis. 1, 2. . 

SlIlMIZU, S. 

1932. On the Lower Triassie Ammonites of Iwai, Oguno-Mura, Nishi- 
tama-gun, Tokyo-to. Jour Geogr., Tokyo, vol. 44, p. 97. (In 
Japanese.) 



1960 AMMONITES FROM IWAI FORMATION, JAPAN 11 

Smith, J. 1'. 

1932. Lower Triassic Ammonoids of North America. U. S. Geol. Sur- 
vey, Prof. Paper 167, pp. 1-199, pis. 1-81. 

Spath, L. F. 

193-4. Catalogue of the Fossil Cephalopoda in the British Museum 
(Natural History). Part -t — The Ammoiioidea of the Trias. 
British Museum Publication, London, 521 pp., 17 pis. 

TOZEK, E. T. 

1958. Triassic Faunas from the Queen Elizalieth Ishnuls, Arctic 
Canada. Al)stract, Bull. (Jeol. Soc. Amer., vol. (i9, pp. 1653-1654. 

Welter, O. A. 

1922. Die Ammoniten der unteren Trias von Timor. Paiiiont. von 
Timor, Lief. 11, Alili. 19. i)]i. 82-160, pis. 155-171. 

Yehaba, Shingo 

1928. The Lower Triassic Cephalopod and Bivalve Fauna of Shikoku. 
Jnp. Jour. Geol. and Geog., vol. 5, p]i. 135-172. 



EXPLAXATIOX OF PLATE I 

The specimens illustrnted on tliis jilate ;ne frcmi the lower fossiliferous 
bed of the Iwai formation, Kaizawa Valley, IIino;le-muia, X'isliitama-gun, 
Tokyo-to, Japan, and are preserved in the Musevmi of Comparative Zoology. 

Fig. 1. DUnei-ocerus sp. in;let., MCZ .5283, X 2. 

Fig. -2. Jiivrniirs sp. indrt., ^K'Z .5284, X 3. 

Figs. 3-5. Dirurrorrrds iivaioisc (Sakagami"), topotypes, MCZ .5282a,h,c, 
X :;, 




PLATE 1 



EXPLAXATIOX OF PLATE 2 

The speeinienw illustrated on this plate are t'vom the Iwai formation, 
Kaizawa Valley, Iwai, Ilinode-mnra, Ni.shitania-gun, Tokyo-to and are 
preserved in the collections of the Geological and ^Nlineralogical Institute, 
Tokyo University of Education. 

Figs. 1, -. Parandiinites sp. indet., fi-oni lower fossiliferous bed of Iwai 
formation, TUE 5261, X 2. 

Figs. 3, 4. Aspenites sp. indet., fi'om upper fossiliferous bed of Iwai 
formation, TUE .1263, X 1. 

Figs. 5, 6. Oivcnites sIiiDihui (tSakagami; from lower fossiliferous bed of 
Iwai formation, TUE 5262, X 2. 

Figs. 7-9. Dicncroccras iicaiensc (Sakagami) from lower fossiliferous 
bed of Iwai formation. Fig. 7, i)aratype, TUE 5254:, X 2; figs. 8, 9, holo- 
type, TUE 5251, X 2. 

Fig. 10. Dieneroccrus sp. indet. from lower fossiliferous lied of Iwai 
formation, TUE 5260, X 2. 



«r.)P'^5^ 











EXPLAXATIOX OF PLATE 3 

Figs. 1-4. Dieneroceras dieneri (Hyatt and Smith), from MeeTcoceras 
beds in Wood Canyon, 9 miles east of Soda Springs, Aspen Eidge, Idaho. 
Figs. 1, 2, holotype, USX:\r 75260, X 1..5; 3, 4, paratype USXM 75260a, 
X 2. 

Figs. 5-7. Owenites Icocneni Hyatt and Smith, from Meekoceras beds. 
Union Wash, Inyo Range, Inyo County, California. Paratype USXM 
7526111, X 2. 

Fig. 8. Aspcnitrs sp. imlet. from lower fossiliferous bed of Iwai forma- 
tion, Kaizawa Valley, Iwai, Hinode-mura, Xishitama-gun, Tokyo-to, Japan, 
MCZ 5285, X 4. 

Figs. 9-10. Faranannitcs pertenuis Smith, from ileeloceras beds in Wood 
Canyon, 9 miles ea.st of Soda Springs, Aspen Ridge, Idaho. Holotype, USXM 
74960, X 1.5. 















^«#' 






V JJs 




,■% 






1^ 







8 



10 



PLATE 3 



BREVIORA 



Mmseiuijni of Compsirative Zoology 



Cambridge, Mass. December 19, 1960 Number 127 

NOTES ON THE CRANIAL ANATOMY OF 
NECROLEMUR 

By 

E. L. Simons ^ axd D. E. Russell - 

INTRODUCTION 

The large number of Avell-preserved skulls of Necrolemur 
antiquus of the late Eocene Quercy phosphorites of south central 
France allow for much more detailed study of cranial anatomy in 
this primate than is possible for most early members of the order. 
In spite of the fact that cranial osteology can be studied in great 
detail, views as to the taxonomic position of this primate, and 
of the allied genera Microchoerus, Nannopitliex, and Pseudoloris 
show considerable variance. 

Although not all of the same provenance, little dental varia- 
bilit}" is evidenced in specimens of Necrolemur antiquus examined 
by us. In the course of this study, however, a number of differ- 
ences in position and size of basicranial foramina have been 
observed, which are in line with mutability of cranial foramina 
(in individuals of the same species) reported by other authors 
(see Edinger and Kitts, 1954). Also, in Necrolemur the proba- 
bility remains that known specimens differ considerably in age 
(from early Bartonian to late Ludian provincial ages, at least), 
but locality data are inadequate for precise age determinations. 
Comparison of upper dentitions in the M.C.Z. and Paris skulls 
has failed to show anj- dental basis for species distinctions 
among them. 

To date, the most detailed studies of the cranium of Necrolemur 
have been by Stehlin (1916) and by Hiirzeler (1948). Stehlin's 
thorough and excellent description can scarcely be improved on, 

^Zoological Laboratories, University of Pennsylvania. 
^ Museum National d'llistoire Naturelle, Paris. 



2 BREVIORA No. 127 

l)iit copies of this work ai'e not as >ieiierally available as could 
he "wished. Also, tlie imi^licatioiis of some of his early observa- 
tions seem to have been neglected in latei- literature. In some 
rather significant points, recent examination of more and dif- 
ferent skulls ])ermits comments su|)pleiiientary to his work. 

ACKXO WLEDi ; e:\ie xts 

The authors Avould like to take this o|)portunity to thank Drs. 
J. -P. Lehman, Curator of Fossil Vertebrates at the ^Museum 
National cVIIistoire Naturelle in Paris, and A. S. Romer of the 
Museum of Comparative Zoology at Harvard for generously 
giving permission to pul)lish on the specimens in their respec- 
tive charges. Preparation of the figures, by ]\Iiss Ellen Cole, 
was supported by a grant from the AV(Miiiei--(iren Foundation for 
Anthropological Pesearch. 

ABBREVIATIONS 

In the absence of specimen numbers, the Paris Museum skulls 
of Nccrolcmur have been numberetl 1 through 5 for convenience 
of reference. Abbreviations used in this paper are as follows : 
M.C.Z., Museum of Comi)arativ(^ Zoology at Harvard College. 
Montauban, Natural History Museum (Geological Collection), 
Montauban, France. Paris, National Museum of Natural History, 
Paris. 

CRANIAL CHARACTERS 
I. AFDITORY REGION 

Most of the information, published to date, regarding the com- 
ponents of the auditory bulla in Necrolemur comes from Mon- 
tauban 9, which has been discussed by both Stehlin and Hiirzeler. 
Even though this skull was prepared with considerable skill, 
the crystalline calcite filling was apparently confused with the 
very similar appearing bone in the region of the epitympanic 
recess. Consequently, the route of the stapedial artery across the 
tympanic cavity of the middle ear was lost just anterior to the 
fenestra ovalis. The extrapetrous portion of the Fallopian aque- 
duct was lost as well. Preparation of Paris 2 has revealed more 
details of the epitympanic region (Fig. 1). 

Exposure of the inside of the bulla in Paris 2 and 5 indicated 
primarily the lack of a free annular tympanic ring. Moreover, 
studies by Simons (in press) on a specimen of Necrolemur at the 



lOfiO 



CRANIAL ANATOMY OF NECROLEMUR 




«( 33 



o <:<: 

&K5 



1; Rh t 

ex iD ^ 

i i ^ 



i; P rt 



OQ 



-. o 



X ;r o 



!<; rt 9 

^ -H ft 

-r '^ cs 

-^ <» ;h 

— a -^^ 

— =4-1 

t£ ^ O 

•n ^- ^ 

=M CC ^ 



O ^H ^ • 



CI ^ b 

— ^ +- 

-: ^ ^ 

* p o 

5 .2 - 

^ ■> P 

1^ *— — * 



'*^ ■^ a5 ^ 



4 BREVIORA No. 127 

British IMiiseuni (Natural History) doiiionstrate that the ecto- 
tympaiiic element is tubular, and mediall}' fused to the ventral 
bulla wall. Iliirzeler's evidence (1946:353; 1948:28) of the 
presence of a free ring-, therefore, can no longer be accepted. 
The bone he identified as such had to be removed during prepa- 
ration to expose the carotid canal and thus cannot be re-exam- 
ined. In M.C.Z. 8879 the meatal tubes are reasonal)ly well pre- 
served but one of the Paris skulls shows an even more complete 
osseous meatus. Together, these indicate that the ectotyrapanic 
(external to the bulla) is about as long as the transverse diam- 
eter of the foramen magnum, curves slightly backward, and 
may be broadest at the external aperture. None of the fossil 
or recent Lemuroidea have this sort of meatus. 

Iliirzeler (1948:27) cites M.C.Z. 8879 as not showing any 
evidence of a fused tympanic ring. Nevertheless, four and pos- 
sibly more transverse struts are exposed on the ventrolateral 
face of the right bulla of the Harvard skull. These bars are 
su]iports for the internal rim of the tubular ectotympanic. 

Also of interest is the fact that the anterior route of the 
])romontory artery (true entocarotid) is apparently variable. 
In Montauban 9. it curves sharply anteromedially shortly after 
leaving the promontory of the petrosal. In Paris 2, this curving 
is much less accentuated. Some crushing is to be allowed for in 
the tympanic region of the latter specimen but the amount of 
curvature illustrated by Hiirzeler (1948, figs. 30 and 31) for 
Montauban 9 is not indicated in the Paris skull. It should be 
further noted that this l)ony tube does not lie in a horizontal 
plane in Montauban 9 and Paris 2, but slopes anterodorsally at 
an angle of about 45°. 

The stapedial artery, like the promontory artery, remained 
enclosed in a bony tube throughout its route within the bulla. 
Branching from the promontory artery just inside the carotid 
foramen, the stapedial artery curved dorsally, lying on and 
following the form of the petrosal promontory. It then passed 
anterior to the fenestra rotunda to the bottom of the fenestra 
ovalis. There it diverged laterally, crossing the fenestra ovalis, 
and continued anteriorly nearly parallel to the promontory 
artery (Fig. 1). The groove mentioned by Hiirzeler (1948:31) 
is surely a remnant of the stapedial tube, as he suggested. The 
exit of the stapedial artery appears to be at the dorsoanterior 
base of the external auditorv meatus. 



iy()0 CRANIAL ANATOMY OP NECR0LE:\IUR 5 

The extrapetrous portion of the Fallopian a(iiieduct is also 
enclosed up to, or nearly up to, its exit at the stylomastoid fora- 
men. Its route lies lateral to the fenestra ovalis, just above the 
stapedial artery and continues posteriorly under the external 
semicircular canal. At this point, the tube forms a "T"' giving 
rise to the small anterolateral opening and a larger posterome- 
dian branch. The former is a natural foramen. Damage to the 
latter region makes it impossible to say whether or not the more 
posterior branch continued as an enclosed tube to the stylo- 
mastoid foramen. In Paris 2, this foramen is single, in Paris 1, 
double. Without exposing the interior of the bulla in Paris 1 
a possible connection between the branching Fallopian aqueduct 
and the double stylomastoid opening cannot be confirmed. That 
the anterolateral foramen of the "T" in Paris 2 could have given 
passage to the chorda tympani seems likely. 

Regarding the foramen designated FX, "Foramen von un- 
liekannter Bedeutung, " by Iliirzeler (1948 :fig. 28), it seems 
probable that this represents the opening of the inferior petrous 
sinus, as originally stated by Stehlin (1916:1355). 

Contrary to the views of a few students, we tind little distinc- 
tion between Tarsius and Xccrohmur in the major carotid rela- 
tionships, both inside and outside the l)ulla. The two genera 
appear to agree in those features of the carotid circulation 
which distinguish tarsiers from Malagasy lemurs, adapids and 
lorises (see Le Gros Clark, 1959:151). Location of the internal 
carotid foramen in Tarsius on the ventral surface of the bulla 
(instead of on the median wall) constitutes a slight difference 
from Xco'ohiintr, but it should be stressed that placement of 
this foramen in the fossil species does approximate the situa- 
tion in Tdfsius, being more ventral than in most, if not all, 
other prosimians. Furthermore, Kecrolemnr and Tarsius are 
alike in having both stapedial and promontory branches within 
the ])ulla, encased in bony canals or tubes, with the promontorj^ 
division the larger. In typical Lemuriformes (Malagasy lemurs, 
adapines and notharctines), the carotid foramen has a quite dif- 
ferent location at the posteroexternal angle of the bulla and, inside 
it, the promontory division is A'ery small (Gregory. 1920 :174:-180) . 
Lorises and the cheirogaleine lemurs differ also, in that the 
carotid divides outside the entotympanic and the main branch 
enters the skull through the foramen lacerum medium instead 
of going through the bulla. A middle lacerate foramen is not 
present in Tarsius and Nccrohmur. The few differences be- 
tween these two genera to be observed in the auditory region 



6 BREVIORA No. 127 

seem best understood with reference to the effects of the anterior 
shifting of the foramen magnum and greater inflation of the 
anterointernal part of the bulla in Tarsius. Some primitive fea- 
tures are also to be seen in the Eocene form. For instance, the 
canal for the promontory artery in Nccrolemur is only slightly 
thicker than that for the stapedial. In Tarsius this difference 
is more pronounced. What is of general significance is that when 
Necrolemur differs from Tarsius it is usually intermediate be- 
tween the latter and yet more primitive prosimians. One could 
hardly expect an Eocene tarsioid to be otherwise. 

II. BASICRANIUM 

Stehlin (1916:1351) mentioned that the alisphenoids partici- 
pated in the composition of the anterior wall of the bulla. How- 
ever, Hiirzeler (1948:26) has pointed out that, although the 
l)ulla is overlapped b}^ the alisphenoids, this does not permit 
the definite statement that the alisphenoids constitute a part of 
the true bulla wall. In Montauban 9, it is possible to follow the 
suture between the ])ulla and its neighboring elements from the 
carotid foramen around the anterior end to the squamosal. The 
diverticulum D 2 (of Hiirzeler) appears to lie outside this 
suture. If then, as Hiirzeler suggested, the alisphenoid forms 
no part of the anterior bulla wall, this diverticulum (D 2) is 
extra-buUar. The broad overlapping of the external pterygoid 
plate of the alisphenoid onto the anterolateral bulla wall in 
Necrolemur (Fig. 2) is a feature of some interest in relating the 
Quercy form to the modern Tarsius. Cope (1885:467) long ago 
stressed the distinctiveness of this region of the tarsier basicran- 
ium when comparing it with the then newly discovered skull of 
an American Wasatchian prosimian, Tetonius homunculus. In 
both Necrolemur and Tetonius these external pterygoid plates 
overlap the bullae, as in Tarsius. Gregory (1920:227) gives the 
following as a general character of lemuroid Primates : ' ' The 
elongate pterygoid plates of the alisphenoids extended back to 
the auditory bullae, whereas in the Anthropoidea they are w^ell 
separated from them." A further distinction here is possible 
in that the posterior extremities of the external pterygoid plates 
in lemurs and lorises, including such fossil forms as SmUodectes, 
Nofharctus, Adapis, and Frouycticchus, typically (although not 
in all cases) reach back to the anterior tip of the bulla, but the 
area of contact is very small and cannot be described as over- 
lapping. Necrolemur, Tarsius, and Tetonius differ in this respect 



1960 CRANIAL AXATOMY OF NECROLEMUR 7 

from lemuriform, lorisiforin, platyrrhine, and catarrhine Pri- 
mates. 

Stehlin could not find the stylomastoid foramen in his speci- 
mens: Hiirzeler (1948) shows it in figures 27 and 28 at the 
posteroexternal angle of the bulla, but does not label it (see 
Fig. 1). Just anterior to this foramen is a fossette, probalily 
for reception of the stylohyal, if, as in Tarsius, the tympanohyal 
was not distinct (van der Klaauw, 1931:239). This foramen and 
fossette lie in the same depression and have a somewhat variable 
degree of separation. In M.C.Z. 8879 the external appearence 
is as a single oblong foramen, while in Paris 1 the two are more 
distinctly set off (Fig. 2). A specimen at the British Museum 
is intermediate in this regard. 

Contrary to Stehlin 's suggestion (1916 :1348) that a true post- 
glenoid i)rocess does not exist in Nccroloiiur, Paris 1 and 5 ex- 
hibit a process that can justly be termed postglenoid. Also a 
l)ostglenoid foramen is present (M.C.Z. 8879, Paris 1, 2 and 5) 
median to this process and between the posterior limit of the 
glenoid area and the external auditory tube. 

Two large foramina, one on either side of the alisphenoid 
pterygoidal wing, were described hy Stehlin (1916:1353-1354) 
as the inner and outer openings of the canalis civinninii (or 
foramen pterygospinosum, Stehlin 1912:1205). He named a 
smaller opening situated anterodorsally in the same region the 
foramen ovale. The position of this latter small foramen is vari- 
alilc. l)ut is always anterior to the glenoid fossa instead of being 
appi-oximately on a line with it, as is the foramen ovale in 
Primates generally. In those specimens in which this foramen 
is relatively large, a groove extends laterally and slightly pos- 
teriorly from it, which would probably not be the case if it were 
the foramen rotundum. Removal of matrix from Paris 1 and 2 
in the region of the external pterygoid plate or wall has re- 
vealed a foramen opening directly into the cranial cavity, lying 
within the wall at the juncture of canals from the three foramina. 
Given this information, probably not known to Stehlin, we sug- 
gest that the posteroexternal foramen (outer opening of Stehlin "s 
canalis civinninii) is the foramen ovale. 

In a footnote, Stehlin (1916:1354) cited Gregory (1915:430) 
as confirming his identifications in the region of the foramen 
ovale. Gregory does this only partially, committing himself no 
further than to say that the foramen ovale is on the external 
(as opposed to the internal) side of the external pterygoid Avail. 



8 



BREVIORA 



No. 127 




FIGURE 2 
Necrolemur antiquus x 3.5 



1. Foramen for branch of internal 6. 
maxillary artery 7. 

2. Foramen ovale 8. 

3. Postg'lenoid foramen 9. 

4. Stylomastoid foramen and fos- 10. 
sette for ? stylohyal 11. 

5. Foramen for auricular branch 
of pneumogastric 



Hypoglossal foramen 
Alisphenoid canal 
Opening of eustachian tube 
Internal carotid foramen 
Inferior petrous sinus 
Posterior lacerate foramen 



1960 CRANIAli ANATOMY OF NECBOLEMUR 9 

He did not definitely say that Stelilin's small foramen is the 
foramen ovale. However, he identified as the foramen rotundum 
the smaller foramen (Stehlin's f. ovale) regarded by us as being 
for the internal pterygoid branch of the internal maxillary 
artery. 

III. ORBIT 

Along its median Avail the orbit is composed principally of the 
frontal and the maxillary. Careful search of Paris 1 revealed 
no OS planum present in the orbital wall (Fig. 3). The lacrymal 
forms a narrow band within the orbit along the anterior rim. 
Frontal, parietal and alisphenoid comprise the posterior wall. 
Xo anterior sutures between the small orbitosphenoid, frontal, 
and palatine, respectively, could be made out in Paris 1, in which 
this region is entirely undistorted. Only a small palatine com- 
ponent is present in the orbit, and this does not separate the 
frontal from the maxillary. 

Stehlin (1916:1345) noted that his material was not adequate 
to allow determination of the maxillo-malar suture. This led 
him to suggest that the malar might reach the lacrymal. Paris 1 
and 2 show that this is not the case ; the maxillary makes up part 
of the orbital rim. AVhen discussing the Eocene lemuroid Xoih- 
arctus, Gregory (1920:227) remarked "... the malar if not 
in actual contact Avith the lacrymal certainly came very close 
to it, whereas in tarsioids and anthropoids it becomes widely 
separated from the lacrymal and limited to the outer side of 
the orbit." Consequently, yccrolcmur resembles the higher 
Primates in this regard, and not the majority of prosimians 
other than Tarsius. 

The absence of an ethmoid component in the rostral orbital 
wall of Necroleniur has been taken by some students as an indi- 
cation of a lack of affinity betAveen it and Tarsius, since in the 
latter the os planum is large. To the Avriters this distinction 
does not seem to have much significance. A primitiA'e prosimian 
condition, Avhere the ethmoid has no orbital plate, is retained in 
such forms as Necroleniur, Pronycticehus, SmUoclectes etc., in 
Avhich, perhaps, there has not been enough orbital expansion to 
effect an expression of this bone in the orl)it by impinging on 
the anterior part of the interorbital septum. In Tarsius and 
some Lorisif ormes the interorbital septum is A'ery narroAv — evi- 
dently an accommodation for relatively large eyes, and in both 
groups an os planum occurs. Moreover, only in Cheirogaleinae, 



10 



BREVIORA 



No. 127 



among living and fossil Lemuriformes, is the os planum present. 
In these small lemurs also, the occurrence of an os planum is 
coupled with large orbits (compared to body size) and a thin 
interorbital septum. For the Anthropoidea, a similar origin for 




FIGURE 3 
Necrolemur antiquus x 3.5 approx. 



1. Sphenopalatine foramen and 
posterior palatine canal 

2. Foramen rotiindum and ante- 
rior lacerate foramen ( ?coal- 
esced) 



3. Ethmoid foramen 
Al)breviation,s: 

as, alisphenoid 

fr, frontal 

7)(.r, maxilla 

OS, orbitosphenoid 

pi, palatine 

pr, parietal 

this orbital element may be considered. In most Ceboiclea the 
orbits restrict the interorbital septum to a thin plate on which 
the OS planum is exposed laterally. Although in Old World 
Anthropoidea the rostrum between the orbits is occasionally 



1960 CRANIAL ANATOMY OF NECROLEMUR 11 

rather broad, it is possible to posit that such breadth is secondary 
and tliat they descend in common from a form in which rela- 
tively large eyes impinged on the interorbital area enough to 
induce the appearance of an orbital ethmoid component. This 
hypothesis is strengthened by observed interorbital narrowness 
in the only known part of an Oligocene catarrhine skull (Simons, 
1959 :8). If the foregoing suggestions apply, then it is not neces- 
sary to expect the presence of an os planum in the stock from 
which Tarsins may have arisen. 

In Nccrolcmur (Paris 1), a small venous foramen can be seen 
situated near and beneath the median dorsal rim of the orbit. 
As in Tarsius, but apparently not in other Primates, below this 
foramen a deep groove curves posteroventrally and (in both) 
lies at a juncture between the plane of the lateral wall of the 
rostrum and that of the hack of the orbit. This is another un- 
usual feature (occurring in both Nccrolcmur and Tarsins) 
which has to be attributed to independent acquisition, by those 
who doubt that any known fossil prosimians have a close phyletic 
relationship to Tarsius. 

A cranio-orbital foramen exists in Paris 1, l)ut could not be 
found in Paris 2, 3 or 5. Running anteroposteriorly and slightly 
above the optic foramen is another groove. A small opening near 
its anterior end appears to be the ethmoid foramen. The sphen- 
opalatine foramen and the posterior palatine canal in Paris 1 
are combined to open posteriorly through a common large fora- 
men in the suture between maxillary and palatine. 

Apparently coalescence of the foramen rotundum and the 
anterior lacerate foramen occurs in Paris 1 but they are separate 
in Paris 5. In both cases, however, the foramen rotundum lies 
within the orbit, as Stehlin noted (1916:1353), and not lateral 
to the postorbital part of the alisphenoids, as Gregory (1915 :430) 
suggested. 

The orbital region of Paris 1 is entirely undistorted and shows 
that the postorbital opening was small. It is of some interest that 
neither lemurs nor lorises, nor any other known fossil prosimians 
of similar size possess a smaller aperture here. As with so many 
cranial characters the primate showing the most interesting re- 
semblance to NecroUmur in respect of the structure of the post- 
orbital region is Tarsius. However, because of the huge flanges 
that encircle the orbit in mature specimens of Tarsius, and the 
greater degree of postorl)ital closure seen in such adults, much 
more revealing comparisons can be made between skulls of 



12 BREVIORA No. 127 

Nccrolcmnr and those of juvenile tarsiers. In the latter, the 
C'ircuniorbital flanges are not yet very pronounced and resemble 
the slight flanges seen in Nccrolcmnr (Fig. 3) . It seems necessary 
to assume, first, that these flanges in Tarsius are concerned pri- 
marily with the support of the enormous eyes, and second, be- 
cause of their very uniqueness, that they were not so developed 
in Eocene forerunners of the living genus. Nccrolemur clearly 
has such flanges in an incipient stage. 

Closure behind the orbit in Tarsius (on the outside) can be 
seen to proceed in successively older juveniles from three main 
centers : 1, ventrolaterally, by an upgrowth of the posterior mid- 
region of the orbital plate of the maxilla; 2, laterally, by an 
anteroposterior spreading of the middle of the postorbital bar ; 
and 3, dorsolaterally, by growth of a flange from the frontal, 
which arises beneath the frontal insertion of the postorbital bar. 
Ossification proceeds downward as this flange, or plate, grows 
alongside the postorbital l)ar, and it eventually fuses with the 
bar, leaving no sutural indication. In most specimens of adult 
Tarsius, the frontal and jugal components of the postorbital wall 
can be distinguished by the fact that the region where they fuse 
is much thinner and consequently more translucent. In Ncc- 
rolemnr, at least two of these components effecting closure ap- 
pear to be partially developed. Paris 1 has an uncrushed post- 
orbital bar which shoAvs an anteroposterior spreading at the 
middle, much as in Tarsius. The evidence is less clear because 
of breakage, but the posterior part of the orbital plate of the 
maxilla also bears a flange in Nccrolcmnr. The third center of 
closure seen in Tarsius, the frontal element, is indicated in 
Nccrolemur only by a distinct angulation along the lateral wall 
of the cranium betwen the orbital and temporal fossae. Although 
such characters as tlie greatly reduced paraconids and the lo.ss 
of certain anterior lower teeth eliminate Nccrolemur from the 
direct ancestry of Tarsius'^, the incipient circumorbital flanges 
and characteristics of postorbital closure in Necrolemnr are sug- 
gestive of a stage to be expected in the Tarsius ancestry. Perhaps 
Pscudoloris or Nanvopifhex are nearer the actual line leading 
to the modern form, but incompleteness of known specimens pre- 
vents the sort of comparisons here made with Nccrolcmnr and 
leaves this possibility insoluble at present. 



^Even if this were not so it would be almost irrelevant, in the absence of 
intermeiliates, to urge an ancestor-desoendent relationship for forms so separate 
in geographic distribution and in time. 



1960 CRANIAL ANATOMY OF NECROLEMUR 13 

It may be noted in passing-, that the manner of postorbital 
closure in Tarsius (insofar as the malar and frontal are con- 
cerned) is distinct from that seen in catarrhines and platyrrhines. 
Closure in this area in Tarsius is chiefly effected by an outward 
and downward growth of a flange of the frontal (with relatively 
little malar expansion) while in higher Primates the greater part 
of the dorsolateral area of enclosure is contributed by the de- 
velopment of an orbital plate of the malar. These differences 
strongly imply that the partial postorbital closure of Tarsius only 
parallels that of the Anthropoidea and is not a character of their 
common inheritance. If it be agreed that some postorbital clo- 
sure arose at least twice among Primates, the possibility that this 
feature also was independently acquired in the ancestral platyr- 
rhine and catarrhine stocks can be more seriously entertained. 

EXPLANATION OF FIGURES 

The lateral and ventral views of the slaill of Necrolemnr are 
based on Paris 1, to which details of missing regions have been 
added from other specimens, principally M.C.Z. 8879. Mislead- 
ing stains and fractures in Paris 1 are largely omitted from 
these illustrations. Certain details of the anterior dentition and 
mandible are drawn from specimens figured by Stehlin (1916). 

REFEEENCES 

Cope, E. D. 

1885. The Lemuioidea and the Iiiseetivora of tlie Eocene period of 
Xortli America. Amer. Naturalist, 19: -l:o7-471. 

Edinger, T. and D. B. Kitts 

1954. The foremen ovale. Evolution, 8: 389-404. 

Gregory, W. K. 

1915. I. On the relationship of the Eocene lemur Notliwctus to the 
Adapidae and to other Primates. II. On the classification and 
phylogeny of the Lemuroidea. Bull. Geo. Soc. Amer., 26: 419-446. 

1920. On the structure and relations of Notharctus, an American 
Eocene primate. Mem. Amer. Mus. Xat. Hist., (n.s.) 3: 49-243. 

HURZELER, J. 

1946. Zur Charakteristik, systematischen Stellung, Phylogenese und 
Yerbreitung der Xeerolemuriden aus dem europiiischen Eocaen. 
Eclogae geol. Helvetiae, 39: 352-354. 

1948. Zur Stammesgeschiehte der Xeerolemuriden. Schweiz. palaont. 
Abh., 66: 3-46. 



14 BREVIORA No. 127 

KlAAUW, C. J. VAN DEK 

1931. The auditory l)ull;i in sonic fossil iiiamiiials. Bull. Aiiier. Mus. 
Xat. History., G2 : 1-352 

Le Gros Clark, W. E. 

1959. The antecedents of man. 374 pp. Edinbiiro-h University Press. 

Simons, E. L. 

1959. An antliropoid frontal bone from the Fayum Oligocene of 
Egypt: the oldest skull fragment of a higher iirimate. Amer. 
Mus. Xovit., no. 1976: 1-1(5. 

1960. Notes on Eocene tarsioids at the British Museum. Bull. Brit. 
Mus. Xat. Hist., Geol. Ser., in press. 

Stkhlin, H. G. 

1912. Die Siiugetiere des sehweizerischen Eocaens. Abh. Schweiz. 

Palaont. Gesell., 38: 1165-1298. 
1916. Die Saugetiere des sehweizerischen Eocaens. Ihid., 38: 1299- 

1552. 



BREVIORA 



Mitaseimim of Coimparative Zoology 

('A.Mi'.iui)(ii;, ^Iass. I )k(kmhkr lM), nXil) Ximi'-f.r 12s 

SIZE OF EXDOCEUOID CEPIIALOPODS i 
By Curt Teichert- and Bernhard KrM:MEL3 

The maximum size of fossil animal groups, ^vhether mammals, 
reptiles, or invertebrates has always been a fascinating subject 
of inquiry'', because phyletic size increase is one of the important 
trends that dominate the evolution of living things. In the case 
of large animals, tlie evidence is often hard to assemble because 
their remains are difficult to obtain, to transport, and to store. 
Squids are the largest living invertebrates and a tradition has 
been handed down in paleontological literature that the largest 
fossil invertebrates likewise arc to be found among the cephalo- 
pods, but few accurate data are to be found in published sources 
which are now readily available. 

Among the nautiloid cephalopods, it has long been suspected 
that the Endoceratida furnished the real giants, but no accurate 
measurements in support of this statement are available. 

Clarke (1897) stated that entire shells of Canicroccras pro- 
tfiformf, 10 to 15 feet long (3 to 5 uK^ers). had been found in 
the ]\Iiddle Ordovician of Minnesota, hi the same publication, 
Clarke figured an internal cast of part of a siphuncle, from the 
base of the body chamber to the adapical end of the spiess, 
which Avas 3 feet and 3 inches long. Miller and Kunnnel (1944) 
described and illustrated additional species of these :\Iiddle 
Ordovician endoceroids from ^Minnesota, which are deposited in 
the Carnegie Museum. One of their paratypes of Endoccras 
clarkei measured 750 mm long, is sejitate throughout and is 
not complete, adapically or adorally. The holotype of Endoiunts 
gracillimvm Miller and Kunnnel (1944) measured 070 nun in 

U'ublicatioii authorizea by the Pirectdr, V. S. Geological Survey. 
-U. S. Geological Survey, Denver, ('olo. 
= Museum of Comparative Zoology. 



2 BREVIORA No. 128 

length, again an iiicoiiiplctc six'ciiiicii consisting only of phrag- 
niocone. These same authors (IcscimIxmI a new species, Endoccras 
(lccora]i(nsf, on two ])ortions of tlic internal mold of the phrag- 
nioeone from the Deeorah formation, Winneskiek Connty, Iowa. 
The larger portion is ahout 62.") mm long and the length of the 
smaller measures about :>2() mm. They estimated the interval 
between the two pieees as about 11.1 mm, so tlie total length of 
this phragmoeone was about 1, ()()() mm. These authors also men- 
tioned that there is on display in the Chicago Natural History 
Museum a larger endoceroid that measures 6 feet in length. 

Teiehert (1927) noted the occurrence, in Middle Ordovician 
limestones of Estonia, of endoceroids as iiuicli as 5 meters long, 
but gave no further details. Flower (19.3.")) stated that specimens 
12 feet in length had been collected and added that he was "not 
wholly inclined to discredit a report of an endoceroid found 
in a ([uarry n(>ar Watertown, New York, which was measured 
before it was broken up and found to attain a length of .30 feet." 
As far as we have been able to ascertain, these somewhat vague 
statements are all that is presently available in the ])ublished rec- 
ord on the subject of the maximum size of endoceroid cep- 
halopods. 

It does not seem to Ix' gcurrally kno\\n that the Museum of 
Comparative Zoology at Harvard laiivei-sity possesses Avhat a])- 
pears to Ix' the largest fragment of an eiuloceroid cephalopod 
on display anywlicrc^ in the world. As Flower (1955) has stated, 
"the removal of even reasonably coiu])lete sjiecimens involves 
something vei-y close to ([uarrying oj)erations, storing them is 
anothei- ])roblem." The specimen in the collections of the Mu- 
seum of Comparative Zoology is, therefore, pi-obably unique 
in nniseums of the Avorld. 

The specimen measures 3,000 nnn in length but is not com- 
plete, adorally or adapically. In general the preservation is fair, 
but as a result of weathering and crusliing the full diameter 
of the conch is preserved only in one jilane, and in the other 
])lane the outer shell is removed exposing traces of septa and 
in places the siphuncle. The tirst recognizable septa are 500 mm 
from the adoral end but the whole s])ecinien could well ho 
phragmoeone as this adoral 500 nnn is slightly crushed and 
weathered and one cannot tell whether septa are present or 
absent. The adoral diameter of the specimen is 2(S() nnn. The 
couch ta|)ers at a uniform rate and the adai)ical diameter meas- 
ures 120 nnn. 



1960 



SIZE OF EXDOCEROID CEPIIALOPODS 




Figure 1 — Large endoeeroid on exhibit in the Museum of Comparative 
Zoology. 



BREVIORA No. 128 



rii I 



he s('i)ta sl()|)c adapieally at an an^lc of about 45° and in 
the mid-part of tlic spcciiiicn are spaced 17 to 20 mm apart. Tlic 
siphunele is visible only on tlie adapieal lialf of tlie specimen. 
About 1,000 mm from the adoral end of tlie shell the siphunele 
has a diameter of about 0" mm; at l.ToO mm from the adoral 
end of the shell the siphunele has a diameter of 75 mm. The 
first endocones ap])ear 2.000 nnu behind the adoral end of the 
shell. The spiess measures 510 mm in length. The surface of the 
shell bears faint annulations that are spaced approximately 10 
to 12 mm a]iart. 

SUMMARY OF MEASUREMENTS 

Length 3,000 iimi 

Adoral dianieter 280 mm 

Diameter 1,00(1 mm from adoral end 220 mm 

Diameter 1,750 mm from adoral end 170 mm 

Adapieal diameter 120 mm 

Diameter of siphunele 1,000 nun from adoral end 95 mm 

Diameter of siidium-le 1,75(1 mm fiom adoral end 75 mm 

Spiess length 510 mm 

A graphical reconstruction of the sludl on the basis of these 
measurements shows that the entire fossil from its presently pre- 
served adoral end to the a])(^\ may have measured about 
5.S00 nnn. 

The total length of the body chamb'er is a matter of guesswork. 
There are tew ])ublished and illustrated records of any straight 
fossil cephaloiiod shells, complete from apex to aperture, which 
are more than a foot or so long. In short shells the ratio of body 
chamber to i)hragmocone may l)e high, even larger than 1:1. 
With increasing total hmgth of conch, however, ratio of body 
chamber to ]ihragm()cone is likely to decrease, although no defi- 
nite figures can be stated, in a specimen of Act{)ioccras beloifoisc 
(Foerste and Teichert, 11)80, pi. 28), which was 450 mm long, the 
ratio of body chandicr to phragmocone was about 1 :2. Leith 
(1942) described a specimen of Ldinlxoccras lamhii (Whiteaves) 
which was 45.5 in. (1,155 mm) long. lie estimated the total 
length of the shell at 1,405 mm. The body chamber was almost 
wholly preserved and not more than 250 mm long. Ratio of 
body chamber to iihragmocone was thus 1 :4.6 in this specimen. 

It shotdd be noted, however, that both Acfi)toc€ras beloitensc 
and Laiiibcoccrds hinthii have body chambers with constricted 



1960 SIZE OF ENDOCEROID CEPHALOPODS 5 

apertures, whereas no endoceroids Avitli constricted apertures 
are known. It seems physiolop'ieally plausible that in large 
straight ccphalopod shells the animal shonld have a Ijetter 
"grip" on a body chamber with constricted ai)erture than on 
one with an unconstricted aperture; therefore, in shells which 
expanded uniformly from the ai)ex to the apertui-c. like the 
endoceroids, the animal itself, and thus its body chamber, should 
have been relatively larger. 

In a juvenile specimen of a straight ammonoid, Baculitcs 
o vat us, Trueman (1941) determined the ratio of length of body 
chamber to phragmocone as 1 :0.7, but in adult shells this ratio 
becomes much smaller. If we assume the ratio of length of body 
chamber to phragmocone in endoceroids to be more like that 
of Actinoceras hdoitense we arrive at a length of the body 
chamber for the Harvard Enduceras of 2,650 mm and for the en- 
tire shell of 8,150 mm, or 28 feet. This is a conservative esti- 
mate, yet close to the possible maximum figure of 30 feet men- 
tioned by Flower. 

Add to this the length of the tenacles which must have ex- 
tended a considerable distance in front of the aperture, cer- 
tainly no less than half the length of the l)ody chamber, and 
we have an invertebrate animal considerably longer than 30 
feet — a truly imposing size. Today's giant squid, Architeuthis, 
rivals and slightly exceeds in length the largest extinct endocer- 
oids. Sparck (1928) records specimens of Architeuthis dux 
from tlie North Atlantic, washed ashore on the Norwegian Coast, 
that have body lengths of up to 2 meters and tentacles as much 
as 10 meters long. The largest specimen to our knowledge is 
that of Aychiteuthis harveyi? recorded by Yerrill (1879, p. 196) 
which measures 624 inches (17 meters). There is a model in 
the ]Museum of Comparative Zoology of a specimen of Archi- 
t(itthls j)rl)u-(ps. which was washed ashor<' in Newfoundland, 
which measures about 15 meters in length. ^Nlore recently. Lane 
(1960, pp. 198-227) has critically reviewed a larger number of 
reports of tinds of and encounters with giant squids. He is in- 
clined to believe that individuals of Aychit( tffhis or some other 
genus, as yet undescribed, may reach overall hnigths of some 

70 feet. 

While the Harvard specimen represents by far the largest 
nautiloid cephalopod on which accurate data are now available, 
it is interesting to compare it with the largest aiuinonoid on 
record. This is Pachydiscus seppenradense Landois from the 



6 BREVIORA Xo. 128 

Upper Cretaceous of western (!ei-iiiaiiy (Landois, 1895, 1898). 
lu 1895, Landois fii-st described this fossil amiiiouoid whose shell 
was 1,800 iiini in diameter and in which tlie last camera was 
550 mm high. Landois' reconstruction provitled the animal with 
a body chambei- e(jnivai(Mit to only one-fonrth of a complete 
whorl. From this he concluded that Uie total diameter of the 
complete specimen of his ammonoid had been about 2,550 mm. 
From later studies (Tnieman. 1941) it is. however, likely that 
Landois' estimate of the len<ith and bulk of liA'ino- chamber was 
too low. If the body cham])er of PacJiydiscii.s .s( ppoiradciise was 
ecjuivalent, as is more likely, to three-fourths or one full volu- 
tion of the shell, the diameter of the adult shell of this ammonite 
would have l)een of the order of 8,500 nnn. oi- moi'(^ than 10 feet. 
A very approximate graphic plot of a shell of this kind shows 
that the total length of the shell of I'ncJn/disciis seppenradense, 
when unrolled, would have been o1' the order of 00 feet, or 
roughly- twice as long as that of the largest endoceroid. 

In another paper Landois (1898) attempted to estimate the 
weight of these giant cepludoixxis. ()n the basis of his (estimated 
measurements he arrived at a total weight of the ammonite 
as 1,455 kg, or 750 kg for the weight of the animal its(df. and 
705 kg for the weight of the shell. 

We shall abstain from any attempt to indicate exact weights 
of the large endoceroids. The order of magnitude Avas almost 
certainly the same as that iid'ei'red by Landois for the giant 
Pachydiscus, something of the order of 1 ton (about 1000 kgL 
It must be assumed that the weight of shell aiul siphuncle, 
which for a h^ngth of over 5,000 nun was entirely tilled with 
calcareous dei)osits. balanced tlie buoyancy provided by the 
empty camerae and confine;! the animal to a strictly beiithonic 
existence. Few, if any. fossil invei-tebrates ever surpassed them 
in bulk weight and si/e. 

One other point deser^■es attention : Phyletic size increase 
is a trend that as a rule continues until the end, or very close 
to the end of the evolutionary life of a particular group of 
organisms, as, for example, in tlie anuuonoids. The endoceratids. 
however, reached their maximum size long before the time of 
extinction, in fact relatively early in their evolution. In North 
America, as well as in northern Europe, endoceroid cephalopods 
survive to the end of the ( )rdovician period, but reach their 
maximal size during ]Middle Ordovician time (Teichert, 1930, 
pp. 235-236). 



1960 size of exdocekoid cel'iialopods 7 

References 

Clarke, J. ^f. 

1807. Tlu> Lower Si!uri;iii ( 'ciili.-ilopocUi of ^liinu'.sota. Geology of 
Minnesota, vol. o, pt. 2, Paleontology, jip. 7(il-812, pis. 47-60. 

Flower, li. IT. 

]955. Status of eniloccroiil classification. Jour. I'aleoutology, vol. 
•29, no. ?,, lip. 3119-:;71. 

FoERSTE, A. F., and Teickert, Curt 

1930. The actinoeeroids of east-central Xorth America. Denison Univ. 
Bull., Sei. Lab. Jour., vol. 25, pp. 201-296, pis. 27-59. 

Laxdois, H. 

1895. Die Riesenannnoniten von Sepjienrade, Pachj/disou!^, Zittel, Sep- 

penradense H. Landois. Westfal. Provinzial. Ver. Wiss. und 

Kimst. f. 1894/95, vol. 23, pp. 99-108, 2 pis. 
1898. Gewichtsverluiltnisse der Riesen-Anmioniten. Ih'ul., vol. 26, pp. 



Laxe, F. W. 

1957. Kingdom of the Octopus. Javrolds, London. 286 pp. 

Leitit, E. I. 

1942. Xotes on the cephalojiod Ldmbcoccras lamhii from Manitoba. 
Jour. Paleontology, vol. 16, no. 1, pp. 130-132, 1 text-fig., pi. 22. 

Miller, A. K., and Ktmmel, Berxiiakd 

1944. Rome large straight Ordovician cephalopods from Minnesota. 
Annals Carnegie Museum, vol. 30, pp. 19-38, 4 pis. 

Sparck, Ragiiar 

1928. Xordens Pyrcverden. Henrik Koppel, Copenhagen, 658 pp. 

Teichert, Curt 

l!t27. Der estliindische Glint. Xatur u. ]\ruseum, vol. 57, pp. 264- 

272, 7 figs. 
1930. Biostratigraphie der Porauiboniten. Xeues Jahrb. f. Mineral, 
etc., Beil. Bd., Abt. B, pp. 177-246, 8 tigs., 4 pis. 

True.max, a. E. 

1941. The ammonite liody-chaniher, with special reference to the 
buoyancy and mode of life of the living ammonite. Quart. Jour. 
Geol. Soc. London, vol. 96, pp. 339-378. 

Verrill, a. E. 

1879. The cephalopods of the northeastern coast of America. Trans. 
Conn. Acad. Sci., vol. 5, pp. 177-476, pis. 25-56. 



BREVIORA 



Miaseiim of Compsirative Zoology 



Cambridge, Mass. December 21, 1960 Number 129 



TYPE AND TYPE LOCALITY OP THE GULF COAST 

SPINY SOFTSIIELL TURTLE, 

TRIONYX SPINIFER ASPER (AGASSIZ) 

By Robert G. AVebb 

Museum of Natural History, The University of Kansas 

The currently accepted type locality of Trionyx spinifer asper, 
Lake Concordia, Louisiana, is in an area of intergradation be- 
tween three subspecies of Trionyx spinifer. Of the nine available 
syntypes of asper, none has been designated as a lectotype, and 
only one of the syntypes of T. s. asper is recognizable as belong- 
ing to that taxon. 

Abbreviations of names of museums from which specimens 
are mentioned are : KU, Museum of Natural History, University 
of Kansas ; MCZ, Museum of Comparative Zoology, Harvard 
College ; SM, Strecker Museum, Baylor University ; TU, Tulane 
University ; and USNM, United States National Museum. 

Agassiz (1857:406) described Aspidonectes (= Trionyx) 
asper as having : 

"... very coarse and large tubercles of the front and 
hind part of the carapace, which extend, behind, even over 
the bony shield, and are there supported by prominent 
warts of the bony plates. These bony warts exist in no other 
species with which I am acquainted: their form is very 
irregular, sometimes oblong and sometimes orbicular ; they 
also project more or less. Another marked peculiarity of 
this species consists in the greater bluntness of the extremi- 
ties of the jaws, which are more rounded than in Asp. spini- 
fer. The jugal arch is also broader. The difference between 
the males and the females is more striking in this species 
than in any other, the males being regularly oval, whilst 



2 BREVIORA No. 129 

the females are almost circular in their outline ... in 
younger specimens of Asp. asper there are . . . two or three 
l)lack lines separating the pale rim of the posterior mar- 
gin 
Several of the syntypes show "prominent warts of the bony 
plates" {supra), which appear posteriorly and principally on 
the seventh pair of pleurals toward the midline on the bony 
carapace (Plate 1). I have seen bony elevations, which are 
circular or elongate resembling short ridges, on the carapaces 
of specimens of Triomjx fcrox (USNM 4373, 55316, 62217) ; a 
photograph of a fcrox (Stejneger, 1944 :pl. 7) clearly shows bony 
prominences on the posterior part of the bony carapace. These 
bony elevations also occur in populations of T. spinifer (SM 
2552, 2558, Texas; USNM 54731, Iowa; USNM 100396, 100404. 
Louisiana) . Bony warts and ridges on the posterior part of the 
carapace are not diagnostic for T. s. aspcr. To my knowledge, 
the subspecies of Trionyx spinifcr are not distinguishable by 
characteristics of the skull, bony carapace or plastron. 

T. s. aspcr closely resembles the subspecies hartwcgi and 
spinifcr but differs in usually having two or more blackish lines 
paralleling the rear margin of the carapace, and usually in 
having the postocular and postlabial stripes united on the side 
of the head. Other characters mentioned above liy Agassiz seem 
not to be of taxonomic worth. 

Agassiz (op. cif .-.4:05-06) did not designate a type, mention 
l^rceise localites, or state the number of specimens that formed 
the basis of his description of Aspidonectes asper. Baur (1893: 
220) restricted the type locality of aspcr to Lake Concordia, 
Louisiana, but did not mention any specimens ; presumably 
Baur's action was based upon an examination of the bony cara- 
pace of USNII 012349, which shows the prominent bony ridges 
described and considered diagnostic by Agassiz and has "Lake 
Concordia, Louisiana" written in ink on the underside of the 
carapace. Barbour and Loveridge (1929:225) listed MCZ 1597 
and 1622 as cotypes. Stejneger (op. cit. :56-58) discussed some 
of the syntypes of asper, and also regarded the type locality 
as "Lake Concordia, La.," designating USNM* 12349 ( = 
012349) and MCZ 37173 as "cotypes." The type locality of 
T. s. asper is currently accepted as Lake Concordia, Louisiana 
(Schmidt, 1953:109). 

T. s. asper intergrades w-ith T. s. hartwcgi and T. s. spinifer 
in the lower Mississippi Valley (Conant and Coin, 1948:11). 



1960 TRIONYX SPINIFER ASPER 3 

The softshell turtles inhabiting the Mississippi River and its 
tributaries in Louisiana (including Lake Concordia) and Mis- 
sissippi represent an intergrading population of spmifer and 
hortwegi, and, to a lesser extent, asper. Most turtles from the 
Pearl River drainage and rivers that drain into Lake Ponchar- 
train adjacent to the east are typical of asper. Lake Concordia 
is a large oxbow on the west side of the Mississippi River in 
Concordia Parish, Louisiana. 1 was a member of a field party, 
from Tulane Uiiiversit}^ which collected three specimens (TU 
16524, 16524.1, 16524.2) of Trionyx spinifer from Lake Con- 
cordia on August 1-3, 1954. Because none of these turtles has 
the postocular and postlabial stripes united on the side of the 
head or any indication of more than one marginal line parallel- 
ing the rear margin of the carapace, none is considered referable 
to asper. 

Some of the nine syntypes discussed below were mentioned 
by Stejneger (op. cit.:51-58). 

(1) USNM 012349 (Plate 1) is represented by a bony cara- 
pace and may be considered the present type (lectotype), 
although never designated as such. ''Trionyx Ferox?, Lake 
Concordia, Louisiana, BLC Wailes, 1851" is written in ink in 
the same handwriting on the underside of the bony carapace. 
Suljsequently, "Ferox?" has been crossed out and "asper" 
added in pencil, and "012349" inked on tlie second pleural. A 
gummed label is pasted on the fifth rib, right side, and bears 
the inscription "asper Ag. (Type)." There is also an attached 
metal tag bearing the number 22676. The carapace has a maxi- 
mal length of 14.9 cm. and width of 12.7 cm. ; its size suggests 
that it is that of a female. On the seventh pair of pleurals and 
in line with tlie longitudinal sutures of the neurals are two, 
short, longitudinal, elevated, bony ridges. 

(2) USNM 01086 is represented by an intact bony carapace 
(19.8 cm. in length and 16.5 cm. in width), disarticulated parts 
of the bony plastron (epiplastron and preplastra lacking), and 
a skull including the lower jaw. There is an attached paper tag 
labeled "cotype." "Miss." is written in ink on the underside 
of the carapace. There are small, slightly-elevated bony warts 
on the sixth pair of pleurals (with few, less conspicuously de- 
veloped, on the seventh pair), and a semblance of ridging as 
seen on the carapace of USNM 012349. "\Vashington Lake" 
(crossed out in pencil), and "1086" are written in ink on the 
skull, which has a basicranial length (occipital condyle to tip 



4 BREVIORA No. 120 

of upper jaw) of 62.8 mm. Tlie locality "Washington, Adams 
Co." is written in pencil on the eard for USNM 01086 in the 
card file in the USNM. The specimen is certainly a female 
judging from the size of the skeletal parts. 

(3) USNM 01084 is represented only by the skull (lacking 
lower jaw) of a female, which has a basicranial length of 63.1 
mm., and the number "1084" and "Wash." inked on it. The 
skull is presumably from the same locality as USNM 01086. 

(4) MCZ 46621 is represented only by the skull (lower jaw 
present) of a female having a basicranial length of 63.0 nnu. ; 
it was received from B.L.C. Wailes. 

(5) MCZ 1622 has been photographetl by Stejneger (op. cit.: 
pi. 16), who mentioned a label in lead pencil, "No. 1622. Ti/pe 
Amyda asper (Agassiz) Lake St. John, Miss. W. Sargent leg. 
et don." (which I have not seen) {op. cit. :5S). Stejneger sug- 
gested that Lake St. Jolm was in Louisiana (op. cit. :footnote 
2) ; however, tlie caption for Plate 16 reads Lake John, Florida. 
Coin (1948:304), commenting on the last -mentioned locality, 
wrote that MCZ 1622 "... is listed in the museum catalogue as 
having come from Lake St. John, Mississippi." Lake St. John is 
a few miles north of Lake Concordia occurring in Concordia 
and Tensas jiarislies. Louisiana, in the Mississip]ii Uiver drain- 
age. 

MCZ 1622 is a young alcoholic specimen, probably a female, 
having a plastron 5.0 cm. in length, and a carapace 6.8 cm. in 
length and 6.0 cm. in Avidth. The postlabial and postocular 
stripes do not join on either side of the head. The undersurface 
is more heavily pigmented than that which 1 judge to be "nor- 
mal" for T. .s\ a^pcr. The most significant character is the lack 
of more than one dark marginal line posteriorly on the carapace ; 
there are small blackish dots posteriorly but these are Avidely- 
separated and not closely-set in a linear fashion to suggest a 
second marginal line. Neill (1951:19-20) believed this specimen 
to show "... some approach to spinifera.''^ I do not regard 
this specimen as representative of T. s. asper. 

Schwartz (1956:15), however, referring to MCZ 1622, stated 
that it ". . . differs in no way, so far as head and carapace pattern 
is concerned, from similarly sized specimens of T. s. ogassizi 
(= asper) from South Carolina and Georgia." It would be 
surprising to find specimens from South Carolina and Georgia 
having a pattern on the carapace resembling that of MCZ 1622 : 
Schwartz's photograph of a juvenile from South Carolina (op. 
cit. -.14:, pi. 3") is certainly asper. 



1960 TRIONYX SPINIFER ASPER 5 

(6) MCZ 46633 bears the locality '' Washington, Mississippi" 
written in ink on an attached paper tag. In a different hand, 
the data ' " AVashington, Miss. Pres. by B. L. C. Wailes" and the 
number "9" is written in ink on an unattached, folded piece of 
stiff, brownish paper that is held in place l)y a metal rod against 
the plastron. The data suggest that this specimen is from the 
same locality as USNM 01084 and 01086. 

MCZ 46633 is a stuffed, adult male having a plastron approxi- 
mately 11.0 cm. in length and a carapace 16.0 cm. in length. 
The postlabial and postocular stripes fail to join by a narrow 
segment on the left side of the head, but appear to be in contact 
on the right side. Bony, longitudinal welts, resembling those 
on the bony carapace of USNM 012349, appear posteriorly on 
the seventh pair of pleurals. The carapace is darkened laterally 
and sprinkled with widely -spaced, black dots ; there is no indica- 
tion of a second marginal line. The pattern on the carapace 
most closely resembles that of T. s. hartwegi. Therefore, I do 
not consider this specimen to be representative of T. s. asper. 

(7) MCZ 46615 bears an attached label having the following 
data written in ink: "Triouyx ferox SOUTHERN SOFT- 
SHELLED TURTLE Natchez, Miss. H. Wheatland coll'n." 
Subsequently '"Trionyx ferox" has been crossed out and 
''Amyda aspera" written in ink; the name ''spinifera" also 
has been added in pencil. The reverse side of this label bears 
the number "8" and the name "B. Chase" written in pencil. 
A small piece of paper also bearing the number "8" is pasted 
on the left side on the bony carapace. Presumably this specimen 
has some relationship to MCZ 46633 that bears the number 9. 

MCZ 46615 is a large, stuffed adult female having a plastron 
that measures approximately 33.0 cm. and a carapace 43.0 cm. 
in length ; the anterior edge of the carapace is studded with 
conical tubercles. Scattered, oblong, bony elevations adorn the 
seventh pair of pleurals. There is no pattern evident on the 
carapace, and the striping on both sides of the head is obscure. 
This large female might represent any of the subspecies spinifer, 
hartwegi or asper; there is no character that identifies the speci- 
men as T. s. asper. 

(8) MCZ 37173 is the stuffed specimen that Agassiz mentions 
liaving received from the University of Oxford (at Oxford, 
Lafayette County), Mississippi (op. cit. AOo) ; the specimen is 
discussed by Stejneger (op. cit.-.ol). The plastron is approxi- 
mately 23.5 cm. in length; the length of the carapace, wrinkled 



6 BREVIORA No. 129 

posteriorly, measures (in a straight line) approximately 29.5 cm. 
Although the tail extends noticeably beyond the posterior edge 
of the carapace and is presumably the basis for Stejneger refer- 
ring to MCZ 37173 as an adult male, its large size indicates that 
it is a female. A red paper label bearing the writing "A. asper 
Ag. Cotype" is pasted on the plastron. Elevated bony knobs 
appear toward the posterior margin of the bony carapace on 
the seventh pair of pleurals. There are well-defined stripes on 
the head, but the relationship of the postocular and postlabial 
stripes is obscure. There is no evidence of more than one dark 
marginal line paralleling the rear margin of the carapace. Hence 
the siDccimen is not recognizable as T. s. asper. 

(9) MCZ 1597, a large alcoholic female, is considered repre- 
sentative of T. s. asper (Plate 2). A paper label on the left 
foreleg bears the inscription "Natches, Miss W. Sargent." The 
carapace, measuring approximately 43.0 cm. in length and 37.0 
cm. in width, has more than one dark marginal line and several 
ocelli. Inner marginal lines in the posterior right and left 
quadrants are mostly continuous, but are obscured by the wrink- 
ling and scratching on the posterior part of the carapace. The 
tubercles on the anterior edge of the carapace are worn and 
resemble rounded knobs ; more lateral tubercles are equilateral. 
The seventh pair of pleurals are in contact medially behind the 
seventh neural. There are indications of raised bony welts on 
the last pair of pleurals. The head is partly extended, but does 
not show the relationship of the stripes on the side of the head. 
The pattern on the snout is obscured, and blackish marks are 
evident on the dorsal surface of the limbs. The plastral surface 
lacks dark markings and measures approximately 32.5 cm. in 
length. MCZ 1597 is discussed by Stejneger, who mistakenly 
refers to the specimen as a male {op. cit.:58). MCZ 1597 (Plate 
2) is herewith formally designated as lectotype of Trionyx spiyii- 
fer asper (Agassiz), as it alone, of the nine specimens hitherto 
considered syntypes of Aspidonectes asper Agassiz, is recogniz- 
able as referable to the subspecies asper. 

Agassiz (1857:405) mentions having received specimens from 
Mr. AVinthrop Sargent of Natchez, Mississippi (MCZ 1597, 
1622, 46615), Dr. L. Harper of the University of Oxford, Mis- 
sissippi (MCZ 37173), and Professor B. L. C. Wailes of Wash- 
ington, Mississippi (MCZ 46621, 46623, USNM 01084, 01086, 
012349). These localities suggest places of residence and are not 
necessarily localities at which the specimens were captured ; at 
least two syntypes (MCZ 1622, USNM 012349) are from locali- 



1960 TRIONYX SPINIFER ASPER 7 

ties different from those mentioned immediately above. Al- 
though forwarded to Agassiz from Natchez, the specific locality 
from which the lectotype (MCZ 1597) was captured is unknown. 
Natchez, Oxford and Washington, Mississippi, are in the drain- 
age basin of the lower Mississippi River, w-hich is inhabited by 
softshells that are intergrades between T. s. spinifer and T. s. 
hartwcgi, although few specimens from there are typical of 
T. s. asper. It is possible, but unlikely, that MCZ 1597 was 
captured at Natchez. Those syntypes having a discernible pat- 
tern on the carapace, which is not that of asper, probably came 
from tributaries in the lower Mississippi River drainage. 

The lectotype, MCZ 1597, probably came from the Pearl River 
drainage (adjacent eastward from the Mississippi River drain- 
age), where most individuals are representative of T. s. asper. 
Occasional specimens of asjJer from the Pearl River drainage 
have only one dark line paralleling the rear margin of the 
carapace, and resemble softshells occurring in the Mississippi 
River drainage. An adult male given to me by William E. Brode 
was stated by him to have come from the Pearl River ; this 
turtle (KU 47120) has only one dark marginal line, and Stej- 
neger (op. cit. :64) mentions another from the Pearl River drain- 
age. Schwartz (op. cit. -.16) and Crenshaw and Hopkins (1955: 
20) write that some specimens from Georgia have only one 
solid line at the margin of the carapace. However, most of 
the softshells inhabiting the Pearl River drainage are typical 
asper, and the Pearl River drainage is probably the provenance 
of the lectotype, MCZ 1597. 

The type loealitj^ of Trionyx spinifer asper (Agassiz), repre- 
sented by the lectotype, MCZ 1597, is herewith designated as 
the Pearl River at Columbus, Marion County, Mississippi. The 
geographic range of T. s. asper (Platypeltis agassizi Baur con- 
sidered a synonym) is the southeastern United States except 
peninsular Florida from the Florida parishes of Louisiana east 
to southern North Carolina ; in streams of the Gulf Coast drain- 
age including that of Lake Ponchartrain, Louisiana, eastward 
to the Apalachicola River system, and those of the Atlantic 
Coast drainage including that of the Altamaha River in Georgia 
northward to the Pee Dee River drainage in South Carolina. 



8 BREVIORA No. 129 

LITERATURE CITED 

Agassiz, L. 

1857. Contributions to tlie natural history of the United States of 
America. A^ol. I. Part II. North American Testudinata. Little, 
Brown and Co., Boston, pp. 233-452d. 

Barbour, T. and A. Loveridge 

1929. Tyijical reptiles and amphibians. Bull. Mus. Comp. Zoo!., 
69(10) :205-360, 

Baur, G. 

1893. Notes on the classification and taxonomy of the Testudinata. 
III. The genera of the Trionychidae. Proc. Amer. Phil. Soc, 
31:213-221. 

Conant, E. and C. J. GoiN 

1948. A new subspecies of soft-shelled turtle from the central United 
States, with comments on the application of the name Amyda. 
Occas. Pap. Mus. Zool., Univ. Michigan, 510:1-19, 2 pis. 

Crenshaw, J. W., Jr. and M. N. Hopkins, Jr. 

1955. The relationships of the soft-shelled turtles Trionyz fcrox ferox 
and Trionyx fcrox aspera. Copeia, 1955, No. 1:13-23, 4 figs. 

Goin, C. J. 

1948. The occurrence of Amyda spinifcra aspera in Florida. Copeia, 
1948, No. 4:304. 

Neill, W. T. 

1951. Taxonomy of North American soft-shelled turtles, genus 
Amyda. Publ. Res. Div. Ross Allen's Rept. Inst., l(2):7-24, 
1 fig. 

Schmidt, K. P. 

1953. A check list of North American amphibians and reptiles. Sixth 
edition. Amer. Soc. Ichth. Herp., LTniv. Chicago Press, pji. 
viii +280. 

Schwartz, A. 

1956. The relationships and nomenclature of the soft-shelled turtles 
(genus Trionyx) of the southeastern United States. Charleston 
Mus. Leaflet, 26:1-21, 1 fig., 3 pis., 2 maps. 

Stbjneger, L. 

1944. Notes on the American soft-shell turtles with special reference 
to Amyda agassizii. Bull. Mus. Comp. Zool., 94(l):l-75, 30 
pis., 10 tabs. 
Transmuted January 19, 1960. 




Plate 1 (X appiox. Vs). Bony carapace of Trionyx spinifer, USNM 
012349, from Lake Concordia, Louisiana; parts of ribs that extend beyond 
carapace on right side are detached. Top. — Dorsal view showing elevated 
prominences posteriorly that Agassiz considered diagnostic for Aspidonectes 
asper. Bottom. — Ventral view showing inscriptions. 




Plate 2 (X appiox. Yo). Dorsal view of lectotype of Trionyx spinifer 
asper, MCZ 1597 ; locality designated as the Pearl River at Columbus, 
Marion County, Mississippi. 



BREVIORA 

Mnaseiam of Compsirative Zoology 



Cambridge, Mass. December 22, 1960 Number 130 



THE MECHANISMS OF CAEAPACIAL AND 
PLASTRAL HINGES IN CHELONIANS 

By R. V. Shah 

Department of Zoology, University of Baroda, Baroda, India 

Deraniyagala (1930) described the carapace and plastron of 
Lissctnys and reported that there are two movable corselets in 
the carapace and four in the plastron. In the carapace the pre- 
nuchal is looseh^ connected to the anterior border of the nuchal 
by a thick ligament. This anterior corselet is moved downwards 
and acts as a valve. The last two marginals form a posterior 
movable valve in the carapace. In the plastron, the epiplastral 
lobe acts as an anterior plastral valve, and a small portion of the 
posterior part of the plastron demarcated by a transverse groove 
forms the posterior plastral valve. Besides these, there is a pair 
of cutaneous flaps which are used as valves to cover the posterior 
limbs after they are retracted under the shell. These flaps are 
known as the femoral valves. Presence of the cutaneous femoral 
valves is a distinguishing character of the subfamily Cyclan- 
orbinae, in which the three genera Lisscmys, Cyclanorhis and 
Cycloderma are grouped. Hasan (1941) worked out the shell- 
closing mechanism and described the principal muscles respon- 
sible for the movements of the six movable parts in Lisscmys. 
George and Shah (1955), while working on the musculature of 
Lisscmys, gave pos.able homologies of the muscles described by 
Hasan. 

In the present study an attempt is made to describe the closing 
mechanism of tlie shell in most ^ of the chelonians in which part 
of the plastron or carapace is movable and acts as a valve. Care- 

1 Xo adult was available of Kotochcli/s platynota in which an indistinct trans- 
verse hingp has been reported. Similarly there is only a juvenile at hand of 
Pyxis arachnoidcs. Siebenrock has described in this species a mobile anterior 
plastral lobe in which the entoplastron becomes involved. Since hinges only develop 
with age, it has not been possible to investigate hinge mechanisms in these species. 



2 BREVIORA No. 130 

ful dissections of the muscles involved in the movements of the 
plastral lobes or the carapace parts were made of the following 
chelonians: Cuora amhoinensis, Emys orbicularis, Emydoiden 
hlandingi, Terrapene mexicana yucatana, T. Carolina Carolina, 
T. c. haurii, T. c. triunguis, T. ornata, Testudo graeca, T. 
hermanni, T. kleimnayini, Kinixys erosa, Kinosternon iaurii, K. 
cruentatum, K. flavescens spooneri, K. herrcrai, K. leucostomum, 
K. scorpioides scorpioides, K. suhruhrum, Sternotherus carinatus 
minor, 8. odoratus, Cycloderma frenatum, Pelusios adansoni, P. 
carinatus, P. castaneus, P. nanus, P. suhnigcr. All specimens 
examined are in the collection of the Museum of Comparative 
Zoology. 

Only two genera of the chelonians listed above, viz., Cycloderma 
and Kinixys, show some part of their carapace movable ; the others 
have some part or parts of their plastra movable. In this regard 
all the chelonians used for the present study could be classified 
in four different groups : 

The first group has only the anterior plastral lobe hinged and 
capable of valvular movements. The hinge is usually situated at 
the joint of hyo- and hypoplastral plates. The species of emy- 
dines examined show this condition. 

The second group includes those that have only their posterior 
plastral lobes movable. The posterior plastral lobe is hinged at 
the level of the joint of the hypo- and xiphiplastral plates. This 
group is composed of the species of Testudo cited above. 

The third group includes those which have both anterior and 
posterior plastral lobes movable. The kinosternines examined 
belong here. 

The fourth group consists of the members of the subfamily 
Cyclanorbinae in which besides the movable anterior and pos- 
terior plastral lobes there are the two cutaneous femoral valves. 

MOVEMENTS OF CARAPACE VALVES 

In Cycloderma and Cyclanorhis, as in Lissemys, the prenuchal 
is movable. Hasan (1941) has described as the M. nucho- 
prenuchalis the muscle which is responsible for the movements of 
the prenuchal. This muscle arises from the under surface of the 
nuchal and is inserted on the posterior half of the under surface 
of the prenuchal. When this muscle contracts, the prenuchal is 
pulled downwards and acts as a small valve to close the shell after 
the head and the fore limbs are drawn under the shell and the 
anterior lobe of the plastron is lifted upwards. George and Shah 



IHGO TURTLE HINGES 3 

1,1955) hoinologised this muscle with that part of the spinalis- 
semispinalis system occurring in this region. In no other chelon- 
ians are there such movable prenuchals. 

The last two marginals in Cydoderma and Cyclanorhis, like 
those in Lissemys, are movable and act as a supracaudal valve. 
The )na)-cjino-i)ifracaudaIcs are responsible for the downward pull 
(Hasan 1941). These muscles arise from the undersurface of the 
marginals and are inserted on the flexible subcaudal valve of the 
plastron. AVhen these muscles contract the two movable marginals 
are pulled downwards and at the same time the subcaudal valve 
jf the plastron is pulled upwards. The joint action of these two 
valves, i.e. the subcaudal and supracaudal valves, close the shell 
from behind and protect the tail and surrounding soft parts. 
These muscles are homologised with part of the flexor caudac 
supcrficialis by George and Shah (1955). 

Tn Kinixys erosa an entirely different situation is present. 
Siebenrock (1916) has described the morphological features of 
the hinged carapace of this species in great detail. Briefly, how- 
ever, it may be mentioned that the hinge is at the level of the 
fourth and fifth costal plates, the posterior part of the carapace 
being moved up and down like a valve at this point. This move- 
ment of such a large part of the carapace is unique to this genus. 

The closing movement is brought about in the following man- 
ner. First the head and the neck are pulled under the shell as a 
result of the contraction of the rctrahens capitis coUiqiic, the 
rectus capitis cervico-plastralis and rectus cervicis muscles. Of 
these muscles the retrahens capitis coUiqiie is the principal muscle 
responsible for the withdrawal of the head and the neck. The 
main part of this muscle in Kinixys erosa arises from the body of 
the seventh and eighth dorsal vertebrae and the undersurface 
of the costal plates near these vertebrae. The muscle inserts on 
the cervical vertebrae and on the base of the skull at its ventral 
side. After the head and neck are retracted fully, the muscle 
contracts still further and this additional contraction pulls the 
hinged posterior part of the carapace downwards and closes the 
shell from behind. In addition, the pelvic girdle muscles, i.e. the 
attrahens pelvium, also help in pulling the hinged part of the 
carapace downward. The attrahens pelvium is very highly devel- 
oped in this genus and when it contracts the entire pelvic girdle 
is pulled forward, and since the girdle is firmly attached to the 
carapace the carapace is pulled with it. Although this pull is 
not so great as the one exerted by the retrahens capitis collique 



4 BREVIORA No. 130 

it does contribute to the movement. A few muscles of the legs, 
i.e. the ilio-tihialis and ilio-femoralis which in part arise from the 
costal plate, also exert a pull on the carapace after the legs are 
fully retracted under the shell. Thus a combined pull resulting 
from the contraction of all these muscles is responsible for bring- 
ing about the valvular movements of the posterior hinged part 
of the carapace. 

MOVEMENTS OF THE PLASTRAL LOBES 

Anterior plastral valve : The anterior plastral lobe is hinged 
at the level of the hyo-hypoplastral joint in Emys orbicularis, 
Emydoidea 'blandingii, Cuora amhoinensis, Terrapene mexicana 
yucatana, T. Carolina Carolina, T. c. haurii, T. c. triunguis, T. 
ornata, Pelusios suhniger, P. castaneiis, P. carinatus and P. 
adansonii. This anterior plastral lobe can be moved upwards like 
a valve which shuts off the shell in front after the head, neck and 
the fore limbs are retracted. This upward movement of the an- 
terior plastral lobe is brought about in the following manner : 
After the head, neck and the fore limbs are retracted, the clavic- 
ular part of the deltoideus muscle contracts and pulls the plastral 
lobe upwards. The anterior part of the 'pecioralis muscle, which 
has its origin from the plastral lobe, also on contraction pulls the 
plastral lobe upward. Some part of the scalenus muscle which is 
inserted on the border of the plastral lobe helps in puULng the 
lobe upward. The precoracoid cartilage is attached to the ento- 
plastral plate by a thick ligament and so when the whole girdle 
is rotated forward with the scapula acting as a fixed point the 
precoracoid and the coracoid which normally lie in a horizontal 
plane come now into an inclined plane with the precoracoid lifted 
upwards. Along with these the plastral lobe also gets pulled 
upwards and thus the rotation of the girdle helps in lifting the 
plastral lobe. These combined actions bring about the valvular 
movement of the anterior plastral lobe. 

Normally the anterior plastral valve has a straight transverse 
hinge. This disposition of the hinge is very necessary for free 
valvular movements of the plastral lobe. But to this normal 
condition there is an exception in Pelusios adansonii. In this 
animal the hinge is not a straight transverse one but is V-shaped 
with the apex directed posteriorly. Because of this type of hinge 
the valvular movements of the anterior plastral lobe in this animal 
are very restricted compared to those possible with a straight 
transversely placed hinged. 



1960 TURTLE HINGES 5 

Posterior plastral valve : A hinged posterior plastral lobe 
occurs in Tcsfiido Ideinmanni, Testudo gracca and Testudo her- 
manni. The hinge is at the level of the hypo-xiphiplastral joint 
and is also transversely placed. The posterior plastral lobe is 
pulled upward as a result of the contraction of the retrahens 
pclvium muscle of the pelvic girdle. This muscle arises from the 
spine of the pubis and is inserted on the entire inner surface of 
the xiphiplastral plate. In these animals the muscle is more 
highly developed than in forms which do not have a hinged 
posterior plastral lobe. Besides the action of the retrahens pel- 
vium muscle the skin connecting the legs and the plastron exerts 
some pull on the plastral lobe which helps in pulling it upward. 

Anterior and posterior plastral lobes : Kinosternon subrubrum, 
K. flavescens spooneri, K. leucostomum, K. scorpioides scorpi- 
oides, K. baurii, K. crucntatum, K. herrerai, Sternotherus carin- 
atus minor, and Sternotherus odoratus possess anterior as well as 
posterior plastral lobes which are hinged and are capable of 
valvular movements. The upward movements of these two plas- 
tral lobes are brought about in the same w^ay as described sepa- 
rately in the first two groups which had only one lobe, whether 
anterior or posterior, movable. 

In Kinosternon, as far as the anterior plastral lobe is con- 
cerned, there is an additional factor pulling the plastral lobe 
upward. There are thick tendons which loosely connect the 
fourth, fifth and sixth cervical vertebrae wdth the plastral lobe. 
The point of attachment of these tendons on the plastral lobe is 
at the border of the epiplastral plates. When the head and the 
neck are pulled under the shell as a result of the contraction of 
the retrahens capitis collique (the main retractor of the head 
and the neck), an indirect pull is exerted on the plastral lobe 
because the latter is connected by thick tendons with the cervical 
vertebrae. Such a type of tendinous connection between the 
cervical vertebrae and the plastron is not met with in any of the 
other chelonians studied. 

The Terrapene species examined have been placed in the first 
group in which only one anterior plastral lobe is hinged and 
movable like a valve. These species, however, require special at- 
tention since, although there is no true posterior plastral hinge, 
the posterior lobe is capable of being pulled dorsally and can 
close the shell posteriorly as efficiently as in those which have the 
posterior plastral lobe hinge. Here the bridge between the cara- 
pace and the plastron is very short and a large portion of the 
posterior part of the plastron is left unsupported. The retrahens 



BREVIORA No. 130 

pelviuni muscle in this animal is very highly developed and so 
when it contracts the posterior plastral border is lifted upward. 
This lifting occurs because the plastral plate bends under the pull 
exerted by the retrahens pelvium muscle. In this way, although 
there is no hinge at the level of the hypo- and xiphiplastral joint, 
the plastral plate closes the shell from behind, after the hind 
limbs and the tail are retracted within the shell. 

Four plastral valves : The fourth group of the chelonians in- 
cludes those which possess four plastral valves capable of valvular 
movements — the members of the subfamily Cyclanorbinae — 
Lissemys, Cyclanorhis and Cycloderma. Here the anterior plas- 
tral lobe is not as well demarcated and hinged as in the animals 
described above, but since the plastron is soft the movement is 
effected at the level of the posterior border of the entoplastral 
plate and the anterior border of the fused hyo-hypoplastral 
plate. This anterior plastral valve is moved upward by the con- 
traction of the specially developed muscle — the nucho-plastralis 
(Hasan, 1941), described as the trapezius by George and Shah 
(1955). This muscle arises from the nuchal plate, runs along 
the base of the neck, and is inserted on the epiplastral plate. 
When the trapezius {—nucho-plastralis) muscles contract the 
anterior plastral lobe is lifted upward to close the shell. In addi- 
tion, other muscles are also taking part in the closing mechanism 
of the anterior plastral lobe ; these are : the clavicular part of the 
deltoideus, the rectus capitis cervico-plastralis, and the part of 
the pectoralis muscle which has its origin on the movable part 
of the plastron. The skin connecting the legs and the plastron 
also exerts some pull after the legs are retracted under the shell. 
The trapezius muscle is found only in the Trionychidae. Thus, 
though the anterior plastral lobe cannot be completely closed in 
Trionyx gangeticus, to a certain extent it can be lifted up to pro- 
tect the retracted head, neck and fore limb. In this animal, as in 
the Cyclanorbinae, the trapezius muscle is present and works in a 
fashion similar to that in Lissemys (George and Shah 1955). 

The infracaudal valve in all the members of the subfamily 
Cyclanorbinae is a small posterior part of the plastron which is 
demarcated by a transverse groove and which lies behind the 
xiphiplastral plates. Here also there is no regular hinged con- 
dition. Part of the flexor caudae superficialis muscle described for 
the closing mechanism of the supracaudal valve in the carapace 
of these animals is used for moving the infracaudal valve upward. 
Thus, when this muscle contracts the supracaudal as well as the 



1960 TURTLE HINGES 7 

infracaudal valves are moved in opposite directions towards eacli 
other to close the shell posteriorly. 

For the protection of the hind limbs in these animals an extra 
pair of cutaneous flaps is developed. These are known as the 
femoral valves. Each femoral valve is more or less hinged with 
the side of the xiphiplastral plates and is capable of valvular 
movements. The closure of the femoral valves is brought about 
when the ligament connecting the shank of the leg and the valve 
is tensed, as the legs are retracted under the shell. The skin 
connecting the legs and the valves also exerts some pull on the 
valves when the skin is drawn in as the legs are pulled under the 
shell. 

Finally, it may be said that some chelonians have developed 
certain features like movable plastral or carapace parts by which 
they can close themselves in a box to protect their soft parts. 
Besides this, wherever the posterior plastral lobe or the posterior 
part of the carapace is movable, this capacity for movement also 
serves to facilitate the egg-laying process by widening the gap 
between the carapace and the plastron. 

ACKNOWLEDGMENT 

I am grateful to Dr. A. S. Romer for giving me all possible 
facilities to carry out this work at the Museum of Comparative 
Zoology at Harvard Universit}'. I am also grateful to Dr. E. E. 
Williams for his valuable suggestions in preparing this paper. 
This work was done during the tenure of a Fulbright Smith- 
Mundt scholarship. 

EEFERENCES 

Debaniyaqala, p. E. P. 

1939. The Tetrapod Reptiles of Ceylon, Vol. 1. Testudinates and Croc- 
odilians. xxxii-l- 412 pp. Colombo. 

George, J. C. and E. V. Shah 

1955. The myology of the chelonian trunk and tail. J. Anim. Moiph. 
Physiol., vol. II, no. 2, pp. 49-64. 

1957. The myology of the chelonian limb. 1. The forelimb of Lisseviya 
punctata. Ibid., vol, IV, no. 2, pp. 81-95. 

1958. The myology of the chelonian Ihnb. 2. The hind limb nmseula- 
ture of Lissemys punctata. Ibid., vol. V, no. 1, pp. 21-33. 



BREVIORA No. 130 



Hasan, S. I. 

1941. The shell and the mechanism of its closure in the Indian pond 
terrapin, Lissemys pimctata punclata (Bonaterre). Proe. Ind. 
Acad. Sci., vol. XIV, sec. B, no. 3, pp. 235-249. 

Hoffmann, C. K. 

1890. Schildkroten in Reptilien. Bronn's Klassen und Ordnungen des 
Thier-Reichs, Bd. 6, Abt. 3, pp. 1-442. Leipzig. 

SlEBENROCK, F. 

1906. Schildkroten von Ostafrika und Madagaskar, in Voeltzkow, Reise 
in Ostafrika in den Jahreu 1903-1905, Bd. 2, pp. 1-40. 

1916. Schildkroten aus dera nordlichen Seengebiet und von Belgisch- 
Kongo. Annalen des K. K. Naturhistorischen Hofmuseums, 
vol. XXX, pp. 1-12. 

ABBREVIATIONS 

At. P. Attrahens pelvium 

At.P.' Area of origin of attrahens pelvium 

Car. Carapace 

C.Carp. The position of the posterior hinged part of carapace in Kinixys 

when shell is completely closed 

C.Dl. Clavicular part of deltoideus 

C.DL' Area of origin of clavicular deltoideus 

C.S.Isc. Cut surface of ischium 

C.S.Isc' Area of cut surface of ischium where it is fused with f)lastron 

C.S.P.Isc. Cut surface of pubo-ischial joint which is fused with plastron 

C.S.P.Isc' Area of pubo-ischial attachment on plastron 

F.C.S. Part of flexor caudae superfieialis 

F.C.S.' Area of insertion of F.C.S. on plastron 

F.V. Femoral valve 

H.A.P. Hinged joint of the posterior plastral lobe 

H.A.P.' Place of valvular movement of the anterior plastral lolie in 

Lissemys 

H.P.P. Hinged joint of the posterior plastral lobe 

H.P.P.' Place of valvular movement of the subcaudal valve of plastron 

in Lissemys 

J.Car. Hinged joint in carapace in Kinixys 

P. Pectoralis 

P.' Area of origin of pectoralis 

P.C.Ca. Precoracoid cartilage 

P.G. Pelvic girdle 

P.G.' Shifted position of pelvic girdle in Kinixys when the shell is 

completely closed 



1960 TURTLE HINGES 9 

PI. Plastron 

R.Carp. The position of the posterior hinged part of the carapace in 

Kvnixys when the shell is fully opened 

E.C.C. Retrahens capitis collique 

E.C.C.P. Rectus capitis cervico-plastralis 

B.C.C.P/ Area of origin of rectus capitis cervico-plastralis 

Rt.P. Retrahens pelvium 

Rt.P.' Area of origin of retrahens pelvium 

S.Dl. Scapular part of deltoideus 

Sk.C. Skin connection between the two hinged parts of tin- (ara]i;ue 

of Kinixys 

Tu. Tendons connecting cervical vertebrae with plastron 

Tn.' Place of insertion of the tendons on the plastron 

Tr. Trapezius 

Tr.' Area of insertion of trapezius 



10 



BREVIORA 



No. 130 



Car. 



J, Cor, 



R.C.C. 







Fig. 1. Kinixys erosa: Side view showing extended neck and the position 
of the muscles, the hinged posterior part of carapace nnd the pelvic girdle 
when the shell is open. 



Car, 



J. Car. R.C.C. 




RG: At.P Rt.^^^G. 



Fig. 2. Kinixys erosa : Side view showing the retracted head and neck, 
the contracted muscles and the shifted position of the pelvic girdle when the 
shell is closed. 



I960 



TURTLE HINGES 



11 




/ 

tVVV 










<^^ -C.S.PISC. 



C.DI.' 

R.C.C P' 




C.S.RIsc' 



C.S.Isc' 



Fig. 3. Pelusios suhniger : Left, the superficial ventral muscles in the 
shell region exposed. Eight, the inner surface of the plastron showing the 
area of origin of various luuscles. 



12 



BREVIORA 



No. 130 



H.A.R 




Fig. 4. Terrapene Carolina Carolina: Left, the superficial ventral muscles 
in the shell region exposed. Eight, the inner surface of the plastron sho-wing 
the area of origin of various muscles. 



1960 



TURTLE HINGES 



13 



R.C.C.R 




Fig. 5. Testiido herma>nni: Left, the superficial muscles in the shell region 
exposed. Eight, the inner surface of the plastron showing the area of origin 
of the various muscles. 



14 



BREVIORA 



No. 130 












Fig. 6. Sternotherus carinatus minor : Left, the superficial ventral muscles 
in the shell region exposed. Eight, the inner surface of the plastron showing 
the area of origin of various niusuk-s. 



I960 



TURTLE HINGES 



15 



H.A.P 




Fig. 7. Lisscmys punctata: Left, superficial ventral muscles in the shell 
region exposed. Bight, the inner surface of the plastron showing the area 
of origin of various muscles. 



BREVIORA 



Mmsemim of Comparative Zoology 



Cambridge, Mass. December 30, 1960 Number 131 



A SECOND RECORD OF THE FOSSIL RODENT 
PALUSTRIMUS WOOD 

By Craig C. Black 

Carnegie Museum, Pittsburgh 

111 July of 1959, while working in the Goshen Hole area in 
eastern Wj^oming, Laura McC4rew took Sabra B. Black and me 
to a microfaunal locality about six miles west northwest of Fort 
Laramie National j\Ionument, just south of the North Platte 
River. A small surface .sample of isolated teeth and about twenty 
pounds of fiiie, gray channel sand were collected at that time. 

The following genera have been recognized in the assemblage 
to date: Prosciurus, Promylagaulus, Hcliscomys, Palustrimus, 
and Palaeocastor. The presence of Heliscomys and especially 
Prosciurus would indicate an earlv Miocene age. Until recently 
Proscii{7-ns was not known to occur after the late Oligocene, but 
it has now been recorded from a new basal Miocene forma- 
tion in South Dakota (Macdonald, personal communication). Its 
presence in this assemblage together with the definitely early 
Miocene Promylagaulus and Palustrimus would lead me to be- 
lieve that the fauna is lowermost Miocene. A detailed report on 
this fauna is deferred in the hope that a larger sample can be 
secured in the near future. The purpose of the present note is 
to call attention to this new early Miocene locality and to record 
the presence of Palustrimus in the assemblage. 

Palustrimus was described by AVood in 1935, from a single 
tooth, LM \ in the Yale Peabody Museum collections from the 
Upper John Day. The present specimen represents the second 
record for the genus and corroborates "Wood's original determi- 
nation. This occurrence is doubly important since the type of 
Palustrimus lewisi, Y.P.M. No. 10572, cannot be found in the Yale 
collections. There is a note with the label for the specimen which 
states that it has been lost since 1950. 



BREVIORA 



No. 131 



I take this opportunity to tliank Laura McGrew for showing 
us the locality, and Dr. J. T. Gregory for allowing me free access 
to the John Day rodent collections then in his care. The drawing 
is by Mr. James 0. Parley and was made possible by a grant from 
the Gulf Oil Corporation. 




Figure 1. Palustrimus sp., LM^, X30. 

Abbreviations used : 

Y.P.M. — Peabody Museum of Natural History, Yale Uni- 
versity 

M.C.Z. — Museum of Comparative Zoology. Harvard Uni- 
versity 



Order RODENTIA 
Family MURIDAE 
Palustrimus sp. 

Referred Specimen. M.C.Z. No. 7353, LM ^ 

Horizon and Locality. Lower Miocene. S. 15, T. 26 N., R. 65 
W., Goshen County, Wyoming. 

Description. The tooth is composed of three transverse lophs 
joined lingually by a high ridge. The valleys between the three 



1960 THE FOSSIL RODENT PALUSTRIMUS WOOD 3 

lophs are extremely deep and pass directly across the tooth to 
the lingual ridge. Each loph is composed of three cusps, the cen- 
tral cusp in each case being dominant and the marginal ones 
lower. The lingual cusps of each loph are joined together by the 
lingual ridge, which carries several smaller cuspules along its 
length. The lophs increase in width from front to back, giving 
the tooth a triangular appearance. The transverse valleys are 
much deeper than the notches between the cusps. The notch 
between the buccal and central cusps is quite deep on the an- 
terior loph, shallower on the central loph and almost completely 
absent on the posterior loph. There is a small accessory cusp 
rising from the floor of the posterior transverse valley just in 
front of the shallow notch between the buccal and central cusp 
of the last loph. The first and second crests are slightly convex 
anteriorlv, while the last crest is slightlv concave anteriorlv. 

The measurements (in mm.) are: anteroposterior, 2.45; width 
anterior loph, 1.35; median loph, 1.60; posterior loph, 1.77. 

Discussion. There is little doubt that this specimen is con- 
generic with Palustrimus. That it is conspecific with P. lewisi 
seems highly doubtful, however, but the present material does 
not warrant the erection of a new species. It differs from P. lewisi 
in the possession of a high, lingual ridge connecting the trans- 
verse lophs and bearing several accessory cuspules and in having 
convex rather than concave anterior and median lophs. In other 
respects the tooth is extremely similar to that of P. lewisi. Both 
have the three transverse crests composed of three cusps each. 

Unfortunately, until more material of this peculiar genus is 
known its taxonomic position eannot be definitely determined. 
It is like nothing else known among the rodents from the North 
American Tertiary. Wood's {op. cit.) assignment of the genus 
to the Muridae would seem to be the most likely one at present, 
although the possibility of a geomyoid relationship (Wilson. 
1949, p. 126) should not be overlooked. 

REFERENCES 

Wilson, R. W. 

1949. Early Tertiary rodents of North America. Carnegie Inst. Wash- 
ington, Publ. no. .584: 60-164, figs. 1-13. 

Wood, A. E. 

1935. Two new rodents from the John Day Miocene. Amer. Jour. Sci., 
(5) 30: 368-372, figs. 1-3. 



BREVIORA 



Miisenjnm of Cooiparative Zoology 



Cambridge, Mass. Dkcembeh '40, 1900 Number 132 



THE STATUS OF SFHAERODACTYLUS PICTUS, WITH 

COMMENTS ON THE DISTRIBUTION OF 

S. SPUTATOR AND S. SAB ANUS 

By Wayne King 

Departiiiont of Biology and Florida State Museum, 
Universitv of Florida 



Garmaii (1887:20), in his description of Sphaerodactyliis 
pictus, lists its range as the island of St. Christopher's (= St. 
Kitts), West Indies. Barbour, in his monograph of the genus 
(1921:263) and in his first elieeklist of Antillean amphibians 
and reptiles (1930:85) lists the range as St. Kitts. In his second 
(1935:10-1:) and third (1937:115) Antillean checklists, Barbour 
includes the island of Nevis in the range of picins, and states 
that it (pictus) is possibly a synonym of S. sputator. 

In his monograph, Barbour (1921:226) separates jnctus from 
sputator on the degree to which the dorsal scales are keeled 
(rather weakly in pictus, and strongly in sputator), and on the 
number of dorsal scales equal to the distance from the tip of the 
snout to the center of the eye (9 in pictus, and 10 in sputator). 
Examination of the types of pictus (MCZ 6071) from St. Kitts, 
and of a large series of sputator (MCZ 16598—16633, 16635— 
16641) from St. Eustatius (= Statia) indicates that the tirst of 
the two characters used by Barbour to separate these species is 
extremely subjective and of doubtful value. The second charac- 
ter, which involves allometric growth of the head and dorsal 
scales, varies from 7 to 10 scales in sputator. The number of 
dorsal scales in the "standard distance" of pictus is thus in- 
cluded. Further examination reveals no character of scutellation 
which will separate the two forms. 



2 BREVIORA No. 132 

(laniiau (1887:20) describes the color aiul pattern of pictus 
as: 

''Greyish with tliree or four roAvs of brown spots on 
each side. On the snout there is a brown band from 
each eye around the end ; a median band meets these 
on the rostral. Behind the eyes, on the head, there are 
six longitudinal bands of brown, four of which join to 
form two on the occiput, and these meet the laterals on 
the neck forming two which are continued above the 
shoulders. A light line across the forehead from one 
orbit to the other. Two or three light streaks, across the 
])ack of the head and neck, appear in some. On a very 
young one there are five narrow, transverse, dark-edged 
streaks of white between the eyes and the base of the 
tail. There are traces of brown blotches on the lower 
surface. ' ' 
A series of sputator in the Museum of Comparative Zoology 
indicates that the juvenile pattern of this species consists of 
white crossbands, with dark browai edges, on the neck, trunk, and 
tail. There are 5 to 8 of these crossbands between the level of the 
eyes and the base of the tail. The crossbands alternate with 
areas of dull brown ground color. The adult has a pattern of 
2 to 8 white crossbands, usually with dark brown edges, on the 
neck and shoulders. The white markings on the trunk are always 
edged, at least in part, with dark brown. The pattern may be in 
the form of crossbands, or may be broken into wavy lines or 
spots. The white markings may fade to a light tan. In a few 
individuals the dark brown edge of the dorsal spots tends to 
form longitudinal rows or stripes on the light ground color. Both 
juveniles and adults have a dark brown stripe on the canthus 
i-ostralis, and a white stripe connecting the orbits across the top 
of the head. From the above description it is evident that the 
variation which occurs in the color ]iattern of sputator includes 
the pattern thought to be characteristic of pictus (see Fig. 1). 
Since their scutellation and patterns are identical, Sphaero- 
dactylus pictus Garman should be considered a synonym of 
Sphaerodactylus sputator (Sparrman), and the range of sputa- 
tor should be extended to include the island of St. Kitts. 

A confusing factor in establishing the status of S. sputator is 
Barbour's reference (1923 :2) to its color pattern. lie states that 
sputator is: 

"... One of the dichromatic forms, as are so many of 
the large-scaled species — and perhaps others as yet 



19G0 



SPIIAERODACTYLUS PICTUS 




= "C s 

« I 

o ^ 

S5j CO .r 

■Id ''H 



cS 


-a 


K 


a 


*^^ 


cS 


CJ 




^ 


Q 


-♦- 




^ 




c 


OC 




CO 


«H 


o 




o 


O 


1—1 






CS 


^ 


^~, 


p 



c 

(5 
O 

1= 



0} 



CC 



■r T. ^_ o 



O OS > ^ 






s 









S o OQ 



o 



p 









(M -4J 



CS rH 05 -^- 



N 



"3 



■3 rt 



be 






^ '• o 



o 



4 BREVIORA No. 132 

little known. Tlie t.y])es are females evidently. The 
males are much smaller than the females, uniform grey- 
ish brown through life, or at the most with a few fine 
scattered dots usually on the head. The females are 
large, bulky and with a great variety of broken bands, 
blotches and spots of varying size. ' ' 
Although Barbour does not list the catalogue numbers of the 
specimens he refers to in this paper, he does state that they were 
collected on Statia in 1922 by James L. Peters. This series is 
MCZ 16598—16633, 1 6635— 16641. At the time of his writing, 
this series contained both (S". sputator and S. sabanus. This is 
evident in Barbour's reference to the large ''females" {= sputa- 
tor) with blotches, bands and spots, and to the small ''males" 
(= sahanus) of a uniform brown. ^ Examination of the present 
series of sputator shows little or no sexual dichromatism, and 
the snout-vent length of the males (31-35 mm., with a mean of 
32.8) is slightly greater than that of the females (28-35 mm., 
with a mean of 31.7) . 

On 10 July 1958, Dr. Walter Auffenberg and I collected an 
adult specimen of S. sputator in Basseterre, St. Kitts, thereby 
confirming the existence of this species at the type locality of the 
synonym, pictus. On 15 July 1958, we collected two adult 
sputator on St. Martin, 21/2 miles west, and 14 ™il6 north of 
Philipsburg, near Devil's Hole. 

The range of S. sputator, as indicated by the specimens avail- 
able to me at this time, includes the i-slands of Statia (MCZ 
16598—16633, 16635—16641 (29); IJMMZ 57010), St. Kitts 
(MCZ 6071 (3); UF 10038), and St. Martin (UF 10039 (2); 
PWII 474A, 606 (5)). Future collecting Avill probably estab- 
lish its existence on Nevis and Saba, the two remaining islands 
of the Saba to Nevis chain of islands. 

The range of 8. sahanus includes all of the islands of the Saba 
to Nevis chain of islands. It is found on Saba (MCZ 45215 —  
45217; USNM 103985—103993, 103995—104003), Statia (MCZ 
54010—54015 (158) ), St. Kitts (UF 10041 (9), 10042 (9), 10043 
(9), 30044 (9), 10045 (10); PWH 422; ITMMZ 83317), and 
Nevis (UF 10040 (3); PWII 414; UMMZ 83316 (2): MCZ 

1 The Hfihaitiis were later sfjuirateil from th<' series and recatalogued under tlie 
luiiiio I'lrgantiiliis as MCZ .54010 - .')401.'5. That they are not elegantuhis is now- 
dear since in addition to other features tliere is no crossbandinff in the juveniles 
iif this .series. Cochran (IJKiS) pointed out tiiat the juveniles of sgbanus are 
\iuniarl;ed lilie the adults, while Harbour (1921) descril>ed the erfwsbanded 
juveniles of flegaiitiiliis. Tliis confusion of elegantuhis and sabanus is not sig- 
uiticant for the present pai)er ; a study of tiiese and other Lesser AntiUean 
sphiierodactyls, including a redetinition of all the species, is in preparation and 
will be presented later. 



1960 SPHAERODACTYLUS PICTUS 5 

38374). Barbour's record of pictus {= sputator) on Nevis 
(1935:104 and 1937:115) seems to be based on the Museum of 
Comparative Zoology specimens of sahanus listed above. 

The fieldwork during the summer of 1958 was supported by 
National Science Foundation Grant G-3896 and the Florida 
State Museum. Specimens collected on this trip have been placed 
in the University of Florida Collections, Gainesville (= UF). 
I would like to thank Dr. Ernest Williams (Museum of Com- 
parative Zoology, Harvard = MCZ), Dr. Doris Cochran (U. S. 
National Museum, Washington = USNM), Dr. Norman Hartweg 
(University of Michigan Museum of Zoology, Ann Arbor = 
UMMZ) and Dr. P. Wagenaar Hummelinck (Der Rijks-Uni- 
versiteit Zoologisch Laboratorium, Utrecht = PWH ) for the loan 
of specimens in their care, and Dr. Walter Auffenberg, Dr. 
William Riemer, Andrew Arata, and Charles Myers for their 
criticism and interest. 

LITERATURE CITED 

Barbour, Thomas 

1921. Sphnerodactylus. Mem. Mus. Comp. Zool., vol. 47, no. .3, pp. 

217-278. 
1923. West Indian investigations of 1922. Occ. Papers Mus. Zool. 

Univ. Michigan, no. 132, pp. 1-9. 
1930. A list of Antillean reptiles and amphibians. Zoologica, vol. 11, 

no. 4, pp. 61-116. 
193."). A second list of Antillean reptiles and amphibians. Zoologica, 

vol. 19, no. 3, pp. 77-141. 

1937. Third list of Antillean reptiles and amphibians. Bull. Mus. 
Comp. Zool., vol. 82, no. 2, pp. 77-166. 

Cochran, Doris M. 

1938. Reptiles and amphibians from the Lesser Antilles collected by 
Dr. S. T. Danforth. Proc. Biol. Soc. Washington, vol. .51, pp. 
147-156. 

Garman, Samuel 

1887. On West Indian Geckonidae and Anguidae. Bull. Essex Inst., 
vol. 19, pp. 17-24. 



BREVIORA 

Museum of Comparative Zoology 



Cambridge, Mass. Fkbiu'ary 27, 19()1 Xu.mbek 138 

OX THE GENERIC LIMITS IX THE FAMILY PJLIDAE 
(PROSOBPvAXClIlA: MOLLUSCAji 

By Edward H. Michelsox 

Department of Tropical Public Health, Harvard School of 
Public Health, Boston, Massachusetts 



Members of the moUuscan family Pilidae have b?eii known to 
science since pre-Linnaean times. Although the family has been 
defined, the generic limits — and particularly generic relation- 
ships — require further clarification. Morphologic investigations 
have been conducted on representatives of individual species, but 
only rarely have these studies been of a comparative nature. 
An attempt has been made in the present paper to review and 
collate the available information upon which generic limits may 
be established. Studies on the comparative morphology of the 
kidney and the ])enial complex are also presented. 

In the following discussion the family Pilidae will be con- 
sidered to consist of seven genera. Included in the genera Pila, 
Lanistes, Afropomns, and Saitha are the Old AVurld species: 
members of Poniacea, Marisa, and Asolenc constitute the N^ew 
World species. These genera have been erected primarily upon 
conchological characteristics based on such criteria as color, size 
and shape of the shell, types of sculpturing (if present), and the 
presence or absence of an umbilicus. 

11 

Attempts have been made by others to divide the family into 
two major groups, the Old and New World forms, on the pres- 
ence or absence of a calcareous operculum. Newer knowledge 

1 This study was supported (In part) by a research grant (E-513-C) and a 
training grant (2E-4f)( from the National Institutes of Allergy and Infectious 
Diseases. National Institutes of Ilealtli, I'ublle Health Service. 



2 BREVIORA No. 133 

has shown that this criterion is unsound, since only PiJa has a 
calcareous operculum. However, the phylogenetic significance 
of the operculum even in Pila is limited since calcification is a 
secondary process which occurs after the snail hatches (Ranjah, 
1942). 

The longisiphonate or brevisiphonate nature of the respiratory 
siphon has also been used as a criterion for the separation of 
Old and New World species, and appears to have some validity. 
Pila and Lmiistes are brevisiphonate, and the present study in- 
dicates that the siphon of Afropomus is similar. Saulea can only 
tentatively be accepted as a valid genus since no description of 
its anatomy exists. The New AVorld genera, Pomocea and Marisa, 
are longisiphonate; however, AiioJene has been reported (Scott, 
1943) to have an aberrant siphon (brevisiphonate ?), and may 
be closely related to the Old World genera. It has been suggested 
that the morphology of the siphon is not of phylogenetic signifi- 
cance, but reflects an adajitation to ecological conditions 
(Prashad, 1925). 

The radulae of the Pilidae are all taenioglossate and have the 
formula 2 :1 :1 :1 :2. Intra-specific variations, however, reduce 
the value of radular morphology at levels below the family. The 
genus Turhinicola w^as erected on the basis of radular morphology 
(Annandale and Prashad, 1921; Prashad, 1931) ; however, Pils- 
I)ry and Bequaert (1927) consider this group to be no more than 
a subgenus of Pila. 

The eggs of the various species provide several promising and 
characteristic differences which may aid in arriving at taxonomic 
limits of the genera. Lipochromes, which color the eggs or egg 
shells, occur in species of Pomacea, but are absent in the eggs of 
Pila and Lanistcs (Comfort, 1947). Eggs of Marisa cornnarietis 
are peculiar in possessing an orange pigment when first depos- 
ited, but this soon disappears (Michelson, 1956). 

The presence of an egg-shell is biologically significant and may 
be of phylogenetic importance. Both Pila and Pomacea produce 
such eggs, and in both cases the eggs are deposited out of water. 
Marisa is completely aquatic and its eggs are gelatinous. Eggs 
of Lanistes were initially reported as membranous by D'Ailly 
(Pilsbry and Bequaert, 1927), but recent- observations have 
demonstrated that they are gelatinous and are deposited below 
the water-line (McMahon et al., 1957). Information concerning 
the eggs of Asolcne is limited to a report by von Ihering {vide 
Pilsbry, 1933) in which it is stated that the eggs are gelatinous. 
The eggs of Saulea and Afropomus have not been described. 



1961 GENERIC LIMITS IN THE PILIDAE 



ITI 



The soft parts of 8 species of Pilidae (representing the genera 
Pilo, Lanisies, AfropoinKs, Pomorra, and Mnn'sa) were examined 
for characteristic anatomical differences. Only two structures, 
the kidney and the penial complex, appeared promising in this 
respect. 

The identity and sources of the material used in this study 
are presented in Table 1. Specimens of Pila, Lanistcs, Afro- 
pomus, and the South American Pomacra were obtained from 
the collections of the Museum of Comparative Zoology, Harvard 
University, through the courtesy of Dr. William J. Clench. 
These specimens were fixed either in Bouin's solution or 70% 
ethyl alcohol and sul)sequently stored in TO*^! alcohol. Speci- 
mens of Marisa and Pomacra palndosa obtained from laboratory 
colonies were first relaxed in boiled water and subsequently fixed 
in Bouin's, Zenker's, or Newcomer's solution. Since not all 
specimens were fixed in the same way only gross and micro- 
anatomical features were studied. For micro-anatomical study, 
tissues were embedded in paraffin, sectioned at 0-12.5;;., and 
stained with Lillie-Mayer hemalum and eosin. A total of 82 
snails were examined including at least 5 specimens of each 
species. 

A. The Kidney. The kidney of members of the Pilidae is com- 
posed of two distinct regions, an anterior and a posterior cham- 
ber. The anterior chamber is a discrete tubular structure that 
partially extends into the mantle cavity opening into it through 
an excretory pore. The posterior chaml)er is embedded in its 
entirety in the visceral mass. This chamber is not compact, but 
consists of a large vacuolated area surrounded laterally and 
ventral ly by a thin, transparent membrane, and bounded dorsally 
by a large shield-like mass of tissue. The posterior chamber is 
further limited anteriorly Ijy the pericardial membrane, although 
access to the pericardium is provided by the renopericardial pore. 
Descriptions of the kidney of specimens representing the various 
genera follow : 

Pila. In P. gloho.sa (Fig. IE) the dorsal surface of the pos- 
terior chamber is broadly rectangular and measures approxi- 
mately 1.8-2.0 times the length of the anterior chamber. The 
dorsal surface is broAvn and blood vessels are not prominent. The 
anterior chamber is triangular and is so oriented that its main 
axis is continuous with the axis of the posterior chamber. The 
kidney in the species from Siam was morphologically similar. 



& 



BREVIORA 



No. 133 



Lanistcs. Tn L. bolfancanus (Fig. 1 C) the dorsal surface of 
the posterior ehaniber ditt'ers from that of PUa in being liroader 
anteriorly. The ratio of the length of the j)osterior elianiber to 
the anterior chamber is approximately 0.8 :1 to 1.1. The anterior 
chamber is considerably longer than in Pila and is triangular 
in shajie. The main axis of the anterior chamber in Lanistcs is 
also oriented so that it is continuous with the axis of the ]iosterioi' 
chamber. 




Figure 1. Senii-diagraninuitic sketch of the dorsal surface of the kidney 
in five species of Pilidac: (A) Afropomus halanoideus, (B) Pomacca 
paludosa, (C) Lanistcs holtaneaniis, (D) Maiisa cornuarieiis, (E) Pila 
fjlobosa. The posterior chamber (p) and the anterior clianilier («) of each 
kidney illustrated arc oi'iented as in Figure lA. 



1961 OENERIC LIMITS IX THK IMLIDAF, 

Afropomns. Tlie kiclm-N' in ^1. balanoidciis {Fig. 1 A) differs 
most radically from those of the other genera. The posterior 
ehamher is distinctly triangular in shape and its apex is reflected 
to the right. Furthermore, it is the only form in which blood 
vessels are prominent on the dorsal surface of the kidney. The 
anterior chamber is irregularly rectangular in shape, and its 
axis lies at an obtuse angle to that of the posterior chamber. 

Pomacea. The morphology of the kidney was similar in the 
four species examined. In P. paludosa (Fig. 1 B), the posterior 
chamber is broadly rectangular and has a prominent protuber- 
ance situated anteriorly on its left margin. The anterior cham- 
ber is irregular in shape and its long axis lies at an obtuse angle 
to the axis of the posterior chamber. 

Marisa. The kidney of M. cornuayidis (Fig. 1 D) is very 
similar to that found in species of Pomacea, thus further 
strengthening the suggestion that Marisa, should be considered a 
subgenus of Pomocea (Baker, 1930; Pain, 1950). The lateral 
protuberance of the posterior chamber in Marisa is larger and 
fartlior anterior than in Pouiacca. In addition, the posterior 
chamber surrounds the anterior chamber more completely in 
Marisa than in Pomacea 

B. The Pcnial Complex. The penial complex in the members 
of the Pilidae arises from the mantle as a finger-like projection. 
It consists primarily of a large outer penial sheath Avhich en- 
folds the true penis. Since there is no direct connection betAveen 
the i^enis and the vas deferens, sperm must be transmitted from 
the latter organ to the former, and thence to the female. Sach- 
watkin (19201 first described the presence of an internal sperm 
canal in the penis of Ampullaria gigas. Prashad (1925), how- 
ever, found that in Pila glohosa an external sperm canal was 
present. Our study indicates that both workers were correct 
and that an internal canal is characteristic of the New World 
species and an external canal characteristic of the Old AYorld 
species (Figs. 2-4). 

TV 

Although additional studies will be needed to establish the 
generic limits in the family Pilidae, there appears now to be 
sufficient information to separate the Old from the New World 
genera, as shown in Table 2. 



BREVIORA 



No. 133 



It is apparent from the foreg'oinfi: that sufficient data are not 
available to permit a critical interpretation of the phylogenetic 
relationships within the family. Nevertheless, there appears to 
be an evolutionary trend towards the establishment of the family 
in the terrestrial biotope. The presence of a respiratory sac, in 
addition to gills, suggests a mori^hological adaptation for the 
transition from the aquatic to the terrestrial habitat. Species in 
two genera {Pila and Poniacca) are highly amphibious and 
even deposit their egg masses out of water ; the presence of a 
calcareous egg shell further reduces the dependence of Pila and 
Pomacea on an aquatic habitat. If the premise is accepted that 
the family Pilidae is evolving towards a terrestrial mode of 
life, then we must conclude that both Pila and Pomacea repre- 
sent evolutionary advances in that direction. 

Table 1 

The Specific Identity, Origin, and Number of Specimens 

Examined 

No. Specimens 
Species Origin Examined 



Pila gJohnsa Swainson 

Lanitites bolt an (anus (Eodiiig) 
Afropomus halanoidcus (Gould) 
Marisa cornuariclis (Linne) 
Pomacea paludosa Say 
Pomacea interrupta Soweiby 
Pomacea columellaris Eeeve 
Pomacea nitilia Eeeve 



Cakntta, India, 

Bangkok, Thailand 
Cairo, Egypt 
Liberia 

Rio Picdias, Puerto Eico 
Miami, Florida 
Chonta anticline, Peru 
Huanuco, Peru 
Iluanuco, Peru 



/ 

5 
10 

8 
15 
12 
10 
10 

5 



Table 2A 

Diagnostic Characteristics of the ]\Iajor Genera of the Family 





Y 


'ilidae i 








Shape of 




Eespiratory 


Sperm 


Genus 


Shell 


Operculum 


Siphon 


Canal 


Pila 


Dextral; sub-ovate 
to globose 


Calcareous 


Brevisiphonate 


External 


Lanistcs 


Sinistral; sub-ovate, 
turbinate, or 
earinate 


Corneous 


Brevisiphonate 


External 


Afropom us 


Dextral ; globose 


Corneous 


Brevisiphonate 


External 


Saulea 


Dextral; sub ovate 


Corneous 


? 


? 


Pomacea 


Dextral; sub-ovate, 
ovate, or globose 


Corneous 


Longisiphonate 


Internal 


ilarisa 


Dextral ; secondarily 
planorboid 


Corneous 


Longisiphonatc 


Internal 


Asolene 


Dextral; sub-ovate, 


Corneous 


Longisiphonate 


? 



ovate, or neritoid 



1 The morphological characteristics ol' the kidney of each genus are presented 



1961 



GENERIC LIMITS IN THE PILIDAE 



Table 2B 

Dia<inosti(' f'liaraeteristics of the major Genera of the Family 

Pilidae^ 





Nature of 


Oviposition 




Geographical 


Genus 


Eggs 


Site 


Behavior 


Distribution 


Pila 


Calcareous shell ; 


In banks or 


Highly am- 


Africa and 




non-pigmented 


mudflats 
near water 


phibious 


Asia 


Lanisies 


Lacking sliell : 


On submerged 


Aquatic or 


Africa 




gelatinous; 


vegetation, 


slightly 






non-pigmented 


etc. 


amphibious 




Afropomus 


? 


? 


f 


Africa 


Saulea 


? 


? 


? 


Africa 


Pomacra 


Calcareous shell ; 


On emergent 


Moderately 


South America ; 




pigmented 


parts of aqua- 
tic vegetation 


amphibious 


Central 
America; 
West Indies 
Southern U.S. 


Uarisa 


Lacking shell ; ge- 


On submerged 


Aquatic 


South America ; 




latinous and pig- 


vegetation, 




West Indies 



Asohne 



mented when etc. 

tirst deposited 



Lacking shell ; 
gelatinous; 
pigmented ( ? ) 



On submerged 
vegetation, 
etc. 



'? Aquatic 



South America 



1 The morphological charaLteristics of the kidney of each genus are presented 
iu Fig. 1. 



BREVIORA No. 1^^ 



Figure 2. Cross-section tlirough tlie penial she;itli (ps) and penis (2') 
of a specimen of Pomacca paliulo.'in. The location of tlie internal sperm 
canal is indicated by the arrow. The anatomy of the penis demonstrated 
in this figure is similar to that observed in all siiecies of romnrea exam- 
ined. Ilemaluin and eosin, X47. 

Figure ?>. Cross-section through the penial sheath and penis of a specimen 
of Mnrisa cornunrictis. The arrow points to the internal sperm canal, 
iremalnm and eosin, X29. 

Figure 4. Cross-section through the penial sheath and penis of a speci- 
men of lAinistcs holtaneanus. The arrow points to the external sperm canal. 
A similar type of sperm canal was found in the penes of specimens of 
Pila glohosa and Afropomus halanoideus. Hemalum and eosin, X43. 



l')61 



GENERIC LIMITS IX THE PILIDAE 




X 



x. 



7} 




•"5 



> ^ 



,^t 



5i; 








]0 BREVIORA No. 133 

LITERATURE CITED 

AriNANDALE, N. niirl B. PRASHAn 

1921. Materials for a generic revision of tlic fresh-water gastropod 
molluscs of the Indian Empire. No. 4. The Indian Ampulla- 
riidae. Reo. Indian Mus., Oalcutta, 22:7-12. 
Bakeir, II. B. 

1930. The mollusea collected by the University of ^lichigaii-Williani 
son Expedition in Venezuela. Occ. Pap. Mus. Zool., Univ. 
Michigan, No. 210, 94 pp. 

Comfort, A. 

1947. Lipochronies in the ova of Fila. Nature, 160:333-334. 
McMahon, p., T. von Brand, and M. O. Nolan 

1957. Observations on tlie polysaccharides of aquatic snails. J. Cell, 
and Comp. Physiol., 50:219-240. 
MiCHELSON, E. H. 

1956. Studies on the liiology of the genus Crrafodes (Mollusca: 
Pilidae). Doctoral Dissertation, Harvard Univ., 171 pp. (Pub- 
lished in part.) 
Pain, T. 

1950. Pomacea (Ampullariidae) of British Guiana. Proc. Malac. Soc 
London, 28:63-74. 

PiLSBRY, II. A. 

1933. Zoological results of the Matto Grosso expedition to Brazil in 

1931, II. Mollusca. Proc. Acad. Nat. Sci., Philadelphia, 85:67- 

76. 
PiLSBRY, H. A. and J. Bequaert 

1927. The aquatic niollusks of the Belgian Congo, with a geographical 

and ecological account of Congo malacology. Bull. Amer. Must 

Nat. Hist., 53:69-602. 
Prashad, B. 

1925. Anatomy of the couuuon Indian apple snail Pila ylobosa. Mem, 

Indian Mus., Calcutta, 8:91-151. 

1931. Some noteworthy examples of parallel evolution in the molluscan 
faunas of South-eastern Asia and South America. Proc. Roy. 
Soc. Edinburgh, 5:42-53. 

Ranjah, a. R. 

1942. The embryology of the Indian apple-snail, Pila globosa (Swain 
son) Mollusca, Gastropoda. Rec. Indian Mus., Calcutta, 44:217- 
322. 

Sachwatkin, V. 

1920. Das urogenitalsystem von Ampullaria gigas Spix. Inaugural 
Dissertation, Univ. Zurich, Alb. Bonniers Boktryckeri, Stock- 
holm, 64 pp. 

ScoTT, M. I. H. 

1943. Sobre la organizacion de Ampullaria (Asolene) megastoma 
Sowerby. Notas del Museo de la Plata, 8, Zoologia, No. 70, 
pp. 269-280. 



BREVIORA 



laseimi of Comparative Zoology 



Cambridge, Mass. February 28, 1961 Number 134 

ENZYMATIC CONSTITUTION OF ALSOPHIS SALIVA 
AND ITS BIOLOGICAL IMPLICATIONS 

By George Hegeman 

University of California 
Berkeley, California 

I. Introduction 

As long as fift^'-eight years ago (Alcock and Rogers, 1902) it 
was noted that the saliva of certain ostensibly harmless colubrids 
had toxic properties. Recently, members of the genus Alsophis 
have been suspected to have a toxic saliva. A. S. Rand (MS.) 
has detailed accounts of Alsophis portoricensis feeding behavior 
and of the effects of bites in man. Neill (1954) relates an in- 
stance of toxic effects in himself from an A. anguilifer bite. 

The purpose of this paper is two fold: 1) to attempt to pro- 
vide enzj^matic confirmation for the tentative conclusion that the 
colubrid Alsophis portoricensis has a truly active salivary secre- 
tion (i.e. with venomous properties) ; and 2) to characterize the 
activity found in terms of similar enzymes represented in 
another colubrid and in a crotalid, thus providing a comparative 
basis for evaluation of the biological implications of this activity. 

Natrix sipedon from Florida and Crotalus atrox from Texas 
provided the other preparations. Commercial crystalline Crota- 
lus venom was made available by Dr. W. R. Sistrom. The 
Natrix and Alsophis preparations were collected by a straight- 
forward method in the laboratory. 

II. Collection of Saliva 

A small tablet of washed, fine-pore cellulose sponge, previous- 
ly moistened with distilled water, was inserted in the reptile's 
mouth with stainless steel forceps. Alsophis readily grasped and 
chewed on the sponge, but with Natrix swabbing was necessary. 



2 BREVIORA No. 134 

When relinquished, the sponge was squeezed and rinsed into a 
tared weighing bottle. The bottle was placed in a chilled desic- 
cator and evacuated until dry over potassium hydroxide at 5°C. 
This procedure was repeated over the course of several weeks 
until a quantity deemed sufficient for subsequent tests was col- 
lected. 

Ten "milkings" coming from two individuals provided 22 
milligrams of Alsophis preparation over a period of three weeks. 
Six Natrix, each milked once on four occasions equally spaced 
over the same three wrecks, gave a total of 26 milligrams of 
preparation. 

III. Estimation of protein content 

Since all known enzymes are proteins, the first step in gaining 
a fair comparison of activities depends on the total protein con- 
tent of the saliva. This was estimated for small redissolved 
duplicate samples of preparation by the Folin-Low^ry method 
(Lowry et al., 1951). Bovine serum albumin was adopted as the 
reference protein standard. Protein by percent weight for the 
three preparations is given below (Table 1). 



'.%) 



Alsophis is here seen to lie between the harmless colubrid and 
overtly venomous crotalid with respect to potentially active 
protein content. These results were used to adjust the dissolved 
preparations used in the later experiments to equivalent protein 
content. 

IV. Spreading factor estimation 

The reports of Neill and Rand (above) indicate the presence 
of a "spreading factor," seen in many natural venoms (Kella- 
way, 1939). In the case of snake venoms this is probably a 
complex mixture of proinvasins and hyaluronidase (Zeller, 
1948). These enzymes respectively inhibit destruction of the 
hydrolytic agent of the venom and destroy the mucoid structural 
material of the cell lattice by a hydrolytic action. The resultant 
loosening of the matrix favors the passive transport of the venom 
components of the tissue. 





TABLE 1 




Preparation 




Jo protein ( : 


Alsophis 




43.0 


Crotalus 




99.5 


Natrix 




16.5 



1961 ALSOPHIS VENOM 3 

The method followed here has been modified from Fanilli and 
McClean (1939). It has the disadvantage that specificity is lost 
and the results are ill-defined. It is well suited to demonstrating 
low activities however. 

Two dilutions of the preparation in question, adjusted to equal 
protein content, were mixed with equal parts of india ink and 
0.9 saline. Of each dilution 0.025 ml. was injected intradermally 
on the shaved back of an etherized rat. Bovine serum albumin 
of approximately equal protein content was substituted for the 
saliva preparations in five otherwise similar control injections. 
Sites of injection were placed within the shaved area so that 
each was straddled by two controls. The diameter of each weal 
was read at one-half hour intervals. Since the progress of the 
spots was a nearly constant percentage of the original diameter 
for the first hour and one half, the observations below are for 
that range (Table 2). Rates fell off sharply after that period; 
the rate for Crotalus fell off sharply after the first half hour. 







TABLE 2 










mgm preparation 


% 


increase in diameter 






protein injected 




per hour (± 10%) 


Crotalus 




0.125 
0.068 




80 
55 


Natrix 




0.125 
0.063 




20 



Alsophis 




0.130 
0.065 




50 
20 


Controls 


1 


0.000 









2 


( ( 




14 




3 


( ( 









4 


( 1 








5 •' 17 

Again, according to this index, Alsophis shows a definite 
tendency toward "venomous" aetivit3\ 

V. Proteinase 

Another type of enzyme frequently active in venoms (Zeller, 
1948) is proteinase. Work on the genus Dromicus, a relative of 
Alsophis, has demonstrated the activity of this enzj'me type in 
extracts of the parotid gland (Donoso-Barros and Cardenas, 
1959). Proteinases act hydrolyticly to reduce the chain length 
of proteins and large polypeptides, proAdding necrotic pre- 
digestion of the food object and destruction of the tissues. The 
fortuitous release of histamine from damaged and dissolved 



4 BREVIORA No. 134 

tissue might be expected to enhance auj- neurotoxic activity in 
the venom too. 

Proteolytic activity was estimated (Northrop et al., 1948) by 
measuring the residual material giving the Folin-Lowry reaction 
in an incubation mixture of enzj^me and bovine serum albumin 
after precipitation with 0.1 molar trichloroacetic acid. Quanti- 
ties of 0.25 mgm protein of preparation and 2.5 mgm of bovine 
serum albumin per milliliter were incubated in M/15 phosphate 
buffer at pH 7.2 and 37° C. Saliva preparations alone and 
albumin alone were also assayed for autogenous proteolysis ; the 
added rates of release in these provided a subtractive correction 
for the incubation mixtures of preparation and protein. 

Proteolysis rate is best taken over the first half hour where 
first order kinetics appear to obtain. The relative activities are 
given below, the average of two experiments. 





TABLE 


3 




'reparation 






^Lgm BSA hydrolyzcd/ 

mgm preparation protein/ 

hour (±2%) 


Crotalus 






28.3 


Alsophis 






12.3 


Natrix 






0.3 



\\. Hemolysis 

The lysis of red l)lood cells is a conspicuous part of the action 
of snake venoms (Kellaway, 1939). The major part of this effect 
is due to the action of phospholipase A on tissue phosphatides 
to produce lysolecithin. This substance renders erythrocytes 
extremely fragile (Zeller, 1948). The subsequent action of 
proteoh'tic enzymes results in the disruption of the cell wall 
(lysis). Other substances in venoms besides lipases promote 
lysis without first acting on an intermediate, and it is these which 
were measured here. 

Following Bernheim (1947), aliquots of washed horse erythro- 
cytes were incubated with measured quantities of the various 
preparations in buffered isotonic saline at 37° C. A blank was 
run to correct for autolysis in the buffered saline, and one aliquot 
was lysed by repeated freeze-thawing to give 100% lysis. At the 
end of the incubation period the cells were centrifuged down 
and the supernatant liquid was immediately read for absorbance 
at 579 miJ., an absorption peak of hemoglobin. This quantity 
when related to that of the completely lysed portion gave a 



1961 ALSOPHIS VENOM 5 

percentage lysis for the cells. Hemolysis rates are given below 
as percent lysis per ten micrograms preparation protein per 
milliliter per hour. 

TABLE 4 
Preparation lysis/hour 

Crotalus 4.42% 

Alsophis 1.20% 

Nairix — 0.73% 

The negative value for the Natrix preparation indicates a lysis 
rate less than the control, i.e. a protective effect. 

VII. Cholinesterase 

The typical elapine venom, noted for neurotoxicity, is ex- 
tremely high in a unique type of cholinesterase (Zeller, 1947). 
Presumably, since it is present in high concentrations, the 
curare-like effect of this venom is due to this enzyme's hydro- 
lytic action on acetylcholine. This blocks transmission at neuro- 
muscular junctions, and hence paralyzes the prey. However, 
inhibitors of this enzyme do not protect animals as fully as 
might be expected from the effects of cobra venom, including the 
neurotoxic effects (Zeller, 1948). The situation here is not as 
clear as might be hoped. 

The assay method was essentially that of Hestrin (1949). The 
preparation was incubated with M/lo phosphate buffer at pH 
7.2 and 30° C ; acetylcholine concentration was four micro- 
moles per milliliter. From the known protein content of the 
preparation and the measured disappearance of acetylcholine 
the activity of the preparation was established. Under the con- 
ditions described, accuracy was one tenth of a micromole. This 
was also the minimum amount of hj'drolysis detectable. 





TABLE 5 




Preparation 




Units esterase 


Crotalus 




0.0 


Alsophis 




0.2 


Natrix 




7.4 



The unit of activit}^ is conventionally defined as micromoles 
acetylcholine hydrolyzed per hour per milligram preparation 
protein under the conditions of the assay. Cholinesterase is 
normally low or non-detectable in crotaline venoms. The hio-h 
activity of the Natrix saliva is surprising. Subsequent tests 
showed that the Natrix preparation was 50 per cent inhibited 



fi BREVIORA No. 134 

by 10"^ molar liexamethonium (1,6-bis hexaiie trimethylam- 
moiiium) ; completelj' inhibited by 10"'* and 90 per cent inhibited 
by 10~*^ molar concentrations of neostigmine (prostigmine), re- 
spectively. This preparation did not appear to hydrolyze either 
benzoyl choline or acetyl /3-methyl choline. Characteristically, 
elapine venom cholinesterase does not act on benzoyl choline, 
but will hydrolyze acetyl ^-methyl choline. It was unfortunately 
impossible to treat the Ahophis preparation to this sort of analy- 
sis due to its low activity and a waning suppl}^ of preparation. 

VIII. Discussion 

The statement that a particular animal is or is not "venom- 
ous" is outwardly a qualitative judgment, but it conceals a 
quantitative .statement. The demonstrated presence of enzymes 
characteristic of venom in Ahophis' saliva constitutes necessary 
but not sufficient proof for the postulated function of the secre- 
tion in subduing prey. 

Alsophis' saliva protein content and the relatively greater 
activity of that protein are superior to those found in the "non- 
venomous" Notrix. Two large teeth surrounded by a fleshy 
ridge in the rear of the upper maxilla appear to be modified for 
administration of the saliva. Although these rear teeth are not 
grooved, the total picture for Alsophis indicates a degree of 
specialization trending in the direction of the adaptations of the 
truly venomous species. However, these morphological features, 
like the relative potency of the salivary secretion, are only an 
indication of the possible function. Critical evidence with re- 
gard to the use and thus the significance of the "venom" will 
probably come from the careful study of feeding behavior. 

Animals killed by .snake venom decompose at a much more 
rapid rate than similar individuals who have died peacefully 
(Zeller, 1948). Vipers, kept from injecting venom by fang 
removal or ligature of the duct, take from two to three times the 
normal time to digest a food object of constant size. Apparently 
the digestive function of venom is a large part of its adaptive 
significance, especially in an animal unable to mechanically 
masticate its prey. It may be that at the level of development 
seen in Alsophis, the effect of the saliva in subduing prey is 
secondary and gratuitous to the digestive function. At higher 
levels of specialization it is almost impossible to disentangle the 
two roles. The enzymes of snake venom are most easily inter- 
preted as a digestive complex (Zeller, 1948). 



1961 ALSOPIIIS VENOM 7 

The cliolinesterase of Natrix saliva, unlike the usual elapid 
type (Zeller, 1947), will not hydrolyze acetyl y8-methyl choline. 
This is especially interesting in light of the problem of the rela- 
tionships between the cokibrids and elapids (Johnson, 1956). 
An enzyme study of the analogous secretions of these two groups 
in a larger range of genera could prove valuable. It would, of 
course, first be necessary to establish the stability of enzvme 
type as a character wathin a genus or genera for which the rela- 
tionships are relatively clear. 

Acknowledgments 

I would like to acknowledge the guidance and suggestions of 
Dr. William R. Sistrom, who cheerfully permitted a small plague 
of snakes in an otherwise orderly bacteriology laboratory. I am 
also indebted to Mr. A. S. Rand and Dr. E. E. AYilli'ams for 
suggesting the problem, supplying valuable advice and informa- 
tion, and seeing the paper through several revisions. Dr. Karl 
Kofford generously supplied the Alsophis portoricensis used. 

PAPEES CITED 

Alcock, a., and L. Eogers 

1902. On the toxic properties of the saliva of certain "non-poisonous" 
colubrines. Proc. Roy. Soc. London, 70:446. 
Bernheim, a. W. 

1947. Comparative kinetics of hemolysis induced by bacterial and 
other hemolysins. Jour. Gen. Physiol., 30:337. 
Doxoso-Barros, R. and S. Cardenas 

1959. Estudio del veneno de Dromieiis chamissonis (Wiegmann). Inv. 
Zool. Chilenos, 5:93-95. 
Fanilli, G., and D. McClean 

1939. A spreading factor in certain snake venoms and its relation to 
their mode of action. Jour. Exptl. Med., 69:81. 
Hestrin, S. 

1949. The reaction of acetylcholine and other carboxylic acid deriva- 
tives with hydroxylamine, and its analytic application. Jour. 
Biol. Chem., 180:249. 
Johnson, R. G. 

1956. The origin and evolution of venomous snakes. Evolution, 10: 
56. 
Kellaway, C. H. 

1939. Animal poisons. Ann. Rev. Biochem., 8:541. 
LoWRY, O. H., N. J. Rosenbrough, A. C. Farr and R. J. Randall 

1951. Protein measurement with the Folin phenol reagent. Jour. 
Biol. Chem., 193:265. 



8 BREVIORA No. 134 

Neill, W. T. 

1954. Evidence of venom in snakes of the genera Alsophis and Rhadi- 
naea. Copeia, 19-i4, no 1:59. 
Northrop, J. H., M. Kunitz and E. M. Herriot 

1948. Crystalline Enzymes. Columbia Univ. Press, New York. 
Phisalix, M. 

1922. Animaux Venimeux et Venins (Vol. 2). Masson et Cie, Paris. 
ZELr.ER, E. A. 

1947. Uber das Vorkommen und die Natur der Cholinesterase der 
Schlangengifte. Experimentia, 3:375. 

1948. Enzymes of snake venoms and their biological significance. 
Advances in Enzymology, Volume 8, p. 459. F. F. Nord ed., 
Interscienee Pub., New York. 



BREVIORA 



Miaseiuiini of Comparative Zoology 



Cambridge, Mass. April 7, 1961 No. 135 

NOTES ON IIISPANIOLAN HERPETOLOGY 

2. A REVIEW OF THE ANOLIS SEMILINEATUS 

GROUP WITH THE^DESCRIPTION OF 

ANOLIS COCHRANAE, NEW SPECIES 

By Ernest E. Williams 
and A. Stanley Rand 

Introduction : Recent investigations in Haiti and the Domin- 
ican Repnblic by expeditions from the Mnseum o± Comparative 
Zoology have added considerably to our knowledge of the dis- 
tribution of the so-called grass-anoles of the semilineatus group 
and have resulted in the discovery of a third member of the 
series in the Cordillera Central of the Dominican Republic. In 
addition to describing the new species just discovered we here 
attempt a summary of the information now available on this 
group. 

Acknowledgments: The new species was collected in the 
summer of 1958 by Clayton E. Ray and A. Staule}- Rand during 
an expedition partly supported by a grant from the Societj* 
of Sigma Xi and enjoying the cooperation of the University of 
Santo Domingo and of the government of the Dominican Re- 
public. The essential and generous aid of Dr. Eugenio de Jesus 
Marcano, who accompanied the expedition, and the use of a 
car and driver furnished by the University of Santo Domingo 
are very gratefully acknowledged. In August 1959, E. E. 
Williams and A. S. Rand collected in the vicinity of Port-au- 
Prince with the support of National Science Foundation Grant 
NSF G-5634. In 1960 A. S. Rand and J. Lazell collected aroiind 
Port-au-Prince in northern Haiti and near Aux Cayes on the 
southwest peninsula, aided by a grant from the American Philo- 
sophical Society. In both Haitian trips the letters provided by 
M. Gerard Philippeaux, Minister of Agriculture, and the willing 
assistance of M. Leonce Bonnefil fils. zoologist in tln' Department 



2 BREVIORA No. 135 

of Agriculture and Natural Resources, were indispensable ele- 
ments in the success of the venture. 

In addition to the specimens collected by these expeditions and 
those already present in the Museum of Comparative Zoology 
(MCZ), material was obtained on loan from the United States 
National Museum (USNM) and the American Museum of Natu- 
ral History (AMNH). The assistance of the curators of these 
collections is gratefully acknowledged. Dr. P. S. Humphrey 
made available for study material collected by him for Yale 
Peabody Museum (Yale) and the University of Florida (UF). 

The previously known species : In order to provide a frame 
of reference for the new species, we first summarize the knowl- 
edge now at hand for the previously known species : Two species 
of "grass anoles," A. semilineatus Cope and A. olssoni Schmidt, 
have long been known in Hispaniola. Both are small {ca. 40 mm 
snout-vent length), slender-bodied forms with elongate heads, a 
dorsal zone of about 10 rows of enlarged keeled scales as large 
as the strongl}^ keeled belly scales, tail only slightly compressed 
and with no clear demarcation of the breaking zones or verticils. 
They are thus a morphologically strongly marked group within 
the Hispaniolan anoles. In habits they are also distinctive, being 
associated characteristically with grass and low bushes. 

Morphology: (See also table, below). Structurally the two 
species differ, very little — they differ in size of scales and 
slightly in body size, tail length and shape of head but in none 
of these regards so strikingly that instant identification can be 
confidently made even by the experienced worker. Body colora- 
tion differs also but not without some puzzling cases. The dew- 
lap in males, both as to color and squamation, is the diagnostic 
difference easiest to employ. The dewlap skin in A. semilineatus 
is white and the gular scales about the same size as the ventrals. 
In olssoni the dewlap skin is red or orange (darkly pigmented 
in alcohol) and the gular scales up to three times as large as 
the ventrals. In life the iris of semilineatus is steel blue, that 
of olssoni dark brown. This difference is not determinable in 
alcoholics. 

Ecology: There exists a real ecological difference between 
the two species but again there is overlap and the species do 
occur side by side. 

Mertens (1939) describes A. semilineatus as "eurytop" occur- 
ring in both dry and wet areas. He found it in mangroves (Puerto 
Plata, Sabana de la Mar), corn fields (Moca), dry open brush 
(Barahona, San Pedro de Macoris, Ciudad Trujillo), meadows 



1961 ANOLIS COCHRANAE 3 

iu the lower drier pine woods (Jarabacoa, Monciou), and in 
damp lush vegetation (Samana, the top of the pass between 
Santiago and Puerto Plata). He, in fact, states that it is 
absent only in the cactus- steppe. Rand (1958, field observations) 
saw it in the Dominican Republic primarily in open situations 
and along- roadsides and in pastures but found also a single 
specimen sitting on a rock in a muddy trail through heavy 
forest (Bejucal), and a dense population living in low vegetation 
in an area of rather dense bamboo along a stream bank (nr. 
Sabana de la Mar). Hassler (field notes for November 4, 1929) 
reports finding this species at Laguna near Samana on "leaves 
in damp woods and in fields nearby." 

Near Port-au-Prince, scmilineatus occurs in the hills to the 
south of towTi and up to the vicinity of Furcy at 4000-5000 ft. 
It is absent from Port-au-Prince itself and from the Cul de 
Sac Plain. In one place it has been observed to overlap with 
olssoni (see below). 

Mertens has described A. olssoni in contrast to A. semilineatus 
as "stenotop," confined to open dry areas. It seems, indeed, to 
be more limited than semilineatus, but in 1959 Williams and 
Rand found it in the moderately dense vegetation of a Port-au- 
Prince garden and also in the irrigated areas of the experi- 
mental farm at Damien, Haiti. It is present in open thorn 
scrub of the Cul-de-Sac Plain both in Haiti and in the Domini- 
can Republic. It is known to occur with semilineatus at several 
localities. Mertens records both forms from the vicinity of 
Ciudad Trujillo, and at Moncion, Sabana de la Mar and Bara- 
hona, all localities in the Dominican Republic. Cochran reports 
both species at San Michel du Nord, Haiti. In 1960 Rand and 
Lazell obtained both species at Gros Morne in northern Haiti. 
They found "A. olssoni on grass and low vegetation along sunny 
roadsides, A. semilineatus on vegetation at the edge of forest." 

In August 1959 Williams and Rand studied a contact area 
between semilineatus and olssoni on Bontillier Road which climbs 
the foothills south of Port-au-Prince. A. olssoni occurred only 
on the lower reaches of the road, A. semilineatus only on the 
portion of the road which parallels the crest of the hill. No 
striking vegetation or habitat difference was evident between 
the two portions of the road, both of which traverse very dis- 
turbed, cut over, areas. The distance between the places at 
which SI inilineatus and olssoni were found closest to one another 
was a matter of a few A^ertical vards. Rand and Lazell returning 



4 BREVIORA No. 135 

to the same area in 1960 found the same general pattern but in 
one instance a semilineatus was taken about 20 feet from an 
olssoni and at the same level. 

Habits: The two species differ very little in habits. Both 
are commonly seen on grass stems and the slender twigs of 
small bushes and plants. Sometimes they occur on fence posts 
and stands of barbed wire and occasionally on the ground. None 
have been seen on the trunks or in the crown of even small trees. 
Numbers of A. olssoni were observed by Williams and Rand at 
Damien, Haiti. Individuals of all sizes were found in the tall 
grass, while on the fence posts most of the animals were large 
males, and in the short 4-8 inch grass most were juveniles. They 
are frequently found facing head downward on a vertical perch 
with the neck bent so that the head is almost horizontal. Both 
sleep with the hind legs fully extended. Both escape by jumping 
off their perch into grass cover nearby. In one area near Port- 
au-Prince a few individuals were chased out of the grass into 
a rock pile. Here they did not go deep into the rock pile in 
contrast to juvenile A. cybotes but hid close to the surface and 
could usually be chased out by poking into the holes with a 
short stick. A. semilineatus is perhaps shier than A. olssoni and, 
according to Mertens, it is less pugnacious. Mertens reports that 
freshly caught olssoni carried out biting battles with each other, 
raising the nuchal crest and displaying their dewlaps. 

Very little is known of the reproductive habits of these forms. 
There is one observation by Rand (field notes, 1958) on A. 
semilineatus in the Dominican Republic : "In one locality 
(Rancho La Guardia, San Rafael Province) in a coffee plantation 
about twenty yards from o])en pasture, a number of eggs of 
this species were found. These were discovered in three of 22 
rotten logs examined in a small area. One log contained 12 eggs, 
another 7 eggs, and the third 4 eggs. These three logs differed 
from the others examined in that they contained nests of a 
large black stinging ant. The eggs were mostly in the loose 
soil just under the logs but some were in the galleries of the ant 
nest. When I picked up the first egg an ant stung me and my 
iuA^oluntary jump sent the egg flying several feet. The other 
logs in the area contained a variety of invertebrate life but the 
ones with the lizard eggs had only the black ants and a single 
centipede nest. The ants provided an effective protection for the 
lizard eggs and it seems possible that the lizards had sought 
out these nests in Avhich to lay their eggs. 



1961 ANOLIS COCHEANAE 5 

The eggs were ovoid with a white flexible skiu. They ranged 
in size from 12 by 9 nun to 7 by G mm. Collected on August 
13 fourteen of the twenty-two eggs hatched, the first on August 
If) and the last on September 17.'* 

Distribution : Both species are very widely dispersed in 
Ilispaniola. Because of their ecological differences the two dis- 
tributions coincide only in limited and scattered areas, though 
in broad terms they overlap widely. No olssoni are at present 
known from the Saniana peninsula and the adjacent areas in 
the north of the Dondnican Republic north of La Vega and east 
of Puerto Plata, or from ths southwestern peninsula of Haiti, 
but none of these areas is so well collected that the absence of 
olssoni from collections can be taken as a demonstration of a 
real absence in the field. Except perhaps in the southwestern 
peninsula, the distribution can be interpreted better in terms of 
present ecology than in terms of any other factor. We list all the 
verified localities^ for the two species below": 

Semilineatus. HAITI: Dcpt. du Nord, Cap Haitien (MCZ, 
USNM), Citadelle (MCZ), Dondon (MCZ); Dept. de I'Arti- 
bonite, Gros Morne (MCZ), San Michel (USNM); Dept. de 
Oucst, Basin Bleu nr Furcy (MCZ), Bontillier Road nr Port- 
au-Prince (MCZ), 5 km south of Dufort, south of Leogane 
(MCZ), Furcy (MCZ, AMNII), Obleon nr Furcy (MCZ); De- 
partment du Sud, Miragoane (MCZ), Les Platons north of Aux 
Caves (MCZ), Place Negre near Jeremie (MCZ). DOMINICAN 
REPUBLIC: Prov. Sa7i Rafael, Rancho La Guardia (MCZ); Prov. 
Barahona, Barahona (AMNH, Senckenberg), Palo (AMNH) ; 
Prov. Benefactor, 7 km north of Carpintero (MCZ) ; Prov. San- 
tiago Rodriguez, Moncion (Senckenberg) ; Prov. Santiago, top 
of pass between Santiago and Puerto Plata (Senckenberg) ; 
Prov. Puerto Plata, 8 km north of Pena (MCZ), Balneario Colon, 
Puerto Plata (Senckenberg), Rio Munoz, 7 km from Puerto 
Plata (Senckenberg) ; Prov. La Vegn, Jarabacoa (Senckenberg); 
Proi'. EspaiUat, Moca (Senckenberg), Rio San Juan (USNM); 
Prov. Duarte, Las Bracitas (AMNH); Prov. Trujillo, nr San 
Cristobal (MCZ) ; Prov. San Pedro de Macoris, San Pedro de 
Macoris (Senckenberg) ; Prov. Seibo. Boca del Inferno (USNM) ; 
San Francisco, 6 km east of Hato Mayor (MCZ) : Rio Yabon 



1 The record of sonUincatus for the island of Narassa is doubtful. A parutype 
of A. olssoni was recorded by Schmidt (3 919) as probably from this island. In 
1921 Schmidt redetermined the si>ecimen as A. semilineatus and cited the species 
without qualification as a member of the Navassa fauna. No additional speci- 
mens of A. semilineatus have been collected, and the record is unconfirmed. 



6 BREVIORA No. 135 

(MCZ) ; Prov. Samana, Laguna (AMNH), Samana (MCZ, 
Senckeiiberg), Sanchez (MCZ) ; Distrito de Smito Domingo, 
Ciudad Trujillo (Seuekenberg). 

Olssoni. HAITI: Dcpt. da Nord Guest, Bombardopolis (MCZ), 
Jean Rabel (MCZ, AMNH), Mole St. Nicolas (MCZ); Dept. 
du Nord, Cap Haitien (USNM) ; Dept. de I'Artibonite, bridge 
over the Artibonite (MCZ), south end of Etang Bois Neuf (MCZ), 
Gros Morne (MCZ), St. Marc (USNM, AMNH); Departmeiit de 
Quest, Boutillier Road nr Port-au-Prince (MCZ), Carrefour 
(AMNH, Yale, UF), Cabrite Id (AMNH), Damien (MCZ), 
Delmas (MCZ), Diquiui (MCZ, USNM), Eau Gaillee (Yale, 
UF), Etang Saumatre (MCZ), Fond Parisien (AMNH), Hatte 
Latham (MCZ, USNM), Manneville (MCZ), Mon Repos 
(USNM), Morne Decayette (MCZ), Petionville (Yale, UF), 
Port-au-Prince (MCZ, USNM, AMNH, Yale, UF), Ste. Philo- 
mene (USNM), Thomazeau (MCZ), between Thomazeau and 
Manneville (MCZ), Trou Caiman (USNM), Trou Forban 
(MCZ), Gonave Id, Anse a Galets (MCZ), Encafe (MCZ, USNM, 
Yale), Pointe-a-Raquettes (Yale, UF). DOMINICAN REPUBLIC: 
Prov. Monte Cristi, Monte Cristi (AMNH, Senckenberg) ; Prov. 
San Rafael, Banica (MCZ); Prov. Independencia, has Baitoas 
(AMNH), Duverge (AMNH); Prov. Barahona, Barahona 
(AMNH), Senckenberg), Cabral (MCZ) ; Prov. Santiago Rodri- 
guez, Monciou (Senckenberg) ; Prov. El Seiho, Sabana de la 
Mar (Senckenberg); Distrito de Santo Domingo-. Ciudad Tru- 
jillo (Senckenberg). 

A THIRD SPECIES DISCOVERED: On September 7 to 8, 1958, col- 
lecting in the vicinity of Constanza in the high interior of the 
Dominican Republic, C. E. Ray and A. S. Rand of Harvard 
University and Sr. Eugenio de Jesus Marcano of the Uni- 
versidad de Santo Domingo obtained 20 specimens of a new 
species of "grass anole." The greater number of these speci- 
mens were collected at night, sleeping on grass stems. A renewed 
effort to collect the same form the next morning obtained very 
few individuals, the lizards being then very wary and difficult 
to see or catch. 

Examination of these specimens reveals that they differ from 
the two previously known species in just the ways cited by 
Doris Cochran (1941, pp. 139-140) for a single specimen from 
Constanza which she then referred hesitantly to A. olssoni. Her 
remarks are quoted in full : 



1961 ANOLIS COCHRANAE 7 

"With some doubt I place with Anolis olssoni a single adult 
male (USNM No. 62103) collected by Dr. W. L. Abbott in the 
hills 5 miles south of Constauza. This individual has much 
smaller scales ou the gular fan than does typical olssoni from 
San Michel, Haiti. It does not approach, however, semilineatus 
in fineness of scales. In fact, while the gular scales are finer, the 
dorsal and ventral scales of the Constanza lizard are actually 
perceptibly coarser than they are in the San Michel specimens. 
The color pattern of this Constanza specimen shows none of the 
definite black markings that so often appear on true olssoni. It 
is lilac gray above, tinged with china-blue on the supraocular 
region, the dorsal tone shades into drab above the lateral light 
stripe, which is very sharply marked anteriorly but less so after 
it passes the shoulder, back of which it fades out almost com- 
pletely. The only definite head marking is a black diagonal bar 
across the temporal region which does not occur in olssoni but 
is found in every specimen of semilineatus. A series of examples 
from Constanza will be needed to determine whether these 
characters are stable and definite, meriting specific separation 
or whether they represent an aberrant or hybridized olssoni with 
some of the seniilineatus characters." 

Dr. Cochran has excellently characterized the new species, 
which may therefore be appropriately named : 

Anolis cochranae new species 
Type. MCZ 57660, an adult $ collected at Constanza, Dom- 
inican Republic, September 7-8, 1958 by C. E. Ray, A. S. Rand, 
E. de Jesus Marcano. 

Paratijpes. MCZ 57661-79, same data as above : USNM 62103, 
hills 5 miles south of Constanza, collected by Dr. AY. L. Abbott, 
April 29, 1919. 

Diagnosis. An Anolis allied to semilineatus and olssoni, dif- 
fering from the first in the much larger dorsal, ventral and 
supratemporal scales, from the latter in having a white rather 
than a red or orange dewlap in the somewhat larger dorsals and 
in having the gular scales little if at all larger than the ventrals, 
differing from both in having the ventrals nearly as large as the 
enlarged dorsals. 

Description. Head: All head scales multicarinate rather than 
smooth or singly keeled. Five to eight scales across head between 
second and third canthals (usually six to seven). Frontal de- 
pression very shallow, the scales in it nearly or as large as the 
posterior frontal or anterior supraorbital. 



8 BREVIORA No. 135 

Supraorbital semicircles in contact (five specimens) or sep- 
arated by one scale row (fifteen specimens), wholly or partly 
separated by one scale row from the supraocular discs. Supra- 
ocular disc consisting' of two to five large keeled scales separated 
from the elongate supraciliaries by at least two rows of scales. 
Canthus distinct, canthal scales four (five in one specimen), the 
second largest diminishing gradually forward. Naris anterior 
to canthal row\ The anterior nasal scale in contact with rostral. 
Loreal rows four to five (three on both sides in one specimen). 
Temporal scales subgranular. Supratemporal scales larger, 
keeled, grading into the large keeled scales surrounding the inter- 
parietal. Interparietal larger than ear, separated from the supra- 
orbital semicircles by one to three scales (usually two). 

Posterior frontal as large as anterior supraorbital. One scale 
nearly as large as the posterior frontal between the latter and 
the canthals. 

Suboculars in contact with supralabials. One scale intervening 
between subocular series and canthals. Five to six (six on one 
side, seven on the other in one specimen) supralabials to the 
center of the eye. 

Mentals wider than long, one to two scales inserted between 
the tips posteriorly. One sublabial on each side in contact with 
the infralabials. Central throat scales keeled, elongate. Gular 
fan in males. 

Trunk : About ten longitudinal rows of much enlarged keeled 
middorsal scales, broader than long, as large as the ventrals (10 
to 12 ill standard distance), grading laterally into the smaller 
flank scales wliicli in some specimens are keeled imbricate, in 
others nearly granular. Ventrals in longitudinal rows, keeled, 
imbricate, mucronate. Postanal plates present in males. Scales 
of gular fan moderate, not extremely elongate, hardly larger 
than ventrals, not clearly arranged in lines. 

Limhs and digits: Hand and foot scales raulticarinate, about 
17-19 lamellae under phalanges 2 and 3 of fourth toe, about 
26-31 under whole toe. Largest arm and leg scales unicarinate. 
about as large as ventrals. 

Tail: Tail subcircular in section, very long, more than 21/2 
times snout-vent length ; verticils not apparent. 

Sisc : Largest 3 41 nnn in snout-vent length ; largest 9 
38 mm snout-vent length. 



1961 



ANOLIS COCHRAN AE 



Color: Essentially as in semilineatus. 

The more significant characters of cochranae may be compared 
with those of semilineatus and olssovi in tabular form: 



semilineatus 
skill of tjular fan white 

flank stripe short 

;;ular scales ea. = 
ventrals 

14-17 enlarged dorsal 
scales in distance snout 
to middle of eye 
(standard distance) 

median rows of enlarged 
dorsal scales about as 
broad as long 

17-21 ventrals in standard 
distance 



olssoni 

skin of gular fan orange 
to red 

dank stripe long 

gular seales>> 
ventrals 

11-13 enlarged dorsal 
scales in standard 
distance 



median rows of enlarged 
dorsal scales mostly 
longer than broad 

11-14 ventrals in 
standard distance 



cochranae 
skin of gular fan while 

flank stripe short 

gular scales ca. = 
ventrals 

10-12 enlarged dorsal 
scales in standard 
distance 

median jows of enlarged 
dorsal scales about as 
broad as long 

11-14 ventrals in 
slandard distance 



Other differences have been listed by Mertens or Cochran, but 
they are at best modal differences or they alter markedly with 
age. These species are indeed close, and females and juveniles 
are sometimes difficult to distinguish. 

Discussion: Anolis cochranae combines in new ways charac- 
ters of ^1. semilineatus and .1. olssoni. It is in no sense an 
intermediate; its characters are either those of one or the other 
or are somewhat exaggerated versions of a trend present in one. 

It is necessary to admit that we know very little about 
this species beyond its existence. Its distribution would appear, 
on present evidence, to be extraordinarily limited. It may well 
be confined to the high interior, but its real range is surely more 
extensive than known at present. The area from which it comes 
is remarkable for certain peculiar forms : Celestus darlingtoni 
and Audantia shrevei in the higher elevations, Anolis aliniger 
(described as a subspecies of A. chlorocyanus by Mertens in 
1939 but in reality a full species) in the vicinity of Constanza 
itself, Anolis clarlimjtoni both from Constanza and from higher 
elevations. The region merits extensive and systematic collect- 



ing. 



The biological relation of A. cochranae to the other two mem- 
bers of the semilineatus group is equally unknown. We do not 



10 BREVIORA No. 135 

know its contacts with either form. Its relationship to A. 
semilineatus in particular is puzzling. In squamation it differs 
strongly enough that we have called it, as a matter of judgment, 
a distinct species. The scale differences from both semilineatus 
and olssoni are as great or greater than the differences between 
other closely related sympatric fully valid species. But in other 
Anolis such, (or lesser) diff'erences are correlated with color and 
dewlap differences that are evident visual cues to species recog- 
nition. In color and dewlap A. cochranae exactly resembles one 
of the neighboring species — semilineatus. If A. cochranae is 
indeed a full species that is at some point in contact with semi- 
lineatus, it is necessary to suppose that there is some unlcnown 
behavioral difference that maintains the distinctness of the 
population in the absence of color cues. 



EEFERENCES CITED 

Cochran, D. M. 

1941. The herpetology of Hispaniola. Bull. X. S. Nat. Mus., 177; 
1-398. 
Mertens, R. 

1939. Herpetologische Ergebiiissc einer Reise naeh der Insel His- 
paniola, Westindien. Abh. Senckenberg. Naturf. Ges., 449: 1-84. 
Schmidt, K. P. 

1919. Descriptions of new amphibians and reptiles from Santo 
Domingo and Navassa. Bull. Amer. Mus. Nat. Hist., 41 : 519-52.3. 
1921. The herpetology of Navassa Island. Bull. Amer. Mus. Nat. 
Hist., 44: 555-559. 



1961 



ANOLTS COCHRANAE 



11 




l-H 
O 

•a 

as 
A 

.a 



Pi 

O 

Si 



OB 
IS 
O 



o 



& 



o 

M 



O 

o 

El 

-S 

Q 






BREVIORA 



Musemnri of Comparative Zoology 



Cambridge, Mass. April 8, 1961 Number 136 

NOTES ON HISPANIOLAN HEKPETOLOGY 

3. THE EVOLUTION AND RELATIONSHIPS OF THE 

ANOLIS SEMILINEATU8 GROUP 

By Ernest E. Williams 

The discovery of a third species of the Anolis semilmeatus 
group, confined apparently to the high interior of the Dominican 
Republic, poses problems in the distribution, biology and evolu- 
tion of the group. 

The distributional data for the semilineatus group has been 
given in Williams and Rand (1961) and need not be repeated 
in detail here. A. semilineatus and A. olssoni are both widely 
distributed north of the Cul cle Sac Plain but occupying eco- 
logically somewhat different situations and thus with but limited 
actual contact or overlap ; only A. semilineatus at present is 
known south of the Cul de Sac Plain in the southwest and 
Barahona peninsulas. 

A. cochranae is found in the center of Hispaniola in the 
Cordillera Central — geographically in the midst of the other 
two species though its contacts with these others are not known. 
The biological peculiarity in the relation of A. cochranae to A. 
semilineatus has also been pointed out in Williams and Rand 
(1961). Thus, though differing strongly from the closely related 
A. semilineatus in certain scale characters, A. cochranae is iden- 
tical in body and dewlap color. This phenomenon is highly 
unusual in the genus Anolis in which body and dewlap color dif- 
ferences are important cues in species recognition. (There are, 
for example, strong body and dewlap color differences between 
A. semilineatus and A. olssoni.) A. cochranae, if it is in contact 
with A. semilineaius, as A. semilineatus and A. olssoni are in 
contact with one another, would seem to be a most anomalous 
case in which it would be necessary to provide some ad hoc 
explanation — such as some unknown behavior difference — for 
the maintenance of the species distinction. 



2 BREVIORA No. 136 

The problem is thus to provide an explanation of the central 
geographic position of Anolis cochranae in Hispaniola and of the 
curious absence in cochranae of the usual anoline species recog- 
nition characters contra a related species that occurs literally 
on every side of it. 

I propose below a suggested history of the semilineatus group 
that appears to solve this problem. It must be admitted that 
this proposed history depends upon taking at face value the 
distributions of the three species as they are known at present. 
This is patently unsafe, but it provides a useful starting point. 

On our present knowledge of distribution it is simplest to 
suppose that the postulated biological problem has not arisen, 
that cochranae and semilineatus are nowhere in contact. This 
is at the moment only a brave hypothesis. Anolis cocliranae is 
known from only two collections; our more extensive knowledge 
of the distributions of semilineatus and olssoni is by no means 
good enough to prove contact or absence of contact with coch- 
ranae. 

Critical to the proposed history is the supposition — uncon- 
tradicted by the available evidence — that olssoni is really absent 
from the southwest and Barahona peninsulas. It does appear to 
be absent from the moist coastal zone at Aux Cayes (observations 
by A. S. Rand and J. Lazell in 1960) and Rand did not collect 
it in the dry area of Oviedo on the Barahona peninsula in 1959. 
It is not present in Hassler's collections from these two areas. 

Let us then take the present distributional evidence at face 
value. Let us assume then that semilineatus is the only grass 
anole of the southwest and Barahona peninsulas and that olssoni 
just touches this area at the southern edge of the Cul de Sac 
Plain. 

The soutliAvest and Barahona peninsulas taken together are 
just that portion of the island which was cut off from the mass 
of Hispaniola by the Pleistocene seaway through what is now 
the Cul de Sac Plain. Residual salt lakes and coral rocks still 
testify to this former seaway. 

The division of Hispaniola into two parts which resulted from 
this seaway provides two suitable theatres — a main island and 
a southern counterpart — for the classic pattern of speciation 
during separation, and intensification of species difference 
("character displacement") during renewed contact. 

On this hypothesis semilinedius is the "autochthonous grass 
anole of the southern cut-off "portion -of " Hispaniola Mid' alss'om 



1961 ANOLIS SEMILINEATUS GROUP 3 

and cochranac autoehtlions of the northern main mass of the 
island. Scmilhieatus has infiltrated the northei-n island all but 
completely, while olssoni is not known to have invaded the south- 
ern island. 

The spread of sonilhicatus through much of the northern 
island is not too surprising in view of its eurytopic ecology 
(Mertens, 1939, AVilliams and liand, 1961). Though character- 
istic of a specialized open habitat, it seems to be sufficiently 
tolerant of forests that these would be less efficient barriers to 
its spread than they would to stenotopic olssoni. It is somewhat 
more surprising" — if it is true — that olssoni has not spread 
along the dry north coast of the southwest peninsula or the east 
coast of the Barahona peninsula, but it would be stopped easily 
by discontinuities in suitable habitat and would for this reason 
be unlikely to reach localities otherwise quite suitable to it on 
the southern island. 

The different coloration in olssoni, including the dewlap color, 
and the large size of the dewlap scales may Avell have developed 
after olssoni came into secondary contact with semilineatus dur- 
ing the latter 's invasion of the northern island fragment. In 
suggesting this we assume that the features in common of 
cochranac and semilineatus are primitive and that modification 
in these features took place exclusively or almost so in olssoni. 
(Surel}' the lack of enlargement in the gular scales is primitive 
ill semilineaius and cochranae; this leaves only color in ques- 
tion.) 

What, however, about the origin and relationship of cochranae 
and olssoni? It must first be noticed that there is some plausi- 
bility in considering these two more closely related to each other 
than to semilineatus. In body squamation (i.e. scale size), coch- 
ranac and olssoni are very similar. This is a feature which, 
unlike the characters of the dewlap or of body pattern, is un- 
likely to be a matter of intra- or inter-species recognition. AVe 
do not know that it is per se adaptive: the difference in scale 
.size between semilineatus, on the one side, and cochranue-olssoni, 
on the other, is more likely to be the external expression of more 
fundamental genetic divergencies. 

No. physiographic barrier, however, will account for the divi- 
sion of the grass anole population of the northern or main 
Ilispaniolan island into two species. It is necessary to suppose 
that the barrier was an area of unsuitable ecology, i.e. moist 
dense forest. Olssoni may then be supposed to have arisen in 



4 BREVIORA No. 136 

the arid coastal lowlands while cochranae arose in the open areas 
of the high pine woods' of the interior valleys of the Cordillera 
Central. (We note that Wetmore and Swales, 1931, p. 24, 
describe the natural vegetation of the Valle Constanza as "forests 
of open pine mingled with areas of dense rain forest.") 

The knowni habitat of cochranae — Valle Constanza — is a 
high interior valley of the Cordillera Central. Though the floor 
of this valley is not very high {ca. 3000 feet) it is surrounded 
by some of the highest peaks in Hispaniola and ingress to it at 
moderate elevations is someAvhat narrow and limited. In such 
an area a grass anole population might indeed enjoy a measure 
of isolation from other grass-bush populations — the more so if 
Vv-e suppose that the separation of olssoni and cochranae dates 
from a period in which the density of the hardwood forest of 
intermediate elevations was at a maximum. 

Relationshipy of the semilineatus group. 

There are no other anoles in Hispaniola which either very 
much resemble or seem very closely related to the semilineatus 
group. A search for close relatives and ancestors takes us at 
once outside Hispaniola. 

Two Greater Antillean groups of Anolis are structurally simi- 
lar — tlie alutaceus-clivicolus-cyanopleurus-spectrwni group in 
Cuba and the krngi-pidchellus-ponccnsis series in Puerto Rico. 
(None of the anoles of Jamaica or the Bahamas are similar either 
ecologically or structurally.) 

Both the Cuban and the Puerto Rican series share with the 
semilineatus group the middorsal zone of enlarged scales (least 
developed in krugi of Puerto Rico). All except alutaccus-clivi- 
colus have keeled ventrals. 

The Cuban anoles are all forest species, A. alutaceus occurring 
in rather deep shade, A. spectrum in less deep shade. But, 
though in this regard they differ from the Hispaniolan species 
which are fonder of open areas, they are closer to the semilinea- 
tus group in structure than are the Puerto Rican species. Like 
the semilineatus group they are small, usually under 40 mm 
snout-vent length, slender, with large dewlaps and well developed 
postanal scales in the males. In color, however, they differ in 
never possessing the flank stripe so characteristic of the semi- 
lincaius group, tending instead to emphasize the light middorsal 
stripe. 

1 I'ine in Hispaniola. in contrast to e.g. Cuba, is confined to higher elevations. 



1961 ANOLIS SEMILINEATl'S GROUP 5 

Of the alutaceus series, clivicolus, which may be a subspecies 
of alutaceus, has the least slender habitus and the least specialized 
squamation. It is easy to envision this as representing the primi- 
tive stock of this series. 

The Puerto Rican series is, on the other hand, more similar 
to the Ilispaniolan species in habits. Two of the three species 
— pulchellus and poncensis — are "grass anoles" or at least 
anoles of open reaches. The third species — krugi — is an anole 
of denser brush. All are larger than any species of the senii- 
lincatus group — nearer 50 mm than 40 mm snout- vent length. 
They are perhaps not as slender as their parallels in Hispaniola 
(though this is a character difficult to estimate objectively) ; the 
dewlaps are relatively small and the postanal scales poorlj^ 
developed. All three have a flank stripe passing forward through 
the eye more or less well expressed. 

In both Cuba and Puerto Rico the series exhibit a wider range 
of structure than do the Ilispaniolan forms. In each series there 
is a species with the middorsal zone of enlarged keeled scales 
less developed than in any Ilispaniolan species (in Cuba — 
clivicolus-alutaceus, in Puerto Rico — krugi) and one Avith 
this zone much more strongly developed than in any Hispaniolan 
species (in Cuba — spcctrutn, in Puerto Rico — poncensis). One 
difference appears in this regard : in all the Cuban forms the 
width of the zone of enlarged dorsal scales is about the same 
{ca. 8 scale rows as compared with ca. 10 in Hispaniolan forms), 
while in the Puerto Rican forms concurrently with increase in 
the size of the middorsal scale zone, there is an increase in the 
number of rows enlarged {ca. 4 in krugi, ca. 12 in pulchellus, 
1.5+ in poncensis). 

The evaluation of these resemblances, which are in each case 
beset with significant differences, is difficult. Parallelism is very 
probable, and it is especially likely that the Puerto Rican series 
is an independent radiation witliin Puerto Rico from the same 
stock that gave rise to A. crisfatellus, A. stratidus, A. guncllachi 
and A. evermanni. The primitive member of the Puerto Rican 
series, A. krugi, is not very different from cristatellus and 
guncllachi and would certainly be classed with them except for 
its obvious position at the base of a small grass anole radiation 
on Puerto Rico. 

The Cuban anoles which display a strong structural affinity 
in spite of some hahilat difference are more probably close 
relatives of the Hispaniolan series. There is in fact no substan- 
tial reason for doubting the relationship. 



6 BREVIORA No. 136 

It must be pointed out that the squamation pattern with a 
strongly developed middorsal zone of enlarged keeled scales, 
smaller laterals, and strongly keeled ventrals as large or larger 
than the middorsals is common in mainland Anolis, particularly 
so in Central America. This pattern occurs also in the Greater 
Antilles in three species which, though certainly anoline, are 
currently referred to other genera: the Cuban species {ophio- 
lepis) to Norops, and a species from Navassa (harboiiri) along 
with one from Hispaniola (wetmorei) to ChamaeUnorops. 

The mainland forms exhibit a whole spectrum of conditions 
in regard to the distinctness, number of scale rows, size of 
scales involved in the dorsal zone, etc. No described form seems 
close enough to the Hispaniolan or Cuban grass anoles to be 
worth serious consideration as representing the ancestral stock. 

Norops opliioh'pis, which occupies the grass anole habitat in 
Cuba, does not seem related either. It has some features peculiar 
to itself —  the reduction of the canthal ridge scales to two, the 
small number of scales in the loreal area {ca. 10-12), the very 
elongate scales betv/een the nostrils, the large mental scales — 
that are unlike not onh- the semilineatus-alutaceus groups but its 
supposed congeners on the mainland. The relationships of 
ophiolcpis are probably with Anolis sagrei and more remotely 
with the homolechis complex ; there are certainly no grounds for 
postulating close affinity to the scmilineatiis-ahitaceus set. 

ChamaeUnorops harhouri and C. wetmorei are even more dis- 
tinct. The basic pattern of squamation is quite heterogeneous 
and yet upon this has been imposed a second pattern of enor- 
mously enlarged keeled dorsals and hugely enlarged keeled 
ventrals exaggerated beyond that seen in any other forms. 

This picture, like the apparent radiation of forms on Puerto 
Rico and the extraordinarily varied array of forms on the 
mainland, suggests strongly that the pattern — enlarged mid- 
dorsal zone, enlarged keeled ventrals — is one of several stereo- 
types that the anoles have again and again produced, that this 
is one of a limited set of squamation patterns possible to the 
anolines and therefore produced in parallel fashion in many 
times and places. 

It is this parallelism that contributes to the notorious "dif- 
ficulty" of the genus Anolis. Narrow groups are rather easy to 
recognize (though the specific and infraspecific structure within 



1961 ANOLIS SEMILINEATUS GROUP 7 

the group may be puzzling in the extreme) but wider relation- 
ships (at least when externals only are considered) are problem- 
atical, becoming- obscurer with each step more distant from the 
species group. 

Origin of the semilineatus group 

The species of the semilineatus group are more uniform than 
the related Cuban series. They most resemble cyanoplcurus, the 
middle term in the morphological series of Cuban forms. This 
species has its range in extreme eastern Oriente and is thus 
geographically closest to the Hispaniolan group. It therefore 
seems probable that the semilineatus series on Hispaniola has 
been rather recently derived from a cyanoplcurus-VikQ Cuban 
ancestor but has been on Hispaniola long enough to achieve 
island-wide dispersal and moderate dilferentiation at the specific 
level. 



Acknowledgments 

I have had the advantage of discussions with A. Stanley Rand 
and with Dr. Hicliard Etheridge. The latter 's osteological evi- 
dence for species groupings within Anolis (unpublished thesis, 
University of Michigan) has in part confirmed, in part guided my 
own thinking on the wider relationships of Anolis species. 

The map-diagram was prepared by Patricia Grubbs. 



REFERENCES CITED 

Mee.tens, R. 

1939. Herpetologischer Ergebnisse eiiier Reise nach der Insel His- 
paniola, Westindien. Abhandl. Senckenberg. naturf. Ges., 449: 
1-84. 

Wetmoee, a. and B. H. Swales 

1931. The birds of Haiti and the Dominican Republic. Bull. U.S. 
Nat. Mus., 155: 1-483. 
Williams, E. E. and A. S. Rand 

1961. Notes on Hispaniolan Herpetology. l'. A review of the Anolis 
semilineatus group with the description of Anolis rochranae, 
new species. Breviora, No. 135 : 1-11. 



BREVIORA 



No. 136 





^ ^ 3 S --S 
. o 



a; 



o 



o 

03 



CO g^ 



c 

m 
Ol 

a 

Ol 



3 

c3 
I- 

o 



o 

s 



05 






^ 



o 



•^ .s 



CO o 

52 OJ 



o 



in : 



ft 



> ^ 
.S « 
-^3 e 



05 



10 



15 J2 






S o 



O 

ft ^ 

O 



d 



t/) J- "^ o .2 

«. V =H 'J^ '■« 



*^ s 

o :;: 

£ c 

;« -rl 

o _ 

«H ft 

el 2 

"^ S 

<^ en 

(D 
;-> 

:§ 1 

? ^ 

■73 •-* 

A en 

o -^ 
ft 

05 



Jh ^ 



o 



OP 






-si 

o -o 

.a 2 



O 



03 '^ 



® 0) 



3 

o ^ 
^ §'^ 

" g =2 

*j o *^ 

O >» 13 

'^ S o 

CO ,^ 

5:; S » 
o 3 fc- 



BREVIORA 



MusenaiM of Comparative Zoology 



Cambridge. Mass. April 10, 19(31 Number 137 

NOTES ON HISPANIOLAN HERPETOLOGY 

4. ANGUS KOOPMANI, NEW SPECIES, FROM 
tllE SOUTHWESTERN PENINSULA OF HAITI 

By a. Stanley Rand 

Biological Laboratories, Harvard University 

Among the reptiles and amphibians collected for the Museum 
of Comparative Zoology in Haiti, with the support of an Ameri- 
can Philosophical Society grant, during the summer of lfJ60, 
are seven specimens that appear to represent an undescribed 
species of Anolis. 

These lizards are small, moderately proportioned, dull colored 
in life, as well in preservative, and rather nondescript animals. 
In life the adult males possess a dark gray gular fan and an 
orange-pink chin and throat that distinguish them immediately 
from any otlier Hispaniolan Anolis. When this chin and throat 
color disappears in alcohol, the combination of scale characters 
distinguishes the species, but there are no unique characteristics. 

Dr. Karl Koopman, Assistant Curator of Mammals, Chicago 
Natural History Museum, provided financial assistance and 
encouragement that helped to make the collection of these 
specimens possible. In recognition of his aid the new species is 
called : 

Anolis koopmani new species 

Type. MCZ 62541, adult male. 

Type locality. Carrefour Canon, 350 m. altitude, near Ducis, 
N. of Aux Caves, Haiti. 

Collector. A. S. Rand and J. Lazell, 4 August 1960. 

Paratypes. Adult males, MCZ 62542-3; adult females, MCZ 
62544-5 ; young males, MCZ 62546-7. All from Les Platons, 750 



2 BREVIORA No. 137 

in. altitude, above Carrefour Cauoii, Haiti, A. S. Hand and J. 
Lazell, 5 August 1960. 

Diagnosis. The presence of a zone of much enlarged middorsal 
scales, keeled head scales, and keeled, imbricate, and pointed 
ventrals distinguish this species from all the Ilispaniolan Anolis 
except those of the seynilineatus group {stmilineatus, olssoni, and 
cochranae) . It is distinguished from the latter in having 6-8 (not 
10) enlarged middorsal rows, 5-8 (not 3-5) loreal rows, and 3-5 
(not 1-3) scales separating the interparietal from the supraorbi- 
tal semicircles. It differs also in coloration, the males having 
pinkish-orange chin and throat. 

Description. (In the following description variations occurring 
in the paratypes follow, in parentheses, the condition in the 

Head. Head scales strongly keeled. Frontal dejiression mod- 
erate. Scales across snout between second and third canthals 8 
(7-9). Nares anterior to canthal ridge; separated from rostral 
by 1 scale. Canthal ridge distinct, not exaggerated, composed of 
4 (4-5) large scales preceded by 2 (1-3) small ones. Second 
canthal largest, third next in size, first and fourth subequal. 

Posterior frontal subequal to (slightly smaller than) anterior 
supraorbital ; separated from canthals by 2 (1-2) scales. Anterior 
supraorbital separated from canthals b}' 3 (2-3) scales. Supra- 
orbital semicircles separated by 3 (1-3) scales; separated from 
supraocular disc by one row of small scales (occasional narrow 
contact). Supraocular disc of 5-6 (5-7) enlarged keeled scales; 
separated from superciliarjr by 5-6 rows of granules. A single 
elongate superciliary. Interparietal scale slightly smaller than 
ear (lA to slightly smaller) ; separated from supraorbital semi- 
circles by 4-5 (3-5) scales. 

Scales in center of supratemporal area, granular, smaller 
than flank scales, smallest in center. Scales over temporal bar 
not (very slightly) enlarged. Temporal scales like supratem- 
poral scales. Suboculars broadly in contact with supralabials, 
separated from canthals (very narrow contact), not continued 
behind eye as a series of large scales. Supralabials to center of 
eye 5 (5-6). Loreal rows 5 (5-8). Loreal scales subequal in size. 

Mentals broader than long, in contact with 6 (5-6) throat 
scales posteriorly. Xo series of enlarged sublabials. Central 
throat scales small, elongate, keeled. 

(hilar fan: Gular fan small. Scales slightly smaller than (sub- 
equal to) ventrals, keeled. 

Trunk: Middorsal scales much larger than flank scales, grad- 



i:»g: 



ANOLIS KOOPMANI 



iug into them. Rows of enlarged middorsals 6-8. Veutral scales 
larger than middorsals, imbricate, keeled. 

Limbs and digits: Scales on arms and legs larger tlian ven- 
ti-als, multiearinate. Hand and foot scales multicarinate dorsally. 
Lamellae under phalanges 2 and 3 of fourth toe 18 (17-19). 
Interdigital pads narrow. 

Tail: Tail round in cross-section. Verticils indistinct. En- 
larged postanal scales present in males. 

MEASUREMENTS 







Snout-vent 


Total 


Head 


Tibia 


Hvnd ley 


Sex 


MCZ # 


length 


length 


length 


lengtli 


length 


male 


62541 


34 mm 


114 mm 


9 nmi 


11 mm 


28 mm (type 


< 1 


62542 


39 


135 


10 


12 


33 


( 1 


62543 


38 




10 


12 


31 


i i 


62546 


23 


68 


6 


7 


18 


( I 


62547 


21 




6 


7 


17 


female 


62544 


33 




7 


9 


26 


i t 


62545 


33 


109 


8 


10 


26 



In life a low nuchal and dorsal crest seems permanently raised 
(absent in females and young males). 

Color in life: Male, MCZ 62542, uniform gray-brown above; 
a whitish line, black edged above, from over shoulder to hind 
leg, indistinct for the posterior half of its length, below this line 
the flanks with scattered dark spotting. Belly light brown, chin 
and throat pale pinkish-orange, with a few scattered black spots, 
gular fan scales colored like the chin, but the skin dark gray; 
iris blue. 

Female, MCZ 62544, plain brown above, a middorsal stripe 
with a scalloped margin outlined with darker brown, a yellow 
stripe from below eye to over shoulder, continued faintly to 
hind leg; venter yellov/ish with faint dark spotting on throat; 
iris blue. . •..,. , 

Habitat: The type was taken in a bush along a trail "git the 
edge of a coffee grove. It was found at dusk among small twigs 
about three feet above the ground where it probably had climbed 
to spend the night. 

Of the six paratypes, three adults were found in heavily 
shaded coffee groves. All were on the ground among damp leaf 
litter. One juvenile was found six inches up in low herbaceous 
vegetation at the edge of the coffee grove. The others were 
purchased from a small boy for two cents each. 

Relationships: The relationships of this species are obscure; 



4 BREVIORA No. 137 

it does not seem to be particularly close to any species now 
known either from Ilispaniola or elsewhere. 

The only Anolis outside of Hispaniola that are at all similar 
to this species are the alutaceus, clivicolus, spectrum, cyanopleu- 
rus groups of Cuba, and this similarity lies primarily in the 
presence of a zone of enlarged middorsals and does not extend 
to other details. I interpret this as parallelism. 

Within Hispaniola, A. koopmani is superficially most like 
semilineatus and olssoni. A zone of enlarged middorsals, 
keeled imbricate, pointed ventrals, keeled head scales, a lateral 
stripe, narrow digital expansions and small size occur also in 
.semilineatus and olssoni and suggest a relationship to them. 
However, most of these characters are not unique to semilifieatus 
and olssoni even in Hispaniola. The nature of the zone of 
enlarged middorsals (fewer rows that decrease in size laterally), 
the strong sexual dimorphism in color, the less attenuate body 
shape, and generallj' smaller scales all argue against this rela- 
tionship. 

Anolis ricordi, clistichus, cybotcs, armouri, shrevei, chloro- 
cyanuSyCoelestinus and Chamaelinorops wetmorei all have special- 
izations that seem to exclude them from close relationships with 
koopmani. 

The remaining species, Anolis monticoJn, darlingtoni, chr'sto- 
phei, heridcrsoni, hahanicocnsis and Xiphoccrcns darlingtoni are 
poorly known. It is possible that some if not all of them are 
closely related to one another and that koopmani's relationships 
lie with these. However, until more information is available 
for this assemblage of species, particularly in regard to color in 
life, behavior and ecology, this hj-pothesis must remain very 
tentative. 

Acknowledgments: I am grateful to Dr. Ernest E. Williams 
for his advice and to Mr. James Lazell and M. Luc Whiteman for 
their assistance in the field. I wish also to thank M. Leonce 
Bonnfil fils and the other members of the Department of Agri- 
culture of Haiti who helped us in so many ways. 



BREVIORA 

Mmseiuiioi of Comparative Zoology 



("A:Mi{Hir)(;E, ]Mass. June 14, 19G1 Number 138 

PFEIFFER'S UNFIGURED SPECIES OF 
STROPHOCHEILVS {MEGALOBULIMUS ) 

By T. E. Crowley and T. Pain 



This paper is a supplement to J. C. Bequaert's "Moiioy,Taph 
of the Strophoeheilidae, a Neotropical Family of Terrestrial 
Mollusks" (Bull. Mus. Comp. Zool., Harvard, 100: 1-210, 1948). 

When Dr. J. C. Bequaert published his monograph on the 
Strophoeheilidae he was unable to deal in detail with two species 
described by Pfeiffer from specimens in the Cumino' collection, 
now in the British Museum (Natural History), London. Neither 
of these had ever been figured, and the types are so far the only 
specimens known. Opportunity has, therefore, been taken to 
complete Dr. Bequaert's monumental work with figures of 
Pfeiffer 's almost unknown species. In addition we are describ- 
ing and figuring S. (M.) capiUaccus (Pfeiffer), so far unfigured, 
and S. indigcns Fulton. 

The authors wish to express their grateful thanks to the Brit- 
ish IMuseum authorities for permission to examine and photo- 
graph the types for reproduction herein, to Dr. R. Zischka for 
specimens of S. (M.) nuligens Fulton, to Drs. W. Blume and 
W. Weyrauch for the loan of material, and to Mr. S. P. Dance 
and Mr. J. A. AYillson for their generous assistance in photo- 
graphing specimens. 

Strophocheilus (Megalobulimus) hector (Pfeiffer) 

Plate 1, Figure 1 

BuUmris hector Pfeiffer 1857, Malak. Blatt., 4, p. 157 (Brazil); 1859, 
Monosr. Helie. \iv., 4, p. 367; 1868, Op. cit., 6, p. 11; 1876. Op. rit., 
8, p. 15; 1877, Op. cit., 8, p. 604. Paetel, 1889, Cat. Conch. Samml., 
4tli ed., 2, p. 212. 

BuUmus (Bonis) hector von Martens 1860, in Albers, Die Heliceen, 2nd 



2 BREVIORA No, 138 

ed., p. 192; 1876, Xovit. Com-hol., Abt. 1, 5, pts. 50-51, p. 21. Pfeiffer, 

1879, Nomeiicl. Helie. Vic, p. 224. 
Strophocheihis (Thaiimastus) hector Pilsbry 1895, Man. of Conch., (2) 10, 

p. 50. 
ThaumasUis hector Pilsbry 1902, Man. of Conch., (2) 14, Classification, p. 

xxi. 
Strophocheilits {? MegalohuUmus) hector Pfeiffer, Bequaert 1948, Bull. 

Mus. Comp. ZooL, Harvard, 100: 118. 
Original description: "T. subimperforata, elongato-ovata, 
solidula. sub epidermide decidua, fulvida alba; spira conica, 
apice rotundata; anfr. 6 convexiusculi, summi conferte capillaceo 
striati, ultimus spiram vix superans, plicato-striatus et obsolete 
decussatus ; columella leviter arcuata, nou plicata ; apertura sub- 
verticalis, acumiuato-ovalis, intus albida, nitida ; perist. album, 
marginibus callo albo junetis, dextro subiuerassato, brevissime 
expanso, columellari superue dilatato, adnato. — Long. 71, diam. 
35 mill., Ap. 36 mill, longa, 191/2 lata." 

New Measurements of Adult Holotype 





Greatest 


Aperture 


Aperture 




Length 


Width 


Length 


Width 




71 mm. 


39 mm. 


35 mm. 


26 mm. 


6 whorls 



Specimen examined: Brazil (Miers Coll.), holotype (Brit. 
Mus., Nat. Hist.). 

Remarks. Bequaert (11)48, p. 118), who had seen no speci- 
mens, was inclined to exclude hector from the Strophocheilidae, 
and Pilsbry (1895, p. 50) placed it in Thaiimastus. Von Mar- 
tens (1876), however, suggested that it might be related to 
Stropliocheilns (M.) ohlongus (Miiller). 

Careful examination of the type has con\dnced us that it is 
indeed correctly referred to the Strophocheilidae, being by 
reason of its nepionic sculpture a member of the subgenus Mega- 
lohulimus. It does not appear, however, to be in any way 
related to S. (M.) ohlongus, the shell being longer and narrower 
in proportion, much paler and with a white lip. It is further- 
more much thinner, the apical sculpture finer, and is covered 
with a brown periostracum. 

S. (M.) hector would appear to us to be quite distinct from 
any other species of Megalohulimus so far known. 



1961 STROPHOCHEILUS 3 

Strophociieilus (Megalobulimus) cocapatensis (Pfeiffer) 

Plate 1, figure 2 

Bulimus cocapatensis Pfeiffer 1855 (August), Proc. Zool. Soc. London, 
(for 1855), p. 115 (Coeapata, Bolivia); 1859, Mouogr. Ilelic. Viv., 
4, p. 367; 1868, Op. cit., 6, p. 11; 1876, Op. cit., 8, p. 15. Paetel, 1889, 
Cat. Conch. Samnil., itli ed., 2, p. 209. 

Bulimus {Bonis) cocapatensis Pfeiffer 1856 (January), Malak. Blatt., 2, 
(for 1855), p. 1-17. Von Martens, 1860, in Albers, Die Heliceen, 2nd ed., 
p. 192; 1876, Xovit. Conchol., Abt. 1, 5, pts. 50-51, p. 9. Pfeiffer, 1879, 
Nomencl. Helic. Viv., p. 224. 

Strophociieilus (Borus) cocapatensis Pfeiffer, Pilsbry 1895, Man. of Conch., 
(2) 10, p. 20. 

Strophocheilus (Borus) cocopatensis Pfeiffer, Pilsbry 1895, Man. of Conch., 
(2) 10, p. 12; 1902, Op. cit., (2) 14, Classification, p. v. Misspelling of 
cocapatensis. 

Bulimus corapatensis Pfeiffer, Paetel 1889, Cat. Conch. Samml., 4th ed., 2, 
p. 10. Misspelling of cocapatensis. 

Strophocheilus cocopatensis Pfeiffer, Pilsbry 1930, Proc. Ac. Xat. Sci. Phila- 
delphia, 82, p. 355. Misspelling of cocapatensis. 

Strophocheilus (Megalohulimiis) cocapatensis Pfeiffer, Bequaert 1948, Bull. 
Mus. Conip. Zool., Harvard, 100, no. 1, p. 126. 

Original description: "B. testa imperforata, ovato-oblonga, 
solida, minutissime decussata. sub epidermide virenti-fulvida 
violaeeo-earnea ; spira convexo-euiiiea, apiee obtusa; sutura al- 
bida, irregulari ; anfr. oYo superis radiatim costatis et minutis- 
sime granulatis, sequentibus peroblique descendentibus, parum 
couvexis, ultimo spiram sub-aequante, basi rotundato ; columella 
recedente, leviter arcuata; apertura subverticali acuminato- 
ovali, intus margaritacea ; perist. iiicrassato, breviter expanse, 
marginibus eallo nitido junctis, eolumellari dilatato, adnato. 
Long. 67, diam. 30 mill." 

Measurements of Adult Shells 





Greatest 


Aperture 


Aperture 






Length 


Width 


Length 


Width 






67 mm. 


33 mm. 


31 mm. 


21 mm. 


5^4 whorls. 


Holotype 


67 


33 


31 


20 


514 whorls. 


Para type 


66 


33 


32 


20 


5i/> whorls. 


Para type 



Specimens examined: Coeapata, Bolivia (Bridges Coll.), 3 
types from the Cuming Collection (Brit. Mus. fXat. Hist.]). 

Reynarlxs. As pointed out by Bequaert (1948. p. 127), the 
radially ribbed and minutely granulated nepionic whorls, men- 
tioned in Pfeiffer 's original description, are characteristic 



BREVIORA 



No. 138 



of Mcgalohulimus, to which subgenus S. cocapateusis un- 
doubtedly belongs. Pilsbry (1930, Proe. Ac. Nat. Sei. Philadel- 
phia, 82, p. 355), in describing his S. carrikeri, infers that it may 
be related to 8. cocapatensis, but we are unable to see any justifi- 
cation for this assumption. Bequaert suggests that it is not im- 
possible that cocapatensis may be the same as S. intcrtextus 
Pilsbry, but comparison of a specimen of the latter with Pfeif- 
fer's type has convinced us that they are in no way related. 

The shell of cocapatensis is imperforate, long, thin and deli- 
cate, brown in color, with a very streaky, pink-fiushed appear- 
ance. The spire is attenuated, the apex pointed, the mouth 
long and narrow, the aperture brown within, and the outer lip 
thin, white, slightly reflected. Columella and callus white. 

Pfeiffer compared cocapatensis with S. rosaceus, but, as pointed 
out by Bequaert, the nepionic sculpture is typical of Megalohuli- 
inus and is not found in CliiUhoi'us, to which subgenus S. 
rosaceus belongs. Pfeiffer later 1856) placed it between "S. 
matthewsi" {— leucostoma) and *S'. capillaceiis but, as he does 
not show it as being- then in his collection, this opinion would 
seem of little value. 

Strophocheilus (Megalobulimus) capillaceus (Pfeiffer) 

Plate 1, fig-ure 3 

Bulimus capillaceus Pfeiffer 18.35 (July), Proc. Zool. Soc. London, (for 

1855), p. 93. 
StropJiocheilus (Bonis) capillaceus Pfeiffer, Pilsbry 1895, Man. of Conch., 

(2) 10, p. 31, PI. 14, fig. 69. 
StrophocJtcilus {Mcgalohulimus) capillaceus Pfeiffer, Bequaert 1948, Bull. 

Mus. Conip. Zool., Harvard, 100, p. 120 (full synonymy;, PI. 14, fig. 5. 
The type of *S'. (M.) capillaceus is in the British Museum 
(Nat. Hist.), from the Cuming Collection. It consists of three 
syntypes, of which that figured herein is now chosen as lectotype, 
no holotype having been selected by Pfeiffer. 

Measurements of Adult Shells 





Greatest 


Aperture 


Aperture 




Length 


Width 


Length 


Width 




64 mm. 


38 mm. 


37 mm. 


24 mm. 


5 whorls. Lectotype 


60 


39 


35 


30 


5 whorls. Sjnitype 


S3 


33 


23 


19 


5 whorls. Syntype 


67 


40.5 


40.5 


22 


5% whorls. Huanaco 


69 


42 


41 


21.5 


5% whorls. Santa Ana 



1961 



STROPHOCHEILUS 




Plate 1 



Fig. 1. Strop1(0chcihtf; {McgalobuUmm) hector (Pfeiffer). Holotypt". 
iiat. sizt". Fig. 2. Sirophoclteilits (Mcgalohulimus) cocupatensis (Pfeiffer). 
Holotyi)o, nat. size. Fig. 3. Strophoeheihis (Mer/alobulimus) capiUaceus 
(Pfeiffer). Lectotype, nat. size. Fig. 4. Slrophochcilus (ilegaJohuJiinus') 
maximus indigens Fulton. Apical aspect of immature shell, much enlarged. 



6 BREVIORA No. 138 

Sprciyucns examined : "Banks of the Kiver Solimoes, " Peru 
(Cuming Collection). Near Santa Ana, Kio Urubamba, Peru, 
3500 ft. (W. Weyrauch Coll., in Pain Collection). Huanaeo, 
Peru (Pain Collection). 

Remarks. Bequaert (1948, p. 120) has dealt at length with the 
probable relationships and position in the subgenus of S. capil- 
laceus, and gave an excellent figure of it, together with a com- 
plete synonymy. To this very full account there is nothing 
which we can profitably add. 

Strophocheilus (Megalobulimus) maximus indigens Fulton 

Plate 1, figure 4; Plate 2, figures 5, 6 

Bulimus Jcremnoicus d 'Orbigny 1837, Voyage Am6r. Meridion., 5, pt. 3, Moll., 

p. 300 (in part only: some specimens from Yuraeare, Bolivia, the 

locality given in Explanation of Plates, p. 695, for fig. 3), PI. 35, fig. 3 

only. Not Helix kremnoica d 'Orbigny 1835. 
Stroplwcheilus (Bonis) maximus ? var. Icremnoicus d 'Orbigny, PUsbry 

1895, Man. of Conch., (2) 10, p. 16, PI. 5, fig. 28 (copy of d'Orliigny's 

fig. 3); 1902, Op. 0(7., (2) 14, Classification, p. iv. 
Strophocheilus (Borus) indigens Fulton 1914, Proc. Mai. Soc. London, 11, 

pt. 3, p. 165, fig. (Peru). 
Strophocheilus (Megalobulimus) iyidigens Fulton, Bequaert 1948, Bull. Mus. 

Comp. Zool., 100, No. 1, p. 98, PI. 24, fig. 1 (copy of Fulton's 1914 fig.). 
Strophocheilus indigens Fulton, Blume 1952, Arch. f. Mollusk., Frankfurt, 

81, pts. 4-6, p. 105. 
Strophodheilus (Borus) l-remnoieus subsp. vcstitus Pilsbry 1926, Proc. Ac. 

Nat. Sci. Philadelphia, 78, p. 6 (Bolivia, probably in Dept. Cochabamba), 

PI. 2, fig. 7. 
Strophocheilus (Megalobulimus) maximus vestitus Pilsbry, Bequaert 1948, 

Bull. Mus. Comp. Zool., Harvard, 100: 94, PI. 19, fig. 4. 

Original description: "Shell ovate-oblong, yellowish brown, 
moderately solid; spire about 13 mm. longer than the aperture; 
whorls 61/2? apex smooth, the second and third whorls with prom- 
inent oblique plications, last two volutions polished and appar- 
ently smooth, but under the lens are seen to be finely granulated, 
the granulations being strong on the middle whorls and gradu- 
ally becoming weaker towards the aperture ; the lower whorls 
have also some irregidar and almost obsolete plications ; aperture 
sub-oval, whitish within ; peristome thickened and very slightly 
expanded, white, margins joined by a moderately thickened 
white callus. Alt. 110, Diam. Maj. 47 mm. The nearest species 
to this is *S'. (Borus) huascari Tschudi, which is broader, has a 
wider aperture, a rougher and duller surface, and its apical 
plications are much finer and closer together than in indigens." 



1961 strophocheilus 

Measurements of Adult Shells 

Length Greatest Aperture Aperture 

Width Length Width Whorls 
135 mm. 57.5 mm. 60 nun. 41 mm. 7 Bolivia: Sacha, Yungas, 

800-1500 m. 
(Bavarian State Mus.) 



132 


63 


59.5 


37 


7 


127 


58 


56 


39 


7 


128 


58 


55 


31.5 


6Y2 



118 


58 


56 


24 


— 


116 


48 


47 


31 


6% 


101 


•16 


47 


26 


51/0 


100 


49 


48 


27.5 


51/2 



Bolivia (W. Weyraudi 
Collection) 
118.5 61 57.5 40.5 6 Bolivia: Chapare, 400 m. 

(Bavarian State Mus.) 
Type of vestitus Pilsbry 
Holotype of indigens, Peru 
Bolivia (T. Pain Collection) 
Bolivia (W. Weyrauch 
Collection) 

Specimens examined: Peru, type of indigens Fulton (British 
Museum [Nat. Hist.], No. 1915-1-5-199). Yungas cle Palmar, 
1200 m., Bolivia (R. Zischka Coll., in W. Blume, T. Pain, and 
W. Weyrauch Collections). 

Remarks. Comparison of the type of indigens, together with 
the shells from Bolivia, with Bequaert's (1948) figure of vestitus 
Pilsbr3% leave us in no doubt that they are identical. All shoAV 
the strong, prominent oblique plications on the second and third 
whorls, noticeably absent on both the typical maximus and the 
subspecies huascari. As pointed out by Bequaert (1948, p. 94), 
indigens { = vestitiis) is of considerable interest in that it bridges 
the gap between typical maxim its and subspecies huascari in re- 
spect of its relatively wider spire, narrower body-whorl and 
smaller mouth. 

Fulton, who described indigens from a unique holotype, did 
not apparently connect it with maximus, although he drew at- 
tention to its close resemblance to huascari. 

Bequaert (1948, p. 94), dealing with vestitus, makes no men- 
tion of the prominent sculpture, although this is easily recog- 
nized in his excellent photograph of the shell he selected as 
holotype from Pilsbry 's type set (Ac. Nat. Sci. Philadelphia 
No. 138105). Dr. Weyrauch informs us (in lift.. 1958) that he 
considers the elongated shell from Oxapampa, Peru, referred by 
Bequaert to vestitus, to be a typical maximus on account of the 
aperture being much longer than in ineligens (= vestitus). A 
similar elongated shell from Peru, without more definite locality, 



BREVIORA 



No. 138 



kindly sent by Dr. Weyraudi. sliows traces of a dark periostra- 
cum and, althonjili much worn aliont tlie spire, has the long aper- 
ture characteristic of typical ))ioximiis. 

S. [M.) ntn.ri))U(s ijuligois Fulton has not so far been ob- 
tained in Peru by Dr. Weyrauch, but, from the similarity of the 
fauna of southeastern Peru and northeastern Bolivia, there can 
be little doubt that iiidigcns occurs also in Peru. 




Plate 2 

Fig. 5. Strophocheilus (Megalobulimus) maximus indigens Fulton. Hol- 
otype, nat. size. Fig. 6. Strophocheilus (MegaloiuUmus) maximus indigens 
Fulton. Yungas de Palmar, Bolivia, nat. size. 



BREVIORA 

Mmseium of Comparative Zoology 



Cambridge, ]Mass. JiiXk 15, 1961 Numbef^ 139 



A NEW SPECIES OF SPIIAEEODACTYLVS FKOM 

NORTHERN HAITI 

By Jaaies D. Lazell 

Although the genus Sphaerodactylus on Ilispaniola is suf- 
ficiently diversified and confused to warrant at least a partial 
revision, the species here described is so remarkably different 
from any other form that I am confident in naming it at this 
time. 

The new species is named for Mr. Benjamin Shreve of the 
Museum of Comparative Zoology for his current work on the 
sphaerodactyls of Hispaniola. 

Sphaerodactylus shrevei sp. nov. 

Type: MCZ No. 62548, Mole Saint Nicolas, Haiti. Coll.: J. 
Lazell and A. S. Rand, 16 July, 1960. 

Diagnosis: The combination of the following characters serves 
to distinguish this species from any other found in the Antilles: 
the presence of a highly convex snout as seen from the side 
(loreal region also somewhat convex) ; very large keeled dorsal 
scales beginning at the level of the axilla ; and the paravertebral 
arrangement of these dorsals, producing a middorsal zone not 
of granules but of small and large, irregularly placed scales. 

Description of type. Snout short. Eye nearer tip of snout 
than ear. Snout, as seen from the side, highly convex. Loreal 
region also somewhat convex. Rostral large with a partial 
median cleft. Nostril between rostral, first supralabial, a large 
supranasal (= internasal) and two small postnasals. A single 
scale between the supranasals (= internasals), which border the 
rostral posteriorly. Granular scales on top of snout somewhat 
larger than interoeular or nape scales. Four supralabials, of 



2 BREVIORA No. 139 

about equal length, to the center of the eye, followed by two 
smaller ones. Four infralabials gradually decreasing in size 
followed by two abruptly smaller ones. Mental short, wider than 
long, bordered posteriorly by two postmentals. Supraciliary 
spine small. Squamation of head and neck granular to the level 
of the shoulder; at that level a moderately rapid change to 
large, flattened, heavily keeled dorsals. Twenty-five dorsals 
from the level of the axilla to the posterior level of the hind 
limb. Five dorsals in the standard distance. A very ill-defined 
middorsal zone of smaller keeled scales not forming a continuous 
row but with the large dorsals meeting irregularly along the 
middorsal line. Middorsals subimbricate or not imbricating, the 
laterals more distinctly imbricating. Throat scales smooth, 
granular, juxtaposed. Chest and belly scales larger, smooth, 
cycloid, broadly imbricate, about nine ventral scales in standard 
distance. Scales of anterior surfaces of limbs imbricate, cycloid, 
smooth, somewhat smaller than ventrals. Scales of posterior 
surfaces of limbs granular, smooth. Digital pads approximately 
twice as broad as the subdigital lamellae. Ten infradigital 
lamellae under fourth toe. The type, the only specimen yet col- 
lected, lacks the tail. It is a female and the escutcheon there- 
fore cannot be described. 

Coloration in life. S. shrevei is a dull-colored animal with a 
pattern composed of three basic hues — each tending to be 
unique on an individual scale. There are very irregular dark 
grey-broW'U blotches across the dorsum ; beginning at the back 
of the head there are three such markings to the slioulders. 
There are three more crudely "Y" shaped markings on the 
body, the most anterior of which bifurcates to the right, the 
remaining ones bifurcating to the left. There are two small 
blotches on the right side of the rump and one on the left. The 
second transverse blotch, on the nape, is broken by a light 
middorsal line that continues down through the fourth marking 
and then fades out. The ground color of the dorsum is ash grey. 
There are scattered over the dorsal surface short transverse 
series of white or partly white scales — from two to four in a 
row — that appear to have no correlation whatever with the rest 
of the animal's pattern. The top of the head is ash grey except 
for a very irregular, dark, grey-brown blotch on the parietal 
area. Coming back from the eye are two stripes, one of which 
runs downward across the cheek ; the other nearly connects 
with the first transverse marking on the back of the head. Xot 



1961 SPHAERODACTYLUS SHREVEI 3 

including the stripe across the cheek, there are five vertical dark 
markings across the pale labials, the anterior two of which are 
connected at the edge of the mandible. There are dark streaks 
on the lateral edges of the chin and throat, the underside of the 
hind limbs, and across the venter just anterior to the anus. The 
ventral surface is white ; a close examination reveals that on 
each scale there are tiny black dots. This peppering becomes 
more noticeable laterally and posteriorly. Along each side of 
the animal is a line of partially connected, small, dark, grey- 
brown blotches; just ventral to this row is another composed 
of widely spaced, single, dark, grey-brown scales. All three 
of the animal's hues: white, grey and grey -brown, are simply 
variations in the intensity of speckling on each scale with tiny 
black or brown dots. 

The pattern of the animal bears no resemblance to that of 
the young or females in the species to which it has been com- 
pared, or to any other Sphaerodactylus I have seen. 

Habitat. The type specimen was taken from a large circular 
rock pile about two-and-one-half feet deep ; this sort of rock 
pile is the result of removing the debris from a heap of charcoal 
after burning, and is composed of rocks that vary in size from 
that of a golf ball to nearly the size of a football. This particu- 
lar heap was of some age, for even in the arid terrain of Mole 
Saint Nicolas several fair-sized thornbushes had sprouted up in 
it. Collecting was very difficult, for any animal uncovered 
could generally manage to dart back into the pile before the 
collector could safely ascertain that it was not a scorpion or 
some other unpleasant handful. In order to get best results we 
excavated areas through the pile, dividing it up into more 
manao-eable smaller piles; this system netted Cdcstns, TypJihrps. 
and Tropidophis, as well as the type of Sphaerodactylus shrevei. 
Another specimen of apparently the same species escaped, leav- 
ing only its tail behind. 

Due to the relative inaccessibility of peninsular north-western 
Haiti it may be some time before additional specimens can be 
obtained. 

Comparisons. From S. copei, the only other comparably large- 
scaled Hispaniolan form, S. shrevei differs in the following 
characters: (1) Snout seen from the side highly convex: (2^ 
No middorsal zone of gramdes; (3) Dorsal scales flatter, not 
swollen, apt not to imbricate, especially in the middorsal area ; 
(4) Pattern a series of irregular dark dorsal blotches with 



4 BREVIORA No. 139 

a line of often connected smaller blotches along each side ; dorsal 
blotches broken by a light middorsal line anteriorly. 

From S. scaber of Cuba, S. shrevei differs in all the men- 
tioned characters and in snout length, which averages slightly 
shorter in the Cuban form. 

S. samanensis Cochran of the Dominican Republic resembles 
^■. shrevei somewhat in squamation but the dorsals are smaller 
and the ventrals larger. There are no smaller middorsal scales 
and the pattern is very different. 

>S*. shrevei differs from S. becki of Navassa again in the 
absence of a middorsal zone of granular scales and in the 
flatness of the dorsal scales, which are rounded, swollen, and 
rather tubercular in 8. becki. 

Two Jamaican forms, S. richardsoni and 8. parkeri, occa- 
sionally possess dorsal squamation similar to that of 8. shrevei 
in that while there is no middorsal zone of granules there may be 
small scales irregularly scattered along the middorsal line, but 
m general the arrangement of the scales is much more regular 
and not of a paravertebral nature. From both of these species 
6'. shrevei differs in the following characters: (1) Snout seen 
from the side highly convex; (2) Head granules extending 
posteriorly to level of axilla instead of just onto nape; (3) No 
enlarged, clearly defined canthal scale; (4) Pattern and colora- 
tion. 

From 8. parkeri it differs also in having only a single small 
scale between the internasals. 

There is a ^'aguer resemblance to 8. shrevei in some Lesser 
Antillean forms. 8. vincenti and aS'. microlepis, for example, 
show a tendency towards reduction in size of the middorsal 
scales, but in these forms the pattern of squamation tends to be 
very regular and the scales are much smaller. 

The new species has been compared with these forms largely 
because there are apparently no closer ones, although it bears 
little resemblance to any of them. Its relationships at the 
moment are not at all clear. 

Acknowledgments. My thanks are due to Mr. A. S. Rand 
for help in collecting the specimen described, to Mr. B. Shreve 
for checking the description and com.parisons made and to Dr. 
E. E. Williams for the loan of comparative material and for 
readin'v the manuscript. A grant from the American Pliiht- 
sophical Society supported the expedition which resulted in 
the discoverv of 8. shrevei. 



1961 SPHAERODACTYLUS SHKEVEI 5 

EEPERENCES 

Barbour, T. 

1921. Sphaerodactylus. Mem. Mus. Comp. Zool., 47: 217-277. 
Cochran, D. M. 

1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus., 177: 
1-398. 

BUIBAL, R. 

19j9. a new Sphaerodactylus from Oriente, Cuba. Herpetologica, 
15: 89-93. 

Table 1 

Comparisen of the snout length ratios of six species of SpTiaerodactylus. 
The ratio is obtained by dividing the standard distance (i.e., the distance 
from the center of the eye to the tip of the snout) into the distance from 
the center of the eye to the ear opening. The higher the figure obtained 
the shorter the snout. 





Specimens 


Mean Value 


Range 


S. richardsoni 


5 


.72 


.67- .81 


S. parlceri 


3 


.76 


.73- .78 


S. copei 


10 


.76 


.70- .80 


S. shrevei 


1 


.80 


- 


S. scaber 


6 


.82 


.71- .92 


S. becki 


4 


.90 


.78-1.00 



BREVIORA 



Miiseiuim of Compsirative Zoology 

Cambridge, Mass. June 27, 1961 Number 140 

A PRELIMINAKY REVIEW OF THE NEARCTIC 
SPECIES OF SIEROLOMORPHA (HYMENOPTERA) 

By Howard E. Evans 



In the course of collecting Bethyliclae and examining material 
in various museums, I liave encountered a good many specimens 
of the curious bethylid-like genus Sierolomorpha Ashmead. Some 
of these specimens are so different from the type, ambigua, for 
a long time the only known species, that it seems desirable to 
provide names for them. Krombein (1951, U. S. Dept. Agri. 
Monogr. 2, p. 748) has already pointed out that there appear to 
be several species in North America. The present paper does not 
pretend to be an exhaustive treatment of this genus. Names are 
provided for several of the more distinctive species, but before a 
definitive revision is possible more collecting must be done and 
more detailed studies made of structural details and their vari- 
ation. 

Actually, Sierolomorpha is not a bethylid or even especially 
close to the Bethylidae. Schuster (1949, Ent. Amer., 29: 124) is 
correct in pointing out its close similarity to certain Mutillidae 
and Tiphiidae, and the arrangement in the Synoptic Catalog 
(Krombein, 1951, op. cif.), in which the genus is placed in a 
monogeneric family between the Tiphiidae and Mutillidae, is 
probably the best that can be achieved at present. Sierolomorpha 
is related to the most primitive Scolioidea, and may not be far 
from the stock which gave rise to the Bethylidae. However, in 
virtually all structural details it stands very much closer to the 
Tiphiidae than to the Bethylidae. 

Only one species of the genus, hospes Perkins, has been de- 
scribed from outside the Nearctic region (from Hawaii). Krom- 
bein has suggested that this species may have been introduced 
from North America. In the collection of the U. S. National 
Museum there is a single male of an undescribed species from 



2 BRRVIORA No. 140 

Panama. I have seen no other specimens from outside the United 
States and Canada. 

The most useful structures for the separation of species in this 
genus appear to be the antennae, especially the tyloides of the 
male (Figs. 1-6), the propodeum, and the first two abdominal 
tergites. There seems to be little variation within the genus with 
respect to the mandibles, elypeus, and most details of the thorax 
including the legs and wing venation. I have therefore made 
little mention of these structures in my descriptions. I have 
studied the male terminal ia of selected specimens and found some 
minor variation, particularly in the volsellar cuspis. However, 
the differences are so slight and subtle that further study to 
determine the extent of individual variation did not seem war- 
ranted. I have therefore made no mention of characters of the 
terminalia in the keys and descriptions. 

The following abbreviations are used for the museums and 
private individuals that supplied material for this study: AMNII, 
American Museum of Natural History, New York; CIS, Cali- 
fornia Insect Survey, Berkeley; CNC, Canadian National Col- 
lections, Ottawa; HKT, Henry K. Townes, Ann Arbor, Michi- 
gan ; MCZ, Museum of Comparative Zoology, Cambridge, Mass. ; 
UCD, UniverNity of California at Davis; USNM, U. S. National 
Museum. 

Key to Species 

Females ^ 



. Abdomen with the constriction between the first two tergites 
strong, the first tergite with a weak to strong apical trans- 
verse depression which is longitudinally striate ; propodeum 
with a moderately wide median groove which is irregularly 

margined by carinae 2 

Abdomen with the constriction between the first two tergites 
weak, first tergite smooth, not depressed or striate apically ; 
propodeum with median groove absent or linear and not 
margined as above 3 

. Legs beyond the coxae In-ight yellowish-brown ; basal segments 
of antennae suffused with light yellowish-brown ; notauli 
strongly diverging anteriorly (Florida and Arizona to 

Alberta, Ontario, and Massachusetts) 

canadensis (Provaneher) 

1 The females of two species, apache and hrevicornis, are unknown. 



1961 XEARCTIC SPECIES OF SIEROLOMORPHA 3 

Legs brown except front tibiae and tarsi yellowish-brown; 
antennae brown, sometimes weakly suffused with paler 
brown on sides of basal segments ; mesoscutum rather flat, 
notauli weakly diverging anteriorly (California, Arizona, 

and Colorado to Saskatchewan and Yukon) 

nigrescens n. sp 
3. Front, pronotum, and mesopleura wnth strong punctures 
pro- and mesonota bright rufocastaneous (Arizona) . . 

hicolor n. sp 
Front, pronotum, and mesopleura with only minute, widely 
spaced punctures; thorax entirely black (Georgia to Con- 
necticut and Kansas) similis n. sp. 

3Iales 

1. Abdomen with the constriction between the first two tergites 

strong, the first tergite with a weak to strong apical trans- 
verse depression which is longitudinally striate ; tyloides 
present on antennal segments seven through ten, short and 
rather prominently projecting (Figs. 1, 2) ; propodeum as 
in female (see couplet 1 of key to females) .... 2 
Abdomen with the constriction between the first two tergites 
weak or absent, the first tergite smooth, not depressed or 
striate apically ; tyloides not present on antennal segment 
seven or, if so, very long and low on segments eight 
and nine 3 

2. Tibiae and tarsi mostly or entirely light yellowish-brown in 

most specimens ; temples rather short, as seen from above 
much shorter than eyes (Florida and Arizona to Alberta, 
Ontario, and Massachusetts) . . . ca7ia densis CProyancher) 
Legs brown except front tibiae (sometimes also front tarsi 
and middle tibiae) light yellowish-brown; temples strongly 
developed, as seen from above nearly as wnde as the eyes 
(California, Arizona, and Colorado to Saskatchewan and 
Yukon) nigrescens n. sp. 

3. Antennae extremely short and compact, segment eleven about 

1.2 X as long as thick, segment thirteen about 1.5 X as 
long as thick; tyloides present on antennal segment eleven 
(Fig. 3) ; head extremely broad, about 1.25 X as wide as 

high (South Carolina) hrevicornis n. sp. 

Antennae elongate, segment eleven about twice as long as 
thick, segment thirteen much more than twice as long as 
thick ; tyloides not present beyond segment ten ; head 
1.1-1.2 X as wide as high 4 



4 BREVIORA No. 140 

4. Ocelli distinctly enlarged (diameter of anterior ocelhis about 

.22 X minimum width of front) ; tyloides present on 
antennal segments seven through ten (Fig. 6) ; front with 
a strong median groove in front of anterior ocellus 

(Arizona) apache n. sp. 

Ocelli not enlarged (diameter of anterior ocellus less than 
.2 X minimum width of front) ; tyloides present on seg- 
ments eight through ten (Figs. 4, 5) ; front with a weak 
median impression if any 5 

5. Punctures of front moderately strong ; tyloides rather short 

(Fig. 5) ; middle and hind tibiae and tarsi dull brown 

(Arizona) ? hicolor n. sp. 

Punctures of front very weak; tyloides elongate (Fig. 4) ; 
all tibiae and tarsi light yellowish-brown (Georgia to 
Connecticut and Kansas) similis n. sp. 

SiEROLOMORPiiA BicoLOR new species 

Holotype. — 9 , ARIZONA: Southwestern Research Station, 

5 mi. W. of Portal, Cochise Co., 9 August 1959, 5400 feet eleva- 
tion (H. E.Evans) [MCZ]. 

Description of type female. — Length 5.8 mm., length of fore 
wing 4.3 mm. Head black ; pronotum, mesoscutum, and scutellum 
w^holly bright ruf o-castaneous ; remainder of thorax and pro- 
podeum black ; abdomen very dark brown, approaching black 
basally; mandibles light brown, darker basally and apieally; 
clypeus suffused with reddish-brown apieally; scape black, flag- 
ellum dark brown, outer side of apical segments paler ; legs 
dark brown except front and middle tibiae and all tarsi light 
brown ; fore wing lightly, uniformly infuscated, veins and stigma 
dark brown. First four antennal segments in a ratio of about 
12:5:7:9, segment three 1.4 X as long as thick. Front strongly 
polished, punctures small but rather strong ; spacing of punctures 
rather irregular, those on the sides being mostly rather close, 
often not much more than their own diameters apart, those on 
the middle of the front rather sparse ; temples also with distinct 
punctures. Head subcircular in anterior view, vertex evenly 
rounded off a short distance above eye tops; inner orbits sub- 
parallel, minimum width of front about .9 the eye height. 
Ocelli small, in a broad triangle, the front angle greater than 
a right angle ; postocellar line slightly exceeding oeello-ocular 
line. Thoracic dorsum strongly polished, non-alutaceous ; pro- 
notum with strong, widely spaced punctures; mesoscutum with 



1961 NEARCTIC SPECIES OF SIEROLOMORPHA 5 

a very few punctures on the sides, impunctate medially. Pro- 
podeum strongly polished medio-basaliy, elsewhere slightly 
roughened by obscure punctures ; median line weakly impressed. 
Mesopleurum polished and with well-defined punctures. First 
abdominal tergite without a subapical depressed and striate 
baud, completel}^ smooth ; constriction between first and second 
tergites weak ; abdomen rather strongly hirsute beyond segment 
three. 

Male (assigned here tentatively). — ARIZONA: Cochise 
Stronghold, Dragoon Mts., 4850 feet elevation, oak-juniper zone, 
2 July 1947 (Werner & Nutting) [Univ. Arizona]. 

Description of male. — Length 5 mm., length of fore wing 
4.8 mm. Head and thorax entirelj^ black, abdomen dark brown ; 
apical half of mandibles light brown; antennae wholly dark 
brown; legs wholly dark brown except front tibiae and tarsi 
bright yellowish-brown; wings subhyaline. First four antennal 
segments in a ratio of about 20 :8 :13 :20, segment three 1.5 X 
as long as thick, segment four 2.2 X as long as thick, this segment 
typical of the remaining segments except the last ; segments eight 
through ten each with a short, rather weak longitudinal polished 
ridge (Fig. 5). Front polished, wholly covered with small but 
well-defined punctures which are separated by scarcely more 
than their own diameters ; vertex and temples more weakly 
punctate; front somewhat impressed along the inner orbits and 
beside the posterior ocelli, faintly impressed medially. Minimum 
width of front approximately equal to eye height; ocelli in a 
broad, flat triangle, postocellar and oeello-ocular lines subequal ; 
ocelli of moderate size, diameter of anterior ocellus .18 X mini- 
mum width of front. Pronotum short, shining and w^th dense, 
rather weak punctures. Mesonotura shining, rather sparsely and 
weakly punctate. Propodeum polished, impunctate, and non- 
alutaceous over most of its surface. Mesopleurum polished, 
densely but weakly punctate. First abdominal tergite smooth, 
without a subapical depressed and striate band. 

Remarks. — The type female was taken on the ground beneath 
oak trees in dry, open forest. The male associated with it tenta- 
tively was apparently taken in the same type of forest and at 
nearly the same altitude. 

SiEROLOMORPHA APACHE new spccies 

Holotype. — $ ARIZONA: 3-5 mi. SW of Apache, Cochise Co., 
8 August 1959, about 4300 feet elevation (H. E. Evans, on 
Baccharis glutinosa) [MCZ]. 



6 BREVIORA No. 140 

Description of type male. — Length 4.5 mm., length of fore 
wing 4.0 mm. Entire body dark brown, the head nearly black ; 
mandibles light brown ; antennae uniformly light brown ; legs 
dark brown except tibiae and tarsi light brown, the front tibiae 
a rather bright yellowish-brown ; wings hyaline. First four an- 
tennal segments in a ratio of about 15:9:13:17, segment three 
1.6 X as long as thick, segment four 2.0 X as long as thick, 
segment eleven 2.1 X as long as thick ; segments seven through 
ten each with a weak longitudinal polished ridge (Fig. 6). Front 
polished, with weak, scattered punctures, with a very strong 
median groove extending downward from the anterior ocellus ; 
eyes large and prominent, inner orbits subparallel below; head 
about 1.2 X as wide as high. Minimum width of front 1.07 X 
eye height ; ocelli large, diameter of anterior ocellus .22 X mini- 
mum width of front; postocellar line 1.15 X ocello-ocular line. 
Pronotum short, shining, weakly punctate. Mesonotum strongly 
shining, obscurely punctate. Propodeum in large part strongly 
polished, median area without ridges or other sculpturing except 
for a simple carina on the posterior third. Mesopleurum shining, 
obscurely punctate. Abdomen with scarcely any indication of a 
constriction lietween the first tAvo segments either dorsally or 
vcntrally; first tergite without an apical striate depression. 

Remarks. — • This striking specimen was taken on vegetation in 
tlie daytime, although the large ocelli suggest that the species 
may be nocturnal or crepuscular. Tlie locality was a dry wash 
in an area of desert grassland. 

SiEROLOMORPHA siMiLis new spccies 

Holotypc. — S . MARYLAND: Takoma Park, 22 Sept. 1945 
(II. & M. Townes) [HKT]. 

Description of type male. — Length 4.4 mm., length of fore 
wing 4.1 mm. Body dark brown, head and thorax almost black; 
mandibles light brown ; antennae uniformly dark brown ; coxae 
dark brown, femora medium brown, paler apically, tibiae and 
tarsi yellowish-brown ; wings lightly tinged with fuscous. First 
four antennal segments in a ratio of about 19 :10 :13 :18, segment 
three almost twice as long as thick, segments four and eleven 
about 2.1 X as long as thick ; tyloides in the form of low carinae 
on segments eight through ten, the carina on eight extending for 
much of the length of the segment, the others slightly shorter 
(Fig. 4). Front strongly polished, with small, rather evenly 
distributed punctures which are separated by 1-2 X their own 



1961 NEAKCTIC SPECIES OF SIEROLOMORPHA / 

diameters; median line of front weakly impressed. Head 1.12 X 
as wide as high, subcircular in anterior view, temples only 
moderately developed, contracted immediately behind eyes. Inner 
orbits convergent below, minimum width of front subequal to eye 
height ; ocelli of moderate size, diameter of anterior ocellus .17 X 
minimum width of front; postocellar line 1.1 X ocello-oeular 
line. Pronotum strongly shining, with a great many minute 
punctures. Mesoscutum strongly shining, obscurely punctate, 
notauli strong, nearly reaching anterior margin. Dorsal surface 
of propodeum shining, with a linear median groove. Mesopleurum 
strongly polished and nearly impunctate. First abdominal tergite 
polished, smooth, with no evidence of a transverse apical de- 
pression; second tergite with a narrow transverse basal impres- 
sion, so that there is vague evidence of a constriction between 
the first two tergites. 

Alloiype.— 2, Kearny, New Jersey, 9 Sept. 1935 (C. W. 
Funaro) [AMNH]. 

Description of allotype female. — Length 5.1 mm.; length of 
fore wing 3.6 mm. Head and thorax dark brown, abdomen 
medium brown ; mandibles and apical half of clypeus light 
yellowish-brown ; antennae dark brown except scape and pedicel 
suffused with yellowish-brown and flagellum light brown be- 
neatli ; coxae brownish but legs otherwise wholly bright yellow- 
ish-brown ; wings lightly tinged with fuscous. First four antennal 
segments in a ratio of about 23:10:11:15, segment three 1.5 X 
as long as thick, segment eleven 1.7 X as long as thick. Front 
strongly polished, punctures small and widely separated Head 
subcircular in anterior view, about as wide as high ; inner orbits 
subparallel, minimum width of front subequal to eye height. 
Ocelli small, in a broad triangle; postocellar line subequal to 
ocello-oeular line. Pronotum polished, with scattered small 
punctures. Mesonotum impunctate, notauli strong on posterior 
.8 of mesoscutum, diverging and attenuate anteriorly. Pro- 
podeum mostly smooth and shining, with a very thin median 
groove which posteriorly is paralleled by some irregular carinae. 
Mesopleurum convex, smooth and shining. Abdomen fusiform, 
depressed ; first tergite smooth and polished, with no evidence of 
a transverse apical impression; second tergite barely depressed 
basally. 

Paratypes. — CONNECTICUT: 1 9 , Bank of Conn. Kiver, 

East Hartford, 2 Sept. 1947 (H. E. Evans) [MCZ] : NEW 

YORK: 1 9 , Sea Cliff, Long Island [MCZ] ; MARYLAND: 3 

$ $ , Takoma Park, same data as type [MCZ, HKT] : WEST 



8 BREVIORA No. 140 

VIRGINIA: 1 $, Shaver's Fork, Tucker Co., Oct. 1938 (G. E. 
Wallace) [Carnegie Mus.] ; SOUTH CAROLINA: 1 $, Green- 
ville, Oct. 1952 (L. & G. ToAvnes) [HKT] ; GEORGIA: 1 $, 
Macon, 1 Dee. 1923 (T. II. Ilubbell) [USNM] ; KANSAS: 3 
$ $ , Manhattan, Sept., Nov. (D. A. Wilbur, T. F. Winburn) 
[Kansas State Univ.]. 

Variation. — The two female paratypes are very similar to the 
allotype. The Connecticut specimen has the basal two antennal 
segments bright amber-colored, contrasting strongly to the flagel- 
lum ; the Long Island specimen is without a head. The males 
show a small amount of size variation ; the smallest specimen 
(Manhattan, Kansas) has a fore w4ng measuring 3.6 mm., the 
largest (Takoma Park, Md.) has a fore wing measuring 4.4 mm. 
The Kansas specimens tend to haA^e the body (especially the 
abdomen) slightly paler in color, but otherwise little variation 
in color or body sculpture can be noted. In most specimens the 
postocellar line is subequal to the ocello-ocular line, and in one 
specimen it is somewhat shorter. 

SlEROLOMORPIIA BREVICORNIS UCW SpCCieS 

Holotype. — S , SOUTH CAROLINA: Greenville, 21 Sept. 
1952 (L. & G. Townes) [HKT]. 

Description of type male. — Length 3.4 mm., length of fore 
wing 2.7 mm. Body dark brown, head almost black ; mandibles 
light brown on apical half; antennae dark brown, very slightly 
paler beneath ; legs dark brown except front tibiae and tarsi 
light yellowish-brown ; wings hyaline. First four antennal seg- 
ments in a ratio of about 12 :6 :7 :8, segment three 1.1 X as long 
as thick, apical segment unusually short and thick, about 1.5 X 
as long as thick; tyloides present on segments nine through 
twelve, but rather small (Fig. 3). Front strongly polished, 
punctures minute, shallow; median line strongly impressed in 
front of anterior ocellus. Head very broad, 1.25 X as Avide as 
high ; front broad, its minimum width .61 X width of head, 
1.18 X height of eye ; ocelli small, diameter of anterior ocellus 
.14 X minimum Avidth of front; postocellar line very slightly 
greater than ocello-ocular line. Vertex forming a broad, even 
arc above the eye tops; temples moderately developed, in dorsal 
vicAV about two-thirds as wide as eye. Pro- and mesonota shining, 
obscurely punctate; notauli strong, complete, diverging ant rior- 
ly. Propodeum mostly smooth and polished, med'an line Aveakly 
grooved, ecarinate. Mesopleurum strongly shining, obscurely 



1961 NEARCTIC SPECIES OF SIEROLOMORPHA 9 

punctate. Venation differing from that of other species only in 
having the second recurrent and second transverse cubital veins 
very weakly indicated and the margin cell somewhat more 
rounded apicalh'. First two abdominal segments without a con- 
striction between them, first tergite smooth and polished, without 
a striate depression along its apical margin. 

SlEROLOMORPHA NIGRESCENS new SpecicS 

Holotype.— $, WASHINGTON: Olympia [USNM]. 

Description of type male. — Length 4.4 mm., length of fore 
wing 3.7 mm. Body dark brown, almost black; mandibles light 
brown ; antennae uniformly dark brown ; legs dark brown except 
front tibiae bright yellowish-brown ; wings lightly tinged with 
fuscous. First four antennal segments in a ratio of about 
20:8:11:16, segment three only 1.3 X as long as thick, segment 
four about 1.7 X as long as thick; segments seven through ten 
each with a short but rather strong longitudinal ridge (Fig. 1). 
Front strongly polished, with minute punctures which are rather 
close together below, much more widely scattered above ; median 
line of front weakly impressed. Head about 1.15 X as wide as 
high, rather thick, seen from above with the temples nearly as 
thick as the eyes, the head across the temples nearly as wide as 
across the eyes. Inner orbits subparallel below; minimum width 
of front 1.13 X height of eye ; ocelli small, diameter of anterior 
ocellus .14 X minimum width of front; postocellar line subequal 
to ocello-ocular line. Pronotum strongly shining, obscurely 
punctate. Mesoscutum polished and nearly impunctate, with 
very strong notauli which diverge slightly anteriorly and do 
not quite reach the anterior margin. Propodeum with two median 
carinae between which it is somewhat grooved, disc otherwise 
with weak and irregular sculpturing, somewhat shining. Meso- 
pleurum shining and with very small punctures. Abdomen with 
a strong constriction between the first two segments ; first tergite 
depressed along the posterior margin, second tergite along its 
anterior mars-in, both depressions with longitudinal striations. 

Allotype.— 9, V/ASHINGTON: Seattle [USNM]. 

Description of allotype female. — Length 4.5 mm., length of 
fore wing 3 mm. Head and thorax dark brown, abdomen medium 
bro"v\Ti, somewhat darker apically; mandibles and apical half of 
clypeus light brov\-n ; antennae dark brown, suffused with lighter 
brown on the sides of the basal flagellar segments ; legs dark 
brown except front tibiae bright yellowish-brown ; wings lightly 



10 BREVIORA No. 140 

tinged with fuscous. First four antennal segments in a ratio of 
about 22 :9 :9 :13, segment three 1.2 X as long as thick. Front 
strongly polished, punctures minute and widely scattered. Head 
subcircular in anterior view, very slightly wider than high ; 
inner orbits subparallel, minimum width of front 1.16 X height 
of eye. Ocelli small, in a broad triangle; postocellar line 1.1 X 
ocello-ocular line. Pro- and mesonota polished, obscurely punc- 
tate ; notauli deeply impressed, diverging only slightly anterior- 
ly, terminating well short of anterior margin of mesoseutum. 
Propodeum and mesopleurum as described for male. Abdomen 
fusiform, shining ; first tergite with a transverse apical depres- 
sion which is strongly longitudinally striate. 

Paraiypcs. — WASHINGTON: 1 $ , same data as tvpe 
[USNM] ; 1 9 , Almota, 24 June 1911 [MCZ] ; IDAHO: l' $ , 
Harvard, 24 June 1935 (J. M. Beck) [USNM] ; 2 5 <? , Moscoav, 
June, July [USNM] ; 1 S . Nezperce, 18 June 1935 (J. M. Beck) 
[USNM] : UTAH: 1 S , Uinta Co. (G. E. Wallace) [Carnegie 
Mus.] ; CALIFORNIA: 1 S , Sagehen, nr. Hobart Mills, 21 June 
1954, on Phacelia humilis (P. D. Hurd) [CIS] ; 1 S , Carnelian 
Bay, Lake Tahoe, 17 June 1958 (R. M. Bohart) [UCD] ; ^ $ $ , 
Pasadena, April 1944 (K. W. Cooper) [USNM] ; 1 <5 , Rio Linda, 
Sacramento (^o., 19 May 1958 (Light trap, Jack Fowler) [UCD] ; 
1 5, Donner Pass, f Aug. 1948 (H., M., G. & D. Townes) 
[HKT] ; 1 S , Cisco, 31 July 1948 (H., M., G. & D. Townes) 
[HKT] ; ARIZONA: 1 6 , North Rim, Grand Canyon, 29 July 
1954 (H. E. Evans) [MCZ] ; COLORADO: 1 9 , AYaldo Canyon, 
30 June 1916 (W. D. Edmonston) [USNM] ; SASKATCHEWAN: 
1 S , Saskatoon, 30 June 1950 (A. R. Brooks) [CNC] ; ALBERTA: 
1 9,30 $ S , Onefour, 3 June 1956 (0. Peck) [CNC] ; YUKON: 
1 9 , Whitehorse, 4 July 1948 (Mason and Hughes) [CNC]. 

Variation. — The four female paratypes are very similar to the 
allotype in color and in all important structural details; the 
length of the fore wing varies from 3 to 3.6 mm. The males 
exhibit much size variation, the smallest (Donner Pass, Calif.) 
having the fore wing only 2.2 mm. long, the largest (Rio Linda, 
Calif.) having the fore wing 4.5 mm. long. There is little color 
variation except that the front tibiae are dull brown in a few 
specimens, while in others the middle tibiae are light yellowish- 
brown like the front tibiae. In most specimens the postocellar 
and ocello-ocular lines are subequal. 

Remarks. — This species is very similar to canadensis and may 
represent no more than a western race of that species. However, 
there seems to be some overlap of the ranges in Arizona and in 
western Canada. 



1961 NEARCTIC SPECIES OF SIEROLOMORPHA 11 

SiEROLOMORPHA CANADENSIS (Provanclier) 

Photopsis canodensis Provancher, 1888, Add. Corr. Faune Ent. 

Canada, Hymen., p. 410 [Type: S, Ottawa, Canada (Har- 
rington) ]. 
Sierola (?) amhigua Ashmoad, 1893, Bull. U. S. Nat. Mus., 

4."): 56 [T.ype: c? , Brookings, So. Dakota (Coll. Ashmead) 

(T'SNIVI, no. 56018)]. 
MutiJla tertia Dalla Torre, 1897, Cat. Hymen., 8: 91 (new name 

for canadensis, preoccupied in Mutilla). 
Sierolomorpha anibigua Ashmead, 1903, Canad. Ent., 35: 42. 
Sicrolomorpha canadensis Krombein, 1951, U. S. Dept. Agri., 

Agri. Monogr., 2: 749. 

Remarks. — Provancher 's canadensis was transferred to this 
genus by Krombein on the advice of R. M. Schuster, who stated 
that a specimen in the University of Minnesota collection which 
had been compared with Provancher 's type was found to be the 
same as amhigua Ashmead. I have not seen Provancher 's type, 
but his description does indeed seem to fit this species rather 
well. The following statement in particular makes it clear that 
his name applies to a species (such as this one) with a rather 
marked constriction l)etween the first two abdominal tergites: 
''la suture entre les segments 1 et 2 enfoncee et crenelee." Un- 
fortunately, the abdomen of the type of amhigua is missing, but 
my interpretation of Ashmead 's species is based upon a topo- 
type which agrees closely in all details regarding the head and 
thorax, including the characteristic sculpture of the median line 
of the propodeum. It seems to me best to consider canadensis and 
amhigua synonyms, with the former having priority over the 
more widely used name amhigua. I admit the poSvsibility that I 
may be confusing more than one species here. However, speci- 
mens from the northern tier of states and from Canada show no 
undue amount of variation, and I consider it very doubtful that 
Ashmead and Provancher had different species I)efore them. 

Distrihution. — This species occurs widely over eastern North 
America, ranging from Florida, Texas, and Arizona to Alberta, 
Ontario, and Massachusetts. I have seen specimens from the 
following localities : MASSACHUSETTS: Waltham, June ; CON- 
NECTICUT: Lyme, June; NEW YORK: Oneonta. Ttluua. River- 
head, L. 1., Northwest, L. T., June-Aug. ; PENNSYLVANIA: 
Hunnnelstown. May; MARYLAND: Bowie, Takoma Park, May- 
July; VIRGINIA: Rosslyn, Creat Falls, Fredricksburg, May- 
June; NORTH CAROLINA: Highlands, Wallace, Pink Beds, 



12 



BREVIORA 



No. 140 



June-July; GEORGIA: Tifton, Valdosta, May; FLORIDA: 
Ormond, Osceola Co., Volusia Co., Brevard Co., Dec. -Jan. ; 
OHIO: Columbus, June; MICHIGAN: Ann Arbor, Erie, Liv- 
ingston Co., June-Aug. ; ILLINOIS: Champaiprn ; IOWA: Slioux 
City, Ames, June; ALBERTA: (no further data); NORTH DA- 
KOTA: Bottineau, Aug.; SOUTH DAKOTA: Brookings, June- 
July ; KANSAS: Lawrence, Manhattan, Pottawatomie Co., Geary 
Co.," May-July; LOUISIANA: Tallulah ; TEXAS: Sealy, Victoria, 
May, Nov.; NEW MEXICO: Mesilla Park, Sept.; ARIZONA: 
Springerville, July. 




r 
' — ( 



V— 



U 



z 



Figs. 1-6. Middle antennal segments of males of six species of Sierolo- 
morpha. In each ease the first segment shown (at the Ijottom) is segment 
seven. These figures are somewhat diagrammatic, as the tyloides normally 
form a somewhat twisted series, here shown as a straight series; further- 
more, the tyloides are normally somewhat obscured by setulae, here omitted. 
Fig. 1 — S. nigrescens n. sp., segments 7-10; Fig. 2 — S. canadensis (Prov.), 
segments 7-10; Fig. 3 — S. hrevicornis n. sp., segments 7-13; Fig. 4 — S. 
similis n. sp., segments 7-10; Fig. 5 — S. hicolor (?) n. sp., segments 7-10; 
Fig. 6 — S. apache n. sp., segments 7-10 and 13. 



BREVIORA 

Mmsenam of Comparative Zoology 



C'AMnRiDGi:. Mass. June 28, 1961 Number 141 

THREE NEW TOADS FROM SOUTH AMERICA: BUFO 
3IANIC0RENSIS, BUFO SPINULOSVS ALTIPERUVIANUS 

AND BUFO QUECHUA 

By Jose M. Gallardo 

Museo Argentiiio de Ciencias Naturales 
Buenos Aires 



INTRODUCTION 

In the course of a general study of the Neotropical Bufonidae, 
I received some material from the British Museum (Natural 
History) (BMNH), the American Museum of Natural History 
(AMNH), the Carnegie Museum (CM) and the Museum of 
Zoology at the University of ]\Iichigan (MZUM) which in- 
eluded the new forms described below. 

One of these is a quite different form of Bufo from the State 
of Amazonas, Brasil; another is a subspecies of Bufo spi7iulosus 
from the Department of Oruro, Bolivia, and the last a new 
species of the Bufo ockendeni group from the Department of 
Cochabamba, Bolivia. 

Bufo manicorensis sp. nov. 

Type. BMNH 1898, 3.10.1, adult male. :\Ianicore, Rio Madeira, 
State of Amazonas, Brasil. 

Descrlpiion. Head elongate and sharp. Rostrum nearly ver- 
tical. Nostrils on a prominence, elongate and oblique. Cephalic 
crests well marked, with smooth or somewhat rippled borders. 
Subnasal crests visible. Canthal crests nearly convergent. ]\Iaxil- 
lary crests somewhat expanded. Preorbital crests slightly slop- 
ing posterolaterally. Postorbital crests sloping anterolaterally, 
close to the anterior border of the tympanum. Suborbital crests 
not expanded, rather distant from the lower border of the eye. 



2 BREVIORA No. 141 

Supraorbital crests somewhat raised ; interorbital space very 
narrow and concave, with granules in tlie parietal region. Parie- 
tal crests elongate and obliciue. forming an angle with the supra- 
orbital crests. Orbitotympanic crests short but distinct. Greatest 
diameter of eye, 6 mm. \>rtical diameter of tympanum, 3 mm. 
Tymi)anum on an outward sloping plane. Paratoids subtriangu- 
lar with indistinct borders, with their long axis oblicpie medio- 
laterally, with flat dorsal granules. One subgular vocal sac. 
Dorsum with flat granules. Belly with the larger granules on 
abdominal region. Limbs with dorsal conical granules. Without 
interdigital membrane in the hand ; 1st and 2nd fingers sub- 
equal ; subarticular tubercles double on fingers 2 and 3 ; two 
carpal tubercles, the inner one smaller, elli])tic and somewhat 
salient, the outer larger and rounded. Subarticular tubercle 
double on toe 4 ; interdigital membrane in the toes to near the 
tip of the digits, but in toe 4 basal and prolonged a.s a serrated 
cutaneous fringe; two metatarsal tubercles small but elongati\ 
the inner more salient ; tarsal fringe absent. Dorsal coloration 
light brown, with some darker spots, not well marked. Belly b'ght. 

Dimensions. Head and body 50 mm. Head length 12. .5 mm. 
Head width 17 mm. Head height 5.5 mm. Interorbital space 
2.5 mm. Elbow to the third linger 21 mm. Femur 18 mm. Tibia 
16 mm. Heel to the fourth t(^e 25.5 mm. Foot 17 nun. 

Diagnostic fcafurrs. Biifo niaHicornis's dilfers from all other 
Neotropical toads ; the shape of the head and cephalic crests 
recall certain Asiatic forms. In the Neotropical area the nearest 
species is Bufo int('rni''diHS Giinther, but this is distinguished 
by the well nmrked ])arietals and preurbitals, the interorbital 
space narrower than the upper eyelid, the distinct tympanum, 
and by the paratoids wlrch are not elli])tical and are separated 
from the eye by the orbitotympanic crests. The new species is 
not as close to Bufo rolliceps Wieginann and differs from that 
species in having less prominent cephalic crests, the interorbital 
space narrower, double subarticular tubercles on the foot and 
no lateral granules in a row continuing the paratoids posteriorly. 

Material studied. BMNH 1898, 3.10.1 (1 specimen), Mani- 
eore, Rio Madeira, Brasil, B. Pift'ard. 

Bufo spixulosus alttperuvianus subsp. nov. 

Type. AMNH 14418, adult female, Challapata, Department 
of Oruro, Bolivia. 

Description. Head very short and wide. Loreal region sloping 



1961 NEW TOADS FROM SOUTH AMERICA 3 

outAvard. Rostrum vertical. No cephalic crests, excepting maxil- 
laries; cantlius rostralis thick. Interorbital space granular. Tym- 
panum sloping out. Paratoids well marked and rounded, con- 
tinued laterally by large granules (each granule with many 
horny points). Two types of dorsal granules: the larger with 
one central horny point and many others around it, the smaller 
with only one horny point. Larger granules on abdominal 
region. First finger longer than second ; subarticular tubercles 
on the fingers, double or semidivided ; palmar outer tubercle 
larger and rounded, inner smaller and elongate. Interdigital 
membrane of the foot basal but prolonged as a fringe on the 
toes; subarticular tubercles on the toes, simple or sometimes 
double ; two metatarsal tubercles, the inner more salient ; a 
thick tarsal fringe. 

Dimensions. Head and body 80 mm. Head length 17 mm. 
Head width 30 mm. Head height 11 mm. Interorbital space 6 
nun. Ujijier eyelid Avidth 6 mm. Eye 7 mm. Tympanum 3.5 
mm. Paratoid 9 mm. by 8.5 mm. Elbow to the third finger 38 
mm. Femur 36 mm. Til)ia 30 mm. Heel to the fourth toe 50 
mm. Foot 35 mm. 

Parafype. AMNH 1-1417, Choro, Bolivia, adult female 82 mm. 

Dlstrihution. The two localities of the material studied, Chal- 
lapata and Choro, are in the Department of Oruro, Bolivia ; the 
type locality is at 3700 metres altitude. 

Diagnostic features. According to Vellard (1959), there are 
six subspecies of Bufo spinulosus -. B. s. spinulosus, B. s. are- 
quipensis, B. s. limensis, B. s. trifolium, B. s. flavopictus and 
B. s. orientalis. B. s. altipernvianiis adds a seventh. 

Bufo s. altiperuvianus differs from B. s. spinulosus, the Bol- 
ivian subspecies structurally and geographically closest (De- 
partment of La Paz) in having the head shorter, not so distinct 
from the body ; the loreal region sloping more laterally ; tym- 
painim larger; paratoids larger and more rounded. Capurro 
(1950: 11) has cited B. spinulosus from Tarapaca Province, 
Chile (west of Oruro Department, on the other side of the 
Cordillera Occidental), but specimens of this provenance that 
I examined at the Chicago Natural History Museum are dif- 
ferent from the form here described. 

Remarks. I name this subspecies after the old Spanish name, 
Alto Peru, of the region from which it derives. 

Material stueVed. AIMNLI 14418 (1 specimen) Challapata, Bo- 
livia. AMNH 14417 (1 specimen) Choro, Bolivia. 



4 BREVIORA No. 141 

BUPO QUECHUA sp. IIOV. 

Type. CM 4225, adult t'eniale, liK-aohaca, 2500 m., Department 
of Cochabamba, Bolivia. 

Description. Head triaiioular, widest at the angle of the 
mouth ; loreal region sloping outward. Maxillary border marked ; 
canthus rosti-alis thick; supraorbital crest absent; parietal 
crest visible ; orbitotympanie crest thick ; one rostral-internasal 
crest more or less marked. Tympanum not visil)le. Paratoids 
approximately elliptic, dorsally smooth. Body dorsally with 
sparse large granules, but with abundant small granules. One 
lateral row of granules, continuing the paratoids, each granule 
with a large central papilla and smaller papillae around it ; 
below the row, lateral granules of the same type. Belly with 
abundant conical and simple granules. Elongate limbs, with 
conical granules dorsally. First finger longer than second ; 
fingers free, borders with small conical granules ; subarticular 
tubercles generally simple, but double on the third finger ; outer 
palmar tubercle large and rounded, inner one smaller and 
elongate. Tarsal fringe absent ; two metatarsal tubercles elon- 
gate and approximately of the same size, the outer one more 
salient ; interdigital membrane near the toe tips, but on the 
fourth only a little more than half its length and prolonged as a 
cutaneous fringe ; subarticular tubercles on toes, small and 
simple. Dorsum light l)rown with three large darker triangular 
spots not well marked ; one interocular with base to the front and 
two others on the body with the base to the rear ; a vertebral 
light line divided the last two triangles. Limbs dorsally with 
transverse wide dark bands. Belly yellowish with dark spots 
shaped very irregularly. 

Dimensions. Head and body 50 mm. Head length 12 nnn. 
Head width 17 mm. Head height 7 mm. Eye 5 mm. Upper 
eyelid width 4.5 mm. Interorbital space 5 mm. Paratoid length 
7.5 mm. Elbow to the third finger 22 mm. Femur 20 mm. 
Tibia 17 mm. Heel to the fourth toe 28 mm. Foot 20 mm. 

Parafupcs. CM 4223, 4224, 4226, lucachaca, Dept. Cocha- 
bamba, Bolivia. Head and body: 62 mm., 41.5 mm., 37 mm., 
respectively. 

Diagnostic features. Four other species related to Bufo que- 
chua have been previously described : B. ockendeni Boulenger, 
B. inca Stejneger, B. leptoscelis Boulenger and B. fissipes Bou- 
leiiqer. In the table below their diiferential characters and the 
altitude at which they are found are shown. 



1961 NEW TOADS FROM SOUTH AMERICA 









00 



« -3 



K5 5 o^-e ^^ ^«g 



Qj 



^ W <1 S HO 



2 o h 



«fc +^ 2--— - — ..._; 

Pq to -i-io^ iH-Scur' 






3 
60 be 



o :;3 



SJ. ^ ^ 





-^ 


^M 


^ 


-M 


o 


'^ 


^ 


'5 


■*! 


GQ 


^ 



- ^ *^ bD 



o 



TS 




s 




cS 






a> 


t:! 


»3 


« 


o 


'^ 


G 


> 


• fH 


> 


ft 


p 


05 









,2 


"3 


=H 


a2 




'« 


• ^ 


^ 




tc 


O 


o 




s 


^ 


cS 


r^ 


a 

03 






o 

4^ 




O 


ft 


'a3 


o 




lO 


71 


Ti 


>■ 


^ 


^ 


a 




o 


L. 


f; 








^ 


«l-l 




5- 




O 


-5 


o 


bJD 


13 



be 
o 















d 
















o 














_ 


<u 


a> 






p 




s 


CS 


G 


o 


o 


33 


C3 


c 




'S: 


rt 


-4— 


.^ 


ft 


rt 




1 


i^ 


^ 


ft 


« 


^^ 


^ 


*"* 


1 


^ 


C^ 


o 




''j 


c; 


Cw 








O 




H 


Ph 


H 


t— ( 







o 


oo 


?D 


o 


cq 




o 


m 


f~i 




t-^ 




C>] 




C^l 





^ -^ — "yn -*-)i t^ 

sa, S ^ t:: t: Q> 



a> 



;z; K <J ^ 



o 

o 

00 



ftci.^O-bC.^^ ^og oc 



M rt 



-«-3 






a 













'0 


'^ 





ft 





05 







T— i 


G 










^ 










cc 






^ X ^ CC O K a « ^ 



CO 




CQ 




a> 




r2 




^ 


, , 


3 


a> 




rt 


S 


•^ 


P 


^ 


rt 


3 


be 


-M 


be 


-^j 













l> 




< 



6 BREVIORA No. 141 

From this comparison of t-liaracters it may be deduced that 
Biifo Jissipes, described from Santo Domingo, Province of Cara- 
Laya, Peru, is the nearest relative of B. quechua. But B. fissipes 
differs in tlie absence of parietal crests and the rudimentary 
interdigital membrane. Comparing the altitudes of the five 
species, it is noticed that B. quechua is seen to live at the high- 
est altitudes; in ."ome localities, such as Yungas de Chapare, 
Department of Cochabamba, Bolivia, it coincides with B. 
odiendeni but this last lives also at lower altitudes. It is inter- 
esting to see that B. ockendeni is, in this group of five species, 
the one that lives at the lowest altitude and has the widest 
known distribution, from Central and S.E. Peru to the depart- 
ments of Cochabamba and Santa Cruz in Bolivia. The other 
three previously described species are restricted to S.E. Peru: 
B. in^a (departments of Ayacucho and Cusco), />. leptoscclis 
and B. fssipes (Department of Puno, but at different altitudes). 
B. quechua occurs in the Yungas of the Department of Cocha- 
bamba, Bolivia. 

Material studied (including comparative material). Bufo 
quechua: CM 4223-26 (4 specimens) Incachaca. Department of 
Cochabama. Bolivia, 2500 m.. J. Steinbaeh. 

USN]\I 118704 (1 specimen) Socotal, Yungas del Chapare, 

Department of Cochabamba, Bolivia, J. Steinbaeh, 11-1929. 

MZUM 89414 (1 specimen) Yungas del Chapare, Department 

of Cochabamba, Bolivia. 

MZUM 76075 (1 specimen) Yungas de Cochabamba, 2200 

m., Department of Cochabaml^a, Bolivia. 

MZUM 68163 (3 specimens) Yungas de Cochabamba, 2200 

m.. Department of Cochabamba, Bolivia. 

MZUM 68166 (1 specimen) Cochabamba Valley, 2600 m.. De- 
partment of Cochabaml)a, Bolivia. 

Bufo fissipes. AMNH 6105 (1 specimen) Juliaca, Peril, H. II. 
Keays. Chicago Natural History Museum 64879, 64850 (2 speci- 
mens) Puno, Peru. 

Bufo iuea. USNM 107648 (1 specimen) 1 mile above San 
Miijnel, Avacueho, Peru, 0. F. Cook. V-27-1915. 

MCZ 4758 (1 specimen) Idma. I^ubamba Valley, 6000 ft.. 

Peril, E. Heller, X-1915. 

B}(fo ockeudeni. ]\ICZ 15425 (1 specimen) Chaijuimayo, Peru. 
Chicago Natural History IMuseum 3581-82 (2 specimens) Chaqui- 
mayo, S.E. Peru, H. C. Watkins. 



1961 NEV." TOADS FROM SOUTH AMERICA 7 

AxMXll (mi-lT (16 specimens) Juliaca, Peru, H. H, Keays. 

MZUM 68153 (1 specimen) Yungas del Chapare, Department 

of Cochabamba, Bolivia. 

.MZTM 68152 (2 specimens) Tarata. 1900 m.. liolivia. 

C^l 3806b, 4515 (2 specimens) Cerru Husanu, west of Santa 

Cruz, 1400 m., J. Steinbaeh. 



Acknowledgments 

1 must thank the Consejo Nacional de Investigaciones Cien- 
tificas y Tecnicas de la Kepul)lica Argentina for the fellowship 
given me for investigations on Neotropical amphibians ; I am 
grateful also to Dr. A. is. Komer, Director of the Museum of 
Comparative Zoologj' at Harvard I'niversity and to Dr. E. E. 
Williams. Curator of Reptiles and Amphibians of this Museum, 
for facilities aiforded me during 1959-60 ; to Dr. A, G. C. Grandi- 
son, Dr. D. Cochran, Mr. C. :^.I. Bogert, Dr. R. Inger, Mr. N. 
Richmond and C. Walker for sending me material from their 
respective museums; and to Dr. \\ Vanzolini for the photo- 
graphs illustrating this ]iaper. 



Bibliography 

Barbour, T. and G. K. Noble 

1920. Amphibians and reptiles from southern Peru, collected by 
the Peruvian Expedition of 191-1-1915 under the auspices of 
Yale University and National Geographic Society. Proe. U. 8. 
Nat. Mus., 58: 609-620. 

BOULENGEB, G. A. 

1902. Descriptions of new liatraeliians and reptiles from the Andes 
of Peru and Bolivia. Ann. Mag. Nat. Hist., (7; 10: 39-4-402. 

1903. Descriptions of new batrachians in the British Museum. Ann. 
Mag. Nat. Hist., (7; 12: 552-557. 

1912. Descriptions of new l)atrachians from the Andes of South 
America, preserved in the British Museum. Ann. Mag. Nat. 
Hist., (8) 10: 185-191. 
Captirro, L. F. 

1950. Batracios de Tarapacii. Invest. Zool. Chilenas, 1(1): 9-12. 
GUNTHER, A. 

1858. Catalogue of the Batrachia Salientia in the collection of the 
British Museum, London: i-xvi -f 1-160, pis. I-XII. 



8 BREVIOBA No. 141 

Mertens, E. 

1952. Amphibieii und Eeptilien in Titschack, Beitr. zur Fauna Perus, 
3: 257-266. 
Stejisteger, L. 

1913. Results of the Yale Peruvian Expedition of 1911. Batrachians 
and reptiles. Proc. U. S. Nat. Mus., 45: 541-.547. 
Vellakd, J. 

1959. Estudios sobre batracios andinos. V. El genero Bufo. Mem. 
Museo Hist. Nat. Javier Prado, 8: 3-48, pis. I-XIV. 




Plate 1. Bufo manicorensis, new species. Type: BMNH 1898.3.10.1. 




o 



p^ 




ia§;%^^^0 ill 




m- 



tf^m^ "^mm^ 



H. S^fiSistt 









1i 






^, 



Xi»:; 




Plate 3. Bufo quechua, new species. Type: CM 4225. 



BREVIORA 



Musemnii of Comparative Zoology 



Cambridge, Mass. Jl'xe 30, 1961 Number 142 

AUSTRALIAN CARABID BEETLES VL THE TROPICAL 

AND SOME SUBTROPICAL SPECIES OF PAMB0RU8, 

MYSTROPOMVS, AND NURU8 

By p. J. Darlington, Jr. 

This is the first of four papers describing new flightless 
tropical (and a few related subtropical) Australian rain forest 
Carabidae of zoogeographic importance. However, these special 
papers will be treated as parts VI to IX of my general series 
on Australian carabid beetles. Some of the species now de- 
scribed have been referred to (but not by name) in the preced- 
ing paper of the series, on transition of wet forest carabid 
faunas from New Guinea to Tasmania (1961b). My localities 
are listed, mapped, and briefly described in No. IV of this series 
(1961a). The holotypes of new species described from my ma- 
terial are placed, at least temporarily, in the Museum of Com- 
parative Zoology. Paratypes will be deposited with the Com- 
monwealth Scientific and Industrial Research Organization at 
Canberra (where they can be compared with the Sloane Collec- 
tion) and in most cases in the Queensland Museum too. In the 
descriptions, proportions are usually given as simple fractions 
(%, %, etc.) but are based on actual measurements made under 
the microscope. 

Pamborus 

Banninger (1940) has correctly distinguished the real species 
of Pamhorus known to him, after examining some of the older 
types that Sloane (1904) was unable to see. What I have to 
say now is mostly concerned with tropical forms unknown to 
Banninger. 

Three species-groups of Pamhorus occur in tropical North 
Queensland. Two of them consist of single, very distinct species, 
elegans SI. and punctntus n. sp., respectively. The third, which 



2 BREVIORA No. 142 

I call the tropicus group, consists of a series of slightly dif- 
ferentiated, allopatric forms extending from South Queensland 
to the base of the Cape York peninsula and probably derived 
from a common ancestor that dispersed rather recently. Charac- 
ters given in the following key to distinguish the four species 
( ? or subspecies ) of the tropicus group should be supplemented 
by comparison of descriptions and of specimens if possible. 

Key to tropical species of Pamborus (with some subtropical 

species in parentheses) 

1. Each elytron with 6 to 8 costae (which may be raised or nearly flat) 
separated by narrower crenulate intervals 2 

— Elytron with 15 or more nearly equal costae 7 

2. Tip of aedeagiis dentate near apex; neck constriction deep; prothorax 
subcordate (wet forests of New South Wales and South Queensland) 
(alternans) 

— Tip of aedeagus not dentate; neck constriction shallow; prothorax 
variable in shape 3 

3. Form more convex; sides of ijrothorax not or slightly sinuate; 7th 
and 8th elytral costae usually strongly developed and not much in- 
terrupted (drier woodlands of New South Wales and South Queens- 
land) (viridis) 

 — Lfess convex; sides of prothorax often more sinuate (tropicus group) 
4 

4. Elytra with costae relatively wide and weakly convex, 5th and 6th 
nearly as wide as 4tli on disc, 7th and 8th usually distinct, but variable 

5 

—  Elytra with costae usually narrower and more convex, 7th and 8th 
more often interrupted or disintegrated 6 

5. Intervals between costae slightly vdder and more strongly crenulate; 
larger (31-36 mm.) (South Queensland) {suhtro-picus) 

— • Intervals between costae narrower (1st reduced to a fine irregular 
impressed line) and less strongly crenulate; smaller (26-30 mm.) 
(Eungella Range) transitus 

6. Shining (Mt. Spec to Atherton Tableland, etc.) tropicus 

— Duller (northern Atherton Tableland to Mossman-Daintree area) 
opacus 

7. Pronotum with basal impressions, not punctate; elytral costae not 
much interrupted; elytral margins green (Herberton — ? to v. Cook- 
town) elegans 

— Pronotum without basal impressions, entire surface densely coarsely 
punctate; all elytral costae much interrupted; bluish black (Ather- 
ton Tableland, etc.) punctatus 



1961 AUSTRALIAN CARABID BEETLES 3 

Pa.MBORUS SUBTROPICUS 11. sp. 

Form of alternans but often broader, somewhat variable, 
sliglitly depressed; rather shining black, pronotum with mar- 
ginal channels (narrowly) and posterior impressions bluish or 
greenish, elytra with crenulate intervals and margins green. 
Read: neck constriction weak. P yoihorax obovii y^ (Mt. Jacob) 
or 1/^ (Kenilworth) wider than long at middle; base slightly 
wider than apex ; sides broadly rounded anteriorly, broadly but 
not strongly sinuate posteriorly ; posterior angle moderately 
produced backward; linear basal impressions uniting with mar- 
ginal channels in moderate impressions, with convex areas not 
or vaguely reaching posterior margin; each lateral margin with 
1, 2, or 3 (number variable, sometimes asymmetrical) seta-bear- 
ing punctures near and before middle. Elytra with margins 
not serrate; each with 8 slightly elevated costae wide on disc, 
narrower externally and apically, separated by strongly crenu- 
late intervals of which the 1st is narrowest but plainly crenu- 
late; 7th and 8th costae usually distinct at least to near middle 
of length but not strongly raised, slightly interrupted, 8th some- 
times disintegrated. Aedeagus not dentate. Length 31-33 (36) ; 
width 12 (13) mm. (^ figures in parentheses show probable size 
of an individual of which I have only elytra). 

Holotype i (M. C. Z. Type No. 30,346) and 1 9 paratype 
from Mt. Jacob, c. 45 miles south of Gladstone, South Queens- 
land, March 1958; and 1 6 paratype from Kenilwortli. west of 
Blackall Range. South Queensland, May 1958 ; all taken by 
nivself in or on the borders of rain forest. 1 have also elvtra of 
this species from Mapleton, on the north end of the Blackall 
Range. 

See key for distinguishing characters of this species. 

PaMBORUS TRAXSITUS 11. sp. 

Form almost of rather broad alternans, slightly depressed; 
moderately shining black, marginal channels and basal impres- 
sions of pronotum without or with only slight metallic color, 
margins and crenulate intervals of elytra green or greenish. 
Head: neck constriction weak. Prothorax between % and % 
wider than long at middle ; Ijase slightly wider than apex ; sides 
broadly rounded anteriorly, sometimes vaguely angulate at 
middle, broadly but weakly sinuate posteriorly; posterior angles 



4 BREVIORA No. 142 

moderately produced backward; linear basal impressions unit- 
ing with marginal channels in moderate impressions, with con- 
vex areas sometimes reaching or nearly reaching posterior mar- 
gins; each lateral margin with 1 to 4 (number variable, often 
asymmetrical) seta-bearing punctures near and before middle. 
Elytra with margins not serrate ; each eh^tron with 8 slightly 
elevated costae very wide on disc, narrower laterally and apic- 
ally, 5th and 6th not or not much narrower than 4th on disc, 
7th and 8th narrower, usually distinct but not strong, often 
somewhat interrupted but rarely disintegrated ; crenulate in- 
tervals very narrow on disc, less strongly crenulate than in 
related forms, interval between 1st and 2nd costae reduced to 
a fine line almost without crenulations anteriorly. Aedeagus not 
dentate. Length 26-30; width c. 10.5-11.5 mm. 

Holotype S (M. C. Z. Type No. 30,347) and 67 paratypes all 
from the Eungella Range, c. 40 miles west of Mackay, Queens- 
land, c. 2000-3000 ft. altitude, Nov. 1957, taken l)y my wife, 
my son, and myself, in rain forest. 

For characters and relationships of this geographically iso- 
lated form, see preceding discussion and key. 

Pambortts tropicus n. sp. 

Form almost of alternans, slightly depressed; rather shining 
black, marginal channels and posterior impressions of pronotum 
and margins and crenulate intervals of elytra green. Head : 
neck constriction weak. Prothorax appearing as long as wide 
but by measurement nearly % wader than long at middle ; base 
not or slightly wider than apex: sides broadly rounded an- 
teriorly, sometimes vaguely angulate at middle, broadly sinuate 
posteriorly; posterior angles projecting backward as usual in 
group ; basal impressions uniting with marginal channels in 
moderate impressions, sometimes with vague convex areas reach- 
ing or nearly reaching base ; each lateral margin with 1 to 4 
seta-bearing punctures near and before middle (number vari- 
able, often asymmetrical). Elytra with margins not serrate; 
each elytron wnth 8 costae usually slightly narrower and more 
convex than in preceding forms especially externally, but 7th 
and 8th costae variable, sometimes distinct (but not strong), 
sometimes much interrupted or disintegrated; crenulate inter- 
vals narrow on disc, broader externally, strongly crenulate. 
Aedeagus not dentate. Length (types) 28-31 ; width 10.5-11.5 
mm. 



1961 AUSTRALIAN CARABID BEETLES 5 

Holotype 6 (M. C. Z. Type No. 30,348) and 30 paratypes all 
from Mt. Spec plateau (Paluma Kange), about 40 miles north of 
Townsville, 2000-3000 ft. altitude, Nov. -Dec. 1957, taken by the 
Darlingtons, in or on the edges of rain forest. Additional speci- 
mens, not types: 5, Kirrama Range, 2000-3000 ft., Dec. 1957; 3, 
Millaa Milfaa, April 1932; 2, Longlands Gap, Sept. 1952 (J. G. 
Brooks) ; 6, mountains above (SW of) Atherton, Dec. 1957 and 
Feb. 1958 (this and the 2 preceding localities are on the south- 
central Atherton Tableland) ; 6, Mt. Bartle Frere, west slope, 
2000-3500 and 3000-5000 ft., Dec. 1957; and 1, Davies Creek 
Road, northern Atherton Tableland, May 1958; all specimens 
except Brooks' collected by the Darlingtons, in rain forest. 

This species (or subspecies) is more like suhtropicus of South 
Queensland than like transitus of the Eungella Range. It dif- 
fers from suhtropicus in having elytral costae usually slightly 
narrower and more convex especially externally, with 7th and 
8th costae more often interrupted or disintegrated. 

Pamborus opacus Gehin 

Sloane (1904, p. 702) and Biinninger (1940) have applied 
the name opacus to this species. It differs from tropicus in being 
obviously duller, and it is also slightly more slender, with 
slightly narrower and more convex elytral costae, and on the 
average it has more marginal pronotal punctures, although 
these vary in number and position. 

Although this species and tropicus are not very different 
structurally, I think they probably are real species, for they 
overlap geographically. Of opacus, I have 6 specimens from 
mountains north of Kairi, on the Atherton Tableland between 
the Atherton area and Davies Creek, taken in rain forest at close 
to 4000 ft. altitude, and 18 specimens from Mt. Lewis, near 
Mossman, taken at about 3000 ft. altitude in rain forest. My 
northernmost locality for tropicus is between these two places 
but at a somewhat lower altitude, near the southern end of 
the Davies Creek forestry road. 

Pamborus elegans Sloane 

Sloane (1915) described this species from "scrub" (rain 
forest) east of Herberton on the Atherton Tableland. It should 
occur in the rain forests on the mountains between Atherton 
and Herberton, but I did not find it. Banninger (1940) records 



6 



BREVIOEA 



No. 142 



it from ''Mac Ivor Kiver, " which may be the Melvor Kiver 
north of Cooktown. 

Pamborus punctatus n. sp. 

Form as figured (Fig. 1) ; entirely dull bluish or purplish 
black. Head quadrate ; eyes small ; antennae short ; neck con- 
striction very deep. Proihorax % or more wider than long 
at middle, widest behind middle, narrowed in front and behind 





Fig. 1 . 
Pambokus punctatus n. sp. 



Fig. 2. 
NURUS REX n. sp. 



1961 AUSTRALIAN CARABID BEETLES 7 

but base about % wider than apex ; sides weakly rounded except 
strongly rounded or vaguely angulate behind middle, slightly 
sinuate near base; basal angles only slightly produced back- 
ward; side margins thickened but marginal channels and basal 
impressions obsolete ; dorsal surface almost evenly but not 
strongly convex, with middle line almost obsolete except an- 
teriorly, and transverse impressions obsolete ; whole surface 
coarsely, irregularly punctate with longitudinal, deep punctures. 
Elytra oval; humeri not serrate; each elytron with 15 distinct 
and 2 additional partial costae, all much interrupted, the outer- 
most reduced to tubercles. Abdomen extensively but not uni- 
formly punctate. Tip of last ventral segment subtruncate in 
male, broadly rounded in female. Aedeagus not dentate. Length 
17-19.5; width 7.3-8.0 mm. 

Holotype ? (M. C. Z. Type No. 30,349) and 2 paratyi^es 
from mountains above (8W of) Atherton. 3000-4000 ft. alti- 
tude, Dec. 1957 and Feb. 1958; and additional paratypes as 
follows: 1, south of Ravenshoe, c. 3000 ft., Feb. 1958; 2, east 
side Mt. Bellenden Ker, 3000-4500 ft., Dec. 1957. All speci- 
mens taken by myself in rain forest. 

This species differs from all previously known Famborus 
in obliteration of all pronotal impressions and in heavy punc- 
tation of pronotum. The small size and numerous elytral 
costae suggest a distant relationship with P. guerini Gory of 
South Queensland etc., but (in addition to the other differ- 
ences) guerini has serrate humeri and pimctatus has not. 

In life this small Pamhorus strikingly resembles the heavily 
catenulate species of Notonomus (especially masculinus Darl. 
1953) which occur in the same rain forest areas. I suspect this 
is a case of mimetic convergence. 

Mystropomus 

Two species of this genus occur in tropical eastern Austra 
lia. One, with alternate elytral intervals raised, occurs in rain 
forest on the Eungella Range. It seems to be at most a poorly 
defined subspecies of subcosfaiiis Chd. of South Queensland 
and New South Wales. I do not think it is worth naming. The 
other tropical species, with elytral intervals equal, is N. regu- 
laris Banninger (1940), which occurs probably throughout the 
main (base-of -peninsular) rain forest system of North Queens- 
land. I now have 113 specimens of it from localities north to 



8 BREVIORA No. 142 

Thonitou Peak aiiil south to the ]\It. Spec plateau. There is 
some geographical variation, but specimens from most localities 
can be referred to the typical subspecies. However, the form 
on the Mt. Spec plateau at the southern extreme of the species' 
range seoms worth distinguishing as follows. 

^MysTKOPO-^IU? REOrLAUIS LAEVIS U. Subsp. 

Similar to large specimens of typical Mystropomus regularis 
Bann. but almost lacking the weak granular elytral costae of 
typical regularis. In the latter each elytron has 6 or 7 dis- 
tinguishable (though scarcely elevated) costae or stripes that 
are more shining than the intervening spaces. In the new sub- 
species only about the 4 inner stripes are indicated at all, and 
they are much less distinct than in the typical regidaris. The 
rest of the eh'tral surface, including the lateral and anterior 
declivities, is virtually undifferentiated, dull, and finely granu- 
lar. As compared with typical regularis, laevis also has slightly 
Avider and more reflexed prothoracic margins and a somewhat 
duller pronotum. Length c. 17; width c. 6.5 mm. 

Holotype S (M. C. Z. Type No. 30,350) and 1 i paratype 
both from ^It. Spec plateau (Paluma Range), c. 40 miles north 
of Townsville, North Queensland, 2000-3000 ft. altitude, Feb. 
1958, taken by myself in or on the edge of rain forest. 

NURUS 

This is a group of very large, stout Pterostichini that I can- 
not separate from Trichosternus by any single constant char- 
acter except the relatively heavy build. Nurus atlas Cast, has 
each $ front tarsus slightly dilated, with only 2 segments 
squamulose, and most other Nurus have 6 tarsi narrow and 
without squamae, but both these conditions exist in certain 
Trichosternus. Tschitscherine (1902) was therefore wrong in 
using form of $ tarsi as a primary generic character and 
Sloane (1894) was riglit in not using it. The known species of 
Nurus form several groups that have been called subgenera and 
that are named in the following key, although 1 am doubtful of 
the value of these subgenera. 

Nurus sensu stricto includes about 4 species in northern New 
South Wales, south at least to near Ebor, and an additional 



1961 AUSTRALIAN CARABID BEETLES 9 

species on Mt. Tamborine in southeastern Queensland.' The 
majority of the species inhabit rain forest, but at least one ex- 
tends into savannah woodland. The two species of Pachymelas 
apparently inhabit savannah woodland and perhaps coastal 
habitats in tropical Queensland; they do not seem to enter rain 
forest. The two previously described species of Nuridius are 
localized in South Queensland, probably in rain forest, al- 
though their habitats are not specified. The three new species 
described below, one tentatively associated with Nuridnis and 
the others not assigned to subgenera, are all rain forest species, 
but they are not directly related among themselves and prob- 
ably represent three separate invasions of isolated rain forests 
by different stocks of Nurus. The genus is apparently absent 
in the main (base-of -peninsular) rain forests of North Queens- 
land. 

Key to subgenera and some species of Xurus 

1. Mesosternum not setose {Nurus sensu stricto) c. 5 species 

— Mesosternum setose anteriorly 2 

2. Humeri without teeth {Pachymelas) S species 

— Humeri with margin toothed or thickened (but sometimes only slightly 
so) 3 

3. Elytra without basal margin {Nuridius) 4 

— Elytra with basal margin 5 

4. Margins of pronotum less strongly reflexed; size smaller (31 & 39 mm.) 
fortis, grandis 

— Margins of pronotum strongly reflexed, size larger (41-45 mm.) . . .rex 

5. Wholly black; stout, prothorax transverse, not much narrowed behind 
nox 

— Greenish black ; more slender, prothorax more narrowed behind .... 
mediiis 

Nurus rex n. sp. 

Form as figured (Fig. 2) ; large; black, head and pronotum 
shining, elytra (except marginal intervals) dull, lower surface 
moderately shining. Head rather small (in genus), c. 2/3 
width prothorax; mandibles long, straight basally, strongly 

1 The type locality of N. imperialis (Slonno 18!>4) is given as "North 
Queensland," but the species really Inhabits South Queensland. I have Ke<>n it 
unlv from Mt. Tamborine south of Brisbane. The type locality of \. crami- 
formin (Sloane 1S99, p. .570) is piven as "Cairns." but the specimen was receive<l 
from French, and I know from other evidence that many of French's specimen.'; 
have wronp localities. If c-raxsif(/rinis is a synonym of atlax, as supposed, the 
type presumably really came from northern New Smith Wales. 



10 BREVIORA No. 142 



to 



curved before apex, with irregular row of small seta-bearing 
punctures near lower edge of outer face near base; antennae 
short, extending only slightly beyond basal angles of prothorax ; 
eyes small, genae long, wider than eyes, slightly, sinuously nar- 
rowed to neck ; 2 supra-ocular setae each side ; frontal impres- 
sions subparallel, irregular ; mentum with deeply emarginate 
tooth. Prothorax cordate, 3/5 or slightly less wider than long 
at middle, strongly narrowed behind, less so in front; base 
slightly narrower than apex; latter broadly emarginate, with 
anterior angles abruptly advanced; base emarginate, posterior 
angles produced backward in arc of emargiuation ; sides 
rounded anteriorly broadly but usually not strongly sinuate well 
before base ; basal angles would be right or slightly acute ex- 
cept narrowly rounded; lateral margins wide, very wide pos- 
teriorly, strongly reflexed, each with 1 or 2 seta-bearing 
punctures before middle and 1 or 2 also near base; base mar- 
gined, apex not; disc with well impressed median line and 
transverse impressions; posterior impression and marginal 
channels joining in obliquely flattened baso-lateral areas; sur- 
face of disc slightly transversely wrinkled but not punctate, 
except extreme base at middle rugose-punctate with elongate 
punctures. Elytra broad, widest behind middle ; humeri ob- 
tuse, with margins forming teeth which are strongly raised but 
hardly prominent laterally; base not margined (margins ending 
inwardly about opposite ends 5th striae); disc very convex; 
each elytron with 7 finely punctate striae lying in broad 
depressions between slightly elevated intervals; 9th interval 
with row of minute foveae (the 8th stria) along inner edge; no 
10th interval ; 7th interval scarcely raised at base ; each 3rd 
interval 2-punctate posteriorly, usually near top of and well 
down on declivity, but position of punctures variable. Lower 
surface : prosternal process and anterior face of mesosternum 
setose ; ventral segments 2 to 5 with some irregular setigerous 
punctation. Male tarsi not dilated, without squamae ; o 
usually with 1, 9 2 principal setae each side last ventral seg- 
ment, but additional, usually smaller setae sometimes present. 
Length 41-45, width 15-17 mm. 

Holotype S (M. C. Z. Type No. 30,351) and 12 paratypes 
all from the Elliot Kange, south of Townsville, Queensland, 
taken at about 3000 ft. altitude, March 2, 1958, by my son and 
myself, under logs in mountain rain forest. 

See preceding discussion and key for place of this species 
among other Nurus. 



1961 AUSTRALIAN CARABID BEETLES 11 

NuRUS Nox n. sp. 

Large ; black, moderately shining- above and below except 
elytra duller with narrow bands along 8tli striae (sometimes 
including margins) more shining. Head of moderate size (in 
genus), % or slightly less width prothorax; mandibles rather 
long, strongly curved before apex, without setigerous punctures 
on lower outer face near base ; antennae extending beyond base 
of prothorax by two or more segments; eyes small; genae sub- 
parallel, slightly sinuate but not much narrowed to neck; 2 
supra-ocular setae each side ; frontal impressions weak, sub- 
parallel, irregular; mentum with emarginate tooth. Prothorax 
transverse-subquadrate, slightly more than i/^ wider than long 
at middle, weakly narrowed behind; base about 1/10 wider 
than apex ; latter slightly emarginate, with anterior angles only 
slightly advanced; base broadly emarginate at middle, sub- 
truncate at sides ; base and apex unmargined ; sides arcuate 
anteriorly, then nearly straight and slightly converging pos- 
teriorly for much of length, then broadly and usually rather 
slightly sinuate before base ; basal angles c. right ])ut narrowly 
rounded ; lateral margin moderate, slightly wider posteriorly, 
moderately reflexed, each with seta-bearing puncture near or 
a little before base but without anterior-marginal setae ; disc 
with usual middle line and deeper transverse impressions; 
basal impression and marginal channels joining in what would 
be large baso-lateral impressions except each impression occu- 
pied by a large convex space ; surface of disc slightly trans- 
versely wrinkled but not punctate, wrinkling mueli closer at 
sides and base, radiating from a central point near base. Elytra. 
wide, not much narrowed anteriorly; humeri rounded, minutely 
obtusely toothed or at least slightly thickened at humeri (teeth 
or thickenings easily overlooked in dirty specimens) ; base 
margined almost to scutellum ; disc very convex ; each elytron 
with 7 vaguely indicated, sometimes finely punctate striae in 
depressions between slightly convex intervals; 9th interval 
(marginal channel) with an irregular series of small ocellate 
foveae on inner edge (= 8th stria) ; no 10th interval; 7th in- 
terval only slightly raised at base ; each 3rd interval usually 
2-punctate with punctures near top of and well down on 
declivity, but punctures variable in position and sometimes 
missing. Lower surface : prosternal process and anterior face 
of mesosternum setose; ventral segments 2 to 5 with setigerous 
punctures tending to form transverse rows, ^lale with front 



12 BREVIORA No. 142 

tarsi not dilated, without squamae : male with 1 ur 2, female 
with 2 or more principal setae each side last ventral segment, 
but setae variable, not always distinguishing sexes. Length 
30-37; width 11.5-15 mm. 

Holotype $ (M. C. Z. Type No. 30,352) and 7 paratypes 
(5 whole specimens and 2 represented by shells of prothorax 
and elytra) all from Mount Jacob, about 45 miles south of 
Gladstone, South Queensland, c. 2000 ft. altitude, March 1958, 
taken by the Darlingtons in and on the edges of rain forest. 

This new species approaches Pachymelas, for the humeral 
"teeth" are small, obtuse, and inconspicuous, sometimes only 
slight thickenings of the margin, but the present species differs 
from the two known Pachymelas (both of which are before me) 
in a number of specific characters, including form of prothorax 
and presence of shining sM&marginal stripes on elytra. 

NUBUS MEDIUS n. sp. 

Rather slender (in genus) ; greenish or bluish black, shining 
above except elytra duller with more or less shining sutural 
intervals and marginal channels, rather dull greenish below. 
Head large, 4/5 or slightly less wadth prothorax, mandibles 
rather long, curved before apex, without setigerous punctures 
on lower edge of outer face ; antennae relatively long, passing 
basal angles of prothorax by about 3 segments; eyes small, 
genae wider than eyes, broadly rounded, slightly narrowed to 
neck ; frontal impressions w' eak, subparallel ; mentum with 
emarginate tooth. Prothorax subcordate, between 1/3 and 1/2 
wider than long at middle ; sides weakly rounded anteriorly, 
straight and converging posteriorly behind middle, then broadly 
sinuate before right or slightly acute, scarcely blunted posterior 
angles ; base about equal to or slightly narrower than apex ; 
latter broadly emarginate at middle, slightly rounded at sides, 
with anterior angles (marginal channels) slightly advanced; 
base broadly emarginate, subtruncate at sides ; base and apex 
unmargined; lateral margins rather narrow (in genus) and 
not strongly reflexed, sometimes slightly scalloped, each with 
one seta before middle and another almost on posterior angle ; 
disc w4th usual middle line, moderate anterior transverse im- 
pression, and deeper posterior transverse impression ; latter 
ending in rather vague, irregular baso-lateral depressions which 
include somewhat transverse convexities near base; surface of 
disc slightly wrinkled, more wrinkled at base, the basal wrinkles 



1961 AUSTRALIAN CARABID BEETLES 13 

radiating' irregularly from central point. Elytra rather strongly 
narrowed anteriorly, with obtuse humeri; latter with small but 
distinct teeth ; basal margins entire ; each elytron with 7 fine, 
faintly punctate striae in depressions between low, rounded in- 
tervals ; 7th interval elevated at base but not acute ; 3rd inter- 
val 2- or 3-punctate posteriorly, above and on declivity. Lower 
surface: prosternal process setose; mesosternum rather sparsely 
setose in fresh specimens, but seta sometimes hard to detect in 
old ones; ventral segments 2 to 5 Avith a few (variable) seti- 
gerous punctures tending to form transverse rows near middle 
of segments posteriorly. Male with front tarsi not dilated and 
without squamae; $ with 1. 2 2 setae each side last ventral 
segment. Length 30-37 ; width c. 11.5-14 mm. 

Holotype S (M. C. Z. Type No. 30,353) and 26 paratypes 
all from the Eungella Kange, west of Mackay, Queensland, 
2000-3000 ft. altitude, Nov. 1957, taken by the Darlingtons in 
rain forest. 

This new species resmbles Nurus sensii. stricto in form and 
metallic coloration but differs in presence of setae on the meso- 
sternum. 



EEFEEENCES 

Banninger, M. 

1940. [Pamljoru.s.] Xovitates Zoologieae, 42: 203-205. 
Darlington, P. J., Jr. 

1953. Australian carabid beetles II. Some new Pterostichini. Psyche, 

60: 90-101. 
1961a. Australian carabid beetles IV. List of localities, 1956-1958. 

Psyche (In press.) 
1961b. Australian carabid beetles V. Transition of wet forest faunas 

from New Guinea to Tasmania. Psyche (In press.) 
Sloane, T. G. 

1894. [Homalosoma, inel. Nurus.] Proe. Linn. Soc. New South Wales, 

(2) 9: 417ff. 
1899. [Eomalosovia, incl. Nurus.] Proe. Linn. Soe. New South Wales, 

24: 567ff. 
1904. [Pamhorus.] Proe. Linn. Soc. New South Wales, 29: 701-703. 
1915. [Pamborus elegans.] Proe. Linn. Soc. New South Wales, 40: 

438-439. 

TSCHITSCHERINE, T. 

1902. [Nunis etc.] Horae Soc. Ent. Eossicae, 35: 515-519. 



BREVIORA 



Muasemiini of Comparative Zoology 



Cambridge, Mass. August 20, 19G1 Number 143 



MIOCENE LIZARDS FROM COLOMBIA, 
SOUTH AMERICA 



By Richard Estes'^ 

Numerous fossil vertebrates were collected by the 1949 Uni- 
versity of California field expedition to the upper Magdalena 
Valley, Huila, Colombia, South America. Among- these, the 
lower vertebrates include bony fishes, lepidosirenid lungfish, a 
bufouid frog (identified as ?Leptodactylidae by Savage, 1951), 
turtles, crocodiles, snakes, and lizards. The lizards are described 
in the present paper. 

I am indebted to Dr. AVann Langston for bringing these 
specimens to my attention, and to Mr. C. M. Bogert, American 
Museum of Natural History, Dr. Ernest E. Williams. Museum 
of Comparative Zoology, Harvard University, and to Dr. Alan 
Leviton, California Academy of Sciences, for the use of com- 
parative material in their collections. 

The manuscript has been read by Drs. J. T. Gregory and 
D. E. Savage, University of California. Museum of Paleontol- 
ogy, and Dr. Ernest E. AYilliams, and I wish to thank them 
for numerous helpful suggestions. 

The sequence and distribution of the fossil faunas of the 
Magdalena Valley has been discussed by Stirton (1953, see 
especially figures 1. 2). Fossil lizards have been found only 
in the I'pper Oligocene Coyaima fauna, and the Upper Miocene 
La Venta fauna. 

Ahhreviotions. A.M.. Am.erican Museum of Natural History, 
Dei)artraent of Vertebrate Paleontology. V.C.. University of 
California, ]\Iuseum of Paleontology. 



1 nonia-tiiu'iit (if Bioloirv. Boston Univcrsitv. ami Research Assopiate. Depart- 
ment of Vertebrate Paleontology. :Mnsenni of Comparative Zoology. Ilarvanl 
University. 



2 BREVIORA No. 143 

COYAIMA FAUNA 
Family TEIIDAE 

Cf. TUPINAMBIS Sp. 

Referred specimen. U. C. no. 56303, maxillary fragment. 

Locality. U. C. loc. \^-4411, Coyaima, Colombia, South Amer- 
ica. 

Age. Late Oligoeene. 

Descriptio7i. The fragment is the posterior end of a right 
maxilla, which bears one complete tooth and the bases of three 
others. The complete tooth has a strongly wrinkled crown 
surface, and in occlusal view is circular. No anteroposterior 
crests are present. 

Discussio7i. The tooth closely resembles posterior maxillary 
teeth of Recent Tupinamhis teguixin. It differs from compar- 
able teeth of Dracaena in not being dorsoventrally flattened. 

LA VENTA FAUNA 
Family TEIIDAE 

TUPINAMBIS cf. T. TEGUIXIN 

Referred specimen. U. C. no. 38856, left dentary and splenial, 
with parts of the coronoid, angular and surangular; the sym- 
physis region missing. Collected by Dr. Jose Royo y Gomez. 

Locality. U. C. loc. V-4526, Lower Red Beds, Honda group, 
Huila, Colombia, South America. 

Age. Late Miocene. 

Description. The maximum length of the specimen is forty- 
three millimeters. The dentary is robust, and has a heavy shelf 
external to the tooth row. Three inferior alveolar foramina are 
preserved. Posteriorly a large re-entrant notch contains a 
fragment of surangular. The latter, and part of the dentary 
anterior to it, is prominently depressed for muscle insertion. The 
splenial is large, and deep anteriorly as in most teiids. There 
are two foramina in the splenial, the posterior one smaller. 
The anterior process of the angular inserts between the splenial 
and dentary, and reaches only as far forward as the level of the 
penultimate tooth. The coronoid is represented only by the 
interior dentary process, which is inserted into a notch between 
splenial and dentary. The teeth are bulbous and have slightly 
crenulated surfaces. Immediately posterior to the broken an- 
terior end of the specimen a pit is present for a replacement 



1961 



.MIOCENE LIZARDS FRO.M COLO.MBIA 



tooth. The subsequent teeth (referred to here by numbers 1-8) 
form a slightly curved series when viewed dorsally, and the 
line of occlusion viewed laterally has a slight dorsoposterior 
curve. Tooth one is slightlv bulbous and smaller than tooth two. 





Figure 1. a. Tupinambis of. T. teguixin, U. C. no. 38856, laljial view of 
left dentary, x 2. b. The same, lingual view. 



Two is taller and slimmer than the succeeding teeth and is 
slightly recurved and concave posteriorly. Tooth three is being 
replaced, its crown is broken, and its thin eroded base has been 
crushed ventrally so that the tooth appears lower than it actu- 
ally was; measurement of the replacement tooth indicates that 
it would be intermediate in height between the preceding and 
succeeding teeth. The replacement pits under the first and sec- 
ond teeth are in the interdental position. Teeth four and five 
are subequal : four has a faint anterior and posterior crest near 



4 BREVIORA No, 143 

the crown. The crown of tooth five is worn smooth. The large, 
fully formed replacement tooth below it has a slightly crenulated 
surface near its apex, and its anteroposterior crests are con- 
nected to the central point of the crown. Teeth six and seven 
are subequal and slightly smaller than teeth four and five, and 
the crests on tooth six are slightly worn on the occlusal surface. 
Tooth seven is worn smooth and is being replaced, but the 
replacement tooth is missing. Tooth eight is about one-half the 
size of the rest of the teeth, has anteroposterior crests, and a 
large replacement crypt below it. All of the teeth are essentially 
round, but the last few show a slight tendency to be longer than 
wide. 

Discussion. The jaw is about the same size as that of the 
Dracaena described below, and very slightly larger than that of 
the largest Tupinanihis tegnixin seen by me. It differs from 
Dracaena in having teeth which are higher than broad (or only 
slightly lower than their broadest dimension), and which are 
not as prominently constricted at the base ; in having a de- 
pressed area on the surangular region ; and in having teeth with 
anteroposterior crests. The only characters in which the fossil 
differs from specimens of Recent T. teguixin are the concave 
splenial and the size of the posterior teeth. The splenial is 
concave in Dracaena and in some species of Cnemidophorus, but 
is usually flat or slightly concave to convex in Tupinanihis. In 
the fossil, however, the concavity may in part be the result of 
postmortem shrinkage or crushing and is not conclusive. The 
posterior teeth of Tnpinamhis teguixin are usually relative^ 
slightly smaller than those of the fossil, and are often more 
linguolabially compressed, though this character varies. The 
minor differences between the fossil as preserved and Recent 
specimens do not seem to warrant taxonomic separation. The 
other Recent species, T. nigropunctatus, has higher crowned, 
smooth, and prominent cuspate teeth, and is smaller than T. 
teguixin. 

Rovereto (1914a) described several fossil species of Tupinani- 
his from the Pliocene of Argentina. Their validity is difficult 
to determine at the present time, but they seem to resemble 
T. teguixin most closely. 

Dracaena colombiana n. sp. 

Type. U. C. no. 39643, complete right dentary, collected by 
Dr. R. W. Fields. 



1961 



MIOCENE LIZARDS FROM COLOMBIA 



Referred specimens. U. C. no. 40277, distal and proximal ends 
of a left femur, catalogued as float from ?Cerbatana gravels. 
U. C. no. 37899, sacrum and associated first caudal vertebra, 
collected by Dr. Jose Royo y Gomez from U. C. loc. V-4517, 
Monkey Unit. U. C. no. 38927, posterior half of right maxilla, 
with bases for three teeth, from U. C. loc. V-4528, Upper Red 
Beds. The four localities occur within an area about six kilom- 
eters in diameter. 

Type locality. U. C. loc. V-4536, San Nicolas, near Villavieja, 
Iluila, Colombia, South America. San Nicolas clays, Honda 
group. 

Age. Late Miocene. 

Diagnosis. Differs from the Recent Dracaena guianensis in 
the following characters : dentary larger and more robust ; tooth 
row more strongly curved upward posteriorly; tooth outline 
round or suboval ; tooth number fifteen. From the living D. para- 
guayensis it differs in possessing five more dentary teeth. It 
differs from Tupinamhis tegnixin in having more posteriorly 
expanded cheek teeth, which are larger, basally constricted, and 
wider than high, and in lacking a depression in the exterior 
notch for the surangular. 

Description. The dentary is 60 millimeters long. The Mec- 
kelian fossa is widely open. The robust symphysis has a rugose 




Figure 2. Dracaena coJoinhiana, n. sp., U, C. no. 39643, tjT^e specimen, 
]Miocene, Colombia, South America. Labial view of right dentary, x 1.5. 



articular surface, and is separated from the main body of the 
dentary by a linguolabial constriction. There are five inferior 
alveolar foramina, and the dentary is rugose labially near the 
symphysis, probably for the genioglossus muscle. A broad shelf 



BREVIORA 



No. 143 



is present labial to the tooth row. The labial coronoid notch is 
prominent, and reaches as far forward as the anterior border of 
the last tooth. There are fifteen subpleurodont (or subacrodont) 
teeth, of which all but six are preserved. Teeth one through 
five are missing; six and seven are conical and slightly striated. 
The eighth is missing, but the base shows that it was molariform. 
Teeth nine to fifteen are molariform, and the ninth projects 
well above the preceding and succeeding teeth. Beyond the 
ninth, the teeth are flattened, button-like, and have finely stri- 
ated, round or suboval crowns, which increase in diameter to 
the twelfth tooth and then decrease posteriorly. Most of the 
molariform teeth have occlusal attrition facets, especially tooth 
twelve. Teeth seven through fifteen have replacement pits at 
their bases, and several of these pits (eight, ten, twelve, four- 
teen) contain almost full term replacement teeth. The tooth 
replacement is directly successive, and apparently alternate. 




Figure 3. Dracaena colomhiana, ii. sp, U. C. no. 39643, type specimen, 
Miocene, Colombia, South America. Lingual view of right dentary, x 1.5. 



The fragment of maxilla is robust, externally concave, and 
bears the bases of three large molariform teeth, of which the 
second has a replacement pit containing a large, flattened re- 
placement tooth. 

The femur has been broken, and there is a section of the 
diaphysis missing. It has been restored to a length of 75 milli- 
meters, by analogy with femora of Dracaena guianensis. It is 
large, but not especially robust, and the patellar groove and 
muscle attachment scars are prominent. A small U-shaped notch 
separates the head from the trochanter major. 

The sacrum is composed of two co-ossified vertebrae, which 
have prominent zygosphene-zygantrum articulations. The maxi- 
mum length of the sacrum from center of cup to tip of ball is 



1961 



MIOCENE LIZARDS FROM COLOMBIA 



20 millimeters. The greatest width from the centers of iliosacral 
articulation is 45 millimeters. The first caudal vertebra has a 
perfectly hemispherical cup and ball, and a prominent zygo- 
sphene-zygantnim articulation. It is 16 millimeters long from 
center of cup to tip of ball. 

Discussion. The distinctive specialization of the teeth of 
Dracaena has long been known. Owen (1845; figured a speci- 
men, and Peyer (1929) discussed the form and function of the 
teeth. Amaral (1950) has described another Recent species, 
D. Paraguay ensis, but I have not seen a specimen. 



[O 



o°ooooO 





^ OOOO oo^^o 




oooooooo 



Figure 4. a. Outline of occlusal view of right dentary, Dracaena coloin- 
hiana, U. C. no. 39643, Miocene, Colombia, b. The same, Recent B. 
quianensis. c. The same, left dentary, Recent Tupinambis ieguixin, A. M. 
no. 62545. d. The same, Tupinambis of. T. ieguixin, U. C no. 38856, 
Miocene, Colombia, a.-d. x 1.5. 



The principal differences from the Recent D. guianensis are 
as follows: (1) The fossil is more subpleurodont in tooth im- 
plantation, resembling other teiids, whereas the Recent species 



8 



BREVIORA 



No. 143 



tends to be subaerodont. (2) D. colombiana has fifteen dentary 
teeth, D. paraguayensis has ten {fide Amaral) and D. guianensis 
twelve. (3) The anterior teeth, as far as they are preserved, are 
less bulbous than in the Recent species. (4) The occlusal outline 
of the tooth crowns is round or suboval in the fossil and sub- 
oval or subrhoml)ic in D. guianensis. (5) The crown surface of 
the molariform teeth is more flattened, and the dorsoventral 
compression of the entire tooth is less in the fossil. 

The referred skeletal elements are somewhat larger than 
would be expected for a modern Dracaena with a dentary the 
size of D. colombiana, as the appendicular skeleton is relatively 
small in the Recent species. If correctly referred, the skeletal 
elements perhaps came from a larger individual. The sacrum 
and first caudal vertebra are characteristically teiid, having a 
perfectly round cup and ball, and strongly developed zygo- 
sphene-zygantrum. They resemble comparable bones of Recent 
Dracaena more than those of Recent Tupinamhis. The femur 
has the deep patellar groove seen in many teiids and resembles 
Dracaena in having more separation between head and tro- 
chanter major than that seen in Tupinamhis. The lack of a 
pronounced fossa posterior to the condyles resembles the condi- 
tion in Dracaena. Some of these bones may be referable to 
the Tupinamhis described above, but on the basis of present 
comparisons, all are referred to Dracaena colomhiana. 



Family IGUANIDAE 
Unidentified Genus and Species 

A fragmentary left dentary of an iguanid, U. C. no. 39644, 
from U. C. loc. V-4517, Monkey Unit, shows some similarity 
to the living Polychrus, but lacks the vertical wrinkling of the 




Figure 5. Iguanidae, unidentified genus and species, U. C. no. 39644, Miocene, 
Colombia, left dentary, broken antero posteriorly, x 10. 



1961 MIOCENE LIZARDS FROM COLOMBIA 9 

tooth surface found in the latter. Many iguanid genera were 
compared with this fossil, but close correspondence was not 
found. It may be new or represent one of the forms not seen, 
and further identification was thus not attempted. 

ECOLOGICAL AND ZOOGEOGKAPHICAL 
CONSIDERATIONS 

Fields (1957, p. 394-395, and 1959, p. 428-429) has described 
the upper Magdalena Valley area during the late Miocene as a 
vast floodplain, subject to alternating periods of flooding and 
drying in a wet-tropical climate. Dry seasons resulted in exten- 
sive development of swamps and mudflats, and wet seasons 
were times of periodic, extensive flooding. The fossil mammalian 
fauna has a dominantly savannah aspect, but also includes 
riparian and aquatic types. 

Recent Dracaena giiianensis is particularly abundant in tidal 
marshy sections and swamps of the Amazon Valley, and Recent 
D. paraguayensis (more primitive in scutellation, yet having a 
reduced tooth count) inhabits drier ground on the fringes of the 
swamps in southern Brazil and Paraguay. The fossil Dracaena 
colomhiana is found in the LTpper Red Beds (playa lake de- 
posit), San Nicolas clays (lacustrine deposit), and the Monkey 
Unit (sheet floodplain deposit). Tupinamhis cf. T. teguixin 
occurs in the Lower Red Beds (playa lake deposit). These units 
all reflect a regional marshy floodplain environment. Accord- 
ing to Fields {op. cit.) the Magdalena Valley in this region 
today has a semi-arid climate and is between four and five 
hundred meters above sea level in the area where these sediments 
were deposited. Neither Tupinamhis nor (especially) Dracaena 
usually occur at this altitude, for both are humid-tropical types. 
They are thus in accord with structural and depositional indica- 
tions in the sediments (op. cit.) for a lower altitude and moister 
climate in this area during the late Miocene. 

These fossils constitute a considerable range extension for the 
genera involved. Dracaena is found today from the Guianas 
southward to the Amazon, Tocantino, and Sao Francisco basins, 
and from the southern Matto Grosso region, Brazil, south to the 
District of Chaco, Paraguay (Amaral, 1950). Tupinamhis occurs 
over much of the forested districts of tropical South America 
from Trinidad south through the Guianas to Uruguay. Thus, 
nearly a thousand miles separate the fossils from their nearest 
Recent representatives. It is apparent that these genera were 



10 BREVIORA No. 143 

once much more Avidely distributed over northern South Amer- 
ica than they are at present, and that their habitat has been 
restricted both by uplift and by increasing aridity. 

The presence of Tupinamhis in these late Oligocene and late 
Miocene sediments indicates an early Tertiary origin for this 
genus, and perhaps also for Dracaena. The bufonid frog men- 
tioned in the introduction is closely related to the Kecent 
South American species Bufo alvarius and B. crucifer, and 
further heightens the modern aspect of the late Miocene herpeto- 
fauna of northern South America. It is clear that modernization 
of at least part of this fauna took place not later than the Mio- 
cene, and quite probably much earlier. 

SUMMARY 

The teiid lizards Dracaena colomhiana, n. sp., and Tupinamhis 
cf. T. teguixin, and an unidentified iguanid lizard occur in the 
late Miocene La Venta fauna, Colombia, South America. The 
fossil teiids, found a thousand miles from their nearest modern 
representatives, indicate a once greater distribution for these 
genera in northern South America. Their presence corroborates 
previous interpretation of the La Venta region during the late 
Miocene as a moist, swampy, lowland floodplain. 

A fragmentary specimen from the Coyaima fauna shows the 
presence of Tupinamtis sp. in the late Oligocene of the same 
region. 

BEFEEENCES CITED 

Amaral, Afranio do 

1950. Two new South American lizards. Copeia, 1950, no. 4, pp. 
281-282. 
Fields, Egbert W. 

1957. Hystrieomorpli rodents from the late Miocene of Colombia, 
South America. Univ. Calif. Pub. Geo!. Sci., vol. 32, no. 5, 
pp. 273-404, 35 figs., 1 pi. 
1959. Geology of the La Venta badlands, Colombia, South America. 
Univ. Calif. Pub. Geol. Sci., vol. 32, no. 6, pp. 405-444, 2 
figs., 4 pis., 2 maps. 
Owen, Sir Eichard 

1840- Odontography. Hippolyte Balliere, London, vols. 1 and 
1845. 2. 
Peyer, Bernhard 

1929. Das Gebiss von Varanus nilotious L. und von Dracatna yuia- 
nensis Daudet. Ann. Soc. Zool. Suisse, vol. 36, no. 3, pp. 71-102, 
9 figs. 



1961 MIOCENE LIZARDS FROM COLOMBIA 11 



ROVERETO, C. 

1914a. Los estratos araucanos y sus fosiles. Ann. Mus. Nac. Buenos 
Aires, vol. 25, pp. 1-247, 93 figs., 31 pis. 
Savage, Donald E. 

1951. Report on fossil vertebrates from the upper Magdalena Valley, 
Colombia. Science, vol. 114, no. 2955, pp. 186-187. 
Stirton, Ruben A. 

1953. Vertebrate paleontology and continental stratigraphy in Colom- 
bia. Bull. Geol. Soc. America, vol. 64, pp. 603-622, 13 figs. 



BREVIORA 

Museitim of CoHiparative Zoology 



Cambridge, Mass. August 21. 1961 Number 144 

A LARGE OPHIACODOXT PELYCOSAUR FROM THE 
PENNSYLVANIAN OP THE PITTSBURGH REGION 

By Alfred Sherwood Romer 

The presence in the early Permian of a varied series of pely- 
eosaurs indicates that the beginnings of this synapsid group 
occurred well back in Carboniferous times. Pennsylvanian re- 
mains of pelycosaurs are, however, very rare. Because of this 
paucity of data, the remains described below are worthy of record 
despite their incomplete and fragmentary nature. 

Along the course of McKnight Road, about 6 miles north of 
Pittsburgh, Pennsylvania, there has recently been considerable 
quarrying of sand and shale deposits of Conemaugh age in order 
to level the surface for building constructioii. The materials 
quarried were used as fill in North Park, several miles farther 
north. During the course of this work geology students at the 
University of Pittsburgh noticed three pieces of bone in freshly 
dumped park fill; through the courtesy of ]Mr. Martin Bender 
of the University's geology department they were deposited in 
the Carnegie ]^.Iuseum. Inquiry made it rather certain that this 
material had been excavated from a locality on the east side of 
McKnight Road near the junction of Brown's Lane. The sand- 
stone excavated here, lies just above the Ames limestone, and 
hence pertains to the upper part of the Conemaugh group of the 
Pennsylvanian. It seems certain that all three of the fragments, 
pertaining to a right shoulder girdle and front leg, were parts 
of a single, presumably articulated, specimen, but the state of 
the quarrying operation precluded any attempt to find further 
remains. The specimens are entered as no. 13942 in the vertebrate 
paleontology collections of the Carnegie Museum, Pittsburgh; I 
am indebted to Curator Craig Black for placing them in my 
hands for description. Needed preparation Avas done at the 
Museum of Comparative Zoology under a grant from the National 
Science Foundation. 



2 BREVIORA No. 144 

The most revealing specimen consists of the distal half of a 
right humerus. It is clearly pelycosaiirian in nature and, it would 
seem, equally certainly ophiacodontid, the diagnostic features 
l)eing the greatly expanded entepicondyle, the transversely di- 
rected supinator process and the strong development of the lateral 
margin of the ectepicondyle. In addition, the preserved portion 
of the shaft indicates that proximal and distal halves of the 
bone Avere sharply twisted on one another as in ophiacodonts and 
in contrast to the lesser torsion in other pelycosaur groups. The 
overall breadth is 96 mm. The bone is almost exactly super- 
posable on a mature Ophiacodon humerus in the M.C.Z. collec- 
tions (no. 1426) from the Archer City bonebed (Putnam 
formation) . This Texas element is 120 mm. in length, and pertains 
to a small individual of 0. rctroversus or to an immediate pre- 
decessor of that species. If the proportions of the present form 
were similar to those of Ophiacodon, the animal in life would 
have had a length of head and trunk combined of about 109 em., 
and a total length (including the usual long pelycosaurian tail) 
of about 204 cm. or about 6'8". 

Our Pennsylvanian specimen thus appears to be one of the 
largest of ophiacodonts, exceeded in size only by 0. retroversus 
and 0. majoi- of the later Wichita formations and by StcreorJiachis 
dominans of the European Permian. Post-mortem, the bone has 
lost the hemispherical ventral swelling with which the radius 
articulated; the supinator process is imperfectly preserved and 
there has been some slight damage to the ventral surface at the 
end of the entepicondyle. Otherwise, apart from crushing, the 
bone is well preserved. Notable and possibly significant is the 
fact that ossification had been completed at the time of death. 
This is in contrast to most specimens of Ophiacodon, in which but 
a small fraction of limb bones show complete ossification of 
their ends, even when tlie animal appears to have reached a 
■'mature" size. 

Although the general pattern of the bone is comparable to that 
of Ophiacodon, there are definite diff^erences in detail. The 
present specimen has, dorsally, a prominent muscle scar proximo- 
medially situated near the end of the entepicondyle ; this is not 
present in Ophiacodon. The ridge extending proximally along 
the lateral margin of the ectepicondyle is better developed in the 
present form than in Ophiacodon. The supinator process appears 
to be less developed than in that genus, but the apparent dif- 
ference is due to post-mortem damage in our specimen. 



1961 LARGE PENNSYLVANIAN PELYCOSAUR 3 

A second fragment preserved is the proximal end of the right 
ulna. This has been greatly crushed, so that the sigmoid articular 
surface for the humerus is unnaturally narrow, and identification 
of diagnostic features is difficult. As in the case of the humerus, 
ossification is practically complete so that, in contrast to most 
Ophiacodon specimens, the whole olecranon is represented by 
bone rather than cartilage. 

A third fragment is the upper end of a right scapula. The 
posterior margin is thickened and rounded in typical pelycosaur 
fashion. The low^er posterior portion of the piece preserved is 
somewliat crushed and distorted, but there is an increase in 
thickness and a curvature of this border such as one would 
expect as the supraglenoid region is approached. In many pely- 
cosaur specimens, even when seemingly "mature," the upper end 
of the scapular blade has a thickened distal edge obviously con- 
tinued in cartilage. The present specimen, which appears to be 
nearly intact in the posterior part of its upper end, shows no 
such terminal surface ; ossification was apparently complete. 
More anteriorly the bone has been damaged so that for about 6 
cm. there is a thick broken margin on the preserved portion. 
Beyond and below this the margin as preserved, although im- 
perfect, is thinner where broken oft". Probably the missinii' area 
here had about the extent indicated by the broken line in the 
figure. The anterior margin flares out laterally to a marked 
degree from the general plane of the scapular blade. It seems 
fairly certain that there was considerable constriction in th<' 
width of the blade toward the bottom of the preserved segment. 
This is in contrast with Ophiacodon and Varanosaurus in which 
the scapular blade is broad throughout its height (the scapula 
of Clcpsydrops is incompletely preserved). 

Fragmentary as are the remains here described, they show 
definitely the presence at this horizon, fairly well down in the 
Upper Carboniferous, of a large pelycosaur which is surely an 
ophiacodontoid and nearly equally surely an ophiacodontid, ante- 
cedent although probably not ancestral to Ophiacodon of the 
Permian. Despite its incomplete nature, this fossil deserves 
taxonomic standing because of its importance in the story of 
pelycosaur evolution, and it is herewith made the holotype of 
Clepysdrops? magnum, sp. nov. It will not probably prove, when 
better known, to be generically distinct from Clepsydrops, but it 
appears to be related to that genus, likewise of late Carboniferous 
age, and it may be provisionally included in it. Because of 
imperfect knowledge of comparable anatomical features, we may 



4 BREVIORA No. 144 

for the present rely for diagnosis simply on the size, which is 
approximately half again that of specimens assigned to the 
genotype, C. collettii. 

Advantage may he taken of the present opportnnity to mention 
a few fragmentary remains of Pennsylvanian pelycosaurs found 
in Ohio by Dr. Donald Baird and Mrs. Baird during a 1955 
ex])edition of the Museum of Comparative Zoology. (1) M.C.Z. 
no. 2411 is from the Summerfield limestone of the Conemaugh 
group ; the locality lies in Center Township, Noble County, Ohio, 
on the south side of state highway 78, just west of its junction 
with route 147. Found here were a crushed claw-shaped ungual 
phalanx 24 mm. long, comparable to the unguals of Clepsydrops, 
two complete pelycosaur centra with transverse diameters of 20 
and 14 mm., and a fragmentary centrum with an estimated 
diameter of about 15 mm. (2) M.C.Z. no. 2295 from the Ewing 
limestone of the Conemaugh, in the central part of section 7, 
Noble Township, Noble County, a fragment of a centrum with 
an estimated diameter of 20 mm. or so. (3) M.C.Z. no. 2777, 
from the Lower Uniontown limestone of the Monongahela group, 
along Leitli Run Road 3.8 miles from the junction with state 
route 7, Washington County, Ohio, an ungual phalanx compar- 
able to that mentioned under no. 2411. It may be noted that this 
is the only identified reptilian bone from the Monongahela. 
There is, of course, little in these fragments to allow us to give 
any positive generic determination. But since these specimens 
are within the size range of the Illinois materials of Clepsydrops 
they may be provisionally referred to that genus. 

As mentioned earlier, described pelycosaurian remains from 
the Carboniferous are very rare indeed. Their rarity is pre- 
sumably to be correlated with the fact that most Carboniferous 
fossil localities are from coal swamps in which, apart from am- 
phibious ophiacodonts, pelycosaurs would not be expected. Prev- 
iously described are: (1) Numerous isolated elements of a small 
ophiacodontid, Clepsydrops (Romer and Price, 1940, pp. 212- 
216) from the McLeansboro formation near Danville, Illinois. 
(2) A fragmentary spine of a small Edaphosaurus from the 
Round Knob formation of the Conemaugh group near Pitcairn, 
Pennsylvania (Case, 1908. pp. 237-238: Romer and Price, 1940, 
p. 388). (3) From the late Stephanian of Kounova, Bohemia, 
a single vertebra of a similarly small Edaphosaurus (Romer and 
Price, 1940, p. 388). (4) From the same locality, a number of 
bones which represent a sphenacodont of good size, to which the 
name Macromerion schwarzenbergii is applicable (Romer, 1945, 
pp. 429-431). 



1961 LARGE PENNSYLVANIAN PELYCOSAUR 5 

Little as we know of Pennsylvanian pelycosaurs, a few general 
conclusions can be reached. That our scanty data on Carboni- 
ferous pelycosaurs should include remains of ophiacodonts such 
as Clepsydrops is not surprising. It is believed that the ophiaco- 
dontoids are the most primitive of pelycosaur stocks and would 
hence be expected to appear at an early period in the record. 
Further, it is believed that many, at least, of the ophiacodontids 
were amphibious in habit — essentially piscivorous water-dwellers 
whose ancestors had never abandoned an aquatic existence. In 
consequence the chances of finding ophiacodonts in the coal 
swamp deposits which constitute most of the Carboniferous 
record is much greater than of finding representatives of the 
more terrestrial sphenacodontoids or edaphosauroids. 

Any idea that the finding of Clepsydrops in the Conemaugh 
takes us near in time to the point of origin of the pelycosaurs 
is an illusion. For fragments of edaphosaur spine have been 
recovered not only from the very late Kounova deposit but also 
from the Round Knob formation of the Conemaugh. Edapho- 
saurus is a highly specialized end form. It is obvious that its 
evolution from an ancestral pelycosaur must have begun at a 
time much further back in the Carboniferous than the Cone- 
maugh. The assumption (not an unreasonable one) that the 
major pelycosaur groups evolved independently from a capto- 
rhinomorph stock only begs the question. 

In general, reptilian stocks of any sort begin with forms that 
are of small size (as well as of primitive nature) and become 
increasingly larger as their history progresses. Such a process 
seems clear in the Lower Permian history of the pelycosaurs. The 
earliest dimetrodonts of the Texas Wichita were relatively small, 
and giants developed in the Clear Fork; Edaphosaurus species 
increase greatly in size in later beds ; the caseids grew from small 
forms to enormous Cotylorhyiichus species; even Ophiacodon 
(which became extinct before the typical Clear Fork) developed 
from small forms in the lower Wichita and New Mexican beds 
to the relativel}' gigantic 0. major, in the Clyde Formation. 

Until the discovery of the present specimen of C. magnus the 
known Pennsylvanian pelycosaur material fell in line with the 
belief that all early pelycosaurs were small. The Clepsydrops 
specimens from Illinois and Ohio represent small animals, as do 
both Penns3dvanian scraps of Edaphosaurus. To be sure, Macro- 
ynerion of Kounova is a good-sized sphenacodont, but Kounova 
is close to the Permian boundary and Macromerion could be 
regarded as a slightly premature representative of the Permian 



6 BREVIORA No. 144 

trend to large size. The discovery of C. magtius, however, destroys 
the illusion, for, as noted, this animal was larger than most of his 
later Permian relatives. 

When and if we ever come upon Upper Carboniferous deposits 
with a representative terrestrial, rather than coal-swamp fauna, 
it can be confidently predicted that there will be included a 
considerable number of pelycosaurs which wall vary widely in 
structure and size. It seems certain that for the origin of this 
fauna, of which so far we have but slight traces, we must look far 
back in the Carboniferous, to at least early Pottsville or Namurian 
levels. 



LITEEATUEE CITED 

Case, E. C. 

1908. Description of vertebrate fossils from the vicinity of Pittsburgh, 
Pennsylvania. Ann. Carnegrie Mus., 4:234-241. 
EOMER, A. S. 

1945. The late Carboniferous vertebrate fauna of Kounova (Bohemia) 
compared with that of the Texas redbeds. Amer. Jour. Sei., 
243:417-442. 
EoMER, A. S. and L. I. Price 

1940. Eeview of the Pelycosauria. Spec. Pap. Geol. Soc. Amer., no. 28: 
1-538. 



1961 



LARGE PENNSYLVANIAN PELYCOSAUR 








Figure 1. Clepsydrops magnus sp. nov. A, B, distal end of right humerus, 
dorsal and ventral surface. C, upper part of right scapular blade; probable 
outline when complete suggested by broken line. D, E, lateral and proximal 
views of upper end of left ulna. All x i/^. 



BREVIORA 



Musemnn of Comparative Zoology 



Cambridge, Mass. September 5, 1961 Number 145 



A NEW SPECIES OF THE CETOMIMID GENUS 
GYRINOMIMUS FROM THE GULF OF MEXICO 

By Henry B. Bigelow 

Gyrinomimus parri new species 

Type. U.S. National Museum No. 196180, from north central 
part of Gulf of Mexico, 26° 52' N, 89° 44' W, OREGON Station 
2573. One specimen, 51 mm in standard length. 

Distinctive Characters. G. parri differs conspicuously from 
G. simplex Parr 1946,^ from the Gulf of Mexico, in its relatively 
much longer dorsal and anal fins, from G. nujersi Parr 1934,' also 
from the Gulf, in the simple structure of its lateral line pores, 
and from G. hruni, Rofen 1957-59,^ from the southwestern Indian 
Ocean, in the number of lateral line pores. 

Description. Proportional dimensions in per cent of standard 
length of the type specimen, 51 mm in standard length. 

Snout in front of eye: 51. 

Diameter of eye: about 1. 

Length of head to pectoral fins: 33. 

Snout to origin of dorsal: 62. 

Base of dorsal: 24. 

Base of anal: 23. 

Rear end of base of dorsal to base of caudal: 10. 

Length of caudal: 12. 

Length of pectorals: 6. 

Height at pectorals: 24. 

Height at origin of dorsal: 17. 

Least height of caudal peduncle: 8. 

Height of base of caudal fin : 12 

ICopfia. 1946: 116, PI. 1. 

2 Bull. Binphaiii Oceanogr. Coll.. 4 (6) : 29, Figs. 8, 9, 1934. 

3 Galathea Kept., 1 : 257. 1957-59. 



BREVIORA 



No. 145 



Greatest tvUltJi. hcttcccn rear end of rami of lower jaw: 18. 

Width between upper ends of f/iH .s7(7.v: 14. 

Dorsal rays: 17. 

Anal rays: 17. 

rri)icipal caudal r(iy.s: 17+ (damaged). 

Pectoral rays : 16. 

Dorsal profile sloping- in nearly a straight line from nape to 
tip of snout, the latter narrowly rounded vertically but rather 
broadly rounded transversely ; eye minute, much nearer to upper 
jaw than to dorsal profile ; upper jaw projecting a little beyond 
lower, margins of both jaws weakly concave, the jaws extending 
rearward about to a perpendicular at upper corner of gill open- 
ings. Trunk highest at nape ; dorsal profile of body and tail 
sectors sloping in nearly a straight line from nape to caudal 
peduncle. Dorsal fin a little higher anteriorly than posteriorly, 
its margin very weakly convex ; anal opposite dorsal and similar 
to dorsal in outline. Caudal (somewhat damaged) apjmrently 
truncate or nearly so ; lateral line with 14 large oval pores from 
nape to base of caudal, the anterior margin of the pores without 
rearward extending flap. Teeth along entire length of each jaw 
in 3 continuous rows, those in the row next the gum nuicli the 
shortest and those in the row farthest from the gum mudi the 
longest. 

Color. General ground tint sooty black, the eyes blue, the 
outer parts of the unpaired fins chocolate brown, the teeth ivory 
white, the inside of tlie mouth pale gray. 

Size. The type (and only known) specimen is 51 mm in stand- 
ard length. 

Rouge. So far known only from the north central part of the 
Gulf of Mexico, 26°52' N, 89^44' W, OREGON St. 2573, from 
a trawl haul at 1350 fms. 




Gyrinomimus parri n. sp. Type specimen, USNM no. 196180. 



BREVIORA 

Miiiseiulm of Comparsitive Zoology 



Cambridge, Mass. December 14, U)(il Number 14() 

NEW HODEXTS FKOM THE EAKLY MIOCENE 
DEPOSITS OP SIXTY-SIX MOUNTAIN, WYOMING 

By Craig C. Black 
Carnegie Museum, Pittsburgh, Pennsylvania 

Diirino- the summers of 1980, 1931. and W.VA. Dr. E. M. 
Sclilaikjer working for tlie Museum of Comparative Zoology, 
Harvard University, made a large collection of early Miocene 
mammals in the southern portion of Goshen Hole, Wyoming. 
]\Iost of these were obtained from the sloi)es of Sixty-Six and 
Bear Creek mountains, from beds that Schlaikjer (1935) in- 
terpreted as the Lower Harrison formation. 

On two occasions during July, 19.5!), Sabra B. Black, Laura 
]McGrew, Bryan Patterson, and I made a brief reconnaissance of 
the area on the western slope of Sixty-Six mountain. During this 
reconnaissance, it became evident that Schlaikjer "s Brule-Lower 
Harrison contact was actually a local channel conglomerate de- 
v('l()]ied within the early ]\Iiocene sediments. He states (1935, p. 
112), "'At the northwest end of Sixty-Six mountain in the N.E. 
1/4, Sec. 7, T.20N., R.(^()W. tlie typical Brule clay is overlain by 
a three-foot clay conglomerate above which are nineteen feet of 
clay sands that grade u]nvard into the gray sands with pipy 
concretions." The "typical Brule clay" below the conglomerate, 
however, is the unit from which the rodents described below were 
obtained together with a badly weathered Cyclopidins skull and 
jaws and the posterior portions of two rami of Mesoreodon cheeki. 
These specimens were found in place and they clearly indicate an 
early Miocene, probably Gehring equivalent, age. BeloAv the grey, 
tuifaceous sand and grading into it are bufif clays with some 
sand which are probably equivalent to the Brule. In this area 
deposition appears to have been continuous from the Oligocene 
through the early Miocene with no sharp lithologic or erosional 
break within this sequence. The clay conglomerate is not ex- 
tensively developed either on Sixty-Si.\ or Bear mountains and 



2 BBEVIORA Xo. 14(i 

where it tloes ai)iiear on Sixty-Six mountain it lies within early 
Miocene sediments as is clearly shown by the oreodonts and ro- 
dents described beloAv. A full account of the relationship of the 
Oli^ocene and Mioceiu^ sediments in the southern part of Goshen 
Hole is still in ])roo'ress and will be published at a later date. 

I would like to take this opportunity to thank the Kellam 
family ot Torrino'ton, Wyoming', "svho helped us innneasurably 
durin<i' oui- stay in the Goshen Hole area. I would also like to 
thank Professor Dryaii Patterson for his criticisms and sug'o'es- 
tions. The drawing's are by Mr. Clifford J. Mori'ow and were 
made possible by a g-rant from the Gulf Oil Corporation. 

Abbreviation used: ]\I.C.Z. — ]\Iuseum of Comparative Zool- 
ogy, Harvard University. 

Family CRICETIDAE 

ScOTTIMrs KELLAMORUM^ U. sp. 

Figure 1 

Ti/jx : :\I.C.Z. Xo. 7.!42. a right maxillary witli M^-M-. 
Hiipodif/ni : Type only. 

Horizon aiul locdh'tji: Section 11. T.2()X.. IMilW.. (ioslien Co., 
AYyoming. Arikareeaii, fi-om the supposed Gehring etiuivalent, 
early ^Miocene. 

Diagnosis: Smaller than Scoftinnis lopludus: teeth narrowei' 
in i-elation to length than in Scottiinus cxigutis: antero-jiosterior 
lophs nu)re ])rominent than transverse ; protoloph very weak on 
M^ absent on M- ; metaloph very weak on ]M-. incomplete on 
Ml. 

Description : In general, the ui)per molars of Scofliiiiiis kd- 
Jamorum, like those of ^. lopJiafus. are longer and narrower than 
those of the various species of Euiin/s. The nuire is as strongly 
developed as in N. lopJiatiis, but there are more accessory trans- 
verse crests than in that species. The cusps on M^-M- are high, 
and the crests are at a lower level. The anterocone of M^ is ex- 
tremely large. There is a connection between the paracone and 
protocone on M^, but it is shifted posteriorly and thus does not 
form a transverse loph. A comparable crest is not present on 
M-. The paracone and metaeone are joined by a low crest which 
in turn is joined by the mesolophid. There is a short crest pro- 
jecting lingually from the mure between the protocone and hypo- 
cone on both ]\P and M-. The anterior cingulum on M- is ele- 

1 Named for David ami Ji'aii KeUaiii of Torringtou, Wyomiug. 



irxn 



NEW MIOCENE RODENTS 



vatecl and stronjily clevclopcd. The [xjstcrior luetaeoue arm on 
M^ is directed baekAvard to fuse with the elevated posterior 
einguliini, whereas on M- it passes lingually to the hypoeone to 
form a weak', transverse metaloph. 




Fio-uie 1. Srottimus lellamontm ii. sp., Typo, M.C.Z. Xo. 7.'U2, IRyV-, 
nuterior end to the right, X20. 



Disciissioit : Two species of Oligocene cumyines descrilied by 
Wood (1937) seem referable to Scott i)iuis. These are Einni/s e.r- 
iguus, from the ]Middle Oreodon beds of South Dakota, and 
Lcid}i))iiis r(t)(.s, from the Middle 01i<>'()('ene (Vdar ("r(M'k beds of 
northeastern Colorado. Wood {up. cit., p. 25-1-255). in describin<i' 
Eiiun/s f.riguus, stated that it was transitional between typical 
Kiinins and ScoftiiiiKs. (Jalbreath (1953. p. 72) stated tluit all 
the s|)('cimens referable to Euimjs c.ridjtus in the Tnivcrsity of 
Kansas collections from the Cedar Creek beds were also "closely 
similar to the type of Lcichjiiufs retus Wood. In fact there is no 
(piestion in my mind but that they represent the same species." 
However, he did not synonymize the two, statinti-. '^J think this 
small sjx'cies {Ei())iys e.rl(ji(iis) is <>:enerically distinct from the 
species of Eiinij/s but am not prepai'cd to say whether or not it 
should be referred to Lcidyntys or to anotlun- genus.'" I'nder tiie 
circumstances, it seems propei- to place Eumns cxiguus in 
Scottinnis and to refer L(if}i/)i)iis rifiis to the svnonvmv of this 
species. This brings tlie numhci- of recognized species of Scottimns 
to three; <S'. lopJuitus, S. (.riguiis. and N. kdldiiKninii. 



4 BRET^aORA No. 140 

Tn the (lovelopiiiont of antero-posterior rather than transverse 
lophs ^cotti))U(s kcllomorum is more advanced than »S'. exigims. 
Since S. exigmis is of middle ( )li<iocene age this is, of course, to be 
expv-cted. S. kcUanioriDn, liowever. is not as advanced a species 
as the earlier- N. lopluitus. The transverse crests seen on M^ of 
N. kcUatnoruhi are also present on »S'. lophatus, but these crests 
are not seen on M- of *S. lophatus. This would seem to exclude 
N. lophatus from the ancestry of »S'. kclloinorKni; presumably a 
species similai* to S. cxiguiis was ancestral to both. 

EuMYS sp. 

M.C'.Z. No. 7384, a partial left ramus with tlu^ incisor and 
Mi-Mo cannot be indentified specifically. It does not show an\- 
tendency toward the formation of a stroiiiz' central antero- 
postei'ior lophid at tlie expense of the transverse lophids, and 
hence is certainly not the lower dentition of Scoftinuis kella- 
inoruni. The posterior ])roto('onid ai-m is extremely elongate on 
both Ml and Mj. On ^I^, a short anterior metaconid arm just 
fails to reach the anteroconid whereas a longer jn-otoconid arm 
does so. In size and general pattern the teeth closely resemble 
those of Eiiiuys rlcgatis and Eiiiui/s oblif/iddr iis. I'nfortunately 
the lower teeth of Leidyniys nernatodo)i and lockingionianus are 
unknown. 

Family HETEROMYTDAE 

Heliscomys schlaikjeri- n. sp. 

Figure 2 

Type: M.C.Z. No. 7335. a right maxillary with V^M~. 
Hypodigru : Type only. 

Horizon and locality: Section 11, T.2()X., ii.lilW., Goshen Co., 
Wyoming. Arikareean, from the supposed Gehring equivalent, 
early Miocene. 

Diagnosis: Size larger than that of any other known sjx'cies 
of Heliscomys: intei-nal cingulum undivided; posterioi- cingnluiii 
on P"* extending from hy])ocone to metacone ; ceiiti-al 1rans\('rse 
valley straight, directed somewhat posteriorly. 

Description : No trace of the |)aracone remains on the anle- 
roloph of the premolar. In this I'cspect, the tooth is typically 
perognathine. The posteroloph bears three cusps, the metacone 



1961 



XEW MIOCEXK KODK.V' 



and liypoconc hciiii;' of ('i(ual size with tlu- protocoiic and the 
ciitostylo hoinji' smaller. The eiitostyle curves forward and joins 
tlic internal edji:e of the i)rotoeone at tlie same level as does the 
hypocone. There is a small posterior eingulum running from 
tlie hyi)oeone to the base of the metaeone, and separated from the 
postei-oloph by a small ]iit. 




Figure 2. Heliscomys scJilailjrri ii. sp., Type, M.C.Z. Xo. 7335, RP^-M^, 
anterior end to the left, X30. 



Tlie molars are somewhat more lophate in ai)pearaiiee ihan are 
those of Tldiscoiin/s (jntjoDji (Wood. 1933) and //. Iintcln ri 
(Wood. 1935). although the j)rineipal cusps are still prominent. 
In this respect the teeth r(>semble more closely those of //. 
t( luiicr ps ((lalbreath. 194S). Tlie molars of both agree in having 
anterior and posterior cingula that rise rather steeply to join 
the pi-otostyle and (Mitostyh'. These cusps are closely appressed 
with no gap between them, a further point of resemblance to //. 
f< iiiiia ps. The median va]le\' is straight on both teeth, slanting 
somewhat ]iosteriorly, not sinuous as in II. (/rrf/m-iil and //. 
IkiIcIk )i. The ant(M'ior cingnlum is stronger than the jjosterior 
on both AC and M-. 

Diifciissioii : Although it is somewhal larger and shows a few 
minoi' ditt'erences, Heliscnintis svhhiikjt ri is extremely close to 
//. hiniictps. Roth sjx'cies lia\'e an nndivded internal eingulum 

2 XaiiU'd for K. M. SdilaiUJcT in rccdLrnilinn (if his extensive work in the (ioslicn 
Hole ai-ea. 



6 BBEVIORA No. 146 

Mild i\ sti'ai<rht median valley, charaftprs found neither in //. 
Iiatchcri nor in II. (jrcf/oiifi. In //. scJtlaikjcri, in addition to tlie 
somewhat laroer size, the median valley of the molars is directed 
somewhat more posteriorly and the posterior eingulum on P^ 
has shifted position. The latter distinction seems unimportant 
since the presence or absence and the position of the posterioi- 
cingulum are variable in fI(Iiscoiini.'<. In //. hatcheri this cin- 
gulum varies from absent to present, in some cases extending 
along the entire ])()steri()r margin of the molars; in a specimen 
from Pipestone Springs described by McGrew (1941), it appeai-s 
to be completely absent, while in //. fcnxiceps it connects the 
hypocone and the entostyle. Peeently. Keeder (1960) has de- 
s('ril)ed two new genera of heleromyids from the White River 
formation and lias eiii])hasized their lai'ge size and tlie (juadri- 
cuspidate nature of P4 in A})l(f<if(ni\( \ix and the (juinquicuspidate 
P4 of Akmaloiinis. Thes(^ genera are known only on their lower 
dentitions and hence can not l)e compared with //. schlaikjrri. 
11. srJilnik j( I'i, however, is somewhat largei' tlian eith(M' genus. 

Tabh' of Measurements (in mm. ) 

P4 Ml M- Ml Mo 

Seal ti III ii.s l( Ihi inonuK, 

aiitfiopostcrioi- 2.50 1.85 

transverse protoloph 1.60 1.43 

traiisveise metaloph 1.53 1.35 

Ktuini.s sp. 

anteroposterior 

transverse metalophid 

transverse hypolopliid 

Ill li.scoiinis schJailcjeri 

anteroposterior 0.90 l.lo 1.10 

transverse 1.20 1.4(1 1.40 



REFEEENCES 

(i.M. BREATH, E. C. 

]948. A new species of lieteromjid rodent fioni tlie Middle Oligocene 

of northeast Colorado with remarks on the skull. V. Kansas 

Publ., Mus. Xat. Hist., 1: 285-300, 2 pis. 
1953. A contrilnition to the Tertiary geology and jpaleontology of 

northeastern Colorado. U. Kansas Publ. Paleon. Contiih. Yerte 

brata. 4: 1-120, 2 pis. 



2.10 


2.00 


1.40 


1.90 


1.70 


1.90 



1961 NEW MIOCEXE RODENTS 7 

McGrew, p. O. 

1941. Heteromyids from the Miocene and T.ower Oligocene. Geol. Ser. 
Field Mus. Xat. Hist.. 8: .").")-.'7. 
Keeder, W. G. 

1960. Two new rodent genera from the Oligocene White River forma- 
tion (Family Ileteromyidae ). Fieldiana : Geology, 10: 511-524 

SCIILAIK.JER, E. M. 

1935. Contribution.s tu the stratigraphy and paleontology of the Goshen 
Hole area, Wyoming. Part IV. New vertebrates and the stra- 
tigraphy of the Oligocene .-md early Miocene. P)ull. Mus. ("onip. 
Zool., 76: 9."189, 41 pis. 
Wood, A. E. 

19.3.3. A new lieteromyid rodent from the Oligocene of Montana. .Idur. 
Mam., 14: 134-141. 

1935. Evolntion and relationships of the heteromyid rodents with new 
forms from the Tertiary of western North America. Ann. 
Carnegie Mus., 24 : 73-262. 

1937. The mammalian fauna of the White River Oligocene. Part II. 
Rodentia. Tran,s. Amer. Philos. 8oc., (n.s. ) 28: 155-2(i9, 11 pis. 



BREVIORA 



Museiamii of Comipsirsitive Zoology 



Cambridge, Mass. Decembek 15, 1961 Number 147 



AUSTRALIAX CARABTD BEETLES YITT. LEIRADIBA, 
ESPECIALLY THE TROPICAL SPECIES 

By p. J. Darlington, Jr. 
Museum of Comparative Zoology, Cambridge, Mass. 

This is the third part of this series to be devoted to flightless 
Carabidae of zooo'eographic importance from tropical eastern 
Aii.stralia. Pertinent earlier parts, inclnding a locality list and 
a discussion of transition of carabid faunas in wet forests from 
New Guinea to Tasmania, are listed at the end of the present 
paper. Information on deposition of types and a brief note on 
methods will be found at the beginninjr of Part YI (1961c). 

The present paper is concerned with the pterostichine genus 
Leiradira. Typical Lciradira are very distinctive Carabidae, dis- 
tinguished from (for example) Notonomiis by: deep-bodied form ; 
heavy, sometimes more or less deflexed head, with mandibles 
somewhat longer and more acuminate than in Notonomus; gen- 
iculate antennae ; mentum tooth absent ; and inner lobe of maxilla 
lined with what look like long slender teeth rather than bristles 
(see following key). These characters led Tschit.scherine (1902, 
p. 506) to place Leiradira in the tribe Deliniiui, far removed from 
Notonomus. However, some of my new North Queensland species 
reduce tiie gap between Leiradira and Xotononius and suggest 
tliat tribal separation of the two genera may not be justified and 
that, although the genera are distinct, they (and Dclinius) may 
be derived from a common ancestor. This is something to be de- 
cided in the future, by the next reviser of the generic classifica- 
tion of Australian Pterastichini. 

As I now understand the genus, Lciradira (in a broad sense) 
includes about a dozen species and is discontinuously distributed, 
chiefly in rain forest, along the eastern edge of Australia from 
just below Cooktown in tropical North Queensland south at least 
to the lower edge of the Dorrigo plateau in warm temi^erate 



2 BREVIORA No. 147 

north-central New South Wales. The ^enns divides into three 
well marked subgenera, which have geographical as well as 
structural bases. They are distinguished in the following key, 
which includes Notonomns for comparison. The tropical species 
of Notonomns will be treated in another paper. 

Key to suhgenera and some species of Leiradira 

1. Mentum tooth noi-iiial, long, deeply emargiuate; inner lobe of maxilla 
lined with many coarse setae not in single row ; head not deflexed ; 
mandibles shorter ; antennae not geniculate, 1st segment shorter than 
segments 2 + 3; form usually more depressed (but variable) 

(Notonomns) 

— Mentum tooth broad and short, or absent ; inner lobe of maxilla lined 
with single row of about 6 to 8 slender teeth; head often (not always) 
more or less deflexed, with mandiljles usually (not always) longer and 
more acuminate than in Notonomus; antennae often (not always) 
geniculate, with 1st segment usually (not always) as long as or longer 
than segments 2 + 3; form relatively deep-bodied {Leiradira, scnsti 
laio) 2 

2. Labrum moderately emarginate, 6-setose ; posterior-lateral pronotal 
setae on thickened margin; (mentum tooth present but liroad and short) 
(North Queensland) (subgenus Metadira) .4 

— • Labrum rithcr deeply, almost semicircularly emarginate or only 4- 
setose ; posterior-lateral pronotal setae inside (not on) thickened 
margin ; (mentum tooth present or absent) 3 

3. Labrum moderately emarginate, 4-setose; mentum without tooth; man- 
dibles shorter, but not so short as in Notonomus (northern New South 
Wales and South Queensland) (Leiradira, sensu stricto) 

3 or more full species 

— Labrum semicircularly emarginate, 6-setose; mentum tooth present 
(but very broad and short) ; mandibles longer than usual (Soutli Queens- 
land and Eungella Range) (subgenus Stomimorplnts) 8 

4. Elytral intervals very imequal (odd several times wider than even ones) ; 
1st antennal segment shorter than segments 2 + 3; (color green- 
purple) ignifer 

— Elytral intervals less unequal, or equal; 1st antennal segment usually 
equal to or longer than 2 + 3 5 

5. Elytral intervals subequal and striae normal, not widened except some- 
times near apex 6 

— Either odd elytral intervals wider than even ones or striae partly wide 
and opaque 7 

6. Larger (15-19 mm.) ; dark blue-purple alticola 

— Smaller (11-12.5 mm.) ; black soror 



1961 AUSTBALLVX CARABID BEETLES 3 

7. Odd elytral intervals wider than even ones; olytral striae not nuu-h 
widened; 3rd elytral intervals usuallj' 3- or 4-punetate; scntellar striae 
weak or obsolete (variable) ; form more slender alternans 

— Elytral intervals nearly equal ; elytral striae widened and opaque espe- 
cially behind middle; 3rd elytral intervals always 2-punctate; scutellar 
striae short but deep ; form less slender opacistriatus 

8. Larger (14.7 mm. or more) violacevs 

— Smaller (c. 10-12 mm.) 9 

9. Prothorax wider (W/L 1.14-1.19") : duller Jacohi 

— Prothorax nari'ower (W/L l.OG-1.08) ; more shining tenuis 

Of the three subgenera, Metadira is most like Notonomus. Even 
the least modifiecl Metadira, anrifer, differs from Notonomns in 
having a shorter mentum tooth and a regular row of about 8 
slender tooth-like processes on inner edge of maxilla instead of 
more than 20 thickened setae less regularly arranged as in, for 
example, Xotonomus doddi Sloane, but there are two indications 
besides the general similarity that suggest a real relationship. 
One is the position of the posterior-lateral prothoracic setae, on 
(not inside of) the thickened margin at basal angles in both 
Metadira and the tropical species of Nrjtonomus. The other is 
the fact that some Mctaelira and some tropical Notonomus have 
elytral striae conspicuously widened and opaque. This is an 
unusual character, unlikely to have evolved independently in 
the two genera. 

Metadira subgen. n. 

Form varying from that of a convex XutunoDnts to that of 
typical Leirndira. Head stout, sometimes somewhat deflexed ; 
eyes small, genae prominent (but variable) ; 2 supraocular setae 
each side; mandibles varying (in different species) in length and 
curvature ; labrum moderately emarginate, 6-setose ; antennae 
with basal segment varying (in different species) from shorter 
to longer than next 2 segments together ; frontal foveae short but 
well defined ; mentum tooth very short and broad, broadly emargin- 
ate; inner lobe of maxilla with single row of setae so thickened 
as to resemble long slender teeth ; palpi slender in both sexes. 
Prothorax cordate, with narrow margins : baso-lateral impressions 
small but deeply impressed ; usual 2 marginal setae each side, 
posterior ones on thickened margins at basal angles. Elytra usu- 
ally with (sometimes almost without) basal margin; humeri ± 
dentate; striae entire, narrow or widened (in different species) ; 
intervals equal or unequal (in different species) ; narrow lOtli 



4 BREVIORA No. 147 

interval present or indicated posteriorly- ; 3rd interval typically 
with 2 dorsal punctures (usually only 1 in alticola and soror, and 
usually 3 or 4 in alternans). Last 3 ventral segments weakly, 
variably sub-impressed across base but not sulcate ; mesosternum 
and prosternal process not setose. Male with anterior tarsi 
slightlj^ dilated, 3 segments with squamae ; S with 1,9 2 setae 
each side last ventral segment. 

Genotype: Leiradira aurifer Darlington (below). 

Five species of this subgenus are now known. All occur on 
(diiferent parts of) the Atherton Tableland/or Mts. Bartle Frere 
and Bellenden Ker (see map) and the northernmost of the five 
species extends north nearly to Cooktown. Three of the species 
{alternans, soror, opacistriaius) may be (distinct) geographical 
forms of one original widely distributed stock (see notes under 
alternafis). The other two species are very distinct, localized 
endemics. All the species are usually found (by day) under cover 
on the ground, in rain forest, but I got one opacistriaius in good 
savannah woodland a few hundred feet outside rain forest by 
the Davies Creek Road. 

Leiradira (Metadira) aurifer n. sp. 

Form as figured (Fig. 1), of a rather convex Notonomus, but 
with characters of Metadira as given above ; head and prouotum 
bright green, pronotum with variable purplish reflections, elytra 
dark copper-purple, lower surface and legs black, mouth parts 
and antennae brownish ; surface shining, except bottoms of striae 
opaque. Head Vjq or slightly less wddth prothorax; mandibles 
shorter than usual in Leiradira, rather strongly curved, acumi- 
nate apically; eyes moderate (in genus), genae c. wide as eyes, 
weakly rounded to neck ; 1st antennal segment c. % (by measure- 
ment) length of 2nd + 3rd segments together; middle segments 
nearly 2X long as wide. Prothorax c. %o ( — ) wider than long 
at middle, moderately narrowed behind, more in front ; base c. 
%o wider than apex; apex subtruncate or very broadly emargi- 
nate with angles scarcely advanced, margined only near sides ; 
base slightly sinuously subtruncate, deeply margined toward sides 
but not at middle ; sides broadly rounded for much of length, 
sinuate near base ; basal angles c. right, scarcely blunted ; disc 
moderately convex with middle line distinct and reaching base, 



19(31 



AUSTRALIAN CARABID BEETLK^^ 







.<^ 






^N f 4: \ 



O^"" 
»> 



. ^^^" 



^^Mt.'B.F 



^^^ 



\ 3 .K\^^/ 



Ravenshoe 



Known distribution of Leiiadira, eudoniic suljgt'nus llctadira, in 
North Queensland. The finely dotted line is the approximate eastern 
edge of high land (the Atherton Tableland etc.). No. 1, aurifer ; 
2, alticola; 3, alternans ; ■i,f<oror; 5,opacistriatus, whiehoceurs also 
north beyond the limits of the map almost to Cooktowu. 



6 BBEVIORA No. 147 

transverse impressions almost obsolete, and baso-lateral impres- 
sions short, linear, connected with side maryins by deeply im- 
pressed marginal grooves. Elytra ^/4 (or slightly more) wider 
than prothorax; basal margin entire, strongly raised, slightly 
scalloped, rectangular at subdentate humeri ; striae very wide 
and opaque especially posteriorly; intervals convex, very un- 
equal, odd ones several times wider than even, 7th strongly con- 
vex but not sharply carinate at base ; a narrow 10th interval 
present posteriorly ; each 3rd interval usually 2-punctate just 
behind middle and near apex (an additional puncture present 
near middle on 1 side in 1 specimen). Length 15-19; width 5.0- 
6.2 mm. 

Holotype i (M.C.Z. Type No. 80,382) and 9 paratypes from 
mountains north of Kairi, Atherton Tableland, North Queens- 
land, 3000-4000 ft., Dec. 1957 ; and 3 paratypes from south end 
of Davies Creek road, Atherton Tableland, Dec. 1957 : all speci- 
mens taken by myself, in rain forest. The two localities are not 
far apart in the mountain mass that lies on the Atherton Table- 
land between the Mareeba-Kuranda road on the north and 
Yungaburra-Lake Barrine road on the south. 

See preceding discussion and key for place of this species 
among other Leiradira. Although it has striking specific charac- 
ters (color, extreme alternation of elytral intervals), it is the 
least specialized member of the group in form, mandibles, and 
length of 1st antennal segment. 

Leiradira (Metadira) alticola n. sp. 

Form as figured (l-''ig. 2), pruthoi-ax strongly cortlate, elyti'a 
narrowed anteriorly ; rather strongly convex ; black, usually with 
purplish tinge, margins of elytra blue ; moderately shining. Head 
c. % or slightly less width prothorax ; mandibles slightly longer 
and less curved than in aiirifcr; eyes small, genae usually longer 
and more pronunent than eyes, strongly convex ; antennae very 
short, 1st segment slightly longer than next 2 together (by meas- 
urement), outer segments scarcely longer than wide. Prothorax 
slightly (less than Ym) wider than long at middle ; apex truncate 
with angles scarcely advanced, not margined ; base virtually 
truncate, strongly margined toward sides ; sides arcuate for c. % 
of length (sometimes subparallel near middle), strongy sinuate 
c. Yq or more of length before base, then parallel or diverging to 
base ; basal angles well defined, right or acute ; disc Avith middle 
line well impressed, very deep basally, transverse impressions 



1961 AUSTRALIAN CARABID BEETLES 7 

obsolete, baso-lateral impressions deep, subliuear, eoiiiiected with 
side margins by deep basal grooves. Elytra c. % (or slighth' 
less) wider than prothorax, narrowed basally; basal margins 
entire, scalloped, forming sharply defined right angles at sub- 
dentate humeri; striae narrow, slightly wider apieally, with a 
little dull sculpture at bottom ; intervals subequal, slightly con- 
vex ; 3rd interval with 1 puncture near or behind middle, usually 
without posterior puncture, but latter present on 1 side in 1 
individual (see Fig. 2). Parts of lower surface (sides of meso- 
sternum, first ventral segment, and parts of other ventral seg- 
ments) more or less punctate or subpunctate. Length 15-19; 
width 5.2-6.5 mm. 

Holotype $ (M.C.Z. Type Xo. 30,383) and 3 paratypes from 
Mt. Bellenden Ker, E. side, 3000-4500 ft., Jan. 1958, and 1 para- 
type, Mt. Bartle Frere, W. slope, near 5000 ft., Dec. 1957. These 
2 mountains are close together at the eastern edge of the Atherton 
Tableland south of Cairns, North Queensland. All specimens 
taken by myself, in mountain rain forest. 

Distinguished from other Leiradira as indicated in the preced- 
ing key; unique in form, and notable for very short antennae. 

Leiradira (Metadira) alternans Darlington 

This species (Fig. 3) was described by me in 1953 (p. 90) from 
Malanda etc. It is widely distributed on the central-southern 
Atherton Tableland, the limits of its known distribution being 
Atherton, Lake Barrine, the lower western slope of Mt. Bartle 
Frere, and Mt. Fisher southwest of Millaa ]\Iillaa. It may be 
represented by soror (below) on the eastern slope of Mt. Bellen- 
den Ker (and perhaps elsewhere on the eastern slope of the 
Tableland) and by opacistriaius on the northei-n edge oC the 
Tableland and northward. 

LEiRADmA (Metadira) soror n. sp. 

Small, slender, convex ; shining black with silky lustre but no 
distinct metallic color. Head % (±) width prothorax : mandibles 
relatively weakly arcuate ; eyes small, genae slightly more promi- 
nent than eyes, broadly convex: antennae with 1st segment 
longer than next 2 together. Proilwrax narrowly subcurdate, as 
long as or slightly longer than wide ; apex subtruncate witli 
angles scarcely advanced, not margined at middle ; base sul)- 
truncate except slightly rounded toAvard sides, not margined 



8 BREVIORA No. 147 

except near sides ; sides broadly arcuate, subparallel before mid- 
dle, strongly sinuate about Yi o of length before base ; basal angles 
c. right, scarcely blunted ; disc with middle line rather fine but 
reaching base, transverse impressions weak, baso-lateral impres- 
sions nearer sides than middle, linear, deep, joining lateral 
margins at base. Elytra -Xo (or less) wider than prothorax, 
subparallel, weakly narrowed anteriorly ; basal margin deep, 
scalloped, rectangular at subdentate humeri ; striae fine ; intervals 
convex, subequal ; 3rd interval usually 1-punctate slightly behind 
middle, posterior punctures usually absent (1 present on 1 side 
in 1 specimen). Parts of lower surface (sides of mesosternum, 
1st ventral segment, parts of other ventral segments) variably 
punctate. Length 11.0-12.5 ; width 3.3-3.7 mm. 

Holotype S (M.C.Z. Type No. 30,384) and 2 paratypes all 
from E. side Mt. Bellenden Ker, c. 3000 ft., North Queensland, 
Jan. 1958, taken by myself in rain forest. 

This species most resembles altcrnans (above) but is smaller, 
narrower, with less alternation of elytral intervals, and fewer 
punctures on 3rd elytral interval. 

Leiradira (Metadira) opacistriatus (Sloane) 

Sloane first (1902, p. 319) described this species in Notonomus, 
then (1913, p. 409) ruled it out of that genus. The types were 
said to be from Cairns, collected by Froggatt ; they probably 
really came from the mountains near Cairns. I could not locate 
the types in Australia, but my specimens fit the description 
reasonably well. They are from the Davies Creek road on the 
northern Atherton Tableland ; near Black Mt. about 20 miles 
north of Kuranda (these are probably virtual topotypes) ; Mt. 
Lewis southwest of Mossman ; and Mt. Finnegan south of Cook- 
town. 

Leiradira, scnsu stricio 

Castelnau 1867, p. 72. 

Csiki 1929, p. 500 (see for additional refereuces and list of previously 
described species). 

Genotype, by present designation : Leiradira auricollis Castel- 
nau (Fig. 4) (genus originally' based on this species and latrcillei 
Castelnau ) . 

Leiradira, scnsu stricto apparently occurs only below the 
tro])ics, from the Blackall Range etc. in South Queensland south 
at least to the lower (eastern) edge of the Dorrigo plateau. The 



1961 AUSTRALIAN CARABID BEETLES 9 

subgenus includes at least three full species, perhaps more. I 
have not studied them and cannot establish synonymies. 

Subgenus Stomimorphus Straneo 

Straneo 1953, p. 1. 

Leiradira-like Carabidae with mandibles long, acuminate; 
labrum deeply emarginate, 6-setose ; mentum tooth present but 
short, broad ; antennae with 1st segment longer than 2nd + 3rd 
together; posterior-lateral setae of pronotum inside thickened 
margin. 

Genotype : StomimorpJi us violaceus Straneo. 

This subgenus is apparently confined to southern and central 
eastern Queensland. The type locality of violaceus is simply 
Queensland. I have what may be this species from Maleny, on 
the Blackall Range, and additional, smaller species are described 
here from Mt. Jacob and the Eungella Range. 

Leiradira (Stomimorphus) violaceus (Straneo) 

Straneo 1953, p. 1. 

Described as violaceous; length 14.7 by 4.9 mm. The unique 
type is a <5 from ''Australia, Queensland" in the Straneo Collec- 
tion. Two specimens that I collected near Maleny, on the Blackall 
Range, South Queensland, in rain forest, are greenish violaceous 
and larger than the type, but I hesitate to describe them without 
more material to show extent of variation. 

Leiradira (Stomimorphus) jacobi n. sp. 

Form as figured (Fig. 5) ; labrum, mentum tooth, and other 
characters as described for Stomimorphus; head and prothorax 
greenish black, elytra purplish darker dorsally, lower surface 
and appendages dark ; moderately shining, elytra slightly duller 
and with distinct, fine reticulate microsculpture. Head not quite 
% (c. .72) width prothorax. Prothorax subcordate, slightly less 
than y^ wider than long at middle, width/length 1.14-1.19 ; apex 
subtruncate ; base slightly emarginate at middle and rounded 
toward sides ; base and apex not distinctly margined ; sides 
broadly rounded for most of length, slightly sinuate before base ; 
basal angles slightly obtuse, slightly blunted ; disc convex, middle 
line fine, transverse impressions weak, baso-lateral impressions 
linear, moderately impressed, not punctate. Elytra % (or more) 



10 BREVIORA No. 147 

wider than prothorax, slightly narrowed anteriorly ; basal 
margins entire, forming obtuse-right angles at humeri ; latter 
finely bluntly subdentate ; striae moderately impressed, entire, not 
punctate ; dorsal intervals equal, slightly convex ; 10th interval 
present posteriorly ; 3rd intervals 2-punctate in all specimens, 
near middle and posterior '% (but position of punctures slightly 
variable). Secondary sexual characters normal, i.e. $ with an- 
terior tarsi slightly dilated with 3 segments s(iuamulose beloAV ; 
and $ with 1, $ 2 setae each side apex last ventral segment. 
Length 9.8-12.0 ; width 3.3-4.1 mm. 

Holotype S (M.C.Z. Type No. 30,385) and 9 paratypes all 
from Mt. Jacob, c. 45 miles south of Gladstone, South Queens- 
land, c. 2000 ft. altitude, Mar. 1958, taken by the Darlingtons, 
in rain forest. 

Superficially this species is deceptively similar to Lciradira, 
sensu stricto, but the species' technical characters are as in Sfomi- 
morphus. 

Leiradira (Stomimorphus) tenuis n. sp. 

Form as figured (Fig. 6) ; with characters of StoruiinorpJius 
as here given ; dark purplish, lower surfaces and appendages 
dark; shining, elytra with faint fine microreticulation. Head % 
or slightly less width prothorax. rrothorax narrow, wadth/length 
1.06-1.08 (all specimens) ; apex subtruncate or very broadly emar- 
ginate ; base broadly emarginate at middle, slightly rounded 
toward sides ; base and apex not distinctly margined ; sides 
w^eakly arcuate for much of length, slightly sinuate near base ; 
disc with fine middle line, very weak transverse impressions, 
moderate linear baso-lateral impressions, not punctate. Elytra 
c. Yg wider than prothorax ; basal margin entire, forming slightly 
obtuse (nearly right) angles at humeri; latter bluntly subden- 
tate ; striae entire, moderately impressed, not punctate ; intervals 
slightly convex, subequal ; lOtli interval present posteriorly ; 3rd 
interval 2-punctate, near middle and apical %. Length 10.0- 
11.5 ; width 3.3-3.5 mm. 

Holotype 9 (M.C.Z. Type No. 30,386) and 2 9 9 paratypes 
all from the Eungella Range, west of Mackay, Queensland, 2000- 
3000 ft., Nov. 1957, taken by the Darlingtons, in i-ain forest. 

This resembles and may be related to jacohi (above) but the 
prothorax of the present species is narrower and the elytra more 
shining. 



1961 AUSTRALIAN CARABID BEETLES 11 

EEFERENCES 

Castblnau, F. de 

1867. Notes on Australian Coleoptera : pp. 1-139 (Reprinted in Trans. 

R. See. Victoria, 8: 30-38, 95-225, 1868). 
CsiKi, E. 

1929. Junk-Schenkling Coleoptera Catalogus, Pars 104, Carabidae, 

Harpalinae III. 
Darlington, P. J., Jr. 

1953. Australian carabid beetles II. Some new Pterostichini. Psyche, 

60: 90-101. 
1961a. Australian carabid beetles IV. List of localities, 1956-1958. 

Psyche, 67: 111-126. 
1961b. Australian carabid beetles V. Transition of wet forest faunas 

from New Guinea to Tasmania. Psyche, 68: 1-24. 
1961c. Australian carabid beetles VI. The tropical and some subtropical 

species of Pamboriis, Mystropomus, and Xurus. Breviora, No. 

142: 1-13. 
1961d. Australian carabid beetles VII. Trichostenuis, especially the 

tropical species. Psyche (In press). 
Sloane, T. G. 

1902. A re\'ision of the genus Xotonomus . . . Proc. Linn. Soc. New 

South Wales, 27 : 252-324. 
1913. Revisional notes on Australian Carabidae. Part IV. The genus 

Xotonomus. Proc. Linn. Soc. New South Wales, 38 : 404-449. 
Straneo, S. L. 

1953. Nuovi Pterostichini VII. Doriana, Suppl. Mus. Civico Storia 

Nat. "G. Doria," Genoa, 1, no. 36, 1-12. 

TSCHITSCHERINE, T. 

1902. Notes sur les Platysmatini de I'Australie. Horae Soc. Ent. 
Rossicae, 35 : 502-534. 



12 



nREVrORA 



No. 147 









Fig. 1. Leiradira (Metadiia) aurifer n. sp. 

Fig. 2. Leiradira (Metadira) altieola n. sp. 

Fig. 3. Leiradira (Metadira) alternans Darlington 

Fig. 4. Leiradira {sensii stricto) auricollis Castelnau 

Fig. 5. Leiradira (Stoniiinorplius) jacobi n. sp. 

Fig. 6. Leiradira (Stomimorxjhus) tenuis n. sp. 



BREVIORA 

Mmseimi of Comparative Zoology 

Cambridge, I\Iass. December 18, 1961 Number 148 

AUSTRALIAN CARABID BEETLES IX. THE TROPICAL 

NOTONOMIJS 

By p. J. Darlington, Jr. 
Museum of Comparative Zoology, Cambridge, Mass. 

This is the last of four papers on flightless Carabidae of zoo- 
geographic importance from tropical eastern Australia — but 
these papers are part of my general series on Australian carabid 
beetles, which will be continued. Pertinent earlier parts of the 
series, including a locality list (with maps) and a discussion of 
the transition of wet forest carabid faunas from New Guinea to 
Tasmania, are listed with references at the end of the present 
paper. Information on deposition of types will be found at the 
beginning of Part VI (1961c). In this as in other parts of this 
series, proportions stated as simple fractions are based on actual 
measurements made with a ruled ocular in the microscope. 

The present paper concerns the tropical species of the dominant 
eastern Australian pterostichine genus Nofonomus. The genus 
as a whole ranges along the eastern edge of Australia from the 
base of the Cape York Peninsula (somcAvhere between Daintree 
and Cooktown) to Victoria and Tasmania in the south, extending 
west into the eastern edge of South Australia, with one very 
distinct species geographically isolated in southwestern Australia 
and two species described from New Caledonia. I have dealt 
with some of the tropical Australian species before (1953) but 
several additional species are now to be described and a new key 
is necessary. 

The tropical species of Notonomus are all apparently related 
among themselves and form what may be called the doddi sub- 
group of Sloane's (1913) kiugi group of the genus. The doddi 
subgroup is characterized by posterior-lateral setigerous punc- 
tures of pronotum on (not inside) thickened margin at basal 
angles ; elytra fully and strongly striate ; 3rd interval not more 
than (but sometimes less than) 2-punctate (except that rarely 



2 BREVIORA No. 148 

individuals have extra adventitious punctures) and 5th and 7th 
intervals imi^unetate ; 8th interval (10th in saepisfriatus) nar- 
row and convex (least so in masculinus) ; metepisterna short; 
intercoxal declivity of prosternuni flat, without setae ; tarsi not 
striolate above; posterior tarsi Avith 1st segment rather long (but 
not quite so long as next 2 together) and with claw segment 
glabrous below; and S with 1, 9 usually 2 setae each side last 
ventral segment (1 seta each side in ? montelliis, and single 
anal setae may be either missing or duplicated in individuals 
of other species), and S with anterior tarsi dilated with 3 seg- 
ments squamulose below, $ usually with tarsi unmodified (but 
some [not all] 2 ? of doddi, flos, monforum, and masculinus 
have front tarsi with 1st segment squamulose). Other characters 
common to all members of the doddi subgroup, and therefore not 
repeated in the following brief descriptions, are: prothorax with 
apex subtruncate or broadly emarginate with angles not or not 
much advanced beyond arc of emargination, margined at sides 
but not at middle ; base slightly sinuously subtruncate, margined 
only at sides ; side margins rather narrow, only slightly wider 
posteriorly; disc with middle line nearly entire, transverse im- 
pressions weak or obsolete, baso-lateral impressions long, linear, 
moderately impressed, not punctate ; elytra with basal margin c. 
rectangular at finely subdentate humeri; narrow extra (10th, 
or 12th in saepisiriatus) submarginal interval present at least 
posteriorly. Although they agree in these characters (wath ex- 
ceptions noted), the species of the doddi subgroup differ re- 
markably among themselves in some other characters, given in 
the following key. 

Explanation of Map on Page 3 

Known distribution of Notonomus in tropical Queensland. The finely dot- 
ted line is the approximate eastern edge of high land (the Atherton Table- 
land etc.). No. 1 (arrow) indicates occurrence of Notonomus transitus endem- 
ic on the Eungella Range south of the limits of the map ; 2, A", doddi in the 
Herberton-Atherton area, and also (arrow) south of the limits of the map on 
the Kirrama Range, the Mt. Fox plateau, and the Mt. Spec plateau ; 3, montel- 
liis; 4, dimorphicus ; 5, spurgeoni; 6, flos; 7, montorum with subspecies azul 
on Mt. B(ellenden) K(er) ; 8 (arrow) ellioti, endemic on the Elliot Range 
south of Tbwnsville, beyond limits of map; 9, masculinus; 10, saepisiriatus. 
Note the wide gap between the Atherton Tableland species and those on the 
mountains of the Mossman-Daintree area ; the 3 species on the latter moun- 
tains are apparently derived from one secondary ancestor. 



1961 



AUSTRALIAN CARABID BEETLES 




^ '-V, 
#^^ ^e4 



'''^on 




?'></o 






: ^'rns 



ilT^i 



\\e' 



'•'t£*v<.i *"' 



Af. 



V'ffoc 



I . V5 -^/^vL / 



'r^'^t. 



\""Af: 






Oo 



I 



oe- 



8 



^ 



^^ 



Distribution of tropical Notonomus. 



4 BREVIORA No. 148 

So far as I know, all the tropical Notonomus are confined to 
rain forest, and they may all be derived from a single primary 
ancestor that invaded tropical rain forest from the more south- 
ern part of eastern Australia. N. transitus, on the Eungella 
Range in east-central Queensland, may be a (presumably modi- 
fied) derivative of this ancestor. Of the more northern species 
(see map), doddi of the Dividing Range system from Mt. Spec 
to Atherton, and montellus of Mts. Bartle Frere and Bellenden 
Ker, may be related to each other ; montorum, of Mts. Bartle 
Frere and Bellenden Ker, and ellioti, of the Elliot Range south 
of Townsville, may be relicts of one more widely distributed 
secondary ancestor ; masculinvs and saepistriatiis, with allopatric 
ranges on the Atherton Tableland, may be related to each other 
in spite of their structural differences ; and the three species 
north of the Atherton Tableland (dimorphicus, flos, spurgeoni) 
may be interrelated and may now be in process of radiation from 
one secondary ancestor. All this suggests the probable complex- 
ity of evolution of the group in the rain forests of North Queens- 
land, especially on and near the Atherton Tableland. The radia- 
tion of Trichosternus in the same area (Darlington 1961d) may 
have been even more complex. Incidentally, no species of either 
Notonomus or Trichosternus has yet been found on the heavily 
rain-forested mountains that lie on the northern part of the 
Atherton Tableland between the Yungaburra-Lake Barrine road 
(on the south) and the Mareeba-Kuranda road (on the north), 
although these mountains are the home of a striking endemic 
Leiradira {aurifer Darlington 1961e). 

Key to tropical species of Notonomus (doddi suhg7'oup) 

1. Species with all following characters: elytra! striae not widened on disc, 
and intervals normal in number and not much interrupted, and 3rd 
interval with dorsal punctures 2 

— Either striae wide and opaque on disc (not just on declivity), and /or 
intervals more numerous or much interrupted, and/or 3rd interval with- 
out dorsal punctures " 

2. Sides of ventral segments 4-6 extensively punctate (more so than 1st 
ventral) ; (dull black, pronotum virtually smooth) iransiius 

— Ventral segments not punctate or, if punctate, 1st most strongly so . . 3 

3. Pronotum deeply and closely transversely strigulose 4 

— Pronotum only normally (lightly or faintly) transversely strigulose. . .5 

4. Brownish black, usually larger (12.5-16 mm.); 9 with 2 setae each 
side last ventral segment doddi 



1961 AUSTRALIAN CARABID BEETLES 5 

— Bluish or purplish; usually smaller (10.5-13 mm.); 9 with 1 seta each 
side last ventral montellus 

5. Male slender (Fig. 3) ( 2 less so) ; (bicolored) dimorphimis 

— Both sexes stouter 6 

6. Wholly purplish or bluish; length c. 12-17 mm spurgeoni 

— Bicolored, head and pronotum purple or coppery, elytra black; broader 
and larger, length c. 16-20 mm flos 

7. Elytral intervals not much interrupted (3rd without dorsal punctures) 8 

— Elytral intervals much interrupted (3rd with or without dorsal ijunc- 
tures 9 

8. Form broader ; dull montorum 

a. Greenish or purplish montorum, sensu stricto 

b. Bluish with elytral margins bright blue subsp. azul 

— Form narrower; more shining, purplish ellioti 

9. Elytron with 9 intervals, without dorsal punctures masculinus 

— Elytron with 11 intervals plus narrow submarginal one posteriorly (7th 
interval tripled), and with dorsal punctures saepistriatus 

NOTONOMUS TRANSITUS 11. Sp. 

With characters of doddi subgroup; form as figured (Fig. 1), 
rather broad and depressed (in group) ; black, upper surface 
sometimes slightly aeneous, marginal channels of elytra usually 
cupreous or dull greenish ; most of upper surface with fine iso- 
diametric microsculpture. Head % or slightly less width pro- 
thorax, without noticeable unusual characters. Prothorax c. y^ 
or less wider than long ; sides weakly arcuate for most of length, 
slightly sinuate almost at base ; basal angles slightly obtuse, not 
much blunted ; disc smoother than usual, with transverse strigu- 
lation very faint or absent. Elytra slightly less than % wider 
than prothorax ; striae rather fine, not punctate, not widened 
even on declivity; intervals nearly flat on disc, more convex 
laterally, the discal ones subequal ; 3rd interval 2-puiictate, be- 
hind middle and behind apical % . Lower surface nearly impunc- 
tate anteriorly but much of abdomen finely and closely but 
rather irregularly punctate. Secondary sexual characters nor- 
mal; all ? 5 (16) with simple anterior tarsi. Length c. 13-18; 
width 4.5-6.1 mm. 

Holotype S (M.C.Z. Type No. 30,387) and 26 paratypes all 
from the Eungella Range, west of Mackaj^ east-central Queens- 
land, 2000-3000 ft. altitude, Nov. 1957, taken by the Darlingtons, 
in rain forest. 



6 BREVIORA No. 148 

Superficially, transit us is similar to the common, variable A'. 
nitidicollis Cliaiidoir of South Queensland. There may be a real 
relationship between these two species. However, transitus leads 
toward the North Queensland species of the genus in position of 
the posterior-lateral prothoracic setae, on the thickened margin 
(inside the margin in nitidicollis) , and it differs from nitidicollis 
in other specific characters. For example, transitus is broader 
and more depressed, with flatter elytral intervals than nitidicollis, 
and the extensive punctation of the abdomen of transitus is lack- 
ing in nitidicollis. The northern limit of nitidicollis, incidentally, 
is probably Mt. Jacob, about 45 miles south of Gladstone, South 
Queensland. Of the North Queensland species, transitus is prob- 
ably nearest doddi but differs in being broader and more de- 
pressed, with flatter elytral intervals, without special pronotal 
sculpture, but with abdominal punctation. 

NoTONOMUs DODDI Sloane 

The type locality is the Herberton District, Atlicrton Table- 
land, North Queensland (Sloane 1913, p. 439). Specimens that 
I collected on the mountains south and west of Atherton (be- 
tween Atherton and Herberton) are virtually topotypes. The 
species extends south (discontinuously) along the Dividing Range 
system to the Kirrama Range, the Mt. Fox plateau, and the Mt. 
Spec plateau not far north of Townsville. 

The exceptionally close and deep transverse strigulation of 
the pronotum is apparently always present in this species, but 
the other pronotal microsculpture is dimorphic. Most individuals 
have the head very finely and the elytra more coarsely (but still 
finely) isodiametrieally reticulate and the pronotum longitudi- 
nally roughened between the deep transverse strigae. However, 
my series of 11 specimens from the Mt. Spec plateau includes 3 
S S with pronotum without longitudinal roughening, although 
the 4 other S S and all 4 2 9 have the roughening present, as 
do all specimens from other localities. The length of this species 
is c. 12.5-16.5 mm., with the average size decreasing southward. 

NoTONOMUS MONTELLUS n. Sp. 

With characters of doddi subgroup as given above ; form as 
figured (Fig. 2), small, slightly depressed; bluish or purplish 
black; head and elytra with fine isodiametric microsculpture, 
less distinct on pronotum. Head % or slightly less width pro- 
thorax, without obvious unusual characters. Prothorax ]/-, (±) 



1961 AUSTRALIAN CARABID BEETLES 7. 

wider than long ; sides broadly arcuate for most of length, briefly 
sinuate before c. right, scarcely blunted basal angles; main (cen- 
tral) part of disc with many deeply impressed, close-spaced, 
slightlj^ irregular, transverse strigae ; surface of disc otherwise 
not distinctly punctate. FAytra \{^ (±) wider than prothorax; 
striae well impressed, not punctate, not or scarcely widened even 
on declivity ; intervals moderately convex, subequal on disc ; 3rd 
2-punctate near middle and apical ^/4 (punctures slightly vari- 
able in position). Lower surface with a little scattered (variable) 
punctation especially on sides of mesosternum and 1st ventral 
segment. Secondary sexual characters normal except 9 with 
1 seta (not 2) each side last ventral segment. Length c. 10.5-13.0; 
width 3.5-4.4 mm. 

Holotype $ (M.C.Z. Type No. 30,388) and 5 5 <5 4 9 5 para- 
types from Mt. Bartle Frere, west slope, 3000-5000 ft., Dec. 1957 ; 
and 1 additional $ (not a type) from Mt. Bellenden Ker, east 
side, about 4500 ft., Dec. 1957. These two mountains are close 
together at the east side of the Atherton Tableland, south of 
Cairns, North Queensland. Specimens all taken by the Darling- 
tons, in mountain rain forest. 

This small Notonomus is closest in technical characters to N. 
doddi Sloane, of the Dividing Range system, and may be related 
to it, but differs in smaller size, more depressed form, color, and 
presence of only 1 seta each side 9 last ventral segment. The 
pronotal sculpture resembles that of the exceptional doddi from 
Mt. Spec, with deep transverse strigulation but without longitud- 
inal roughening. 

Notonomus dimorphicus n. sp. 

With characters of doddi subgroup as given above; S (Fig. 3) 
exceptionally slender, 9 (Fig. 4) less so; head and prothorax 
aeneous, elytra purplish black ; moderately shining with fine iso- 
diametric mieroseulpture on head and elytra, not distinct on 
pronotum. Head c. % or more width prothorax, without notice- 
able unusual characters. Prothorax slightly longer than wide in 
slender $ $ , slightly wider than long in stouter 9 9 (but slightly 
variable in both sexes) ; sides rather weakly rounded, in $ $ 
nearly straight and converging both before and behind sub- 
median curve but more regularly arcuate in 9 9; sides briefly, 
variably sinuate just before l)ase ; basal angles right or slightly 
obtuse, not much blunted; surface of disc with faint transverse 
strigae, not punctate. Elytra c. % ( ± ) wider than prothorax ; 



8 BREVIORA No. 148 

striae not widened except slightly so (striae 1 and 2) at extreme 
apex, on declivity ; intervals moderately convex, not interrupted, 
subequal on disc; 3rd 2-punctate, near middle and posteriorly 
(middle puncture sometimes duplicated). Lower surface nearly 
impunctate. Secondary sexual characters (other than dimor- 
phism of form) normal. Length c. 12-15; width 3.6-4.6 mm. 

Holotype 6 (M.C.Z. Type No. 30,389) and 7 £6,1 9 2 para- 
types all from Mt. Lewis, near Mossman, North Queensland, 
c. 3000 ft., Dec. 1957, collected by the Darlingtons, in rain forest. 

Males of this species are unique in form at least among the 
tropical species of the genus, and at first I thought they might 
represent transition toward Leiradira, but the mandibles, an- 
tennae, and setae of the inner edge of the maxilla are as usual 
in Notonomus. The 9 9 are close to N. spurgeoni (below) but 
are more slender and bicolored. All specimens that I refer to 
the present species were taken within a comparatively small area 
of continuous mountain forest, and I feel sure they represent a 
single population. A single 9 that I refer to spurgeoni, although 
labeled from Mt. Lewis, may have been taken outside of and 
below the area occupied by dimorphicus. 

Notonomus spurgeoni Darlington 

This is the most northern known species of Notonomus. I 
described it (1953, p. 98) from a series from Mt. Spurgeon, 
about 12 miles northwest of Mt. Lewis in the same mountain 
system. Three specimens from Thornton Peak, northeast of 
Daintree, near 4000 ft., Dec. 1957, taken by the Darlingtons in 
mountain rain forest, seem to be the same species, although the 
color is bluish rather than purplish. A single 9 from Mt. Lewis 
is also apparently referable to this species (see note under 
dimorphicus, above) . 

Notonomus flos n. sp. 

With characters of doddi subgroup as given above ; form as 
figured (Fig. 5), rather large and broad in group; black, head 
and pronotum bright violaceous or cupreous ; moderately shining, 
elytra slightly duller; fine isodiametric microsculpture very dis- 
tinct on elytra, absent or indistinct on head and j^ronotum. Head 
73 (it) width prothorax, without obvious unusual characters. 
Prothorax large, % to ^ wider than long ; sides broadly arcuate 
for almost entire length, usually minutely sinuate at base (the 
sinuation involves hardly more than widening of the marginal 



1961 AUSTRALIAN CARABID BEETLES 9 

bead) ; basal angles obtuse (sometimes nearly right) ; surface 
of disc with weak transverse strigae. Elytra slightly wider than 
prothorax ; striae well impressed, not punctate, not or not much 
widened on disc but inner ones wider and dull on declivity ; 
intervals convex, subequal on disc ; 3rd 2-punctate near middle 
and posteriorly (exact position of punctures variable as usual). 
Lower surface nearly impunctate. Secondary sexual characters 
normal except some 9 9 (2 of 4) with front tarsi with 1st seg- 
ment squamulose below, although not dilated. Length c. 16-20 ; 
width 5.5-6.4 mm. 

Holotype $ (M.C.Z. Type No. 30,390) and 17 paratypes 
(13 S S , 4: ?2) all from Mt. Lewis, southwest of Mossman, 
North Queensland, c. 3000 ft., Dec. 1957, taken by the Darling- 
tons in mountain rain forest. 

The comparatively large, broad form, color, and characters 
given in the key should easily distinguish this species. It was 
found with dimorphicus on Mt. Lewis, and the occurrence of 
spurgconi too there or nearby raises an interesting problem of 
speciation. I think all 3 species have probably been derived 
from one ancestor, but I do not know how the divergence has 
come about. 

NOTONOMUS MONTORUM n. Sp. 

With characters of docldi subgroup as given above ; form as 
figured (Fig. 6), rather large, broad, and depressed in group; 
greenish or sometimes vaguely purplish with elytral margins 
usually greenish and never bright blue ; rather dull, entire upper 
surface with very fine, isodiametric reticulate microsculpture. 
Head c. % width prothorax, without noticeable unusual charac- 
ters. Prothorax Y^ to % wider than long ; sides broadly arcuate 
for much of length, usually broadly but slightly sinuate pos- 
teriorly ; basal angles usually subprominent, c. right, scarcely 
blunted ; surface of disc with weak transverse strigulae. Elytra c. 
y^ or more wider than prothorax ; striae widened and opaque 
especially posteriori}^ ; intervals moderately convex, subequal on 
disc ; 3rd without dorsal punctures. Lower surface with sides of 
ventral segments (especially 1st) finely and irregularly punctate, 
but sterna nearly impunctate. Secondary sexual characters of 
S S and most ? 9 normal, but exceptional 5 9 with 1st segment 
front tarsi with squamae below. Length c. 15.5-19.5; width 5.2- 
6.9 mm. 



IG BREVIORA No. 148 

Holotype S (M.C.Z. Type No. 30,391) and 76 paratypes from 
Mt. Bartle Frere, west slope, 3000-5000 ft., Dec. 1957, and 12 
additional paratypes with the same data except 2000-3500 ft. 
altitude ; all specimens taken by the Darling-tons, in mountain 
rain forest. Mt. Bartle Frere is at the eastern side of the Ather- 
ton Tableland south of Cairns, North Queensland. A weak sub- 
species (below) occurs on neighboring Mt. Bellenden Ker. 

See key for distinguishing characters of this species, and see 
also note under ellioti (below). 

NOTONOMUS MONTORUM AZUL n. SUbsp. 

Structurally similar to typical montorum (above) but more 
shining and different in color, bluish black with elytral margins 
bright blue in all specimens. Of 5 9 5,1 has and 4 have not 
squamae on 1st segment of front tarsi. Length c. 16-19 ; width 
5.3-6.6 mm. 

Holotype $ (M.C.Z. Type No. 30,392) and 11 paratypes all 
from Mt. Bellenden Ker, east side, 3000-4500 ft., Jan. 1958, 
collected by myself in mountain rain forest. Mt. Bellenden Ker 
is about 10 miles north of Mt. Bartle Frere at the eastern edge 
of the Atherton Tableland south of Cairns, North Queensland. 

NOTONOMUS ELLIOTI n. Sp. 

With characters of doddi subgroup as given above ; form as 
figured (Fig. 7), narrower than montorum, moderately de- 
pressed ; purplish black, rather shining ; fine reticulate micro- 
sculpture present but lightly impressed on head and pronotum, 
slightly more distinct on elytra. Head % or slightly less width 
prothorax, without noticeable unusual characters. Prothorax c. 
y^ or slightly less wider than long ; sides broadly, rather weakly 
arcuate for most of length, then slightly sinuate near base ; basal 
angles c. right, well defined ; disc faintly, transversely strigulose 
(as in most species of genus). Elytra c. ^ or less wider than 
prothorax ; striae narrow anteriorly but widening on posterior 
part of disc and especially on declivity; intervals moderately 
convex, subequal on disc (odd ones slightly wider tlian even) ; 
3rd without dorsal punctures. Lower surface almost impunctate. 
Secondary sexual characters normal in <5 ; 9 unknown. Length 
c. 16-18 ; width 5.0-6.0 mm. 

Holotype $ (M.C.Z. Type No. 30,393) and 5 paratypes (all 
S $) all from the Elliot Range (actually from near the summit 



1961 AUSTRALIAN CARABID BEETLES 11 

of "Sharp Elliot"), c. 3000 ft., Mar. 1958, taken by my son and 
myself in mountain rain forest. 

This species may be related to montorum of Mts. Bartle Frere 
and Bellenden Ker. If so, the two mountain species are pre- 
sumably relics of a once more widely distributed stock. As com- 
pared with moniorum, the present species is more slender, with 
elytral striae narrower, especially anteriorly. 

NoTONOMUS MASCULiNUS Darlington 

N. masculinus (Fig. 8) is a relatively large, broad species with 
elytral intervals normal in number but heavily catenulate (much 
interrupted) and without dorsal elytral punctures. It has an 
extensive distribution on the southern part of the Atherton Table- 
land. I described it (1953, p. 99) from Millaa Millaa, and other 
known localities are indicated below. I have noted elsewhere 
(1961c, p. 7) the resemblance between this species and Pamborus 
punctatus, and have suggested that it may be a case of mimetic 
convergence. 

In 5 5 of niascHlinus the front tarsi are moderately dilated, 
with three segments squamulose below. In some $ $ the front 
tarsi are normal (without squamae) but in others the front tarsi, 
though scarcely widened, have the 1st segment biseriately squam- 
ulose. My single 9 from Millaa Millaa has the front tarsi 
squamulose as described, and so does the single 9 from Ilerber- 
ton. A single 5 from between Millaa and Innisfail has the tarsi 
simple. In a series of specimens from mountains (including Mt. 
Fisher) between Millaa Millaa and Ravenshoe, 5 9 9 have tarsi 
simple, 3 squamulose. Three 9 9 from Longlands Gap all have 
simple tarsi. My single specimen from the western foot of Mt. 
Bartle Frere (the northeastern known limit of the species' range) 
is a 5 . 

NoTONOMus SAEPisTRiATUs Sloaue 

triplicate Darlington 1953, p. 100 (new synonymy). 

Sloane (1907, p. 364) described this species (Fig. 9) from 
Atherton, on the Atherton Tableland, North Queensland. I mis- 
interpreted Sloane 's description and re-described the species 
from Lake Barrine and Yungaburra, on the Tableland not far 
from the type locality. In 1957-1958 my wife, son, and I took 
a total of 8 more specimens of the species at Atherton, Yunga- 
burra, and Lake Eacham. At Atherton, we found it only in 
patches of rain forest on the flat part of the Tableland along 




BREVIORA 



No. 148 








1961 



AUSTRALIAN CARABID BEETLES 



13 






Fig. 1. Notonomus transitus n. sp. 

Fig. 2. Xotononius moiitellus n. sp. 

Fig. 3. Notonomus dimorphicus n. sp. (slender $ ). 

Fig. 4. Notonomus dimorphicus n. sp. ( 9 ). 

Fig. 5. Notonomus flos n. sp. 

Fig. 6. Notonomus montorum n. sp. 

Fig. 7. Notonomus ellioti n. sp. 

Fig. 8. Notonomus masculinus Darlington 

Fig. 9. Notonomus saepistriatus Sloane 



14 BREVIORA No. 147 

the road toward Mareeba. The species, therefore, seems to be 
uncommon, confined to a very limited area (a strip about 10 
miles long) of the central Atherton Tableland. It apparently 
does not occur in the mountains west and south of Atherton, 
where it is replaced by doddi, and does not overlap the range of 
masculinus. In other words, these 3 strikingly different species 
seem to be allopatric. My 3 9 9 of saepistriatus all have narrow, 
unclothed front tarsi. 

REFERENCES 

Darlington, P. J., Jr. 

1953. Australian carabid beetles II. Some new Pterostichini. Psyche, 

60: 90-101. 
1961a. Australian carabid beetles IV. List of localities, 1956-1958. 

Psyche, 67: 111-126. 
1961b. Australian carabid beetles V. Transition of wet forest faunas 

from New Guinea to Tasmania. Psyche, 68: 1-24. 
1961c. Australian carabid beetles VI. The tropical and some subtropical 

species of Pamborus, Mysiropomus, and Xuriis. Breviora, No. 

142: 1-13. 
1961d. Australian carabid beetles VII. TricJiosternus, especially the 

tropical species. Psyche (In press). 
1961e. Australian carabid beetles VIII. Leiradira, especially the trop- 
ical species. Breviora, No. 147 : 1-12. 
Sloane, T. G. 

1907. [Notonomus saepistriatus.] Proc. Linn. Soe. New South Wales, 

32: 364-365. 
1913. Revisional notes on Australian Carabidae. Part IV. The genus 

Notonomus. Proc. Linn. Soc. New South Wales, 38: 404-449. 



BREVIORA 



Museum of Cointiparative Zoology 



Cambridge, Mass. Decembek 19, 19()1 Number 149 



A PRELIMINARY STUDY OF THE SILURIAN 

CERATIOCARIDIDS (CRUSTACEA: PHYLLOCARIDA) 

OF LESMAHAGOW, SCOTLAND 

By W. D. Ian Rolfe 
Museum of Comparative Zoology 

AND 

T. P. BURNABY 

University College of North Staffordshire, Keele, Englaud 

The following ten species of Ccratiocnris have been recorded 
by Peach (1901, pp. 450-451) and Jones and Woodward (1888b, 
p. 72) from beds of uppermost Yalentian age in the Lesmaha- 
gow inlier, Lanarkshire, Scotland : 

C. hiornata M'Coy, 1851 
*C. ancjusta Etheridge, AYoodward and Jones, 1886b 
*C laxa Etheridge. Woodward and Jones. 1886b 

C longa Jones and Woodward, 1885 

C. murcliisoni (Agassiz), 1837 
*C. papilio Salter, 1859 
*C. stijgia Salter, 1860 

C. patula Etheridge, Woodward and Jones, 1888 

C. rohusta Salter, 1860 

C. attenuata Etheridge, Woodward and Jones, 1886b 

[=C. tyrannus Salter in Etheridge and New- 
ton, 1878, nom. mid.] 

Species indicated by an asterisk are based on type specimens from 
Lesmahagow. It is unlikely that ten sympatric species of Cerati- 
ocaris would be ecologically compatible (Gause's principle — see 
also Simpson, 1961, p. 74), and examination of the supposedly 
diagnostic characters of these species suggests that the majority 



2 BREVIORA No. 149 

are mere variants. Thus C. inonwfa is stated by Jones and Wood- 
ward (1885, p. 894; 1886a, p. 343; 1888b, p. 37) to ''agree per- 
fectly in form and proportions [of the carapace] with C. papilio 
from Lesmahago, also in ornament, except that the postero-dorsal 
convergence of the striae is not present." Yet elsewhere they 
state that the C. inorvafa from Tjpsmahagow" (BM 59648) is "near 
to C. papilio in form" but that its "proportions are different 
from those of C. papilio" (1885, pp. 460-461; 1886a, p. 346; 
1888b, p. 49). My own measurements (W.D.I.R.) on this speci- 
men differ from those given by Jones and Woodward, but even 
if their values are taken the proportions are found to be identical 
with the figured specimen of C. papilio provided bj^ Jones and 
WoodAvard (1885, pi. 10, fig. 1; 1888b, pi. 12, fig. 1). The speci- 
men is too poorly preserved to distinguish whether the diagnostic 
striae convergence is present or not. 

C. laxa is a juvenile instar of C. papilio, as Jones and Wood- 
ward first thought (1885, p. 396). C. murchisoni is used to de- 
note moderately large styles of several species, in this case of 
C. papilio. In Etheridge, Jones and Woodward's own words "C 
rohusta, being based on some small caudal appendages without 
carapaces, is troublesome and unsatisfactory to deal with. We 
find some equivalent styles and . . . stylets in C. papilio, stygia, 
ocuminaia. etc., but none of these seem small enough for the 
several little sets of trifid appendages, more or less perfect, 
which we have met with. C j'ohnsta takes in some of these ; 
but Oxford Mus. T is relatively broad, and might be termed 
lata [= patula 1888] ; BM 58878 from Muirkirk has very nar- 
row members (angusta).'^ C. longa was first regarded as a 
variety of C. rohusta (1885, p. 464) although some specimens 
"may well belong to C. papilio or C. stygia" (1886b, p. 458). 
This 'variety' was raised to species level later as the style was 
considered too long for either C. papilio or C. stygia (1888b, 
p. 43). C. attenuata differs in having "narrower and smaller" 
abdominal segments and style and stylets shorter than C. gigas 
or C. murchisoni (1886b, p. 456-457). 

Style length is unsuitable as a character for specific differ- 
entiation since only when cameo and intaglio are available is 
it possible to know if the long, needle-like, distal portion of 
the style is preserved. Furthermore, the style is a hollow 
structure and appears "relatively broad" simply due to flat- 
tening during burial. It is therefore suggested that all the 
above mentioned 'species' belong to either C. papilio or C. 



1961 PRELIMINARY STUDY OF CERATIOCARIDIDS 3 

stygia, which form the subject of the present preliminary 
study. An account of the morphology of the species is given 
elsewhere (Rolfe, in press). 

Although Salter (1860, p. 156) stated that C. stygia had its 
"margine ventrali plus minusve angulato." the diagnostic char- 
acter of C. papilio was "the much shorter body — scarcely longer 
(tail included) than the great carapace — [which] easily distin- 
guishes it." Jones and \Yoodward (1885, p. 392; 1886a, p. 341; 
1888b, p. 36) misrepresented Salter's diagnosis of these two spe- 
cies, thus "as mentioned by Salter, one (C. papilio) has the cara- 
pace more oblong than the other (C. stygia).''^ This revised diag- 
nosis led them to suggest (1885, p. 393 ; 1886a, pp. 341-342 ; 1888b, 
p. 36) that only the first of the three figures "termed C. papilio, 
evidently from oversight" by Salter (1860, p. 154) was in fact 
that species, the remaining two being C. stygia. Salter had al- 
ready suggested (I860, p. 156) that there were "at least two 
varieties of carapace in C. stygius itself." Subsequent workers 
have found it difficult to distinguish the two species, and it has 
been suggested that they are identical (Stormer, 1935, p. 294). 
This difficulty was also encountered in the present study and 
hence the museum collections of material available to Salter 
and Jones and Woodward [i.e. their hypodigm) have been re- 
studied to see if the species distinction could be maintained. 

One complicating factor in the use of the older collections is 
the separation of parts from counterparts. Specimens of iden- 
tical dimensions occur in different institutions, and without 
bringing all the material together it is impossible to be certain 
that part and counterpart of the same individual are not treated 
as two individuals. This does not affect the present study, how- 
ever, since it is argualjle that such duplication Avill be distributed 
evenly throughout the sample. 

Two new collections have been examined to determine if, for 
example, C. papilio and C. stygia were allopatric or even succes- 
sional species (= chrono- or palaeospecies). The first of these, 
the A. Ritchie collection in the Grant Institute of Geology, Edin- 
burgh University, came from the ^Jamoytius Beds', half a mile 
up the Logan Water from Logan House (Ritchie, 1960, p. 647). 
The other was collected by J. S. Jennings from a horizon ca. 700 
feet higher in the succession at the locality near Logan Reser- 
voir known as Shank's Castle (Peach and Home, 1899, p. 573). 
This collection is now deposited in the Geology Department, 
Universitv College of North Staffordshire. 



4 BREVIORA No. 149 

The older material is housed in the several museums listed on 
Table 1. Few of the specimens have accurate localities indicated, 
l)ut the majority probably came from the Shank's Castle region. 
Several other localities are known, however, and these have been 
detailed by Etheridge (1873b, p. 49). The authors wish to thank 
the several curators for access to collections, and Professor H. B. 
Whittington for reading the manuscript of this paper. 

Salter's criterion for distinguishing C. papilio from C. stygia 
by the number of segments protruding from the carapace is arti- 
ficial, since it depends solely on the degree to which the thorax 
and abdomen are impacted into, or drawn out from the carapace 
after death or exuviation. Jones and Woodward suggested that 
"C. stygia was rather larger than C. papilio ; its telson was 
larger ; the carapace was markedly distinct by its trapezoidal 
outline, deep ventral region, and mucronate antero-dorsal angle, 
which was not nearly so often lost in fossilization as the front 
angle of C. papilio' '\l^^rj, p. 395; 1886a, p. 344; 1888b, p. 40). 
Style length has already been criticised as a specific character ; 
thus of 202 styles measured only 64 were complete and associated 
with the last abdominal segment. The ratio of style length/last 
abdominal segment length ranges from 1.5 to 2.9, and a scatter 
diagram of this ratio showed normal uncorrelated variation when 
plotted against the last abdominal segment length. This does not 
confirm Jones and Woodward's assertion that "in C. stygia the 
style is usually rather more than twice, and in C. papilio only 
about twice as long as the ultimate segment" (1888!), p. 39). 

The remaining two characters utilised by Jones and Wood- 
ward are carapace size and shape. We here define a 'size factor' 
P such that 

P = log (LH) 
and a 'shape factor' Q such that 

Q = log (L/H) 
where L and H are the overall length and height of the cara- 
pace in millimeters. Figure 1 shows the relationship between the 
size and shape factors for the 128 intact carapaces listed in Table 
I. The distribution of points is the same as would have been ob- 
tained by plotting L against H on double-log paper and rotat- 
ing the diagram through 45°. The object of performing the 
logarithmic transformation and rotation analytically is two-fold : 
to normalise the distribution of the shape factor Q, and to sim- 
plify investigation of the extent to which growth in Ccratiocaris 
is allometric. 



1961 PRELIMINARY STUDY OF CERATIOCARIDIDS 5 

The overall mean value of the shape factor and standard de- 
viation were found to be 

Q = 0.2775 ± 0.05917 
corresponding to a geometric mean value of the ratio L/H of 
1.894. The regression of Q on P was calculated by the first-mo- 
ment method of Wald and Bartlett. The observed P values were 
divided into five groups and the mean values of Q and P were 
found for each group, the number of points in each group being 
20, 20, 48, 20, and 20. The five mean points are shown on Figure 
i- (Qi-Q.-,)- Their coordinates are (2.8109, 0.3097), (3.0779, 
0.2787), (3.2100, 0.2717), (3.3350, 0.2698), and (3.4830, 0.2646). 
The regression is thus non-linear, although as may be seen 

from Figure 1, the four points Q2-Q5 lie on a practically straight 
line. The deviation of Q^ from this line is scarcely significant, 
and the line itself does not depart significantly from a direction 
parallel to the P axis, if the first group of points is ignored. 

If we interpret the assemblage as a growth series, it follows 
that growth is isometric except in the range of size for which P 
is less than 3.07 (LH less than 1175 mm.^). In this latter size 
range, growth is not isometric, though the degree of allometry is 
small and is barely significant. 

Changes in the allometric constant during growth are well 
known in crustaceans, and commonly separate two distinct in- 
stars or mark a more critical ecdysis such as the prepuberty moult 
(Teissier, 1960; Simpson, Roe, and Lewontin, 1960, pp. 412-415). 

If we discard the 20 smallest specimens and test the marginal 
distribution of the remaining 108 Q values b}^ plotting on prob- 
ability paper, we find that the distribution is an almost perfect 
unimodal normal curve (a straight line plot). There is thus no 
evidence to justify splitting up the assemblage on the basis of 
carapace shape. If, moreover, we regard the assemblage as a 
single homogeneous sample of a single species, we can readily 
calculate confidence limits for Q values, to test whether a given 
specimen may reasonably be regarded as a member of the as- 
semblage. (It will be as well to withhold judgment in the case 
of very small specimens, but otherwise there should be no need to 
worry about allometry.) 

The first writer (W.D.I.R.) has been unable to trace either of 
the syntypes of C. papilio (Salter, 1859, p. 262) and hence speci- 
men GSM 7479, the original of Salter's 1860, p. 154, fig. 1, is 
here treated as the neotype of the species. Jones and Wood- 
ward accepted this specimen as a genuine C. papilio (= ]\I.P.G. 
X 1/15 in 1885, p. 393; 1886a, p. 342; 1888b, p. 36). Calcula- 



BREVIORA 



No. 149 



TABLE I 
Length (L) and height (H) of 123 museum specimens of ceratiocaridid 
carapaces from the Lesiiuliagov/ area, in millimeters 



Specimen number 


L 


H 


Specimen number 


L 


H 


Specimen number 


L 


H 


GSM-165-2 


25 


11 


Ke-09 123 gb 


53 


26 


AR 59-153 


60 


33 


BIVI-24161 


26 


12 


BM-24164 


52 


27 


AR 59-144 


60 


34 


BM-24163 


28 
35 


13 
15 


H -A 1908 
GSM -7479 


52 
57 


27 
25 








BM -59648 


AR 59-106 


57 


36 


BM-16483 


33 


16 


GSE-6647-I 


55 


26 


K-6-2 


57 


36 


H-no number 


28 


19 


BM- 16501 


52 


27 


E -190^30/10 


71 


29 


Ke-09 123 fu 


34 


16 


K-5-3 


49 


30 


E-190;/30/l 


69 


30 


Ke-09 123 eg 


42 


15 


Ke-R28 


49 


30 


K-1 


56 


37 


Ke-Alrdrie Coll. 


34 


19 


BM-24157 


57 


26 


BM-16513 


56 


37 


J-LW8 R2 


34 


19 


Ke-09 123 dn 


50 


30 


Ke-R33 


62 


34 


Ke-09 123 dy 


36 


18 


GSM -87338 


50 


30 


AR 59-M 


68 


31 


Ke-09 123 cb 


36 


19 


AR 59-148 


54 


28 


K-7 


65 


33 


Ke-R22 


37 


20 


E -190^30/9 


51 


30 


AR 59-P 


67 


32 


GSE-6650 


39 


19 


K-6-1 


48 


32 


BM-41894 


64 


34 


Ke-09 123 cd 2 


41 


19 


GSE -6647-2 


57 


27 


J-LW8 B 


59 


37 


BIVI-45161 


39 


20 


H-no number 


52 


30 


H-A 1900-1 


58 


38 


J -27 


44 


18 


Ke-09 123 en 


60 


26 


Ke-R4 


69 


32 


H-SUrl< Coll. 


45 


21 


K-5-2 


51 


31 


BM-16482-I 


67 


33 


BM-24150 


43 


23 


J-LW9 E 


59 


27 


BM-164S2-2 


62 


36 


AR 59-152 


43 


24 


Ke-Rl 


57 


28 


H-A 1900- 2 


66 


34 








GSM-272 


54 


30 
29 


Ke-09 123 ez 
BM-24153 


74 
67 


33 
37 


Ke-09 123 ed 


48 


22 


H -A 1903 


56 


GSM-X 1/13 


41 


26 


BM-no number 


56 


29 


AR 59-147 


66 


38 


Ke-R29 


43 


25 


AR 59-154 


58 


28 








J-LW8 Q2 


45 


25 


K-5-1 


52 


32 


BM-24155 


72 


"i? 


GSM -256 


45 


25 


H -A 1904-1 


4? 


24 


Ke-R9 


78 


33 


AR 59-146 


49 


23 


AR 59-Q 


60 


28 


AR 59-150 


63 


41 


BM-24151 


47 


24 


BM-41896 


60 


28 


AR 59-151 


70 


38 


Ke-R23 


44 


26 


Ke-R2 


56 


30 


EM -58669 


83 


33 


BM-41895 


45 


26 


E-190y30/'7 


."■-3 


32 


Ke-R25 


66 


42 


BM-24160 


45 


26 


H-A190B-3 


53 


32 


BM-24149 


73 


39 


H -A 1901 


47 


26 


BM-14517 


57 


30 


E-1 391/9^6 


80 


36 


Ke-09 123 ex 


47 


26 


Ke-09 123 dz 


58 


30 


Ke-09 123 ea 


68 


43 


BM-24158 


50 


25 


GSM-no number 


55 


32 


Ke-R24 


70 


43 


AR 59-149 


47 


27 


H-A190e-2 


55 


32 


BM-41898 


83 


37 


Ke-R8 


53 


24 


H-T.WIsfi Coll. 


59 


31 


K-3 


75 


41 


Ke-Alrdrle Coll. 


48 


27 


GSM-X 1/22 


60 


30 


BM -16479 


82 


42 


E-190?/30/ll 


54 


25 


Ke-09 123 el 


62 


30 


GSM-X 1/21 


85 


46 


J-LW8-L 


54 


25 


Ke-R3 


62 


30 


E-1865/11/17 


74 


53 


H-Macnair Coll. 


54 


25 


H-A 1904-2 


55 


34 


Ke-09 123 ce 


84 


47 


H -A 1902 


52 


26 


K-8 


64 


30 


BM-45154 
GSIVI-X 1/19 


90 


47 


H-66 


52 


26 


Ke-09 123 ey 


60 


32 


105 


50 


AR 59-108 


52 


26 


E- 190^30/4 


52 


37 


Ke-09 123 dr 


96 


61 


BM-16495 


53 


26 








H-Macnair Coll. 


100 


70 



Ranked according to Increasing size factor P = log (LH); the five groups delineated by 
the heavy lines are those referred to in the text. 

Repositories: AR 59 - Ritchie Collection, Grant Institute of Geology, Edinburgh 
University; BM - British Museum (Natural History), London; E - Royal Scottish 
Museum, Edinburgh; GSE - H. M. Geological Survey, Edinburgh; GSM - H. M. 
Geological Survey Museum, London; H - Hunterlan Museum, Glasgow University; 
J - Jennings Collection, Geology Department, University College of North Staf- 
fordshire; K - Kilmarnock Public Museum; Ke - Kelvlngrove Museum, Glasgow. 



1961 



PRELIMINARY STUDY OP CERATIOCARIDIDS 







10 

X • 



^ 



1- 



•l^ • . • 

• • • \ , » 



'o^ r 



CD 

(0 
0^ 
10 

5: . 

OQ 



10 

d 



6 



h 
d 



d 



o 



R- 



Co 

10 

f1 



^ 

h 



h 

CM 



h' 



o 
h 

(^ 
c\i 

00 

f\j 

N 
(\j 

Co 

10 

c\i 



(H/lJ^o/ = o ^0±OVJ 3=1 VHS 





•-a 

1- 


OQ 

s 
o 

•s 


■1^ 


>-> 





'3 

a; 

a 

CO 


10 

oo 

00 

T— 1 


a; 

ft 

ft 






55. 


fl 


s 


TS 


c3 














t» 


^ 


<V 


" 


be 


'O 


-^ 






e 

■2 


S 

T3 


o 


J3 


1 
01 

5D 





CS 


cS 

g 

CS 












00 









fe 






g 

^ 


C 








<t-i 

13 


^ 


CO 






o 

^M 




o 

5^ 


CO 




CS 


3 






O 


o 


bi 


c; 




0) 


CS 


13 


, 




oo 


O 




^ 


•^ 


73 


CS 


0) 






05 




c 


-4-> 




fc 


•a 






"3 
V 


02 


"u 

-^ 

g 






t-3 












ft 

CS 




o 


-^3 


(m* 


.r-( 


3 


T— 1 

bi 




13 

3 






^tH 


o 


oo 





«c 


3 








o 


o 


• r-l 


a 
o 





I— 1 


be 


0) 

3 



bO 






a 


CO 

• r-l 




<I> 

> 


'El 

00 
00 


^ -2 


a; 
3 




s^ 


o 


o3 


tl 




1—1 




Q 


4> 


II 


a; 
a 

5 


so 

5^ 


c: 


c3 

s 
la 


be 


CS 





4. 


2 

00 

3 




.^ 




-^- 


b£ 


1 






c 


f-H 




00 
cS 


g 

s 




UJ 


1 


00 

o 
o 


o 


> 


0^ 
-♦- 

"ci 
CO 

■2 


C3 
CO 




^-1 


a 

'-^ 

3 

■73 


1-^ 


CO 




CO 

o 

be 


£ 

O 
t-, 




^ 

a. 1-5 

^ 


00 


3 
'zj 


> 

CS 






^ 


%4 


«> 


«^ 


^ 

^ 


13 

3 


ft 


ft 




OJ 
X 




CO 


J3 






3 

be 


Kl 


cS 
«-i 

CS 




5*H 

o 


00 








p. 

+^ 



V5 




c« 

!2; 


00 

to 




"be 




s 


CJ 


& 




s 


£- 


s 
3 

1 

00 


irt 


*r^ 




1h 

be 

-3 


CS 


CO 




-73 

EC 







<4-l 







5^ 
• i-i 

Id 

s 


'S 


'a 


be 
be 

3 


«<-i 



^ 




3 




C^ 



















W 


CO 


o 


-t^ 


1 


s 


+-' 


K 




cc 


O 


'3 

!3 


.s 


01 


g 
ft 

CO 

"3 




be 


CO 

Q 




T-l 




33 


t3 




.S 


«3 


X 




bi 


c5 


SO 

o 

6 




CO 


to 


0" 

rH 


-4-' 






C» 


_g 





S 


3 


"ft 


'E 



8 BREVIORA No. 149 

tion shows that this specimen lies within the 95 percent confidence 
limits for group 3. The 'primary' types of C. stygia are unknown 
but one of the "good specimens of C. stygia" figured by Jones 
and Woodward (1885, pi. 10, fig. 2; 1888b, pi. 12, fig. 2) BM 
45154, is similarly within the 95 percent confidence limits for 
group 5. C. stygia is thus not separable from C. papilio on 
statistical or indeed any other present evidence, and the former 
junior synonym should be suppressed. The morphospecies C. 
papilio and C. stygia form the one biospecies (or transient spe- 
cies) C papilio. It is worth noting that the specimen of C pap- 
ilio figured by Jones and Woodward (1885, pi. 10, fig. 1; 1888b, 
pi. 12, fig. 1) BM 58669 is 'abnormal,' as can be seen from its 
position on Figure 1. 

No significant differences in P or Q can be detected in the 
stratigraphically separated material of the Ritchie and Jennings 
collections. 

There is a notable lack of clustering in terms of the size factor 
P corresponding to the mean size of successive moult stages or 
instars. Attempts to divide the Ritchie or Jennings specimens 
alone into instars were also unsuccessful. Intra-instar variation 
may obscure the limits of successive instars if sufficiently great, 
but not all crustaceans obey Brooks' Law (Needham, 1950, pp. 
10-11), and C. papilio may be another exception. At least part 
of this difficulty is due to sampling and preservation. Thus small 
individuals are only rarely collected (see Fig. 1) and all large 
individuals found have been incomplete. Relatively gigantic 
specimens up to two feet in total length occur both in the Les- 
mahagow and liagsliaw Hills inliers, but they can only be recon- 
structed from fragments and hence do not appear on Figure 1. 
A further complicating feature of the few large specimens avail- 
able is their distinctive dendritic carapace ornament. It can be 
.ijrgued, however, that this ornament is characteristic of adult 
aistars (Rolfe, in press). Such giant individuals must have had 
younger growth stages coincident in size with the specimens of 
Figure 1, and no such dendritic ornament has been observed in 
that size range. It seems preferable to extend the name C. papilio 
to include these large individuals, at the risk of 'lumping,' until 
better sampling has been made. 



1061 PRELIMINARY STUDY OF CERATIOCARIDIDS 9 

SUMMARY 

Of ten species of Ceratiocaris recorded from the Lesmahagow 
inlier, only C. pnpilio and C. stygia are sufficiently well founded 
to demand preliminary investigation. Both Salter's diagnosis of 
these two species and Jones and Woodward's sul)sequent de- 
finitions are artificial. Analysis of 128 carapaces in museum col- 
lections shows that C. papilio is indistinguishable from G. stygia, 
and the latter should be suppressed as a junior synonym. 

Carapaces of C. papilio show isometric growth except in the 
smallest individuals. The material cannot be resolved into a 
series of distinct instars. 



EEFEEEXCES 

Only those references not listed by Van Rtraelen and Sehmitz, 1934, are 
given here. 
Xeedham, a. E. 

1950. Growth and regeneration rates in relation to age in the Crus- 
tacea with special reference to the isopod, Asellxfi aqvai\c\ts 
(Linn.). Jour. Gerontology, 5: 5-16. 
Ritchie, A. 

1960. A new interpretation of Janioufiiis lenroorTi White. Xature, 
188: 647-649. 

EOLFE, W. D. I. 

(In press) Grosser morphology of the Scottish Silurian phylloearid crus- 
tacean, Ceratiocaris inipiUo Salter. Jour. Paleont. 
Simpson, G. G. 

1961. Principles of animal taxonomy. Columltia Univ. Press, Xew 
York. 

SnrpsoN, G. G., A. Eoe and E. C. Lewontin 

1960. Quantitative Zoology. Eevised ed. Hareourt, Brace and Co., 
Xew York. 

ST0RMER, L. 

1935. Dictyocaris, Salter, a large crustacean from the Upper Silurian 
and Downtonian. Xorsk geol. tiddskr., 15: 267-298. 
Teissier, G. 

1960. Eelative growth. In The Physiology of Crustacea, Waterman, 
T. H. ed., vol. 1, pp. 537-560. Academic Press, XeAv York and 
London. 
Van Straelen, V., and G. Schmitz 

1934. Crustacea Phyllocarida (=:Archaeostraca). Fossilium Catalogus; 
pars 64, Berlin. 



BREVIORA 

Museum of Compsirative Zoology 



Cambridge, Mass. January 5, 1962 Number 15U 

THE GENUS BETHYLUS IN NORTH AMERICA 
(HYMENOPTERA: BETHYLIDAE) 

By Howard E. Evans 



Bethylid wasps are predominantly tropical and subtropical in 
distribution, with only a few species of diverse genera penetrat- 
ing temperate regions and virtually none entering arctic or sub- 
arctic regions. The sole exception to this statement, illogicallv. 
is the type genus of the family, Beiliylus Latreille. This rather 
liighly evolved genus is circumpolar in distribution. In North 
America, specimens have been taken close to the Arctic Circle, 
but none have been taken south of New York, Illinois, Colorado, 
and central California. In the Old World there are several 
species of northerly distribution and several others from the 
Mediterranean region. The geiuis is not known from the South- 
ern Hemisphere. 

In North America, four species have been described in the 
genus; these are: casianeus Kieffer, amocnus Fonts, hrachypit lus 
\Vhittaker. and flauicoynis Whittaker. A fifth species, decipiens 
Provancher, has recently been transferred to the genus by 
Krombein (1958, U.S. Dept. Agri., Monogr. no. 2, first suppl., 
p. 98). Examination of types in the U.S. National Museum re- 
veals that two additional species, Arysepyris californicus Brid- 
well and Perisetnus oregonensis Ashmead, properly belong in the 
genus. Since the latter species is the tj^pe of the genus Digoniozus 
Kieffer (1905, In Andre, Spec. Hymen. Eur. Alger., v. 9, p. 245), 
this name can be added to the synonymy of Bethylus. 

One of these seven names can be removed from further con- 
sideration here. I have recently had an opportunity to study the 
type and only known specimen of Bethylus castaneus Kieffer 
(1907, Berlin. Ent. Zeitschr., 51: 295). The wings of this speci- 
men are in poor condition, but enough remains to be sure that 
this species belongs not to Bethylus but to the related genus 
Goniozus (new combination). 



2 BREVIORA No. 150 

Thus there are six specific names aA'ailable for the North 
American Bethylns, three of them ncAvly assigned to the genus. 
The question naturally arises as to how much synonymy is in- 
volved and how many species, in fact, are there? The present 
paper is an attempt to answer that question. 

ANALYSIS OF THE PROBLEM 

Specimens of this genus are not common in collections, but 
by borrowing material from many sources I was able to obtain 
about 80 specimens. One's first impression, on scanning this 
material, is the remarkable uniformity of the specimens in size, 
color, and structure. The only notable color differences are sex- 
ual : the males have yellow mandibles and wholly yellow an- 
tennae, while the females have dark mandibles and the antennae 
more or less infuscated apically. There are no noticeable differ- 
ences in the structure of the mandibles and clypeus, in the sculp- 
turing of the head or thorax, or in the male genitalia. 

There is, however, one character which varies strikingly, and 
that is wing length. The wings vary all the way from small pads 
scarcely larger than the tegulae to wings of normal size. This 
is not unusual in the genus, as several brachypterous species 
have been described and the European fuscicornis is known to 
exhibit much variation in wing length. In the case of the North 
American Bethylus, it was of interest to know whether wing 
length varied in a continuous spectrum or whether there were 
certain wing-length types which might represent different species. 
Following 0. W. Richards (1939, Trans. R. Ent. Soc. London, 
89: 185-344) in his revision of the British species, I first deter- 
mined the relative wing length of each specimen by dividing the 
length of the fore wing by the length of the hind tibia (which is 
much easier to measure accurately than total body length). I then 
plotted the number of individuals exhibiting a given relative 
wing length (Fig. 1). The males fell into two distinct groups. 
Those of the first group (Type A) might be termed subapterous, 
since the wings are exceedingly small, barely surpassing the 
anterior margin of the propodeum. Males of the second group 
(Type B) might be termed micropterous, since the wings are 
still very small, extending about to the beginning of the pro- 
podeal clecliAdty. When one plots the females on this same scale 
he obtains a somewhat different picture (lower half of Fig. 1). 
The subapterous forms (Type A) tend to have slightly longer 
wings, the micropterous forms (Type B) slightly shorter wings, 



19()2 



NOUTH AMERICAN [-.irrn VM'S 



CD 








-ia 



oS 


f-l M^ 


1^ (> 


' 0^ 

cpoo 






. OS 

in in 



1 o» 



o 



lO'^lO<sJ,-,OoCO(^-o>Ol-'^'^'-' 








^ 








^ 












Ji 0^ 








o^o\ 








ijhj 




if 




 o> 








O<0 




.— 




k5 Kj 




•^ 




^?^ 




'_ 




10 K) 




V- 




1 Ov 




=^ 




sO-fi 




c 




i6 «) 

> 




-*— 

r— i 




oS 




-*^ 




-> i-i 




Op 




f^^ 




0) 




•-I ,~^ 




ex 












^ZO 




*r" 




— t ■-) 




if 


• 


in S 




o 


P 






o 


•—I .—1 












+J 




■_ 


r^ 


n 










^^ 


u. 


^ 


'^ 


-« r-t 


_J 


^ 


+- 




"x,^^ 


_j 


c 


' O 








« Jo 




c; 


X 


r-t r-i 


u. 


>■ 


O) 










r-l —1 


_1 




s 



O o 



<i^S^ 


« 


1—* 




-(-^ 


JTi 


1 ov 


s 


S 


«aj 


rr 


--" 


< (S 


^ 


r 


t- P 


^~^ 






tfi 


^ 


4^ 


■^ 

3 


h£ 














is; 


>■ 


r* 




"^ 


-♦— 








I <^ 




t:! 


■<■ ■«- 




fj 




=4-4 


• rH 




o 


p^ 


1 OS 




^ 



«i s 



<— » ^^ "H »^ »-1 ""^ 



5 ^ 






o o 

d o 
Z E 



be 






4 BREVIORA No. 1 30 

SO tliat the two curves overlap slightly. Furthermore, there is a 
class of individuals with wings of moderate length, reaching about 
to the posterior margin of the first abdominal tergite, which 
might be termed brachypterous (Type C), as well as a few in- 
dividuals with wings of normal length (macropterous. Type D). 

The fact that wing length varies discontinuously suggests the 
possibility that several species may be involved, each exhibiting 
a different winff length. Presumablv there would be four such 
species, with males of the two less common ones still to be dis- 
covered. However, in the absence of other characters one cannot 
rule out the possibility of polymorphism. 

I once again turned to Richards' study of the British species, 
and discovered that the most useful character for separating the 
three forms occurring in Britain is the ratio between the distance 
separating the hind ocelli and the distance separating the hind 
ocelli from the occiput. I determined this ratio for all speci- 
mens availalile to me but obtained a unimodal curve, with the 
mean 1.7, the range of variation from 1.3 to 2.2 (close to the 
range for the European juscicornis) . Thus these measurements 
failed to support the po.ssibility of more than one species. How- 
ever, in the course of making the measurements I found myself 
able to recognize "long-headed" and "short -headed" individ- 
uals. The difference was slight, but sufficient to induce me to 
measure the heads and determine the width/length ratio. In the 
case of the males I again obtained two separate curves (Fig. 2. 
top). For the females I obtained a bimodal curve (Fig. 2, bot- 
tom). It was at once apparent that all the subapterous individ- 
uals (Type A) were "long-headed" (left hand curves in Fig. 2), 
all the micropterous individuals (Type B) "short -headed" 
(right hand curves in Fig. 2). The brachypterous females (Type 
C) were all "short -headed", while the fully winged females 
(Type D) were of both types. 

Here was a suggestion that two species might be involved, 
with both species being polymorphic for wing length in the 
female sex. Upon sorting the specimens into two lots represent- 
ing probable species, several other differences previously over- 
looked or discounted in importance were discovered. The most 
important of these involved the sculpture of the propodeum, the 
shape of the male subgenital plate (Fig. 3), and the wing vena- 
tion of the few available fully winged females. Thus I am now 
convinced that two polymorphic species are involved. The name 
(innit lilts Fonts is applicable to the "long-headed" species, while 



1962 



NORTH AMERICAN BETHYLUS 



dccipiens (Provancher) is the earliest name for the "short- 
headed" species. The two species are widely sympatric east of 
the Rockies, but amoenus is not known to occur west of the 
Rockies. The characters separating the two species are summar- 
ized below, as are their synonymy and distribution. 



N 0. of 
Males 



IZ 

11 

10 

9 

a 

7 

5 
4 
3 
Z 

1 




.76 79 80 81 .62 &h .54 .85 .£6 .6 7 .es .89 

wh/lh 




No. of 
Females 




.78 19 .60 62 &Z .63 64 65 .66 .87 .68 .39 .90 .91 .92 

wh/lh 

Fig. 2. Number.s of individuals (ordinate) exhibiting given relative head 
lengths (width of head [WH] divided by length of head [LH]), males at 
top, females at bottom. 



The only good series of decipicns from one locality is the 
series of 16 females and 4 males from Chilliwack, British Colum- 
bia, on which Whittaker based his descriptions of hraehypterus 
and ftavicornis. As discussed further below, I have studied or 
oi)tained the necessary information on this entire series. Twelve 
of the females are micropterous (Type B), three are brachyp- 
terous (Type C), and one is maeropterous (Type D). This 
12 :3 :1 ratio, obtained in a series from one locality, is approached 
rather closely by the ratio for the species throughout its range. 



6 BREVIORA No. 150 

which is 41 :9 :4. Unfortunately, only one reared series of this 
species is available. That is a series of five females and three 
males in the U.S. National Museum reared from Vicia angusti- 
folia at the Lummi Indian Reservation, Washington. All indi- 
viduals in this series are micropterous. 

Unfortunately, no good series of am.oenus is available; the 
longest series consists of three females and a male taken on 
different dates at Bar Harbor, Maine. All of these individuals 
are subapterous, and in fact only one fully winged individual 
of this species is known. This is a female taken by 0. W. Richards 
on the window of an automobile at Buffalo, N. Y., 19 Sept. 1928. 
The ratio of subaptery : braehyptery : macropter^' in the females 
of this species is 23:0:1. It is, of course, entirely possible that 
brachypterous individuals of amoenus may some day be discov- 
ered. It is also quite possible that polymorphism for wing length 
may occur in the male sex. At present only seven males of 
ainocnus are known, only thirteen of dccipiens. Clearly any hy- 
potheses on the genetics of polymorphism in these wasps will 
have to await the day when much more material has accumulated 
in museums. At present it appears that only the females are 
polymorphic for wing length and that the polymorphism arises 
from a very simple genetic mechanism. 

The Nearctic dccipiciiH is undoubtedly closely related to the 
Palaearctic fuscicornis and may well be derived from it. Not 
only are the oeellar measurements similar, as noted earlier, but 
the sculpturing of the propodeum is similar and the male sub- 
genital plate virtually identical. However, there is no doubt in my 
mind that they are specifically distinct. The antennae of fusci- 
cornis are shorter and the scape is black at the base, yellowish 
apically (the scape is wholly yellowish-brown in both Nearctic 
species). Richards has found that the frequency distribution 
of relative wing length in fuscicorni.'i is more or less trimodal or 
((uadrimodal, but less distinctly so than in dccipiens and with a 
much larger proportion of longer-winged individuals. Further- 
more, the males of fuscicornis are typically macropterous rather 
than micropterous as in dccipiens. 

TAXONOMIC TREATMENT 

Key to North American Species of Bethylus 

Propodeum with a median polished ridge, remainder of disc con- 
trastingly alutaceous; head rather short (width/length ratio 



1962 NORTH AMERICAN BETHYLUS 7 

.85-. 91 ill female, .89-. 92 in male) ; wings of micropterous in- 
dividuals reaching at least nearly to middle of propodeal disc 
(relative wing length .57-.91) ; fully winged individuals with 
radial vein curved upward sharply apically, vein arising 
from basal vein barely indicated ; fore tibiae clear yellow ; 
male subgenital plate strongly emarginate, but the side-pieces 
relatively broad and blunt (Fig. 3, a) . .decipiens (Provancher) 
Propodeum somewhat convex dorsally but without a median 
ridge which is set off from the remainder of the disc ; head 
slightly longer (width/length ratio .79-. 84 in female, .85-.86 in 
male) ; wings of most individuals extremely small, reaching 
barely beyond anterior margin of propodeum (relative wing 
lenglli .26-. 54) ; fully winged individuals with radial vein not 
curved upvrard sharply at apex, vein arising from basal vein 
nearly as long as transverse median vein; fore tibiae of female 
usually at least weakly sutfused with brownish ; male subgenital 
plate with a strong emargination, the side-pieces reduced to 
slender, acuminate processes (Fig. 3, b) amoenus Fonts 





Fig. 3. Subgenital plates of (a) Bethylus decipiens and (h) BetJiylus 
amoenus. 

Bethylus decipiens (Provancher) 

Gonatopus decipiens Provancher, 1887, Add. Corr. Faune Eiit. 

Canada, Hj-men., p. 179 [Type: 9, Cap liouge, Quebec (Que. 

Prov. Mus., yellow label no. 1382) ] . — Muesebeek and Walkley, 

1951, U. S. Dept. Agri. Monogr. 2, p. 1038. 
rtrL^icniHs oregonerisis Ashmead, 1893, Bull. V. S. Nat. Mus., 

45: 70 [Type: 5, Portland, Oregon (U. S. Nat. Mus. no. 

40422)]. New synonymy. 



8 BREVIORA No. 150 

Digoniozus ore(joncnsis Kieffer, 1905, Spec. Hymen. Eur. Alger., 

9: 245 [Made type of new genus Digoniozus]. — Muesebeek 

and Walkley, 1951, U. S. Dept. Agri. Monogr. 2, p. 732. 
Arysepyris californicns Bridwell, 1919, Proc. Hawaiian Ent. Soc, 

4: 34 [Type: 9, Parkside, San Francisco Co., Calif. (U. S. 

Nat. Mus. no. 64124)]. New synonymy. 
Bethyltis bracliypterus Whittaker, 1929, Trans. R. Ent. Soc. Lon- 
don, 76 : 385 [Type : 2 (not 6 as stated), Chilliwack, Br. Col. 

(British Museum) ]. — Muesebeck and Walkley, 1951, U. S. 

Dept. Agri. Monogr. 2, p. 732. New synonymy. 
Bethylns fiavicornis Whittaker, 1929, Trans. R. Ent. Soc. London, 

76: 386 [Type: 6% Chilliwack, Br. Col. (British Museum)]. 

— Muesebeck and Walkley, 1951, U. S. Dept. Agri. Monogr. 2, 

p. 732. New synonymy. 
Glenosema calif ornicus Muesebeck and Walkley, 1951, U. S. Dept. 

Agri. Monogr. 2, p. 727. 
Bcthylus decipiens Krombein, 1958, U. S. Dept. Agri. Monogr. 

2, First Suppl., p. 98. 

Remarks on types. — Provancher's decipiens was transferred 
to Bethylns by Krombein upon his examination of the type. Dr. 
Krombein has kindly placed his notes at my disposal, and they 
leave no doubt that Provancher's name applies to this species. 
The propodeum is alutaceous but with a median polished ridge, 
and the wings extend almost to the posterior slope of the propod- 
eum. The type is in good condition. 

The type of Ashmead's oregonensis is also in good condition 
and is a fully winged female of this species. The type of Brid- 
welLs calif ornicus is unfortunately in poor condition, the head, 
abdomen, and legs all being missing. However, the wings and 
propodeum are typical of the micropterous form of decipiens. 
A topotypic female in the collection of the California Academy 
of Sciences is very similar to the type and is in good condition. 

Whittaker 's two names require special discussion. The types 
and most of the paratypes are in the British Museum and I 
have not seen them. However, Mr. G. E. J. Nixon has been good 
enough to examine tliese specimens and send me the critical in- 
formation on them. I have studied one paratype of hrachypterus 
in the collection of Cornell University as well as two of this 
species and one of fiavicornis in the collection of Roliert M. Fonts 
of Laredo, Texas. The characters AVhittaker used for separating 
the two species are color characters which happen to be those 
which separate the sexes, and it happens that all the specimens 



1962 NORTH AMERICAN BETHYLUS 9 

of braclnjpfo'iis are females and all of Jlavicornis are males — 
Whittaker's statements to the contrary notwithstanding. The 
entire series is from Chilliwack, British Columbia; the type of 
hrachypterus is a fully winged female, that of flavicornis a 
mieropterous male. As indicated earlier, three of the paratypes 
of brack ijpte)-us are lirachyjiterous, the remaining twelve mierop- 
terous. 

Specimens cxantincd. — 40 9 9, 10 £ i. ALASKA: 1 9, Fair- 
banks, 25 June ]948 [I^SXMJ ; 1 9, Circle, 2 July 1958 (C. 
Lindroth) [CXC] ; 1 9, Xenana, 17 June 195:3 (K. I. Sailer) 
[USXM] ; 1 9, 2 <3 6, Mile 1476, Alaska Highway (C. Lindroth) 
[CXC]. BRITISH COLUMBIA: 1 9, Mile 290, Alaska High- 
way, 19 June 1951 (W. Mason) [CXC] ; 1 9, Smithers, 12 June 
1958 (C. Lindroth) [CXC]; 1 9, Cranbrook, 12 May 1922 (C. 
Garrett) [CXC]; 1 9, Victoria, 28 Aug. 1923 (K. F. Auden) 
[CXC] ; 3 9 9,1 S, Chilliwack, May-June, Sept. 1927 (0. Whit- 
taker) [CU, Coll. R. M. Fonts]; 1 S, Galiano, 2 Aug. 1929 
[Coll. Fonts]; 1 9, Kaslo (A. X. Caudell) [USXM]; 2 9 9, 
Terrace [MCZ]. WASHIXGTOX : 1 9, Olympia [USXM]; 

5 9 9,4 S S, Red River Rd., Lummi Ind. Res., 1 Aug. 1944 
(Vicia angusHfolia, W. W. Baker) [USXM]. OREGOX : 1 9, 
Portland [USXM] ; 1 9, Forest Grove, 1 Apr. 1919 (A. C. Bur- 
rill) [USXM] ; 1 9, Ashland Loop, Siskiyou Mts., Jackson Co., 

6 Aug. 1950 (Malkin & Thatcher) [CAS]. CALIFORXIA : 1 9, 
Land's End, San Francisco, 11 July 1922 (F. X. Williams) 
[CAS] ; 1 9, Parkside, San Francisco Co., 8 Sept. 1910 (J. C. 
Bridwell) [USXTVE]. UTAH : 1 9, Logan [MCZ]. COLORADO: 
2 9 9, 1 $, Fort Collins, June, Sept. 1895 (C. F. Baker) 
[USXM]. IDAHO: 1 9, Coeur d^Alene (H. J. Rust) [USXM]. 
ALBERTA: 1 9, Edmonton, June 1917 [USXM]; 1 $. Elk- 
water Lake, 19 July 1956 (0. Peck) [CXC]. OXTARIO : 1 9, 
Sudbury, 1892 [CXC]. QUEBEC: 1 9 , Anticosti Island, 9 Sept. 
[MCZ]. XEW BRUXSWICK: 1 9, Penobsquis, Dec. 1927 (C. 
A. Frost) [MCZ]. XOVA SCOTIA: 1 9, Portapique, 23 July 
1929 (C. A. Frost) [MCZ]. MAIXE: 4 9 9, Bar Harbor, July- 
Oct. (A. E. Brower) [USXM]. XEW YORK: 1 9, Grand 
Island, 11 Oct 1922 [USXM]; 1 9, Xorth Fairhaven, 1 Sept. 
1918 [CU].i 



1 The following abbreviations have been employed for the museums involved : 
CAS, California Academy of Sciences. San Francisco ; CXC. Canadian National 
Collections, Ottawa : CU, Cornell University. Ithaca : MCZ, Museum of Compara- 
tive Zoology, Cambridge; USNM, U. S. Nati()nal Museum, Washingttm. 



10 



BREVIORA 



No. 150 




Map showing distribution of North American Bethyhis. Solid triangles : 
Bethylus amoenus, subapterous form ; hollow triangle : macropterous form 
of amoenus. Solid circles: B. decipicns, mieropterous form; half -solid circles: 
braehj'pterous form of decipiens ; hollow circles: macropterous form of 
decipiens. 



Bethylus amoenus Fonts 

Bethylus amoenus Fonts, 1928, Proc. Ent. Soc. Wash., 80: 127 
[Type: 9, Slaterville-Caroline, Tompkins Co., N.Y., 14 Jniie 
1904 (Cornell Univ. no. 934) ]. — Mnesebeck and Walkley, 
1951, U. S. Dept. Agri. Monogr. 2, p. 732. 

Remarks on types. — The type is in good condition. I have 
also studied a male allotype, bearing the same data, in the collec- 
tion of Robert M. Fonts. 

Specimens exammed. — 24. 5 5, 7 $$. NORTHWEST TER- 
RITORIES: 2 9 9, Norman Wells, 3-13 July 1949 (W. Mason) 
[CNC]. ALBERTA: 1 9 , Aspen Beach, 23 Aug. 1944 (0. Peck) 
[CNC]; 1 9, Elkwater Lake, 19 July 1956 (0. Peck) [CNC]. 
SASKATCHEWAN: 1 S, White Fox, 10 July 1944 (0. Peck) 
[CNC] ; 1 c^, Holdfast, June 1946 (W. A. Nelson) [CNC] ; 1 9, 



1902 XORTH AMERICAN BETHYLUS 11 

Assiniboia, June 1955 (J. R. Vockeroth) [CNC] ; 1 i, Saska- 
toon, 15 Sept. 1924 (K. M. King) [CNC]. MINNESOTA: 1 ?, 
Eaglesnest, 26 An^. 1959 (W. V. Baldnf) [TTSNM]. WISCON- 
SIN: 1 9, Cranmoor, 20 May 1910 (C. AV. Hooker) [USNM]. 
ILLINOIS: 1 2, Palos Park, 17 March 1933 (Frison & Mohr) 
[111. Nat. Hist. Survey]. MICHIGAN: 1 9, AVexford Co., 4 
July 1952 (R. R. Dreisbach) [Coll. Dreisbach] ; 1 9, Presque 
Isle Co., 28 July 1952 (P. B. Kannowski) [Coll. Dreisbach] ; 1 9, 
Midland Co., 20 June 1945 (R. R. Dreisbach) [Coll. Dreisbach]. 
ONTARIO: 1 9, Jordan, 25 Sept. 1916 (W. A. Ross) [CNC] : 
1 9, Rondeau Park, Kent Co., 28 June 1936 (G. Steyskal) [Coll. 
Dreisbach] ; 1 9, Prince Edward Co., 10 July 1950 (J. F. Brim- 
ley) [CNC] ; 1 9, Belleville, 2 Oct. 1956 (J. M. Smith) [CNC]. 
NEW YORK: 1 9, Buffalo, 19 Aug. 1928 (0. AV. Richards) 
[MCZ]; 2 9 9, Ithaca, 28 May, 23 June (Babiy, Evans) [CU, 
MCZ] ; 1 9,1 $, Slaterville-Caroline, 14 June 1904 [CI^ Coll. R. 
M. Fonts] ; 1 <5, Caroline-Harford, Tompkins Co., 15 June 1904 
[CU]; 1 S, Gannett Hill, 30 Aug. 1925, 2000 feet [CU]. 
MAINE: 3 9 9, 1 5 , Bar Harbor, July, Sept., Oct. (A. E. 
Brower) [USNM] ; 1 9, Southwest Harbor, 6 Sept. 1922 [CU]. 
NOA'A SCOTIA: 1 9, Portapique, 22 July 1929 (C. A. Frost) 
[MCZ]. 

BIOLOGY OF THE GENUS 

The only specimen of this genus which I have collected was 
taken walking over the ground in a small sand pit. Several speci- 
mens in collections are labeled as having been taken sweeping, 
one while "sweeping Carex," another "while beating for ants." 
Several specimens of both species were taken by A. E. Brower 
at Bar Harbor, Maine, on "Great Heath," one of the female 
decipiens "on flowers of Ilex verticillata.'" A female amoenus 
from Palos Park, Illinois, is labeled "in wet peat sample," 
while a series of decipiens from the Lummi Indian Reservation, 
AYashington, is labeled "Rd Vicia angusti folia." Apparently 
these insects occur in a variety of situations. B. amoenus has been 
collected in every month from March to October, decipiens from 
April to October and also in December. 

Two specimens of amoenus bear host data. One is the female 
listed above from Cranmoor, AVisconsin, which is indicated as a 
probable parasite of Eudemis vocciniana. This name is now re- 
garded as a synonym of Rhopohata naevana (Hbn.), an oleu- 
threutid moth known as the black-headed tireworm. The other 



12 BREVIORA No. 150 

specimen is the female listed from Belleville, Ontario, which is 
labeled as a parasite of Brachypterohis pulicarius L. This is a 
nitidulid beetle introduced from Europe to the United States 
about 1918. 

If the latter record is correct, it is the only kno^\^l instance 
of a Bcthylus attacking a beetle. The European cephalofcs 
Forster and fuscicornis (Jurine) attack various caterpillars, 
chiefly Microlepidoptera but occasionally Noctuidae. The fe- 
male wasps sting and malaxate their rather large prey, then drag 
it to a place of concealment, such as a hollow^ stem. Several eggs 
are laid on the prey and several larvae develop on a single host. 
Further details regarding the biology of these two species may 
be found in the papers of Richards (1932, Trans. Ent. Soc. So. 
England, 8 : 35-40 ; 1939, Trans. R. Ent. Soc. London, 89 : 185- 
344). 



BREVIORA 

Miiseiuim of Compsirative Zoology 

Cambridge, Mass. January 12, 1962 Xr. miser 151 

A NEW PHYLLOCARID CRUSTACEAN FROM THE 
UPPER DEVONIAN OF OHIO 

By W. D. Ian Rolfe 

INTRODUCTION 

Wlieii c'lirating- the collections of non-trilobite arthropods in 
the Musenni of Comparative Zoology, the writer recently found 
a fossil crustacean which had been sent to Professor P. E. Ray- 
mond for determination. The specimen was received from the 
Cleveland Museum of Natural History through Dr. D. H. Dunkle 
in January 1944 ; the late Professor Raymond published no 
description of the specimen and left no manuscript notes with 
it. Recent collecting in the same area by Mr. G. Lammers of 
the Cleveland Museum failed to find further specimens, although 
a fragment of a second specimen from a different locality was 
recently donated to the Museum of Comparative Zoology by Mr. 
R. Pritschan of Cleveland. This specimen will be referred to as 
the MCZ specimen to distinguish it from the original Cleveland 
^Museum specimen. 

During the preparation of this description Mr. Lammers called 
the writer's attention to the fact that H. K. Brooks of the Uni- 
versity of Florida had collected and studied the echinocaridids 
of this region. In correspondence, Mr. Brooks informed the 
writer that he had photographed this specimen some years ago, 
but was kind enough to allow the writer to submit this account 
for publication. 

The writer is indebted to Dr. G. A. Cooper, Professor A. La 
Rocciue and Professor F. G. Stehli for searching through the 
collections at the U. S. National Museum, the Ohio State Univer- 
sity and Western Reserve University, Cleveland, for additional 
material, and to 'Mv. W. E. Scheele. Director of the Cleveland 
Museum of Natural History, for allowing the specimen to be 
retained for description. Professor H. B. Whittington kindly 
took the photograph for Plate 1 and offered helpful criticism of 
the manuscript. 



2 BREVIORA No. 151 

SYSTEMATIC DESCRIPTION 

Subclass MALACOSTRACA Latreille, 1806 
Siiperorder PHYLLOCARIDA Packard, 1879 

Order ARCHAEOSTRACA Claiis, 1888 

Suborder RHINOCARINA Clarke in Zittel-Eastman, 1900 

Family OHIOCARIDIDAE fani. nov. 

Diagnosis. Carapace valves deep, with anterodorsal-medio- 
ventral fold and broad median dorsal plate. Rostral plate and 
number of thoracic and abdominal segments unknown. 

Remarks. The family Rhinocarididae comprises five genera 
which form a compact group characterised by elongate carapace 
valves and a narrow median dorsal })late. The present genus is 
so distinct from the previously described Rhinocarina as to Avar- 
rant the erection of a second family. 

Genus OhiOCAKIS gen. nov. 

Type species. Ohiocaris wycoffi sp. nov. 

Diagnosis. As for the family. 

Ohiocaris wycoffi sp. nov. 
Plate 1 ; Figure 1 

Descripfion. The Cleveland specimen is exposed with the dorsal 
surface of the carapace uppermost in one half of a concretion. 
It is preserved as a very thin film of golden brown fcuticular 
material, but this has been destroyed over much of the .specimen 
so that an internal mould is revealed. The concretion has been 
split apart so that the two carapace valves and median dorsal 
plate are separated by matrix from two complete abdominal seg- 
ments, a fragment of a third, and stylet fragments. The where- 
abouts of the counterpart of the concretion are unknown. 

As may be seen from Figure 1 and Plate 1, the carapace valves 
are deep ; a well-defined fold, semicircular in cross section, runs 
from the anterodorsal region of each carapace valve, immedi- 
ately posterior of the strong carapace horn, and dies out ventrad 
of the centre of the valve. The ventral or free margin of each 
valve is bordered by a narrow reflexed rim except in the mid- 
ventral region, where it continues as a marginal ridge inside the 
ventral margin. 



1962 



NEW DEVONIAN CRUSTACEAN FROM OHIO 



The broad median dorsal plate is separated anteriorly from 
the cephalic region of the carapace by a shallow transverse 
groove. The grooves laterally separating the plate from the main 
area of the carapace valves are narrower and deeper than the 
anterior groove, and are confluent posteriorly with the carapace 



mandibles 



anterior carapace 
horn 



oblique fold 



left valve 
of carapace 




pereiopods 



p osier o median 
groove 



marginal 
ridge 



m e dian dors a I 
plate 



last abdominal 
segment (? = 7th) 




ventral process 
of style head 



stylet 



Figure 1. Outline drawing of Ohiocaris u-ycoffi gen. et sp. nov. showing 
structures visible on Plate 1. Cleveland Museum of Natural History 33241. 
X 1.1. 



rim and a groove marking the posterior edge of the median dorsal 
plate. These grooves doubtless mark the position of marginal 
rims analogous in structure to those at the ventral edge of the 
carapace valves. The plate bears a faint posteromedian groove 
which extends anteriorly for 1.4 mm. 

The carapace valves and median dorsal plate are crazed by 



4 BREVIORA No. 151 

veinlets of a brown mineral ( ?collophane), whereas the abdomi- 
nal segments and the matrix are unaffected, indicating differen- 
tial chemical desiccation. The fragments of test preserved are 
smooth and free from ornament and the few wrinkles present 
are clearly secondary. 

The abdominal segments are inverted relative to the carapace 
and thus the style and stylets are exposed ventral side upper- 
most. The last segment ( ?7th) is 1.3 times the length of the 
preceding segment and has a concave posteroventral margin. 
After the photograph for Plate 1 was taken, the fragmentary 
st^'le and stylets were broken from the matrix. The dorsal head 
of the style thus exposed was found to be of the echinocaridid 
type illustrated by Echinocaris snblevis Whitfield, 1880, figure 6 
(=Hall and Clarke, 1888, pi. 29, fig. 13). Only the bases of 
style and stylets are preserved but the former is triangular in 
cross section and much shorter tlian the stylets. As in all the 
known archaeostracans, the head of tlie style embraces the proxi- 
mal portions of the stylets laterally and ventrally. A small 
denticle, 0.2 mm. long by 0.4 mm. broad, projects posterolaterally 
from the right lateral edge of the ventral style process or plat- 
form. 

The cuticle of the style head, stylets and abdominal segments 
lacks ornament. 

Only the inflated coxal parts of the mandibles are j)reserved. 
and excavation has failed to reveal the toothed gnathal lobes 
which were probably broken off at burial. The mandibles have 
been impressed tli rough the anterodorsal region of the carapace 
valves, and the plane of section show^s the left mandible to have 
had a wall thickness of 0.7 mm. 

At least four recurved ridges on the anteroventral region of 
the carapace fairly certainly mark the position of simple pereio- 
pods. As they are only seen as impressions through the thin cuti- 
cle of the carapace no detail of their structure can be discerned. 

The MOZ specimen is a fragment sliowing the median dorsal 
plate only. 

Dimensions, in millimeters 

Oleveland ^luseuni of Natural History 33241 
Maximum length of undistorted right carapace valve 34.0 
Maximum height of undistorted right carapace 
valve, to right edge of groove bordering median 
dorsal plate 25.0 



1962 NEW DEVONIAN CRUSTACEAN FROM OHIO 5 

Length of median dorsal i)late, along mid-line 21.0 
Maximum width of median dorsal plate, at a point 

7 ram. posterior from transverse groove 7.3 

Length of penultimate abdominal segment 4.3 

Length of last abdominal segment 5.5 

Maximum dorsal width of style head 4.5 

Length of style head to base of style 3.3 

Width of style at base 1.1 

Cross-sectional diameter of stylet 1.6 

Museum of Comparative Zoology 6556 

Maxinnim length of median dorsal plate ca. 21 

Maximum width of median dorsal plate 7.2 

Holotype. Cleveland Museum of Natural History 33241. Col- 
lected by Dale Wycoff, 25th May, 1934, from the Chagrin Shale, 
Upper Devonian. Locality — shore of Lake Erie at mouth of 
Porter Creek, 12 miles west of Cleveland, Cuyahoga County, 
Ohio. 

Other material. Museum of Comparative Zoology 6556. Col- 
lected by Raymond Pritschan and donated to the Museum via 
G. Lammers, July 31, 1961 ; found as float from Chagrin Shale. 
Locality — Painesville, 25 miles northeast of Cleveland, Lake 
County, Ohio ( ? Whitfleld's 1880, p. 37, Leroy locality). 

Remarks. The spread out carapace valves of the Cleveland 
specimen recall the condition in Dithyrocaris, which possibly 
lived with the valves in this attitude. The marginal rim fore- 
shadows the well-developed submarginal wall and doublure struc- 
ture of Dithyrocaris, and is similar to the condition in the ceratio- 
caridids Caryocaris curvilata (Gurley) and CaUizoe hohemica 
Barrande. The anterodorsal-medioventral fold is more anteriorly 
situated than that of Pephricaris horripilata Clarke, whereas the 
"Schragrippe" and "Schnabelfurche" of Silesicaris nasuta 
Ciiirich (1929, p. 29) run closer to the ventral margin. As men- 
tioned above, the style resembles that in Echinocaris rather than 
that in any rhinocaridid, but the long smooth last abdominal 
segment is different from that of every species of that genus. 

The posteromedian groove on the median dorsal plate may 
prove to be of phylogenetic significance as a vestige of the non- 
rhinocaridid simple dorsal hinge, and homologous with the me- 
dian fold of other members of the Khinocarina. Ohiocaris shows 
the greatest development of the median dorsal plate and suggests 
a derivation from the Middle and early LTpper Devonian rhino- 
caridids. Thus the plate width/carapace width ratio is 0.146 in 



6 BREVIORA No. 151 

Ohiucaris, Ijut only O.UI)(i in the specimen of Elyinocaris slliqua 
figured by Beeeher (1902, pi. 19, fig. 8; Yale Peabody Museum 
22410). Hall and Clarke's reconstruction of Mcsothyra oceani 
(1888, pi. 32, fig. 1) is inaccurate in showing the hypothetical 
median dorsal plate too broad. Measurement of the specimen 
upon which this reconstruction was based (New York State 
Museum 4576) shows that the ratio is 0.073, not 0.150 as figured. 
Two other specimens of M. oceani (NYSM 4577, 4581) give com- 
parable ratios of 0.093 and 0.082, and in the Rhinocaris columhina 
figured by Clarke (1893. fig. 4. NYSM 4786) the ratio is only 
0.074. 

The concretion in which the Cleveland specimen occurs has 
the characteristic orange colour of oxidised Chagrin material 
described by Cushing, Leverett and Van Horn (1931, p. 34). 
Fossils do not seem to have been recorded previously from the 
Chagrin west of Cleveland (Cushing ef al., 1931, p. 35). The 
MCZ specimen forms part of a small collection comprising many 
of the same sj^eeies of brachiopods and pelecypods as those listed 
by Cushing et al. (1931, p. 35), and in addition the crustaceans 
Echinocaris multinodosa Whitfield, E. suMevis Whitf. and 
PalacopahKnuni mwlxryiii AYliitf.. 



EEFERENCES 

Beecher, C. E. 

1902. Revision of tlie Phyllocarida from tlie Chemung and Waverly 
groups of Pennsylvania. Quart. J. Geol. Soc. London, vol. 58, 
pp. 441-449. 
Clarke, J. M. 

189.3. On the structure of the carapace in the Devonian crustacean 
Bhinocaris ; and the relation of the genus to Mesothyra and the 
Phyllocarida. Amer. Nat., vol. 27, pp. 793-801. 
Cushing, H. P., Leverett, F., and F. R. Van Horn 

1931. Geology and mineral resources of the Cleveland district, Ohio. 
Bull. U. S. Geol. Surv., 818. 
GURICH, G. 

1929. Silesicaris von Leipe und die Phyllokariden iiberhaupt. Mitt. 
Min.-Geol. Staatsinst. Hamburg, Heft 11, pp. 21-90. 
Hall, J. and J. M. Clarke 

1888. Trilobites and other Crustacea. Natural History of New York, 
Palaeontology, vol. 7. 
Whitfield, R. P. 

1880. Notice of new forms of fossil crustaceans from the Upper Devon- 
ian rocks of Ohio, with descriptions of new genera and species. 
Anier. J. Sci., ser. 3, vol. 19, pp. 33-42. 



1962 



NEW DEVONIAN CRUSTACEAN FROM OHIO 






r 









*%.. 




.■•\ 



# 



r 



^r 



%•• 



Plate 1. Ohiocaris icycoffi gen. et sp. nov., Cleveland Museum of Natural 
History 33241, x 1.7. 



BREVIORA 



useiuinn of Connparative Zoology 



Cambridge, Mass. January 15, 1962 Xtmber 152 

NEW AUSTRALIAN DACETINE AXTS OF THE GENERA 
MES08TRUMA BROWN AND CODIOMYRMEX WHEELER 

(Hymeuoptera — Formicidae) 

By Robert W. Taylor 

Biological Laboratories, Harvard University 

The two new species described below are of considerable in- 
terest as members of tlie Australian ant fauna. The rare genus 
Mcsostruma, to which Mesostruma hrowni n. sp. is added, in- 
cludes two previously described species which have recently been 
revised by W. L. Brown, Jr. (1952). Mesostruma is known only 
from eastern Australia and occupies an important phylogenetic 
position in the subtribe Epopostrunuti, being almost exactly in- 
termediate in character between the major Australian genera 
Epopostruma Forel and Colohostruma Wheeler. Codiomyrmex 
flagellatus n. sp. is the second member of its genus to be de- 
scribed from northern Queensland, and as such is the second 
representative of the important short-mandibulate stock of the 
subtribe Strumigeniti to be recorded from Australia. The de- 
ficiency of this element in the Australian fauna is of considerable 
zoogeographic interest. Its historical absence or scarcity on the 
continent has perhaps been important in allowing adaptive radia- 
tion of the short-mandibulate Epopostrumiti of the genus Colo- 
hostruma (Brown, 1952, 1959; Brown and Wilson, 1959). 

INIesostruma browni new species 
(Figs. 1-5) 

Holotype worker. Synthetic aggregate length (TL)^ 3.6; head 
length (HL) 0.78; head width (HW) 0.71; mandibular exten- 
sion (ML) 0.37; Weber's length of alitrunk (WL) 0.8-t; cephalic 

lAU measurements stated in the present paper are given in miUimeters with 
indices in units. The conventions foUowed in measurements are those estabiisntu 
for the Dacetiui by Brown (195;}a, lySSb). An ocular scale with units of O.OKj.s 
mm. was used for measuring, with correction to the nearest unit, the maximum 
error being estimated as ±0.01 mm. 



BREVIORA 



No. 152 



index (CI) 91 ; mandibulo-cephalic index (MI) 47. Correspond- 
ing precisely with the generic characteristics cited by Brown 
(1948, 1952) for head shape, mandibular and antennal structure, 
l^etiole and postpetiole form, and body sculpturing. Head shape 
as in Figure 1, similar to large M. laevigata workers, but clypeus 
and anterior part of head somewhat more transverse. Eyes small, 
strongly convex as in M. turneri. Humeral angles rounded ; 
dorsum of alitrunk in profile strongly and evenly convex. Pro- 
podeal lamellae as in Figure 2, more extensive than in either 
previously described species. Petiolar node as in Figure 2, similar 
to that of M. turneri, but more massive and less acute above in 
side view. 




Mesostruma irowni new species, Figs. 1 and 2, worker (Holotype). Fig. 1. 
Full-face view of head. Fig. 2, AUtrunk, nodes and base of gaster in 
side view. Figs. 3-5, male (Allotype). Fig. 3. Full-face view of head. Fig. 
4. Alitrunk, nodes and base of gaster in side view. Fig. 5. Forewing. 
Pilosity and sculpture omitted from Figs. 2-4. Scale line: 1 mm. 



1962 NEW AUSTRALIAN DACETINE ANTS 3 

Sculpture of head, alitrunk and petiole consisting of large 
circular umbilicate foveae, more widely spaced than in the 
other described species, rarely separated by distances less than 
their maximum diameter ; the surfaces between them smooth and 
strongly shining. Dorsum of alitrunk less densely sculptured 
than head, with a median longitudinal area almost devoid of 
foveae. Posterior parts of sides of alitrunk opaque, the foveae 
indistinct and mixed with coarse punctures. Foveae of petiolar 
node smaller than those of head and alitrunk ; those of postpetiole 
indistinct, postpetiolar dorsum finely and irregularly sculptured 
and feebly shining. Dorsum of first gastric segment smooth and 
strongly shining, with no trace of longitudinal striae. 

Color rich golden brown ; petiole, base and apex of gaster, legs, 
mandibles and antennae lighter. 

Type locality. Two miles east of Berry, New South Wales (B. 
B. Lowery). 

Worker variation. Thirty-two paranidotype ivorkers, collected 
with the holotype, have the following dimensions : TL 3.2-3.9 ; HL 
0.72-0.83 (mean 0.77) ; HW 0.66-0.74 (mean 0.71) ; ML 0.32-0.38 
(mean 0.35) ; WL 0.77-0.92; CI 89-96; MI 42-49. No significant 
structural variation is indicated in the series. The workers are 
monomorphic with no bimodality in the frequency distributions 
of the dimensions listed, and no perceptible allometric differentia- 
tion between the head and mandibular dimensions, within the 
sample. 

The measurements and indices of ten paratype specimens from 
Riverview College, Sydney (B. B. Lowery), fall within the above 
ranges and have almost identical means, but an eleventh specimen 
of the same series is much smaller: TL 3.1 ; HL 0.69 ; HW 0.62 ; 
ML 0.34; WL 0.74; CI 90; MI 49. The HL of this individual is 
2.80 standard deviation units smaller than the overall mean 
HL of all 45 specimens examined (76.7 ± SD 2.75), its HW is 3.0 
standard deviation units smaller than the overall mean HW 
(70.4 ± SD 2.80). The specimen is thus of extremely small size 
when compared with workers from mature colonies, and is 
perhaps an old nanitic which had survived into the mature colony 
with which it was collected. 

A single paratype worker from Barrington Tops, New^ South 
Wales (T. E. Woodward), has the following dimensions: TL 3.2; 
HL 0.72 ; HW 0.66 ; ML 0.32 ; WL 0.77 ; CI 92 : MI 44. 

Paratype queens. The first series of dimensions are those of 
an alate from Burns Bay, Sydney (B. B. Lowery) ; the second 



4 BREVIORA No. 152 

series those of a dealate from Pymble, New South Wales (C. Mer- 
covich). TL 4.0, 4.3; IIL 0.80, 0.86; HW 0.76, 0.82; ML 0.37, 
0.40; WL 1.07, 1.16; CI 95, 95; MI 46, 46. Differing from the 
workers in the usual characters of full sexuality: larger size, 
presence of ocelli and wings, and unreduced structure of ali- 
trunk. Coloration as in worker, the ocellar area dark brown. 
Wings clear with pale yellow veins, venational pattern similar 
to male. The same diagnostic features as those of the worker 
serve to distinguish queens from those of M. turneri. 

Allotype male. TL 3.0; HL 0.54; HAV including compound 
eyes 0.63 ; WL 0.88 ; forewing length ca. 2.2 mm. Head as in 
Figure 3. Compound eyes large, elliptical, strongly convex, their 
longest diameters about 0.28 mm. Mandibles slender, acute, 
probably not opposable. Antennae robust, thirteen segmented. 
Maxillary and labial palpi well developed; palpal formula ap- 
parently maxillary 5, labial 3, as in the worker (the mouthparts 
have not been dissected from the unique specimen). Body profile 
as in Figure 4. Mesonotum with w^ell developed notauli, their 
posterior portion, forming the stem of the "Y," indistinct. 
Propodeal lamellae smaller than in the female castes. Petiole 
sub-clavate, the node low, sloping gradually back from the 
anterior peduncle ; antero-ventral tooth obsolete. Lateral edges 
of post-petiole lacking aliform appendages, but each with a 
distinct, low, obtuse, longitudinal carina. Gaster broad, some- 
what flattened basally ; the basal edges of its first segment feebly 
carinate longitudinally, as in many epopostrumite workers. Geni- 
talia exposed, enfolded by the parameres. The latter similar to 
those of Oreciognathus (Brown, 1953b) : broad in dorsal view, 
with convex lateral outlines and strongly concave inner faces, 
the apices rounded with their tips turned inwards and opposed 
mesally. Apex of sub-genital plate acute. Cerci short and stout. 
The penis valves and volsellae have not been dissected from the 
specimen. 

Forewing venation (Fig. 5) of the Solenopsis type, as in 
Orectognathus, the apical elements (Rsf 5, Mf 4 and Cu-A) 
feebly developed, and the radial cell open. Hindwing narrow, 
with a broad posterior fringe of microtrichiae, and four well 
developed subapical hamuli; venation as in Orectognathus. 

Head and most of alitrunk coarsely and closely punctate. 
Punctures of the pronotal dorsum and prescutellar area similar 
to the foveae of the workers, but almost contiguous. Sides of 
alitrunk Avith quite extensive shining areas, especially on the 



1962 NEW AUSTRALIAN DACETINE ANTS 5 

median parts of the larger sclerites. Petiole and postpetiole with 
eoarse punctures, those on the latter shallow and irregular. First 
gastric tergite semi-opaque, with very irregular and shallow 
large, flat, piligerous punctures. 

Color blackish-brown ; antennae, mandibles, under-mouthparts, 
and legs yellowish-brown. Wings clear, their veins pale yellow. 

Described from a unique specimen collected with the holotype 
and its associated paranidotype worker series. 

Material examined. Northeastern New South Wales : 2 miles 
east of Berry (type locality) December 28, 1959, holotype and 
32 paratype workers, allotype male (B. B. Lowery). Burns Bay, 
Sydney, February 2, 1959, ex leaf litter, a single alate queen 
(paratype) (B. B. Lowery). Riverview College, Burns Bay, Syd- 
ney, April 19, 1959, eleven paratype workers ( B. B. Lowery). 
Pymble, Sydney, March 18, 1956, a single dealate queen (para- 
type) (C. Mercovich). Barrington Tops, ex leaf mould (Berlese 
funnel sample), a single paratj'pe worker (T. E. Woodward). 

The holotype, with paratypes, has been returned to Father 
Lowery for eventual deposition in the Commonwealth Scientific 
and Industrial Research Organization collection at Canberra; 
the allotype, with paratypes, is in the Museum of Comparative 
Zoology, Harvard University ; the remaining paratypes are in 
the Queensland Museum. 

Biology. The following information regarding the biology and 
ecology of 31. hrowni has been provided by Father Lowery. 

The type locality is about two miles inland from Seven Mile 
Beach, in low hill country behind scrubby alluvial coastal flats. 
The collection was made in a grassed clearing in a heavily tim- 
bered area, with Eucalyptus and turpentine growing in black 
non-sandy soil. 

The Riverview College locality overlooks Burns Bay, Lance 
River Cove, Sydney. The colony taken there was found nesting 
in damp yellow sand beneath a cover of moss and a little grass. 
The site was in a clearing in low scrub about 15 to 20 feet high, 
with Eucalyptus corymhosa, Grevillea, Lantana and Leptosper- 
nuon. The soil near the nest contained a few small termite gal- 
leries and a large nest of the locally dominant ant Acropyga 
australis. The alate paratype queen was collected nearer the 
Burns Bay foreshore, wandering on leaf litter in warm sunshine, 
during the late afternoon. 

Father Lowery has collected three further colonies of M. 
lyrowni within 200 meters of the type nest site. One of these 



6 BREVIORA No. 152 

colonies was nesting about four inches below the surface of 
coarse black sandy soil, under a small rock, but probably not 
in direct contact with it. A maze of termite galleries was located 
immediately beneath the rock, and permeated the surrounding 
soil. 

The new species is dedicated to Dr. W. L. Brown, Jr. of 
Cornell University, a leading authority in ant taxonomy who has 
worked particularly with the Dacetiui and has devoted much 
study to the Australian ants in general. 

The addition of M. hrowni to Mesostruma requires no change 
in the basic concept of the genus, as formulated by Brown 
(1952). Indeed Mesostruma retains its appearance as a compact 
and distinctive genus, with its species abundantly distinct from 
each other. 

Considering the characters of the worker and queen, M. hrowni 
seems to be most closely related to M. turneri Forel, which it 
resembles in the structure of the alitrunk, propodeal lamellae 
and petiole. It has a proportionately narrower head, however 
(see Brown and AYilson 1959, fig. 7), and lacks the longitudinal 
striation of the basal gastric segment seen in turneri. The ab- 
sence of humeral denticles, and the general form of the alitrunk, 
propodeal lamellae and petiole distinguish M. hrowni from M. 
laevigata Brown. The new species differs from both the previ- 
ously described species in its less dense sculpturing and overall 
glossiness, and its more extensive propodeal lamellae and more 
massive petiolar node. The three known species of Mesostruma 
may be separated by the following key (based on the workers). 

1. Ilunieri rounded; dorsum of alitrunk strongly and evenly convex in 
profile; eyes protruding and very eonvex '1 
Humeri acutely subdentate; dorsum of alitrunk not so markedly convex 
in profile; eyes less convex and protruding only slightly. (Victorian 
mallee) ^1/. laevigata Browai 

2. Head broad, CI 98-100; head capsule opaque, the foveae almost con- 
tiguous ; gaster finelj' longitudinally striate over basal half or more of 

segment I (vicinity of Cairns, Queensland) M. turneri Forel 

Head narrovrer, CI 88-96 ; head capsule strongly shining, the foveae 
separated by smooth areas at least as wide as their maximum diameter; 
basal segment of gaster smooth and strongly shining (northern New 
South Wales) M. hrowni n. sp. 

In the discussion above I have not considered the enigmatic 
species f Mesostruma, monstrosa (Viehmeyer), 1925 (Brown, 
1948). The unfortunate circumstances surrounding the original 



1962 NEW AUSTRALIAN DACETINE ANTS 7 

selection of this species, which was based on an apparently ab- 
normal specimen, have been discussed by Brown (1952). Dr. 
Brown now believes (personal communication) that Viehmeyer's 
species was most likely based on a defective Epopostruma speci- 
men. In anj^ case monstrosa seems best ignored, pending the 
location and competent re-examination of the type. 

CODIOMYRMEX FLAGELLATUS UCW SpCcicS 

(Figs. 6-9) 

Holotype worker. TL 1.9 ; HL 0.48 ; HW 0.32 ; scape length 
(SL) 0.23; ML 0.08; WL 0.47; CI 68; MI 19. General form 
much as in C. semicomptus Brown (1959) (Fig. 9), but smaller 
and more lightly constructed. Shape of head as in Figure 6. 
Dorsal surface of cranium convex, sloping towards occiput and 
clypeus as in C. semicomptus, the convexity less pronounced, 
however, and the occipital lobes more broadly rounded when 
viewed from the side (c/. Figs. 7, 9). Mandibles strongly convex, 
rising above the anterior clypeal border ; with five strong, acute, 
conical teeth, each slightly smaller than the one posterior to it. 
Basal lamella normally hidden at full mandibular closure, set 
at a slightly lower level than the teeth and oblique to them; its 
shape as in Figure 8 — roughly right -triangular, the posterior 
edge almost perpendicular to the mandibular axis, and the 
anterior edge diagonal, forming the hypotenuse. The anterior 
lamellar edge rises almost immediately from the base of the 
proximal mandibular tooth so that the diastema is very brief. 
Clypeus almost perfectly plane. 

Body profile somewhat as in C. semicomptus, with which it is 
compared in Figures 7 and 9. Alitrunk narrow, its maximum 
width 0.59 X the HW; its dorsum almost perfectly plane, with- 
out sutures or a median longitudinal carinula. Alitrunkal dor- 
sum in side view evenly arched between the pronotum and the 
propodeal teeth. In dorsal view the sides of the pronotal disc 
are evenly rounded and those of the remaining alitrunkal dorsum 
almost parallel, with the distance between them, at the base of 
the propodeal teeth, slightly more than half the pronotal width. 
Dorsum of alitrunk margined with a fine carina, less distinct 
than in C. semicomptus, enclosing the pronotal disc anteriorly, 
and continuous with the upper edges of the propodeal spines 
posteriorly. The latter with their infradental lamellae similar 
to those of semicomptus. Propodeal spiracle minute, circular, its 
margin not appreciably expanded ; it contrasts with that of C. 



8 



BREVIORA 



No. 152 



semicomptus, which is larger and has a wide and very conspicu- 
ous rini-like margin (Fig. 9). Profile of petiolar node as in 
Figure 7, shorter than that of semicomptus, only about as long as 
high in side view ; seen from above the node is slightly longer 
than broad with rounded sides and a truncate anterior border. 
A full complement of areolate spongiform appendages is de- 
veloped, distributed normally as in Figure 7. Gaster depressed ; 
basigastric costulae reduced to about five feeble lines on either 
side of segment one, the two groups of costulae separated by a 
wide median shining area; the costulae extend back about y^ 
the length of the segment. 

Mandibles shining, with a few scattered punctures. Head 
capsule with a close cover of large, flat, irregular, shallow punc- 
tures, which are almost effaced in a small region in the center 
of the frons. Antennae very finely punctate, their scrobes punc- 
tate-granulose. Alitrunk, both nodes, gaster, legs and anterior 
parts of frons and clypeiis smooth and strongly shining. Petiolar 
l^eduncle coarsely granulate. 




Coflionv/rmi .r ffncjcUatits new species, Figs. 6-8, liolotype -workoT. Fig. C^. 
Full-face view of head. Fig. 7. Alitnuik nodes and liase of gaster in side 
\iew. Fig. 8. ilandiblc. Codiomyrmcx semicomptus Brovrn, paratope 
worker. Fig. 9. Alitrunk, nodes and base of gaster in side view. Scale line : 
O.IG mm., for Fig. 8; 0.25 mm., for Figs. 6, 7 and 9. 



1962 NEW AUSTRALIAN DACETINE ANTS 9 

Occiput with a number of very fine long (0.05-0.08 mm.) hairs 
with clavate tips, somewhat finer than in C. semiconiptus. Hairs 
of clypeus shorter and more distinctly clavate. A very few^ simi- 
lar hairs are present on the anterior part of the pronotal disc and 
the dorsal surfaces of both nodes. The pilosity otherwise consists 
of exceedingly long (0.13-0.36 mm.), fine, tapering hairs, which 
are distributed symmetrically. On the head, five pairs in the 
following positions (see Fig. 6) : on the occipital border, about 
halfway between its midpoint and its lateral extremity on either 
side ; on either side and a little anterior to the median part of 
the f rons ; at the edge of the cephalic dorsum just anterior to its 
widest point; at the edge of the cephalic dorsum at about mid 
head length ; on either side of the posterior extension of the cly- 
peus, above the antennal insertions. On the body, six pairs : two 
pronotal ; one on each node ; two at the base of the gaster — 
distributed as shown in Figure 7. On the forelegs : a single 
flagellum on the outer side of the tibia, near its apex. On the 
middle and hind legs: single hairs on the outer sides of the limbs 
near the bases of the tibiae and basitarsi. These long flagella curve 
upward and inward on the head capsule ; those of the body are 
erect with their apices turned posteriorly ; the anterior pronotal 
and postpetiolar pairs are more tangential, and inclined laterally. 

Type locality. Clump Point (near Mourilyan) Queensland, 
June 3, 1953 (T. E. Woodward), collected from a Berlese fun- 
nel sample of leaf mould. 

Worker paratype variation. Seventeen worker paratypes, col- 
lected with the holotype, show no significant variation in size or 
structure. The specimens have at some time been subjected to 
drying and fungal attack while in alcohol storage, with the 
result that some are fragmentary. A few fungal hyphae are still 
attached to several of the specimens, and a number have had 
the pubescence, particularly the elongate flagella, damaged dur- 
ing the consequent cleaning. 

Among the known Codiomyrmex, C. flagellatus is most closely 
related to C. semiconiptus Brown, the only other known Aus- 
tralian species. The general resemblances between these forms 
are indicated in the accompanying figures. The two species may 
be readily separated by the characters discussed in the above 
comparative dsecription; the difi^erences in size, pilosity, and 
propodeal spiracular structure are especially characteristic. 

The adaptive significane of the peculiarly sparse and elongate 
body pilosity of C. flagellatus is not understood, and deserves 
examination should live material become available for future 
study. 



10 BREVIORA No. 152 

The holotype, with paratypes, has been placed in the collection 
of the Queensland Museum. Paratypes are deposited in the 
Museum of Comparative Zoology, Harvard University, and the 
Commonwealth Scientific and Industrial Research Organization 
Collection, Canberra. 

I wish to acknowledge the generous assistance of Father B. B. 
Lowery of Sydney, and Dr. T. E. Woodward, of the University 
of Queensland, who collected the bulk of the material cited above 
and made it available to me for study. I wish also to thank Dr. 
E. 0. Wilson, of Harvard University, for his assistance and 
advice during the preparation of this paper. 

REFEEENCES 

Brown, W. L., Jr. 

1948. A preliminary generic revision of the higher Dacetini (HjTiien- 

optera: Formicidae). Trans. Amer. Ent. Soc, 74:101-129. 
1952. The claeetine ant genus Mesostruma Brown. Trans. Roy. Soc. 

S. Aust., 75:9-13. 
1953a. Revisionary studies in the ant tribe Dacetini. Amer. Midi. Nat., 

50:1-137. 
19o3b. A revision of the dacetine ant genus Ortciognaihus. !ilem. 

Queensland Mus., 13:84-104. 
1959. Some new species of dacetine ants. Breviora Mus. Comp. Zool. 

Harvard, 108:1-11. 
Brown, W. L., Jr. and E. O. Wilson 

1959. The evolution of the dacetine ants. Quart. Rev. Biol., 34(4): 

278-294. 



BREVIORA 

Museum of Comparsitive Zoology 



Cambridge, Mass. February 15, 1962 Number 153 

AN0LI8 SCRIPTUS GARMAN 1887, 

AN EARLIER NAME FOR 

ANOLIS LEUCOPHAEUS GARMAN 1888 

By A. Stanley Rand 

Garman in 1887 described Anolis scripfus on the basis of five 
specimens in the Museum of Comparative Zoology, giving the 
type locality as "Silver and Lena Keys, Fla." Barbour in 1914 
re-examined Garman 's type series and decided that they were 
identical with A^wlis cristatellus from Puerto Rico and the Vir- 
gin Islands and therefore placed A. script us in the synonymy of 
A. cristatellus. In the course of an examination of the Museum 
of Comparative Zoology anoles referred to A. cristatellus I had 
occasion to study the type series of A. scriptus. I find that the 
series is mixed and none is cristatellus. One, a juvenile, is A. 
homolechis quadrocelifer of Cuba; the other four are conspecific 
with the form from the southeastern Bahamas described by Gar- 
man (1888) as Anolis leucophacus, and apparently subspecifically 
identical with the form from the Turks and Caicos Islands now 
called alhipalpebralis Barbour 1916. 

Clearly the name scriptus can no longer be kept as a synonym 
of Anolis cristatellus, but correction of its status raises certain 
problems. Since the series is mixed a lectotype must be selected 
to fix the name. 

The type series, three adult males and two juveniles, are all 
somewhat faded from their long period of preservation. One of 
the juveniles possesses the scale characters of Anolis JiomoJechis 
and the color pattern, dark spots over the shoulders, is still suf- 
ficiently evident to identify it as Anolis homolechis quadrocelifer. 
This is the specimen labeled as coming from Lena Key, which 
thus would appear to be Cayos de la Lena, near Cabo San An- 
tonio, Cuba.^ I have arbitrarily excluded this juvenile from the 

1 For further information on this form see Rnibal and Williams (1961). 



2 BREVIORA No. 153 

concept of A. scriptus and it therefore needs no further dis- 
cussion. 

The remaining four specimens seem to belong to a single species 
and I herewith designate M.C.Z. No. 65950 as the lectotype of 
Anolis scriptus Garman. 

The labels accompanying these specimens say "Silver Key 
Florida." I, like Barbour, have been unable to locate a Silver 
Key anywhere in the West Indies. There is a Silver Bank near 
the islands from which the types must have come in the south- 
eastern Bahamas but it is completely submerged. 

These specimens are very like cristatellus as both Garman and 
Barbour agreed. Garman distinguished them from cristatellus 
on the basis of the greater size of the two paravertebral scale rows. 
Barbour (1914, p. 274) said, "I can not see, however, that these 
are at all enlarged ; and there is no other character in which they 
vary from true A. cristatcllns." An examination of the type 
series helps to explain this contradiction. Two of the males have 
the two paravertebral scale rows enlarged more than is usual in 
cristatellus, but the third male has the paravertebral scale rows 
scarcely enlarged at all and it is undoubtedly this specimen that 
Barbour examined. 

However, a close comparison shows certain other and more 
constant differences between the type series of scriptus and the 
many specimens of cristatellus examined. In scriptus the dorsal 
scales lateral to the paravertebral rows are larger than they are 
in specimens of cristatellus of similar size. In cristatellus also, 
the frontal ridges are higher and sharper and the frontal depres- 
sion correspondingly deeper than in the type series of scriptus. 
Finally, in cristatellus, there are only 1-3 scales behind the inter- 
parietal and these are abruptly larger than the very small dorsal 
scales. In scriptus there are many more rows of enlarged scales 
in this position and they grade more gradually into the dorsal 
scales. 

In all of these characters the type series of scriptus differ from 
cristntellus and agree with specimens of the species now called 
leucophaeus. So far as I can find, the types of scriptus do not 
show any scale differences from leucophaeus, nor does leucophaeus 
show any additional differences from cristatellus. 

From this it appears that scriptus and leucophaeus are synon- 
ymous and scriptus as the older name must be substituted for 
leucophaeus. 



1962 ANOLIS SCRIPTUS 3 

The species "leucophaeus" is quite widely distributed in the 
southeastern Bahamas and has been divided into four subspecies. 
These races have been described primarily on the basis of color 
pattern, and they are all very similar in scalation. They are 
diagnosed in Table I. 

The types of script us lack the many dark spots characteristic 
of leiicophaeus and the lectotype has a well-developed tail crest 
which is lacking in sularum. Thus the name scriptus definitely 
does not apply to the populations called leucophaeus and sularum. 

Distinguishing between albipalpehralis and mariguanae is more 
difficult. The diagnostic difference between them is the pres- 
ence of a broad dark lateral band in mariguanae. This is absent 
in the type series of scriptus but it is also absent in many of the 
adult males of mariguanae and best developed only in the juve- 
niles and females. Even the small ''type" of scriptus lacks this 
band but this specimen is so faded that one cannot be positive 
that the band was never present. Many of the females of albipal- 
pehralis have dark middorsal blotches which are lacking in the 
small "type" of scriptus but, since they are absent in man}' alhi- 
palpebralis, this is not conclusive. The male scriptus have a com- 
plex mottling along the sides in addition to a light narrow lateral 
line. The light lateral line is found in both albipalpcbralis and 
mariguanae but the mottling in the types of scriptus is most like 
that found in albipalpebralis. Finally, the lectotype of scriptus 
has a dark line running posteriorly from the eye onto the neck. 
This marking is found in some of the males of albipalpebralis 
but in none of the mariguanae examined. So far as can be deter- 
mined there are no useful scale differences between mariguanae 
and albipalpebralis. From this it appears that the "type" series 
of scriptus, while not indisputably assignable to either of these 
races, is most like albipalpebralis and the lectotype most clearly 
so. For this reason it seems necessary to replace the name albi- 
palpebralis by the name scriptus. In accordance with this change 
the type locality of scriptus is restricted from "Silver and Lena 
Keys" to "Silver Key," Turks and Caicos Islands. Further 
restriction seems pointless at this time. The correct names for 
the races of this species now stand as follows: 

Anolis scriptus scriptus Garman 1887 = Anolis albipalpebralis 

Barbour 1916 
Anolis scriptus leucophaeus Garman 1888 
Anolis scriptus mariguanae Cochran 1931 
Anolis scriptus sularum Barbour and Shreve 1935 



BREVIORA No. 153 



■3 op -- 



»-. 


^ 


<u 


s 






ft 


-*— 


a> 


c 


ft 


ft 



O) 



=0 


c 




« 




s 


ci 




3 




•^^ 


3 




be 




Si. 


tx 




c3 




•«* 










t 


^ 








=0 


^ 




a 




■2 










r«» 










o 










s 










^ 






OQ 




«l-l 






o 




o 




cc 


^ 




01 




15 


o 




a> 




tH 








11 


^ 
o 


'^ 




ft 


■r. 


_ft 


C3 


03 


en 
to 


.f- 


"rt 






5 


K 


13 


H 


1— ( 


^ 










H 











S3 bD 



^ -S 





HI 








> 


rM 






o 








o 


rt 




>-i 


cS 


■^ 




-^^ 


ns 


iw 




J5 


a> 


c: 




o 


;« 


o 






O) 






o 


>5 


d 


'M 


,^ 


^ 


p, 



o g 3 0^ •« __ § 

• s- S, r-^ ^ t:? g 

rt ■;; S o ^ c? ^H 

ft 



.^ 


a* 




be 


r^ 






1— • 


bt 


o 


"« 


.5 


IH 






c 




^ 




o 


5 


Is 





m 
ft 



fii 






rt 






c4 


OQ 

3 


I— 


3 
be 


es 


7? 




« 


j^ 


S 


^ 


O 






c: 


1—1 


K 


ft 




O 
o 
3 




o 


3 






Jh 


• p- 


*" 


c 









1IJG2 ANOLIS SCRIPTUS 5 

EEFERENCES CITED 

Cochran, D. 

1931. New Bahaman reptiles. Jour. Washington Acad. Sci., 21: 39-40. 
Barbour, T. 

1914. A contribution to the zoogeography of the West Indies, with 
especial reference to amphibians and reptiles. Mem. Mus. Comp. 
ZooL, 44: 209-359. 
1916. Additional notes on West Indian reptiles and amphibians. Proc. 
Biol. Soc. Washington, 29: 215-220. 
Barbour, T. and B. Shrkve 

1935. Concerning some Bahaman reptiles, with notes on the fauna. 
Proc. Boston Soc. Nat. Hist., 40: 347-366. 
Garman, S. 

1887. On West Indian Iguanidae and West Indian Scincidae in the 
collection of the Museum of Comparative Zoology, Cambridge, 
Mass., U.S.A. Bull. Essex Inst., 19: 25-53. 

1888. Reptiles and batrachians from the Caymans and Bahamas. Bull. 
Essex Inst., 20: 103-116. 

RuiBAL, R. AND E. E. Williams 

1961. The taxonomy of the Anolis homoleohis complex of Cuba. Bull. 
Mus. Comp. Zool., 125: 211-246. 



BREVIORA 

Mmseium of Compsirsitive Zoology 



Cambridge, Mass. April 4, 19G2 Number 154 

NOTES ON HISPANIOLAN HERPBTOLOGY 
5. THE NATURAL HISTORY OF THREE SYMPATRTC 

SPECIES OF ANOLIS 

By A. S. Rand 

The lizard genus Anolis is abundant in species and individuals 
in the Greater Antilles. Quite a number of the common species 
occur together over wide areas. There is thus a special oppor- 
tunity to study the ecological relations of sympatric species of 
a single genus under conditions genuinely favorable for field 
observation. 

Several papers have now been published which have begun to 
exploit this fortunate opportunity: Ruibal (1961) and Collette 
(1961) working on the anoles of Cuba; Oliver (1948) on those 
of Biniini in the Bahamas; Williams and Rand (1961) on the 
scmUincatus group in liispaniola. These and earlier workers — 
Grant (1940) for Jamaica; Stejneger (1904) and Schmidt 
(1928) for Puerto Rico and especially Mertens (1939) for His- 
paniola — have established that in each well studied case there 
are small but definite differences in microhabitat among the 
sympatric Anolis. 

In this paper I document the same point for the three com- 
monest species of Anolis in Hispaniola. I describe also a number 
of behavioral differences which seem to be associated with the 
ecological differences and attempt to assess the adaptive sig- 
nificance of these behavior patterns. This study was made during 
a two-month field trip to the Dominican Republic and supple- 
mented with observations on captive lizards kept in the lab- 
oratory in Cambridge. 

ACKNOWLEDGMENTS 

I wish to thank the government of the Dominican Republic 
that, through the Universidad de Santo Domingo, provided us 



2 BRE\'TORA No. 154 

with transportation and other assistance in that country ; Dr. 
Engenio de Jesus Marcano who helped us in the field and who 
sent me live specimens on our return. I wish also to thank Mr. 
Clayton Ray for inviting me to accompany him on this trip, and 
for his assistance in the field; Dr. E. E. Williams for helping 
to arrange the trip and for his advice and assistance throughout, 
and Dr. A. L. Rand for critically reading the manuscript. Also 
I wish to thank Sigma Xi for providing some of the funds that 
made this trip possible. 

MATERIAL AND METHODS 

The field observations during the summer of 1958 were made 
incidentally to the main project, a survey of Dominican caves for 
vertebrate fossils. Consequently, they include no lengthy ob- 
servations in any one locality but are a synthesis of data col- 
lected from widely separated places. Though I noticed no geo- 
graphical variation in behavior, obviously this is a possible source 
of error that runs throughout the present study. This is par- 
ticularly true as observations are lumped for different subspecies 
in two cases: Anolis clilorocyanus chlorocyanus and A. cliloro- 
cyanus cyanostictus ; and Anolis distichus ignigularis and A. 
distich Its dominice^isis. 

The main areas, where observations were made, were : Santo 
Domingo and vicinity, Santiago and the nearby Sierra Septen- 
trional, Padre Las Casas, Sabana de la Mar and the mountains 
south of there, and the Sierra de Neiba. These field observations 
are least complete on A. cJdorocyonus and uneven on the other 
two species. For example, I found eggs only of cyhotes, while I 
observed copulation only in distichus. 

The laboratory observations made at the Harvard Biological 
Laboratories involved only two species, A. cyhotes and A. chloro- 
cyanus chlorocyanus. These animals, ten of each species, all 
adults and about half of them males, were sent to me by Pro- 
fessor Eugenio de Jesus Marcano. The lizards were released in 
a room 12 x 20 ft. and 9 ft. high. The room was a constant 80° F. 
and lighted by a south window. Eight small tree trunks 7 feet 
tall, mounted on bases so that they stood upright, an 8 foot rub- 
ber tree, a potted pine bush, and some other potted plants w^ere 
arranged around the room. 

The li'-ards were fed meal worms on the floor and in dishes 
taped at various heights above the floor, on the tree trunks. 
Fruit flies were also released in the room. The whole room was 
watered daily. 



] 962 THREE IlISPANIOLAN ANGLES 3 

For a period of about three weeks, daily observations of vary- 
iiiw lengths were made. The lizards quickly came to ignore me 
as I sat quietly in one corner of the room and watched them 
with low power binoculars. 

This was obviously not a natural situation but the observations 
that could be checked against the field notes show a gratifyingly 
close agreement. This method seems to be a useful adjunct to 
field observations though obviously no substitute for it. 

All three of these species are small to medium sized lizards 
possessing the characteristic specializations of the genus : en- 
larged subdigital lamellae and a throat fan or dewlap that is best 
developed in the male. In each species the males grow to a larger 
size than do the females. 

A)iolis disticluis varies in dorsal coloration from green to gray 
to brown, frequently with mottling on the back. The dewlap is 
yellow in A. d. doniinicensis, and red with a yellow border in 
A. d. ignigidaris. The males measure 51 mm in snout-vent length 
(Cochran 1941). 

Anolis cydotes is dorsally gray or brown, sometimes reddish, 
frequently with two indistinct greenish lateral stripes. The dew- 
lap is whitish or yellowish in color. The males measure 67 mm 
in snout-vent length (Cochran 1941). 

Anolis chlorocyanus is usually bright green with the ability 
to change to brown. A. c. cyanostictus also has a rust-red spot 
in front of the shoulder and one on the head. A. c. chlorocyanus 
has a dewlap that is light blue anteriorly and dark, almost black 
posteriorly. A. c. cyanosiicius has the dewlap proximally cad- 
mium yellow, distally sky-blue. The males measure 71 mm. in 
snout-vent length (Cochran 1941). 

OBSERVATIONS 

The observations from both field and laboratory have been 
combined and arranged according to topics. Under each topic 
heading all three species are compared. 

Geographical Distrihutio7i. All three species are widespread 
in the Dominican Republic. Apparently they avoid only the 
high, wet pine forests of the Cordillera Central and are less 
common in the dry regions on the south coast near Asua and 
in the northwest near Puerto Plata. However, A. chlorocyanus 
is replaced by A. coelestinus in the Barahona Peninsula. Geo- 
graphical variation in each of these species has been described 
byMertens (1939). 



4 BREVIORA No. 154 

Hahitat. I found these three species at almost every locality 
I visited in the Dominican Republic. Mertens called distichus 
and cybotes eurytopic forms and chlorocyanus only slightly less 
so. He records both distichus and cybotes from the mangrove 
areas along the coast to the lower pine forests around Constanza 
at some 1200 meters altitude. 

The most significant environmental factor for all three species 
seems to be some sort of vertical perch, a tree trunk, fence post 
or cliff face, but A. chlorocyanus seems to avoid the most open 
areas. 

Anolis distichus lives primarily on isolated trees and fence posts 
and along the edges of woods and trails and in open woods. 
A. cybotes lives in these situations but also occurs in the deeply 
shaded interiors of densely wooded areas that distichus avoids. 
A. clilorocyanus certainly occurs in the edge situations that dis- 
tichus prefers and probably in the heavier woods as well. 

Though there are differences such as these in the extremes 
tolerated, these animals live in the same habitats over most of the 
Dominican Republic and part, at least, of Haiti. Mertens notes 
that in separated localities he observed individuals of all three 
species living in the same tree. My observations agree with this. 

Microhabitat. These three species, though living in the same 
habitat, have differences in their microhabitat preferences. 

A. distichus lives almost exclusively on exposed (i.e. not closely 
surrounded by vegetation) tree trunks, fence posts and similar 
structures, within 10 to 15 feet of the ground. It is seldom seen 
on the ground or in the smaller branches of bushes or trees. 
Though it must descend to the ground to reach isolated trees, 
it does not spend much time on the ground. A. distichus is often 
very common on the palisade fences in small villages. 

A. cybotes is also primarily an animal of exposed tree trunks 
and fence posts within 10 feet of the ground. However, it also 
frequents rocks and fallen logs and smaller individuals are 
frequently seen on the ground. It also avoids, during the day 
at least (see Sleeping), small twigs and foliage. These field ob- 
servations are confirmed by the laboratory data which show 
cybotes spending most of its waking time on perches less than 
five feet from the floor. 

A. chlorocyanus lives also on tree trunks and fences but unlike 
the other two, frequently ranges high in the trees and out among 
the smaller branches. I saw a female fall from the crown of a 
30 foot palm tree. I suspect that this species is one that lives 
primarily up among the branches and ranges down the trunk 



1962 THREE HISPANIOLAN ANGLES 5 

rather than the reverse. However, individuals were seen also 
on fences and once I saw a dozen individuals on a pole frame- 
work nine feet high that with a few bits of palm thatch was all 
that was left of a shed in the middle of a treeless pasture. Like 
distichus, chlorocyanus must occasionally descend to the ground 
to reach isolated trees. 

In the laboratory, chlorocyanus spent most of its time high 
on the perches and ventured out among the leaves of the rubber 
plant much more than did the cybotes. 

Territoriality. As Mertens noted, all three species appear to 
be territorial. He says that for A. cyhotes and chlorocyanus he 
found only a single adult male and one or more females of each 
species on any one tree. When he placed an additional male on 
one of the occupied trees it was immediately attacked. A. dis- 
tichus, he saj^s, defends a smaller territory and the large trees 
may have several males, each with its own territorj^, as well as 
a number of females. 

These observations agree with mine, although it was only in 
areas where distichus were extremely common that I found more 
than one adult male distichus on the same tree, and then only 
on large trees. 

Certainly in the laboratory no male of either cyhotes or chloro- 
cyanus would tolerate another male of the same species in his 
immediate vicinity for long, and even females were chased away 
if they approached too closely. Occasionally a male of one species 
would display to an individual of the other species. 

In addition to this horizontal effect a vertical stratification on 
individual trees or posts is evident in at least two of the species, 
distichus and cyhotes. 

On the smaller trees that were occupied by two individuals of 
distichus, the lizards were usually of opposite sex and the male 
was usually closer to the ground than was the female. On one 
tree there were four lizards of this species : a large male near 
the base, a female above him, a slightly smaller female above her 
and, highest of all, a juvenile who was about 7 feet above the 
ground. During the hour that I watched this tree all of the 
lizards moved a number of times but this stratification remained 
the same for it was actively enforced. When one of the higher 
individuals moved in sight of and within 2 or 3 feet of a lower 
one, the lower animal immediately reacted to it. The male did 
this by bobbing his head and pumping his dewlap, the female 
by a short charge in the direction of the intruder. In every case 
the intruder retreated immediately, around the tree and up. 



6 BREVIORA No. 154 

In cyhotes almost all of the individuals on the trees were large 
and most of them were males. Tlie individuals that were seen 
on the ground were almost always juveniles or females; the 
females are much smaller than the males in this species. This 
spatial distribution seems to be a real phenomenon but I have 
no field evidence to support the hypothesis that it is the result 
of territorial defense. However, there are certain laboratory 
observations that support this contention. In the laboratory 
I saw no vertical stratification but I also saw no evidence that 
the females tended to spend more time on the ground than did 
the males. This may be because the perches were small and nu- 
merous enough so that each lizard could occupy its own, as 
they usually did. However, there was a definite stratification 
horizontally with the large males on the perches near the window 
and the smaller males and females on those farther away. AVhen 
a smaller individual invaded a perch near the window, and this 
happened quite frequently, the resident male would display to 
it and the intruder would retreat. Thus, though there is no 
direct evidence that the distribution seen in the field is due 
to territorial defense, there is evidence that defense could be 
at least a contributing factor : the males taking the most de- 
sirable positions, the elevated perches, and ciiasing away any 
smaller individual that attempted to move in, with the result 
that the juveniles and females would spend most of their time 
on the ground. 

It is of interest to note that this vertical stratification in 
cyhotes and disiichus results in greater difference in size between 
the individuals that occur together. Adult cyhotes are larger, 
and the adult males much larger, than adult distichus. This size 
difference could be reflected in the size of the prey items taken 
and so reduce the competition for food between the two species. 
The young of cyhotes are of course no bigger than adult dis- 
tichus but these are the individuals that live primarily on the 
ground and so do not occur on the tree trunks with the distichus. 
The importance of this in reducing competition for food, if it 
acts in this way at all, is of course unknown. It would be inter- 
esting to have an analysis of the stomach contents of various 
sized individuals of each species. 

In chlorocyanus, I have no clear evidence for vertical strati- 
fication either in the field or in the laboratory. However, the fact 
that most of the individuals caught on fences and tree trunks 
were adult males suggests that the females and the juveniles may 



1962 THREE HISPANIOLAN ANGLES 7 

Stay higher in the trees. This phenomenon may also give maxi- 
mal emphasis to the size difference between this species and the 
distichus with which it comes in contact for it is the large indi- 
viduals of chlorocyanus that descend the tree trunks and they 
meet first and presumably would compete most for food with the 
smallest distich us. 

Posture. Seen in silhouette these three species are usually im- 
mediately recognizable. This is partly because of their different 
proportions but even more because of their very dift'erent postures. 

A. distichus rests facing either up or down or angling across 
the tree with its head and at least the anterior part of its body 
well off' the substrate and with its neck bent so that its head is 
parallel to the substrate but further away from it than are its 
shoulders (Cf. figure 4, plate 2, Mertens). 

A. cyhotes typically rests facing down the tree with the fore 
part of the body off the substrate and the neck bent dorsally so 
that the head is nearly parallel to the ground. In one individual 
seen resting on the underside of a log which slanted at about 45°, 
the neck was bent back well over 90°. This posture is true in 
the laboratory as well as in the field. On the ground the posture 
is much like that of distichus with the neck bent so that the head 
is parallel to the ground and raised above it. 

A. chlurocyanus usually rests with both its head and body 
quite close to the substrate and its neck bent only a little if at all, 
both in the field and in the laboratory. 

While these postures are typical of the normal resting posi- 
tion, both cyhotes and disticltus when mildly alarmed flatten 
against the substrate, and cklorocyanus, when about to display, 
raises itself up on its legs. 

I believe these differences in posture can be correlated with 
feeding behavior as discussed below. 

Mertens says that most Anolis rest with their heads pointing 
toward the ground. I noticed this most commonly in cyhotes and 
less so in the other species. 

Activity. A. distichus appears to be a much more restless 
lizard than either of the other tw'o species. Like them it spends 
most of its time resting quietly but an individual seldom remains 
in one place for more than a few minutes. It then moves quickly 
a few inches away on its tree trunk and rests quietly again for 
another few minutes. 

A. cyhotes, on the other hand, seems to spend much longer 
periods of time resting in one spot. Again, when a change is 



8 BREVIORA No. 154 

made, it is made quickly. Tn the laboratory when a cyhotes 
moved from one tree to another, the lizard frequently ran down 
to the base of the tree, stopped for a few moments, left the tree 
with a jump and ran part way across the floor, paused, ran the 
rest of the way and with a jump started up the new tree, usually 
pausing again before settling down. 

A. chlorocyaniis, though also spending periods of time im- 
mobile, moved quite frequently, going slowly and deliberately 
about in the trees, in the laboratory as well as in the field. In 
the laboratory, moving from one tree to another was a single 
process. The lizard moved slowly down to the base of the tree, 
jumped off, dashed across the floor and with a jump started 
up the new tree. As little time as possible was spent on the floor. 

Climhing. I have no observations on the climbing ability of 
distichus. 

In the laboratory A. cyhotes did not appear to be as sure a 
climber as A. chlorocyanus. I saw several of the former fall to 
the ground from the smooth leaves of the rubber plant while I 
noted that only one of the latter did so even though chlorocyanus 
spent much more time on these leaves than did cyhotes. While 
both these species started their climbs in the lab with a jump 
up on to the vertical surface, and I saw a large male cyhotes make 
a 6 or 7 inch vertical jump to reach a hanging branch, chloro- 
cyanus made a great many more horizontal jumps. Particular!}' 
common was one used to cross the 10 inches that separated the 
two closest perches. 

A. cyhotes is apparently a quickly moving lizard, quite at home 
on the ground but not so much so in the more treacherous footing 
of smooth green leaves and small twigs. A. chlorocyanus, on the 
other hand, is shy of the ground, moving across it only occa- 
sionally and then with as much speed as it can manage, while 
in the less secure footing of more arboreal situations, its de- 
liberate movements help keep it from falling. 

Feeding. On several occasions a distichus was seen to move 
from its resting position on a tree trunk or fence post and snap 
up something small from the bark. Twice I saw a male interrupt 
his displaying to do this. Once I saw a male move up to an ant 
about an inch away, follow it up the tree a couple of inches and 
then apparently lose interest and turn away. Thus there seems 
to be some selection of food and not everything small that moves 
within range is eaten. 



1962 THREE HISPANIOLAN ANGLES 9 

A. cyhofes was never seen to catch anything in the field, though 
on two occasions I saw an individual struggling with a large 
dragonfly ; each time the lizard had the head and thorax in his 
mouth and the wings and abdomen still protruding. In the lab- 
oratory, cyhotes came willingly to the floor to take meal worms. 
Sometimes a lizard would return to a tree to eat the meal worm 
but usually it remained for several additional minutes before 
returning either to its old tree or a new one. I never saw a 
cyhotes moving around on the floor looking for food. On one 
occasion a male left his perch and ran about 10 feet across the 
floor to make an unsuccessful attempt to capture a 2-inch cock- 
roach. 

I have no data on chlorocyavus feeding in the field. In the 
laboratory several individuals were seen snapping at small ob- 
jects on the leaves and twigs. This species only rarely came down 
to the floor to capture meal worms and each time that one did 
so it moved slowly to the base of the tree, then rushed out, seized 
the meal worm, ran back and climbed up the tree before stopping 
to eat its captured prey. 

On the basis of this limited evidence, some tentative generali- 
zations about the relations of feeding, posture and movement 
can be made. 

A. distichus seems to feed primarily on smaller insects that 
it catches on the tree trunk. The posture with the head held 
high above the substrate would enable it to see more of the tree 
trunk than it could if the head was held low. The frequent move- 
ments are necessary if the lizard is to take advantage of the in- 
sects that happen to be on the opposite side of the tree trunk. 
The head is held parallel to the surface in which the lizard is most 
interested. 

A. cyhotes males get at least part of their food from the ground 
after sighting it from their elevated perch. The posture of this 
species keeps the head roughly parallel to the surface in which 
the lizard is most interested, in this case the ground. Since the 
lizard can see a large sector of the ground around him at all 
times, he need not change his position frequently to maintain 
a careful scrutiny of a considerable area. 

A. cJilorocyanus almost certainly gets most of its food from 
the trees in which it usually lives and its slow movements sug- 
gest that at least part of the time, unlike the other two species, 
it goes looking for it instead of lying in wait. The head is held 
parallel to the surface in which it is most interested. 



10 BREVIORA No. 154 

We need more observations on all of these points and par- 
ticularly on the feeding of A. cJilorocyanus. But it is interesting 
that the two species, distichiis and cyhotes, that occur in the 
same microhabitat, on the tree trunks, seem to differ so markedly 
in feeding behavior. 

Mertens remarks, in passing, that most Ayiolis get their food 
on the ground and among the roots of the trees and that their 
usual posture, as he records it, oriented toward the ground, may 
be related to this. I believe that this is true of cyhotes but not 
of the other species. 

Escape. Each of the arboreal species has a noticeably differ- 
ent method for evading herpetologists and presumably other 
predators. 

A. disiichus, when approached, quickly moves around to the 
other side of its tree trunk, usually moving up or down at the 
same time. If followed, the lizard may continue this maneuver- 
ing to keep the tree trunk between itself and the pursuer for 
some minutes. Soon, however, the lizard will either run up the 
tree out of reach or run down it to the base where it is concealed 
by the surrounding vegetation. Only when very hard pressed will 
one leave his tree or post and run out into the grass. Occasionally, 
and this was noted especially early in the morning, distich us 
would hide under a bit of bark or in a hole in the tree. Mertens 
says that distichiis, when approached, runs around to the other 
side of its tree or post and then down to its base to hide. He 
does not mention any of the other behavior described here. 

A. cyhotes showed the same tactics in the field and in the lab- 
oratory. When approached, an individual would remain still 
until I came very close, then suddenly it would dart around to 
the other side of the tree ; it might stop there but only until I 
moved into view again, then, instead of employing evasive ac- 
tion on the tree, it would usually run down and, unlike distichus, 
frequently leave the tree completely. In the laboratory the lizard 
usually ran a few feet away, where it might remain on the floor 
for some moments or might immediately climb a new tree. Mer- 
tens records these same flight reactions for this species. An 
A. cyhotes, discovered on the ground or on a fallen log, frequently 
hid under whatever cover was available. Sometimes a large male, 
when first approached, would display his dewlap to me before 
fleeing. This was observed occasionally in the wild and became 
very common with one male in the laboratory after he became 
accustomed to me. 



]962 THREE IIISPANIOLAN ANGLES 11 

Usually in the field, cJilorocyanns, when approached, imme- 
diately began to climb up the tree. This action was slower and 
more deliberate than that of the other two lizards but, since it 
was started sooner, the lizard was usually carried safely out of 
my reach. In the laboratory and in the field, when on a fence, 
where the lizard could not climb out of reach, it usually climbed 
as high as it could get and then dodged about there. Only very 
occasionally did one run down to the ground, though frequently 
one jumped to a nearby perch, if available, or ran out among 
the foliage and twigs at the ends of the branches. Mertens also 
notes that cJilorocyanns usually climbs up its tree and conceals 
itself in the crown. 

The escape behavior of the three species closely parallels the 
feeding behavior. A. distich us conducts its evasive behavior on 
the tree trunk. A. cyhotes willingly leaves its perch to escape on 
the ground, and A. chlorocyanus retreats whenever possible up 
into the top of the tree. 

Daily Activity. Since I did not spend long periods of time 
watching any one lizard or groups of lizards, I have no detailed 
information on the variations in activity during the day. 

All three species seem to be strictly diurnal. They w^re seen 
sitting in the sun more frequently in the early morning than 
at other times of the day, presumably to raise the body tempera- 
ture to the preferred level. A. (listicJius, at least, seems to feed 
more actively in the morning than at any other time. On several 
occasions I captured one with food in its mouth and three times 
I had one snap at the knot of the thread noose with which I was 
trying to snare it. 

Sleeping. I do not know where A. clisticlnis spends the night 
but, since individuals sometimes use holes in trees as hiding 
places in the early mornings, they may possibly use these during 
the night. 

Two individuals of A. cyhotes Avere found asleep at night, in 
the field. Both of them were males and were asleep on top of 
the foliage of the outermost twigs of small bushes. They were 
plainly visible from my vantage point, though perhaps not to a 
predator, such as a snake, climbing the bush. In the laboratory 
one evening during an examination of the room, I located seven 
cyhotes asleep. Five of them were in the needles at the tips of 
the branches of the pine bush, one on a small vine where it 
stretched away from the tree trunk, and the last on top of the 
topmost leaf of a large-leaved potted plant. Shaking a branch on 



12 BREVIORA No. 154 

which a cyhotes was sleeping woke the lizard but it did not move 
until the branch was shaken vigorously or the lizard touched. 
Then it jumped immediately to the floor and remained there un- 
moving. 

A. chlorocyanus was seen asleep only in the laboratory where 
during the period mentioned above I located six animals. Three 
of them were sleeping between large leaves of the rubber plant 
and the wall, one was between a board and the wall, one inside a 
cold radiator, and only one exposed, on top of its perch. When 
disturbed, the chlorocyanus immediately sought new hiding 
places. 

As Mertens notes, A. distichns and A. cyhotes sleep with their 
liind legs partly flexed, while chlorocyanus usually sleeps with 
them extended along the tail. 

I do not know enough about the real and potential nocturnal 
predators of Anolis to speculate on the adaptive significance of 
these sleeping places. But it is striking that they are so different 
from the situations in which the animals spend the day. Male 
cyhotes, which in the daytime live on substantial vertical surfaces, 
during the night sleep on the small flexible foliage and twigs 
of bushes and vines. A. chlorocyanus during the day is usually 
exposed to view and during the night sleeps under or behind some 
sort of cover. 

Ue'productwn. Since copulation was observed only in distichus 
and eggs found only of cyhotes, it is not possible to compare these 
species with respect to reproductive behavior. I am including 
these data in the hope that eventually comparative data will be 
available. 

Three times pairs of distichus were seen in copulation in early 
to mid-afternoon on tree trunks from four to six feet above the 
ground. In no case was the preceding courtship observed. In one 
case tlie lizards were oriented diagonally up a large tree six 
feet from the ground. The larger male was on the higher side 
and on top of the smaller female. One of his front legs was 
across her shoulders and holding on to the tree in front of her 
front leg; one of his hind legs Avas across the base of her tail, 
the toes resting on her thigh and the trunk in front of it. His 
other legs were spread out holding the tree. His tail was bent 
under hers and his head Avas resting on but not biting her neck. 
Her position was that of any resting lizard except that her tail 
was strongly arched. The position observed in the other cases 
was virtually identical. 



1962 THREE niSPANIOLAN ANGLES 13 

SUMMAKY AND CONCLUSIONS 

These observations on the behavior and ecology of these three 
species of lizards, Anolis distichus, A. cyhotes and A. chloro- 
cyanus in the Dominican Republic are obviously incomplete. 
However, certain tentative conclusions can be advanced. 

These species are sympatric and occupy the same macrohabi- 
tats over much of their ranges. The species differ somewhat in 
their microhabitats but they are not clearly separated spatially 
in this way; members of each species occur on the same tree 
within feet and sometimes within inches of each other. They all 
have the same basic body form, though differing somewhat in 
size, in proportions and in morphological adaptations such as 
development of the enlarged subdigital pads. They are all in- 
sectivorous in diet. However, in the details of their behavior 
they are very different. These details can be fitted together to 
form a picture of three morphologically similar species of the 
same genus living together in the same habitat but living three 
very different lives. 

Reviewing these very briefly, Anolis distichus is strictly an 
animal of the lower tree trunks. Its territorial pattern results 
in .spacing out the individuals living on the same tree with the 
largest at the bottom and the smallest higher up. This species 
feeds on small insects on the trunk, utilizing a posture and pat- 
ern of activity that enables it to forage effectively in this sort of 
place. \Yhen frightened it takes evasive action on the tree trunk, 
and copulation takes place there. 

A. cyhotes, though partially an animal of the tree trunks, is 
also closely associated with the ground below. Its territorial 
behavior results in a spatial distribution with the large indi- 
viduals on the trees and the smaller ones on the ground. Even 
for the indi\dduals on the trees the perches there seem to be pri- 
marily lookouts from which to survey the ground. The postures 
and patterns of activity seem fitted best for this and much less 
so for watching the tree on which the lizards sit. Certainly they 
do go to the ground to capture food spotted from these perches. 
They run to the ground when frightened and they bury their 
eggs in the ground. 

A. chlorocyanus in its behavior seems as closely related to the 
tree tops as cyhotes is to the ground. It seems to be mainly the 
large males that descend the tree trunks. Some feeding un- 
doubtedly occurs high in the trees even in these large males and 



14 BREVTORA No. 154 

chlorocyanus retreats upwards "when frightened. Its slow de- 
liberate movements seem adapted to the more precarious arboreal 
footing among the leaves and twigs. 

These are not just three similar animals doing the same things 
in the same way in slightly different places but three similar 
animals, each with a unicpie set of complex behavior patterns 
which interlock functionally so that each species has its own way 
of life within the same habitat. 

This situation can be an example of the operation of the Gause- 
Volterra hypothesis that closely related species can live together 
only if they differ in ecology. 

Since many of these differences can be correlated with feeding 
behavior they can be interpreted further as serving to reduce 
interspecific competition for food. 

However, the demonstration that these conclusions are valid 
must await the collection of further data, particularly data on 
just what environmental factors act to limit the population densi- 
ties of each species. 

LITEEATURE CITED 

Cochran, D. M. 

194]. Tlie heipetology of Hispaniola. Bull. U. S. Nat. Miis., 177: 1-.398. 

COLLETTE, B. 

1961. Correlations between ecology and morphologj- in anoline lizards 
from Havana, Cuba and southern Florida. Bull. Mus. Comp. 
ZooL, 125: 137-162. 
Grant, C. 

1940. II. The Reptiles. In Lynn, W. G. and C. Grant, The Hcrpetology 
of Jamaica. Bull. Inst. Jamaica, Sci. Ser., 1 : 61-148. 
Ingek, R. F. 

1959. Temperature responses and ecological relationships of two Bor- 
nean lizards. Ecology, 40 (1): 127-136. 
Meetens, R. 

1939. Herpetologishe Ergebnisse einer Reise nach der Insel Hispaniola, 
Westindien. Abb. Senckenb. naturf . Ges., 449 : 1-84. 
Oliver, J. A. 

1948. The anoline lizards of Bimini. Anier. Mus. Novitates, no. 1383: 
1-36. 

RUIBAL, R. 

1961. Thermal relations of five species of tropical lizards. Evolution, 

15: 98-111. 
Schmidt, K. P. 

1928. Amphibians and land reptiles of Porto Eico, with a list of those 

reported from the Virgin Islands. Sci. Survey Porto Rico and 

the Virgin Islands, 10, part 1: 1-535. 



1962 THREE HISPANIOLAN ANGLES 15 

Stejneger, L. 

1904. The herpetology of Porto Rico. Rep. T. S. Nat. Mus., 1902: 
553-724. 
Williams, E. E. and A. S. Rand 

1961. Notes on Hispaniolan herpetology. 2. A review of the Anolis 
semilineatus group with the description of Anolis cochranae, 
new species. Breviora, no. 135: 1-11. 



BREVIORA 

Mmseiiam of Connparsitive Zoology 

Cambridge, Mass. April 12, 1962 Number 155 



NOTES ON HISPANIOLAN HERPETOLOGY 
6. THE GIANT ANGLES 

By Ernest E. Williams 



INTRODUCTION 

Mertens (1939) has called attention to the existence of geo- 
graphic variation in the giant anoles of Hispaniola and has 
distinguished a typical western race, Anolis ricordii ricordii 
Dumeril and Bibron, and an eastern race, A. r. haleafus Cope. 

The distinction between these two forms is sharp and un- 
equivocal ; the situation is, however, more complicated than 
Mertens' limited sample (16 specimens) led him to believe. 
Study of the unreported series of Hispaniolan giant anoles in 
the American Museum of Natural History (AMNH) plus the 
specimens in the Museum of Comparative Zoology (MCZ) and 
the United States National Museum (USNM) (91 specimens in 
all) makes it clear that at least three vicariant forms are recog- 
nizable. The form occurring from Port-au-Prince north to Cap 
Haitien and Port-de-Paix is the one to which Mertens has shown 
that the name ricordii Dumeril and Bibron must be attached. 
Another, occurring in the north and east of the Dominican 
Republic, may be called, following Mertens, by Cope's name 
haleatus (Eupristis taleatus Cope 1864, Proc. Acad. Nat. Sci. 
Phila., p. 168, type locality "St. Domingo"). A third unnamed 
population occurs on the Barahona peninsula. A fourth popu- 
lation inhabiting the southwest peninsula of Haiti may be dis- 
tinct. I list below the distinguishing characters of the three 
well-marked forms. 



BREVIORA 



No. 155 



ricordii 

Very low nuchal and 
dorsal crests 

Nuchal crest scales as 
long as or longer than 
high, and not or but 
little higher than the 
very low dorsal crest 

Head scales small, 
numerous (7-9 across 
snout at level of second 
canthal 2) 

In S S deep hlacTc 
spots above shoulder, 
sometimes also on occi- 
put ; no other evident 
pattern 



Table 1 

Barahona 
population 

Low nuchal and dorsal 
crests 

Nuchal crest scales 
higher than long but 
small, not higher than 
the weakly developed 
dorsal crest 

Head scales larger, 
fewer (4-6 across snout 
at level of second 
canthal) 

Whole of body in both 
sexes with very irregu- 
lar small Matches and 
mottlmg 



haleafvs ^ 

A prominent nuchal 
crest, a variable but 
lower dorsal crest 

Nuchal crest scales 
much higher than long, 
always much higher 
than scales of dorsal 
crest 

Head scales large, few 
(2-5 across snout at 
level of second 
canthal) 

Both sexes with no 
evident pattern or 
transverse handing or 
reticulation 



CHARACTER ANALYSTS 

The scales on the snout are swollen, bosslike in all Hispaniolan 
giant anoles. The differences are solely in the size of these boss- 
like scales. The contrast is extreme in this regard between the 
giant anoles of northern Haiti {ricordii) and those of the north- 
ern Dominican Republic (haleatus) . The animals of the Bara- 
hona and southwestern peninsulas, however, are intermediate, 
overlapping in this regard the north Dominican (haleatus) and 
approaching the north Haitian populations. On this character 
alone it is not possible to separate every specimen of the north 
Dominican and the southern populations though there is a well- 
marked average difference. 

Number of scales across the snout is a measure of a more 
general feature — general scale size — - which is somewhat greater 
in haleatus than in ricordii. Mertens has cited a number of key 
regions, i.e. loreal region, base of the tail, etc., which show this. 

1 A. Salle, who collected the type of Eupristis haleatus worked both in the 
northern Dominican Republic and at the base of the Barahona peninsula (map 
of travels compiled by W. J. Clench). It was therefore necessary to confirm the 
application of the name by checking the characters of the British Museum type. 
Miss A. G. C. Grandison has courteously confirmed that the nuchal crest soales 
of the type are indeed significantly higher than the dorsal crest scales. 

2 I count the canthals forward from the anterior border of the orbit (see Fig. 1). 
The reverse count — from the anteriormost canthal back — is sometimes used, 
e.g. Oliver (1948). This, however, has the disadvantage that the scales here are 
small and variable and do not provide a stable starting point. 



1962 HISPANIOLAN GIANT ANGLES 3 

The snout, however, provides the clearest expression of this 
general feature and one which can be reduced to a simple count 
with an adequate numerical range (2-9). Mertens has used a 
similar transverse count but makes it directly in front of the 
eyes. I have chosen a transverse count somewhat further for- 
ward because this transverse line, anterior to the supraorbital 
semicircles seems to me more suitable as a place for a standard 
count utilizable for all species of Anolis. 





N,S- 

BflLEflTUS RICORDII 

Fig. 1. The two extremes of anterior head squamation in Anolis ricordii 
subspecies. The arrows indicate the place at which the count across the 
snout is to be made. 



The pattern of geographic variation of nuchal crest scales 
is not so simple. Again, the populations of Port-au-Prince and 
northern Haiti and northern Dominican Republic show the 
extremes, the crests of Port-au-Prince and of north Haiti popu- 
lations very low with a long anteroposterior base, while those 
of north Dominican Republic populations are tapering, high, 
short based anteroposteriorly. The southern populations could 
again be described as intermediate but on careful examination 
this is not quite accurate. Both the north Haitian and southern 
populations have a reduced nuchal crest as compared with the 
north Dominican population but the nature of the reduction is 
different. The nuchal crest scales of north Haitian animals are 
long-based, low, rounded scales, not at all tapering; they are 



4 BREVIORA No. 155 

sometimes shorter but usually slightly higher than the very 
low, long dorsal erest scales. In the giant anoles of the Bara- 
hona peninsula tlie nuchal scales are, in contrast, still short 
anteroposteriorly, relatively high, tapered, but very small, not 
appreciably higher than the tallest dorsal crest scales and no- 
tably smaller in area than the largest dorsal crest scales. 

The few specimens from the southwestern peninsula do not 
permit adequate analysis of the populations of this important 
geographic area. The four adults examined all come from the 
vicinity of Fond des Negres halfway along the peninsula. In 
squamation, these are very like the animals of Port-au-Prince 
and north Haiti. The one remaining specimen is a juvenile 
(78 mm snout-vent length) from the foothills of the Massif de 
la Hotte. Although it is less than half-grown, its dorsal and 
nuchal scales are appreciably higher and less broad based than 
those of the adults from Fond des Negres and indeed are very 
comparable to those of the Barahona population ; in addition, 
in this individual these scales are double and even triple even 
on the nape. It thus seems very possible that the populations 
of the middle of the southwest peninsula and those of its extreme 
end differ significantly. (See further discussion below.) 

The nuchal crest scales seem in all populations to be some- 
what variable. HoAvever, within each adequately-sampled popu- 
lation the range of variation is very characteristic and does not 
blur the distinctions tabulated above. Despite statements by 
Boulenger (1885) and by Mertens (1939), I do not find a clear 
correlation with sex ; females, for example, of haleatus do not 
consistently have lower nuchal crest scales than male haleatus 
of the same size. There is, however, clear ontogenetic change ; 
the smallest individual of haleatus at hand (AMNH 28651 from 
San Juan Bay, Samana, 41 mm snout-vent length) has the nuchal 
crest scales only incipiently enlarged and could not be recog- 
nized as a member of the haleatus population on that character. 
However, in haleatus the characteristic, tapering, spinelike nu- 
chal scales develop very early and even specimens little more 
than half grown (e.g. MCZ 57719, Santiago, S , 83 mm snout- 
vent length ; MCZ 5445, Samana Peninsula, S , 88 mm snout- 
vent length) are very readily recognizable. 

Dorsal crest scales are more variable than nuchal crest scales, 
and variable in a peculiar fashion : there is sometimes a regular 
alternation of relatively high triangular single scales and pairs 



1962 HISPANIOLAN GIANT ANGLES 5 

of miicli lower, more quadrangular scales.^ However, such a 
regular alternation occurs — if present at all — on only a por- 
tion of the back and there is present between nuchal crest and 
dorsal crest, on the one hand, and dorsal and tail crests, on 
the other, various irregular conditions with double and single 
crests erratically intermingled and scale types somewhat inter- 
mediate and rather irregularly so. In ricordii itself with a very 
low dorsal crest the double condition with all low scales is most 
frequent ; in haleatus the situation is individually very variable ; 
in the Barahona and the southwest peninsulas the double condi- 
tion of the dorsal crest predominates. 

The body color of live specimens and well-preserved alcoholics 
is probably useful. The usual specimen, however, requires much 
interpretation and its evidence must be received with some 
skepticism. In tabulating pattern, above, I have ignored all 
ill-defined discolorations and have tried to assess pattern on the 
basis of real aggregations of pigment rather than fortuitous 
darkening of random areas by formalin. Even with this qual- 
ification the problem is not simple : these are Anolis and have 
the power of color change ; an adequate discussion of their pat- 
tern would be possible only if the whole repertoire of color 
changes were known. I record my information below by popu- 
lation. 

Ricordii: Females from Port-au-Prince and north Haiti ap- 
pear to be plain green above with no markings of any sort. 
Males from Port-au-Prince are also mainly green (as preserved 
they may show an obscure, very fine brown reticulation) but 
are distinguished by a large, very characteristic patch of black 
above the shoulder but of ill-defined shape and varying extent. 
There may also be irregular black patches extending onto the 
back of the head. There may also be a white patch at the corner 
of the mouth. A Cap Haitien male (Senckenberg 10445) is quite 
similar to Port-au-Prince specimens. Of three males newly re- 
ceived from Ti Guinin a little to the east of Cap Haitien (MCZ 
66147-9) two have almost no black at the shoulder (only small 
and inconspicuous spots) and none at all on the head. The other 
male is as devoid of black spotting as any female. All show fine 
brown reticulation quite like that of similarly preserved Port- 
au-Prince males. 



1 In the tabulation above I have compared the nuchal crest scales with the 
highest crest scales at midbody. 



6 BREVIORA No. 155 

Balcatus: Schmidt (1921, p. 10) records specimens from 
Sanchez and Villa Rivas as "usually green with dark-edged 
transverse bands of light greenish yellow. ' ' Preserved specimens 
which have any definable pattern show this transverse banding 
which is present on dorsum, limbs and tail; it appears to be 
most persistent on the tail. It is very conspicuous in a specimen 
near hatchling size (AMNH 28651). There are frequently num- 
bers of small spots on the sides of the belly and on the venter. 
The MCZ Santiago series, though preserved in a light phase, 
show very irregular light banding much invaded by dark retic- 
ulations, but the bands are very prominent on limbs and tail. 

Barahona Population: I have unfortunately been unable to 
find any description of Barahona giant anoles in Hassler's notes. 
I must therefore rely entirely on the preserved specimens, which, 
however, are unusually consistent, though from several localities 
and with very different collection dates. Collected by Hassler, 
they differ strikingly from the taleatus specimens, most of which 
were also collected by Hassler. All Barahona specimens, and 
most clearly those from Valle de Polo, show irregular small 
blotches, very variable in tint and extent, scattered over the 
entire dorsum. The closest approach to this condition is seen in 
those haleatus which show reticulation. There are never in the 
Barahona specimens the large shoulder spots of ricordii, sensu 
stricto. 

One specimen (AMNH 51241 from Enriquillo) does show 
transverse banding, but more obscurely than in haleatus. The 
small blotches are also very obscure in this specimen, which 
therefore could not in this phase be recognized as a member of 
the Barahona population on its color. The nuchal and dorsal 
crest scales are, however, typical harahonae, and the hypothesis 
which I have adopted is that the blotching so characteristic of 
most preserved specimens is characteristic only of one of the 
phases of the color repertoire possible to this population. 

Population of the Southwest Peninsula of Haiti: Again the 
material is inadequate for proper analysis, the more evidently 
so since the few available specimens suggest that this is not a 
unit sample. The juvenile from the Massif de la Hotte is, as 
preserved, reddish or purplish brown with indistinct broken 
dark longitudinal lines on head and nape and more evident 
narrow brown transverse bands on the back, six between shoulder 
and groin, in pairs with very slightly lighter reddish-brown be- 
tween each pair. The tail is obscurely annulate. 



1962 HISPANIOLAN GIANT ANGLES 7 

111 contrast, the three adult females from the vicinity of Fond 
des Negres are essentially patternless (MCZ 66016 is dark and 
obscurely vermiculate, and USNM 72631, 72633 are plain light 
green except for a subocular half ring of scales that is conspicu- 
ously white). The single male is similarly nearly patternless 
but has also the subocular half ring of white and, in addition, 
a series of small black spots above the shoulder, very like a 
vestige of the large black shoulder spot of Port-au-Prince ani- 
mals and very like those of the two Ti Guinin males. 

TAXONOMIC EVALUATION 

There appear to be at least three distinctive populations of 
giant anoles, all readily separable by nuchal crest development 
and body color, less sharply separable by the size of the scales 
on the snout. These populations are allopatric and thus may be 
species or subspecies. 

The differences between typical haleatus and typical ricordii 
are such that they could well imply specific distinction. How- 
ever, all the populations south of the Cul de Sac Plain, both 
the rather distinctive Barahona animals and the very poorly 
known populations of the southwest peninsula, are to some de- 
gree intermediate between the two northern extremes although 
presenting some features that are their ovm. In thus bridging 
the morphological gap between the extremes, they strongly sug- 
gest that the Hispaniolan giant anoles belong to a single species. 

The giant anoles are scarce and local. It is not to be expected 
that intergradation between the several populations will be easy 
to demonstrate ; specimens from many of the critical intermedi- 
ate areas are conspicuously lacking. 

The Fond des Negres and Ti Guinin specimens, on the other 
hand, may be intergrade populations. The vestigial shoulder 
spots present in the two Ti Guinin males and the single Fond 
des Negres male suggest this conclusion, as does, in the latter 
case, the combination of nuchal and dorsal squamation most like 
typical ricordii with somewhat lower scale counts across the 
snout. No giant anoles are at hand from the northwest Domini- 
can Republic. This is the area in which intergradation between 
ricordii and haleatiis is to be expected. It is thus not at all sur- 
prising to find the first suggestion of loss of characters of the 
typical race just to the east of Cap Haitien. 



8 BREVIORA No. 155 

On the southwest peninsula, the Fond des Negres area is one 
in which such intermediate populations are to be expected, as 
recent collections from the area show. Thus, Dromicus parvi- 
frons parvifrons and D. parvifrons protenus appear to meet in 
this area and several anole races (to be reported later in this 
series) show intergradation at just this point. 

I thus interpret the Fond des Negres giant anoles as inter- 
grades between typical ricordii and a population to the west 
occupying the tip of the southwest peninsula. This western popu- 
lation I infer to be represented at present by the single juvenile 
specimen collected by P. J. Darlington in the foothills of the 
Massif de la Hotte. 

The wide-banded specimen from Enriquillo in Barahona is 
reminiscent of the banded color phase in typical haleatiis, but 
in squamation it is not intermediate and it is geographically 
quite unsuitable as a member of an intergradient population. 
Nevertheless, the presence of this color phase hints at a closer 
relation with ialeafus than otherwise could be inferred. 

The absence of other evidence or hint of intergradation is 
easil}^ accounted for by the gaps in the distributional record, 
and, granted the desirability of further evidence, recognition 
of subspecies status seems justified for typical ricordii, for 
baleatus, for the Barahona population and probably also for a 
population at the west end of the southwest peninsula of Haiti. 
The latter two populations are currently nameless. That from 
the southwest peninsula is at the moment very insufficiently 
known, and it would not now be appropriate to describe it. The 
Barahona population, on the other hand, is well recorded and 
may be formally named: 

Anolis ricordii barahonae new subspecies 

Type: MCZ 43819, Polo, Valle de Polo, Barahona, Dominican 
Republic, an adult female collected by W. G. Hassler, Septem- 
ber 1932, J. C. Armstrong donor. 

ParaUjpes: Valle de Polo, MCZ 56141, AMNH 51036, 51235-6; 
Herman's Finca near Paradis, AMNH 51231-3; Barahona, 
AMNH 50255-6, 50261 ; Halfway between Trujin and Enriquillo, 
AMNH 51230; Enriquillo, AMNH 51241. 

Diagnosis: A subspecies of ricordii distinguished from the 
typical form and from baleatus Cope by the nature of the nuchal 
crest (formed by small but slender tapering scales not or very 



1962 HISPANIOLAN GIANT ANGLES 9 

little higher than the highest scales of the dorsal crest), by the 
size of the scales of the snout (4-6 across snout between second 
canthals), and by a characteristic phase of coloration in both 
sexes in which small blotches are present, irregular in shape and 
of varying intensity. 

List of Specimens Examined: A. r. harahonae (17 specimens). 
DOMINICAN KEPUBLIC. Polo, Yalle de Polo: ]\ICZ 43819; 
Valle de Polo: AMNH 51036, 51235-7, MCZ 56141; Barahona: 
AMNH 50255-6, 50261; Herman's Finca near Paradis: AMNH 
51231-3 ; Halfway between Trujin and Enriquillo : AMNH 51230 ; 
Enriquillo: AMNH 51241; locality uncertain: AMNH 51229, 
51234, 51240. 

A. r. ricordii (12 specimens). HAITI. Port-au-Prince : AMNH 
49501; Diquini: MCZ 8619, USNM 118902, 123988; Source Le- 
clerc, Morne Decay ette, near Port-au-Prince: MCZ 65729-31; 
PetionviUe: MCZ 60013-4; 3Iarcaco: USNM 69437; Port-de- 
Paix: MCZ 63338; Ti Guinin near Cap Haitien: MCZ 66147-9. 
[Records by Mertens : Port-au-Prince, 4 specimens; Cap Haitien, 
1 specimen.] 

A. r. haleatus (57 specimens). DOMINICAN REPUBLIC: 
Pena, near Santiago: MCZ 57713-9; Sayitiago: MCZ 7831; Las 
Bracitas: AMNH 41465-6; El Rio: AMNH 41294, USNM 62104- 
5; Rio San Juan: USNM 74940-1; Samaria Peninsula (various 
localities) : MCZ 5445, AMNH 28651, 39807-15, 39817-23, 39825- 
9, 39837, 40224-30, 40387-90, 42285, 42775, 44841-4; La Romana: 
MCZ 16321. [Records by Mertens: Santiago, 5 specimens; Moca, 
4 specimens; Finca Arhol Gordo near Villa Altogracia, 1 speci- 
men.] 

A. r. subsp. nov. (1 specimen). HAITI: Foothills, Massif de 
La Hottc: MCZ 38217. 

A. r. subsp. nov. x r. ricordii. Pemel near Fond des Negres: 
MCZ 66015-6 ; Fond des Negres: USNM 72631, 72633 ; also 72632, 
skeletonized. 

BEHAVIOR 

The giant AnoUs of Hispaniola is infrequently seen by collec- 
tors and consequently our information on its ecology and habits 
is limited. Such information as is available is summarized below 
by A. S. Rand. Quotation marks refer to Rand's personal ob- 
servations. 



10 BREVIORA No. 155 

A. ricordii ricordii, Morne cle Cayette, near Port-au-Prince, 
Haiti (A. S. Eand and J. Lazell). " A large individual was 
seen about 15 feet up on a branch of a 30-foot high tree on the 
edge of a patch of brush on the hillside. It remained sitting 
quietly in sight for about half an hour until we climbed the 
tree to attempt to capture it. It then climbed out among the 
small branches where, desi3ite an intensive search, we were un- 
able to locate it again." 

Port-de-Paix, Haiti (A. S. Rand and J. Lazell). "A female 
was seen sitting head down on the trunk of a large tree about 
4 feet from the ground. The tree was one of a series forming a 
fence row. The lizard allowed us to approach and noose it." 

A. ricordii haleatus, Pena near Santiago, Dominican Repub- 
lic (C. E. Ray and A. S. Rand). "Local people who brought in 
a series at this locality reported that they lived among the upper 
branches of the larger shade trees in the coffee plantations." 

Finca Arbol Gordo, Dominican Republic. Mertens reports 
that the single specimen collected by him jumped to the ground 
from the trunk of a palm tree. Mertens kept this animal in cap- 
tivity for some time and wrote that it was active only in the 
late afternoon. When approached it squirreled around to the 
other side of its perch. It slept exposed on small branches with 
its hind legs flexed. 

A. ricordii subsp., Camp Perrin, Haiti (A. S. Rand and J. 
Lazell). "A small boy guided Rand to a stand of coffee where 
a crowd of Haitians had treed a large individual. The lizard 
was about 20 feet up in a small tree. It was too high to noose 
and the tree too small to climb so we attempted to scare the 
lizard to a more favorable situation by poking it with a long 
pole. After considerable fuss the lizard ran out along a small 
branch to the next tree, down that to the ground, across the 
ground to hide among the roots of a large Cieba tree. After 
considerable prodding among the roots we dislodged the lizard 
which avoided the herd of assistants successfully and climbed 
the Cieba out of both sight and reach." 

Above Maceline, near Camp Perrin (A. S. Rand and J. Lazell). 
" In a small stand of coffee we saw a single animal about 15 feet 
up in one of the large trees on a large branch. When Lazell 
climbed the tree it squirreled about the branch concealing all 
of itself except one eye. It then climbed up and out among the 
small branches and disappeared from ^•iew." 



1962 HISPANIOLAN GIANT ANGLES 11 

Mertens speculates that the giant anoles of Hispaniola were 
originally inhabitanis of the lowland rain forest and have 
adapted to cultivated areas by living in the crown of the taller 
trees. The observations of Kand and Lazell tend to support this 
conclusion. 

INTRAISLAND ZOOGEOGRAPHY 

The portion of Hispaniola south of the Cul de Sac Plain, in- 
cluding the southwest and Barahona peninsulas and the La Selle 
and La Hotte ranges, has, as Mertens has emphasized, a number 
of endemic species and even genera. I have recently (Williams, 
19(jl) stressed the importance of this area, which I have called 
the "southern island," in the initial ditferentiation of forms 
now widespread in the portion of the island north of the Cul 
de Sac Plain. 

Barahonae as an isolate in "southern island" easily fits this 
pattern. Baleatus and typical ricordii do not. 

This is merely the first of many examples to be discussed in 
this series which will show that not all the patterns of differ- 
entiation in Hispaniola can readily be explained by the simple 
hypothesis of "southern" and "main island" isolates. As in 
the present case so in a considerable number of others, there 
appear to be additional loci of differentiation, e.g. northern 
Haiti, which are not so evidently marked off' by present or 
recently past barriers to distribution. 

These cases of subsidiary loci of differentiation are not at 
present well analyzed. Hispaniola has not been really well col- 
lected, and it is now evident that the differentiation of local popu- 
lations that occurs within it is finer grained than the sporadic 
collecting of the past could reveal. For many of the instances 
of minor loci of differentiation, data are just now being gathered. 

RELATIOXSHIPS OF EICORDII AND THE OTPIER 
WEST INDIAN GIANT ANOLES 

Anolis ricordii clearly belongs to the series of giant anoles 
that includes equestris of Cuba and cnvicri and rooscvclti of the 
Puerto Rican-Virgin Island complex. Aiiolis garmani of Jamaica 
seems, even on externals, to show no features, except size, which 
specially link it to this group, and Richard Etheridge in an 
unpublished thesis (University of Michigan) has shown that 
osteologically it belongs to quite a dift'erent section of the genus. 



12 BREVIORA No. 155 

No full study has ever been made of the Greater Antillean 
giant anoles as a group, though they clearly merit such a study. ^ 
I do not intend to attempt this here, but in order to place the 
races of ricordii within a frame of reference which might add to 
our understanding of them and of their origin I have compared 
all of the species for a number of mostly qualitative characters 
which were simple to determine and to evaluate. Table 2 presents 
the results of this comparison. 

It is sufficiently clear even from the limited evidence presented 
by Table 2 that we are dealing with four good species. The 
Hispaniolan populations are a unit as compared with the forms 
of the neighboring islands and, as in the case of the other poly- 
typic giant anole, A. equestris, the intraisland differentiation 
has involved primarily scale size and color pattern. 

There is one unusual zoogeographical feature in the distribu- 
tion of the giant anoles — the occurrence of A. cuvieri and A. 
roosevelfi on the same island l)ank, cuvieri on the mainland of 
Puerto Rico, roosevelti on Culebra in the Virgin Islands. These 
two are very distinct species. Their differences are at another 
level than those between ricordii and haleatus, and their occur- 
rence on what must at various times during the Pleistocene have 
been a single land mass provides a special problem in accounting 
for their origin and dispersal. 

ACKNOWLEDGMENTS 

This paper was in part made possible by collections made 
under National Science Foundation Grants NSF G-5634 and 
G-16066 and grants from the American Philosophical Society 
and the Society of Sigma Xi. Assistance given by the govern- 
ments and people of the Haitian and Dominican Republics is 
also gratefully acknowledged. I am indebted also to Charles M. 
Bogert and Doris M. Cochran for loans from the collections 
under their care. Dr. Norman Ilartweg permitted examination 
of the second known specimen of Anolis roosevelti Grant. Dr. 
W. J. Clench provided information on the areas visited by A. 
Salle in the Dominican Republic. 



1 For further information on the non-Hispaniolan species see Stejneger (1904), 
Grant (1931), Schwartz (1958). 



1962 



HISPANIOLAN GIANT ANGLES 



13 






S X 



13 



3 
o 



<D 



O 

o 



or! 

'ctf 
02 






cS 



«o 



o •- a 



ai 



Si 

CD 

I - ^ 

o  



-I s ^ s 



r- ^ O 






o 
o 

s 

CD 






03 2: 



M 



35 



bfl 


c3 


'C 


S 


ei 


QJ 


• rH 




cS 


-M 


'S 




Si 


<S 


;h 


<D 


M 


« 


> 


i^ 



CO 



o 
•■o 



AS 






> 



I ^ 



CC 



-G 
GQ 



O) 



a 



lO ^ ^ 



CO 



o 



o 
o 






< — I © 
32 rs 



.9 

cS 



rt ii 



O 



*o 












60 c 



00 

« 



o 

05 






oj c: 



a ^ oj 



1> 



^ i: -S 02 '^ 

p: m « .s a S m 



CO 

CO 



a 

a 

CO 































u 






















^^ 


'-; 










o3 






(M 
















"3 










<u 


"3 






H 




'^ 














^ 








■*^ 


<SJ 






5 

< 






0) 








(D 

'o 


5: 

p 


-^- 




*o 




c5 




33 

o 




H 


••s 


r— < 


a> 


o 


«o 


02 

o 
02 




-*— 
S 


^ 


-M 

fl 


-4^ 


3 


S 

bJO 




"T? 






02 


O 


Ol 


TiH 


3 

02 


3 


o 
o 

i 


CO 

9 


o 

o 

CD 


CD 


S 


> 


0) 



CO 



a 
a 

CO 





a? 

03 












IE 

o 










^ 












O 










^ 


-a 










rS 












M 










QJ 


O 










■3 












3 










^ 


03 








<D 


be 












^ 




















-M 


QJ 




















IB 


O 


'73 




^ 




c8 

'T3 


rH 








a 


■to 


s 

'o 
02 


in 


o 
!5 




'o 

OD 
35 

02 

9 


3 

3 
02 


.2 

s 

s 


o 
o 


•73 


O 
O 

s 

02 

'5 


c3 
3 
cr 


• rH 

13 
be 

'•w 

Eh 

03 


p 

s 

> 






CD 




a 

l-H 



;h 



« S5 S O 










• rH 





g 








3 


OQ 


^ 










05 


03 


=H 


03 








73 














03 

p 






-3 
-4^ 


03 


;-i 












1— H 


<o 


c:! 


^ 

K 




;h 


3 


=s 


+J 


i) 




a 

Sh 

3 
1— 1 


Sh 


.(H 


OS 

»H 

3 


03 


03 

u 
02 


3 
p 




!* 

ft 
3 

03 


CO 



O 



it 

.2 3 
S - 

4— 1 — ' 

O C3 

;ih 02 



O 

O) 



73 O 
tS 05 



CQ 




t-t 




bc 




^ 




03 


''""^ 


3 




a 




3 


d^ 

^ 


<o 






O! 
<B 


73 


3 


s 

X 



;-, 

& 


3 
> 

3 




'rt 


T! 


3 


a 





2 





a 


cS 


cS 


3 


-^ 


02 


a 


iJ 




02 


ci 



14 BREVIORA No. 155 

REFEEENCES CITED 

BOULENGER, G. A. 

1885 Catalogue of the lizards in the British Museum (Natural His- 
tory). London, vol. 2, i-xiii + 1-497. 
Grant, C. 

1931 A new species and two new subspecies of the genus Anolis. 
Jour. Dept. Agr. Puerto Rico, 15: 219-222. 
Mertbns, R. 

1939 Herpetologische Ergebnisse einer Reise nach der Insel His- 
paniola, Westindien. Abhandl. Senckenb. Naturf, Ges., 449: 1 84. 
Oliver, James A. 

1948 The anoline lizards of Bimini, Bahamas. Amer. Mus. Novitates, 
no. 1383: 1-36. 
Schmidt, K. P. 

1921 Notes on the herpetology of Santo Domingo. Bull. Amer. Mus. 
Nat. Hist., 44: 7-20. 
Schwartz, A. 

1958 A new subspecies of Anolis equestris from Eastern Cuba. Herpe- 
tologica, 14: 1-7. 
Stejneger, L. 

1904 The herpetology of Puerto Rico. Rep. U.S. Nat. Mus., 1902: 
549-724. 
Williams, E. E. 

1961 Notes on Hispaniolan herpetology. 3. The evolution and rela- 
tionships of the Anolis semUineatus group. Breviora, No. 136 : 
1-8. 



1962 



HISPANIOLAN GIANT ANGLES 



15 










5D 


03 




10 












^i) 






o 


C> 




^ 


n, 




v 


en 




<) 






;^ 


^^ 




VK- 


6 






ai 




^ 




73 






^<i 


<\> 




P3 


O 






N- 




O 


Tl 




s- 


MO 




+J 








C3 




o 

g 


O 




lO 






<D 




ZD 


V 

8 


1 




CS 


o 


o 






-■c 


o 


g 


|^) 


NU 




<u 


o 


-*^ 










^ 

-4-S 




^ 




r- 


TS 


ca 

•f-5 


1 


'^ 








ci 


<M 


^Kj 






tn 




r 




c 




4) 




o 


'-i 


o 


O 


02 


r^ 


^ 


^< 


n:^ 






-4-' 


y 


N-J 


C3 


"^ 


CT( 


GJ 


i~. 


O 






A 


O 


« 


•O 


^-^ 


O 


O 


n- 


rO 


^) 


^ 


'W 


1=; 








Ci 




o 


rr 










o 




 


 


(0 
C7^ 




o 

-4— 
— 




■S 


^ 


O 


0) 


3 

ft 

O 

Pi 




c: 


en 


o 


"^ 








-^ 


c 












o 


•4^ 




<!i) 


^ 


02 


■*-' 


3 




^ 
^ 


1 


"S 


-t- 


00 


v, 


■;i 




& 


bi 


73 




<o 




tn 


'r^ 


O) 


No 

o 




s 


3 


o 


O 


^ 


V 




O 












^ 




o 




c 








03 


IW 






v 


-^- 




.o 






^ 


"Vl 


5 
o 










o 

Ol 


to 




S 




V! 


, 


1 






^ 


O 


1 






=4-1 


M 


O 




<o 




o 






•-^ 


;> 




J3 


— ' 


b 


Tl 


MJ 




o 


r* 


C! 


t. 


o 




.^ 


o 




8 








03 


!^ 


fO 




1-1 


a 





D O 



fi ►^ 12; 



BREVIORA 



seum of Comparaitive Zoology 



Cambridge, Mass. April 13, 1962 Xumber 156 

THE FOSSILIFEROUS TRIASSIC DEPOSITS OF 

ISCHIGUALASTO, ARGENTINA, AND PRELIMINARY 

DESCRIPTION OF ISCHIGUALASTIA, A NEW 

GENUS OF DICYNODONT 

By Alfred Sherwood Romer and C. Barry Cox 



THE FOSSILIFEROUS TRIASSIC DEPOSITS 
OF ISCHIGUALASTO, ARGENTINA 

By Alfred Sherwood Romer 



Early in 1958, a joint expedition of the Museo Argentino de 
Cieneias Naturales of Buenos Aires and the Museum of Com- 
parative Zoology, Harvard University, was organized to explore 
continental deposits in the western Argentine in search of fossil 
vertebrates. The personnel included, for the Buenos Aires Mu- 
seum, Dr. Guillermo del Corro, geologist, and Mr. Orlando 
Gutierrez, assistant ; from Harvard, Professor Bryan Patterson, 
preparators Arnold C. Lewis and James Jensen, Mrs. Romer and 
the writer. The earlier part of the trip was spent in the general 
region of Mendoza. In late April it was decided to move our 
base northward and explore the valley of Ischigualasto. 

This region lies some 300 miles to the northeast of Mendoza, 
in eastern San Juan Province, close to the border of La Rioja, 
in the department of Yalle Fertil, at a latitude and longitude 
of approximately 68°W, 30°5'S. No adequate maps of this area 
exist; Ischigualasto is shown on the San Juan sheet (no. 37) 
of the 1/500,000 map of Argentina, but the details are highly 
inaccurate. Frenguelli (1948, fig. 1, pis. 1, 2) gives sketch maps 
showing the general topography of the valley and its relation to 
adjacent regions. The name derives from an Indian village once 



2 BREVIORA No. 156 

present at the marofin of the valley. This, however, disappeared 
long since and the valley is oompletely uninhabited. In 1874 
Stelzner (1885, pp. xii, 74-75) rapidly traversed the valley on 
mnle-back, presumably following a trail which runs from Jachal, 
in San Juan Province, through the valley and on eastward to 
Patquia in La Rioja. Bodenbender (1911, pp. 94-113) similarly 
crossed by this trail several decades later. A small coal seam 
had been discovered in the hills west of the valley at a locality 
named los Rastros because of dinosaur-like footprints discovered 
there; Huene (1931) was taken, in 1927, on a hasty trip to see 
these footprints ; he traversed the valley but was unable to stop 
there. Until recent years nothing was known of the geology of 
the region except for such observations as Stelzner and Boden- 
bender could make as they crossed it, and nothing of fauna and 
flora except the footprints just mentioned and a few plants col- 
lected by Bodenbender. 

The reason for this former paucity of knowledge is obvious. 
The valley is extremely arid, and in the days of mule travel, a 
stay there was impossible because of the almost complete lack of 
water and fodder.^ With the coming of automobile transporta- 
tion and, particularly, of vehicles with four-wheel drive, the 
situation has changed radically, and in 1948 Frenguelli was able 
to publish a rather comprehensive account of the stratigraphy of 
Ischigualasto as the result of exploration of the area in 1943, 
primarily in the interests of paleobotany. 

During the course of his work there, three fragments of cyno- 
dont skulls and jaws were recovered, and were described by 
Cabrera (1943). These were of interest as indicating the pres- 
ence in Argentina of Triassic beds similar in fauna and presumed 
age to those of southern Brasil. But since so little reptile ma- 
terial was collected during Frenguelli 's survey of the region, it 
seemed unlikely that work there would be profitable for the ver- 
tebrate paleontologist. 

My attention was first called to Ischigualasto by Professor 
Huene before the results of Frenguelli 's work became known. 
As noted above, Huene had crossed the valley on mule-back 
during his inspection of the los Rastros footprint site and, al- 
though unable to stop and prospect, had been struck by the 



1 Bodenbender (1911, p. 96) breaks off in tbe middle of his description of the 
region to insert a paragraph of emphatic and italicized warning: "NOT A — A 
los futuros exploradores hago presente que no se puede contar . . . con sufl- 
ciente pasta para los anUnales," etc. 



1962 TRIASSIC DEPOSITS OF ISCHIGUALASTO 3 

seemingly favorable nature of the beds for vertebrate explora- 
tion. Like our Argentinian colleaj^ues, I had failed to be im- 
pressed as to prospects by the few materials obtained by 
Frenguelli. A different light, however, was shed upon the possi- 
bilities by a publication by Heim (1949). He had been commis- 
sioned in 1944 to report on the los Rastros coal mine, but in 
crossing the Ischigualasto valley had become so interested in the 
region that a considerable part of his report is devoted to the 
Ischigualasto beds and, incidentally, to the fossils found there. 
He photographed a cynodont skull in situ (pi. VIII, fig. 2) and 
says (p. 22) : "El senor de la Vega me llamo la atencion sobre 
los restos de vertebrados que se han hallado frecuentemente en 
el suelo de las arcillas, especialmente en la parte media de la 
formacion. Se presentan como acumulaciones de fragmentos de 
huesos, de color pardo oscuro. A veces aim se distingue la forma 
del craneo y de la dentadura con restos de dientes negros, pero 
sin que se pueda restaurar esqueletos." 

This report indicated that investigation of the Ischigualasto 
region might prove profitable, and our expedition moved to that 
region. A road of passable quality runs to Valle Fertil, a village 
south of the valley, which serves as departmental capital. Be- 
yond this, settlement is sparse and presently ceases altogether, 
the roads deteriorate into a confusing series of trails, and it was 
necessary to hire a local guide to reach our destination.^ At 
Cerro Morado, about 50 km. north of Valle Fertil, there is 
reached the southern edge of the valley which extends some 20 
km. to the north-northwest. At its southern end its width is 
about 7 km.; it narrows to 21/2-3 km. at the north. Its western 
boundary is a range of rugged sandstone hills, the eastern boun- 
dary an unbroken cliff of red sandstone, "los Colorados," strik- 
ing in appearance, with a height, for much of its length, of 150 
to 200 meters. The valley itself, almost bare of vegetation except 
for scattered thorn bushes, exhibits an expanse of shales of 
varied pastel colors and occasional sandstones. It is drained 
by a series of broad sandy arroyos which converge at about the 
center of its western margin where, at Agua de la Pena, there 
is a small stream of water (non-potable, according to our guide). 
From this point the deep gorge of the Rio de la Peiia runs west- 
ward through the hills to the Rio Bermejo. Northwestward from 

2 Since our work there, a new highway has heen constructed that links Patquia 
with Pogancilla and Chilecito. This passes a little to the northeast of the Ischi- 
gualasto basin, rendering it comparatively accessible. 



4 BREVIORA No. 156 

the main valley extends the narrower valley of the Rio de la 
Chilca, which turns westward to join tlu> Bermejo. 

We made camp near Agua de la Pena, began exploration of 
the valley — and were immediately astounded by the abundance 
of vertebrate remains which it contained. Every vertebrate 
paleontologist dreams of finding, someday, a virgin territory 
strewn with fossil skulls and skeletons. Almost never does this 
dream come true. To our amazement and delight, it did come 
true for ns at Tschigualasto. All about us, in the clays of the 
valley, were exposed specimen after specimen of fossil reptiles. 
In most instances the greater part of the time of a field party 
is taken up in prospecting for specimens. Here little of this 
sort of w^ork needed to be done, and our energies could be de- 
voted to excavating the better specimens chosen from the wealth 
of materials readily available. Approximately six weeks were 
spent here. Our work was hampered by the shortness of the day- 
light hours at this season of the year, and by the fact that a 
number of trips to Valle Fertil, each taking a full day, were 
necessary to carry out fossil blocks and obtain food supplies, 
gasoline and water. Nevertheless, well over 100 specimens, mainly 
plaster blocks containing skeletons, partial skeletons or skulls, 
were recovered. 

The geology of the region has been described by Frenguelli 
and by Heim, and discussed by Groeber and Stipanicic (1952, 
pp. 87-99). Apart from faults of a minor nature, the geologic 
structure is simple, the sediments uniformly dipping gently to 
the east-northeast with, in consequence, a regular succession of 
beds from west to east. Toward the Bermejo, in the western 
hills (an area not visited by us), are beds thought to be of Car- 
boniferous and Permian age — the Paganzo beds of Bodenbender 
— followed by the Estratos de Ischichuca.^ Farther east, the 
hills bordering the Tschigualasto valley are formed by the Estra- 
tos de los Rastros, dominantly sandstones, which conformably 
overlie the Ischichuca and include the coal seam and footprint 
locality mentioned. The shales occupying the valley constitute 
the Estratos de Ischigualasto, with a thickness estimated at 400 
to 500 meters; it is in this formation that nearly all the verte- 
brate fossils of the region have been found. The steep cliffs at 
the east side of the valley constitute the lower part of the Estratos 
de Gualo of Frenguelli, for which Groeber and Stipanicic prefer 
the designation of Estratos de los Colorados. The four upper 

3 The beds beneath Cerro Morado may be the Isehichuca. 



1962 TRIASSIC DKPOSTTS OF TS( HIGUALASTO 5 

formations, at least, of the series — the Estratos de Isehiehuea, 
de Ids Rastros, de Ischiguahisto, de los Colorados — are obviously 
parts of a sinole sedimentary cycle, Avithout evidence of any dis- 
conformity. In the region of Agua de la Pefia a local fault 
(Frenguelli, 1948, pi. II. profile A) obscures the transition be- 
tween los Rastros and Isehignalasto formations. But elsewhere 
(as Frenguelli 's profiles B-D, pi. IV) deposition can be seen to 
have been uninterrupted, and (apart from a conglomerate bed 
which is taken as the upper boundary of the los Rastros forma- 
tion) the transition is marked mainly by a diminution in im- 
portance of the sandstones which are so prominent in the los 
Rastros. The uppermost beds of the los Rastros yielded frag- 
mentary verteV)rate remains which appear comparable to those 
of the Tschigualasto formation. The typical beds of the Estratos 
de los Colorados contrast strongly with those of the Estratos de 
Ischigualasto in color, predominance of sandstones, and re- 
sistance to erosion ; but as Frenguelli notes, a zone of transition 
is apparent at the base of the cliffs. 

Stelzner "lumped" the entire series of beds found in this re- 
gion as "Rhaetic" (in the broad sense in which that term has 
frequently been used in Argentinian geology). Bodenbender 
believed the "Gualo" beds to be Cretaceous, the Ischigualasto 
Jurassic, the los Rastros "Rhaetic." Frenguelli, more correctly, 
considers the series as a whole to be older, the "Gualo" to be 
Rhaetic or lower Liassic, the Ischigualasto to be upper Keuper, 
the los Rastros to be lower Keuper. Groeber and Stipanicic 
(table I) believe the spread in time of deposition of these beds 
to be rather narrower ; the los Colorados are assigned with doubt 
to the Rbaetic, and l)oth Tschigualasto and los Rastros to the 
upper Norian — i.e., the uppermost Keuper. The vertebrate re- 
mains suggest a lower position (Romer 1960a, pp. 1291-1292; 
1960b, pp. 86-87) although full discussion should be postponed 
until the fauna has been more thoroughly studied. The Norian 
stage is one in which is found the typical Upper Triassic dino- 
saurian fauna ; the Ischigualasto fauna is, on the contrary, one 
in whicli there is little evidence of dinosaurs and in which 
gomi^hodont cynodonts and rhynchosaurs are dominant. The 
Ischigualasto formation is essentially comparable to the Santa 
Maria beds of Brasil and the Manda beds of Tanganyika. It is 
surely pre-Norian and not improbably pre-Carnian ; the gompho- 
dont-rhynchosaur faunas would appear to be essentially Middle 
Triassic in age. 



6 BREVIORA No. 156 

By agreement, the entire collection was shipped to Cambridge, 
where it is being prepared. All types and representative speci- 
mens of all forms found will be deposited in the National Museum 
in Buenos Aires. Complete preparation, however, will be a 
lengthy process, not only because of the considerable quantity 
of material but also because of the refractory nature of the mat- 
rix enclosing a large proportion of the specimens. In general, 
publication of the scientific results will not appear for some 
time, since we do not wish to publish before preparation has pro- 
ceeded to the point at which all material of a given form has 
become available for study. In the case of the dicynodonts, how- 
ever, the material included only a few specimens of a single large 
form; these have been prepared, and have been studied by Dr. 
Barry Cox, of King's College, University of London. A prelimi- 
nary description is appended ; it is hoped that a full account 
may be published within the year. 

It is gratifying to us to have been instrumental in opening 
up a new Argentinian area for exploration by vertebrate work- 
ers. Since our trip, several further expeditions to the region 
have been made by the University of Tucuman, under the direc- 
tion of Dr. Oswaldo Reig, with successful results ; certain ma- 
terials collected on these later expeditions have already been 
described (Reig 1958, Casamiguela 1960). There are vast areas 
of late Paleozoic and early Mesozoic deposits in the western Ar- 
gentine which have never been visited by vertebrate paleontolo- 
gists. Although the chances of finding beds as unusually 
productive as those of Ischigualasto are not great, it is highly 
probable that other faunas which will aid in rounding out the 
early history of vertebrates in Argentina await discovery. 

I have elsewhere (Romer 1960a) expressed our thanks to a 
number of friends who aided us in the Mendoza region. We are 
further deeply grateful to various other persons who aided the 
general work of the expedition and our exploration of Ischigua- 
lasto. The cordial cooperation of Dr. Adolfo D. Holmberg, then 
Interventor, and members of the staff of the Buenos Aires Mu- 
seum, was much appreciated. Professor Rosendo Pascual of the 
La Plata Museum accompanied us during the early portion of 
the trip. Much valuable scientific information was given us by 
the geologists of the Yacimientos Petroliferos Fiscales, Comision 
Nacional de Energia Atomica and the Direccion Nacional de 
Mineria, including, among others, Drs. Pablo Groeber, Pedro N. 
Stipanicic, Martinez Cal, Hector de la Mota, Luis A. Barrio- 
nuevo, and Vicente Ferreiro. The Comision Nacional de Energia 



1962 TRIASSIC DEPOSITS OF ISCHIGUALASTO 7 

Atomica aided us in the difficult matter of water and gasoline 
supply at Ischigualasto. Dr. Mario E. Terrugi aided us greatly 
in many regards, and the Harvard members of the expedition 
appreciate very much the hospitality extended to them by Dr. 
and Senora Terrugi and by the late Dr. Bernhard Dawson and 
Senora Dawson of La Plata. Our expedition was made possible by 
grants from the National Science Foundation and Life magazine. 



REFERENCES CITED 

BODENBENDER, G. 

1911. Con8tituci6n geol6gica de la parte meridional de La Rioja y re- 
giones limitrofes (Republica Argentina). Bol. Acad. Nac. Cienc. 
C6rdoba, 19:1-220. 
Cabrera, A. 

1943. El primer hallazgo de terapsidos en la Argentina. Notas Museo 
La Plata, 8, Paleont. no. 55:317-331. 
Casamiquela, R. M. 

1960. Noticia preliminar sobre dos nuevos estagonolepoideos argen- 
tinos. Ameghiniana, 2:3-10. 
Frenguelli, J. 

1948. Estratigrafia y edad del llamado Retieo en la Argentina. Gaea, 
8:159-309. 

Groeber, p., and p. N. Stipanioic 

1952. Triasico, pp. 13-141 in Geografia de la Republica Argentina. 
Buenos Aires, Soc. Argentina Estudios geog. Gaea, 2 (1):1-541. 
Heim, a. 

1949. Estudio geologico del Carbon "Retieo" y del Valle de la Pena 
(Provincias de San Juan y la Rioja). Ministerio de la Industria 
y Comercio, Bol. Dir. General de Industria Minera, 89:1-31. 

HUENE, P. 

1931. Die fossilen Fahrten un Rhat von Ischigualasto in Nordwest- 
Argentinien. Palaeobiologica, 4:99-112. 
Reig, O. a. 

1958. Primeros datos descriptivos sobre nuevos reptiles arcosaurios del 
Triassic de Ischigualasto (San Juan, Argentina). Rev. Assoc. 
Geol. Argentina, 13:257-270. 

ROMER, A. S. 

1960a. Vertebrate-bearing continental Triassic strata in Mendoza region, 
Argentina. Bull. Geol. Soc. America, 71:1279-1294. 

1960b. Explosive evolution. Zool. Jahrb., Syst. Bd., 88:79-90. 
Stelzner, a. 

1885. Beitrage zur Geologie und Palaeoutologie der Argentinischen 
Republik. Geologiseher Theil. Capel & Berlin, xxix -f 329 pp. 



8 BREVIORA No. 156 

PRELIMINARY DIAGNOSIS OP LSCHIGUALASTIA, A 
NEW GENUS OF DICYNODONT PROM ARGENTINA 

By C. Barry Cox 

Department of Zoology, King's College, University of London 

Above is given a brief account of a joint Buenos Aires-Har- 
vard expedition to the Valley of Ischigualasto, San Juan 
Province, Argentina. Among the remains collected from the Ischi- 
gualasto formation, presumably of Middle Triassic age, were 
several skulls and parts of posteranial skeletons belonging to a 
new genus of large dieynodont. A preliminary diagnosis of this 
new genus follows below; it is named Ischigualastia jenseni after 
Mr. James Jensen, who was responsible for the extremely pains- 
taking collection and preparation of this material. 

Ischigualastia jenseni, gen. et sp. nov. 

Holotype of I. jenseni: Number 18.055, Museo Argentino de 
Ciencias Naturales, consisting of skull and partial skeleton. 

Geological Horizon and Locality: Ischigualasto formation 
(Triassic), approximately 100 m. above the base of the forma- 
tion ; about 2 km. north of Agua de la Peiia, Ischigualasto Valley, 
Department of Valle Pertil, San Juan Province, Argentina. 

Genotype: Ischigualastia jenseni Cox. 

Generic and Specific Diagnosis: Large dicjaiodont (type skull 
55 cms. long, 46 cms. broad). No teeth in upper or lower jaws. 
Skull triangular in dorsal view, greatest width across occiput. 
Very wide interorbital region, very narrow intertemporal re- 
gion. Tapering snout, without nasal ridges or bosses. No pineal 
boss, but a slight mound in front of pineal foramen. No post- 
frontal bone. Preparietal bone probably present. Interparietal 
forms whole of posterior half of intertemporal bar, widely sepa- 
rating squamosals from postorbitals. No sharp median inter- 
temporal ridge. Zygomatic arches bowed outward. Sharp 
transition between dorsal and occipital surfaces. Occiput almost 
semicircular in outline. No tabular bone visible. Stapes lacks 
stapedial foramen. Short interpterygoid vacuity. No ectoptery- 
goid bone. Pterygoid broadly meets maxilla. Palatine and pre- 
maxilla meet, excluding maxilla from internal nares. Palatal 
surface of premaxilla bears pair of anterior ridges. Premaxilla 



1962 TRIASSIC DEPOSITS OF ISCHIGUALASTO 9 

extends some way anterior to maxilla. Aseendinji: portion of 
epipterygoid slender, not expanded to form part of lateral wall 
of braincase. No lateral winji: on dentary. Stout retro-articular 
process. 

Five sacral ribs. Acromion })rocess of scapula absent or ves- 
tigial. Coracoid foramen between precoracoid and scapula. 
Sternum constricted halfway along its length; dorsal surface 
bears bosses for attachment of ribs. Ulna has large olecranon 
process, with cartilaginous epiphysial union with rest of bone. 
Femur with well-developed head set off from rest of bone. 

A more extensive and illustrated account of Ischigualastia will 
appear later. 



BREVIORA 



Mmseiuim of Comparative Zoology 

Cambridgk, Mass. May 2S, 1962 Number 157 

A RHACIIITOMorS AMPHIBIAN, SPATHICEPHALVS, 
FROM THE MISSISSIPPI AN OF NOVA SCOTIA 

By Donald Baird 

Department of Geology, Piiiiceton University, 
Princeton, New Jersey 

In the past few years field parties from the Museum of Com- 
parative Zoology at Harvard College, supported in part by funds 
from the National Science Foundation, have discovered several 
deposits of fossil amphibians which date from the earlier half of 
the Carboniferous period. Most of the material recovered comes 
from redbeds of the I'pper Mississippian Mauch Chunk group in 
West Virginia (Romer, 1941) and Pennsylvania; more recently, 
amphibians have been found in beds of equivalent age at two 
sites in Nova Scotia. 

In continuation of the Harvard collecting program, field par- 
ties sponsored jointly by Princeton University and the Nova 
Scotia Museum of Science have collected additional bones and 
trackways of amphibians in Pennsylvanian as well as Mississip- 
pian beds. One of these finds is noteworthy because no member 
of the order Temnospondyli from the Mississippian system has 
hitherto been described. 

This specimen, the right half of a skull table preserved as a 
natural mold, comes from the same horizon and locality as the 
embolomerous lower jaw described as PhoUdcrpeton{1) hrcton- 
cnsc Romer (1958). The matrix is a flaggy calcareous siltstone 
characterized by well-sorted quartz grains and silvery muscovite 
mica. 



2 BREVIORA No. 157 

Su])erorder Labyrinthodontia 
Order Temnospondyli 

Suborder RHACHITOMI^ 

Family LOXOMMIDAE 

Genus SpATHICEPHALUS Watson 1!)29 

[Spathiocephalus Romer 1945, errore] 

Emended Diagiwsis. A loxomniid extremely specialized in its 
expanded cheek and snout, constricted interorbital area and 
shortened skull table, and having numerous small, chisel-shaped 
maro'inal teeth. 



e^ 



SpATHICEPHALUS PEREGER, U. Sp. 

Diagyiosis. Differs from N. minis Watson, the only other known 
species, in its relatively narrower skull table. 

Type. PU 17182, Princeton University Geological Museum. 

Occurreyice. Point Edward formation, Canso group, Upper 
Mississippian (probably early Xamurian). Beach of cove be- 
tween Point Edward and Keating C'ove, 4 miles northwest of 
Sydney, Cape Breton County, Nova Scotia. Collected by Donald 
Baird, William F. Take and Jane McN. Take, 1960. 

DESCRIPTION 

The dorsal surface of the skull table bears a deep, coarsely 
reticular sculpture rather similar to that of Megaloceplialus 
lineolatus (Cope) but with narrower ridges. Aside from the 
separation of prefrontal from postfrontal, the contacts between 
skull elements are normal for a loxommid. 

The narrow frontal l)0iie is Ijounded medially by a deep, 
straight suture which is ridged rather like the edge of a file. 
Anteriorly the frontal is beveled to receive the end of the nasal ; 
anterolaterally it bears a long, striated facet for the articulation 
of the prefrontal. As in other loxommids the prefrontal must 
have swelled laterally to form the antorbital process which dif- 

^ AssigiiiiU'iit of the T.oxoiiiinidno to the Rliac-hitoiiii is now (•(iiifiiiued 1iy 
the association of rhat-hitonious vertebrae with a skull of Mc/jaloeepliahis 
lineolatus (Cope) from Linton, Ohio (cf. Baird, 19.")7). 



1962 



NEW SPATHICEPIIALUS 



ferentiates the orbit proper from its anterior extension, the lacri- 
mal fenestra, and frives the loxommid eye-socket its characteristic 
keyhole shape. Most of the lateral edge of the frontal forms the 
thick orbital rim. At its waist the frontal is 4.3 mm. wide, mak- 
ing- the interorbital distance a mere 8.6 mm. — extraordinarily 
narrow for a skull with an estimated width of 185 mm. 

The short parietal extends laterally into a square lappet ; there 
is no indication that this lappet represents a former intertem- 




Fig. 1. Skull of Spaihicepltalus pereger, n. sp., x %. Eestoration in 
flashed liiU'S based on <S'. minis; dotted lines conjectural. Abbreviations: 
/, frontal; j, jugal ; /, lacrimal; m, maxilla; n, nasal; p, parietal; pf, post- 
frontal ; nm, premaxilla ; po, postorbital ; pp, postparietal ; prf, prefrontal ; 
q, quadrate; qj, quadratojugal; sq, squamosal; si, supratemporal ; t, 
tabular. 



4 BREVIORA No. 157 

poral element. Centered in the anterior half of the inter-parietal 
suture lies the parietal foramen, 2.3 mm. in diameter, Avhieh 
housed the parapineal eye. The postparietal (dermal supraoc- 
eipital) is relatively lartje. 

A thin oval lamina slopes outward from the corner of the 
tabular. A'entromedial to this flano-e lies the hackward-jntting 
tubercle which is characteristic of loxommid tabulars but which 
also occurs in such sjenera as E clops (Romer and "Witter, 1942). 
"Whether this process is a true homoloo- of the tabular horn in 
other labyrinthodonts and perhaps of the tabular lamina in 
Acanthostcga remains to be determined. Unfortunately, the lat- 
eral margins of the tabular and supratemporal are not clearly 
defined in the specimen. The greatly broadened supratemporal, 
however, is evidently embayed by the apex of the otic notch — a 
condition also found in Baphefes and MfgaloccpJwlus hrevicornis 
[Orihoaaurus pochyccpJialus of AYatson. 19261. 

The postfrontal, which in most labyrinthodonts lies postero- 
medial to the orbit, here lies directly posterior to it. In conse- 
quence the postorbital has been thrust outward onto the cheek 
along with most of the supratemporal; its orbital margin is 19 
mm. long and nearly straight. In its new orientation the post- 
orbital might be called exceptionalh' long rather than excep- 
tionally broad. 

Watson's (1929, tig. 22) illustration of the skull of Spathi- 
cephalus viirus, scaled to the interorbital width and post-orbital 
length of S. pereger, forms the basis for the reconstruction in 
Figure 1. The restored skull of S. pereger measures 177 mm. in 
micl-sagittal length as compared to 198 mm. for the type skull 
of S. mirus. As nothing is known of the posteranial skeleton in 
this genus, the dimensions of the Avhole animal cannot be esti- 
mated. 

COMPARISON 

Inconsistencies between the Point Edward specimen and the 
published account of SpatJiicepJwlus mirus might be interpreted 
as generic distinctions. Many of the discrepancies, howcA'er, can 
be attributed to diiferences in the manner and quality of preser- 
vation. For example, the suture pattern illustrated by AVatson 
is that of the ventral surface of the skull roof and can be ex- 
pected to differ somewhat from that of the dorsal surface. 



1962 NEW SPATHICEPHALUS 5 

The single element labeled "postorbital" in Watson's fignre 
of S. ))iirus occupies the j^ositiun of the postfrontal and half of 
the post orbital in *S^. peregcr. Since the suture between jugal and 
squamosal meets the outer corner of this element — as it does 
the corner of the postorbital in all known loxommids — Wat- 
son 's identification nnist be correct. If (as he suggests) the 
postfrontal forms part of the undifferentiated interorbital bar 
in S. viirus, then the two species differ markedly in this respect. 
Watson illustrates a suture v;hich bisects the antorbital process 
at right angles to the lateral margin of the nasal bone, and in- 
terprets this suture as separating the lacrimal from a wing of 
the interorbital bar. New evidence now permits a more conven- 
tional reconstruction of this region. No parietal foramen was 
observed in the Scottish material, but a foramen of normal size 
and position is evident in the Nova Scotian specimen. 

One clear distinction lies in the relatively narrower skull 
table of the new species: in *S'. mirus tlie ratio of interorbital 
width to table width is 1 :7 while in S. peregcr it is 1 :6. Another 
difference is that even when allowance is made for post-mortem 
distortion of the cheek region, the orbits seem to have been more 
obliquely placed in *S'. peregcr than in S. mirus. 

DERIVATION OF SPATHICEPHALUS 

Grotesquely modified though it is, the skull structure of 
S path ice phalus (as elucidated by the new specimen) can be read- 
ily derived from that of more typical loxommids. As it happens, 
one species which can be said to represent a \:)re-Spat]iicepJialus 
morphological stage is also the only older loxommid known : 
Loxomma allmanni Huxley from the Gilmerton ironstone of 
Scotland. The Lower Limestone group in which it was found 
directly underlies the Limestone <.'oal group in which Spathi- 
cephalus mirus occurs. 

In the transition between the mor])hological stages represented 
by Loxomma and Spathicephalus, posteromedial migration of the 
orbits has displaced the supraorbital elements so that the frontal 
bone forms much of the orliital rim — a condition repeated in 
various genera of labyrinthodonts l)ut more commonly associated 
with enlarged orbits. Concomitantly the cheek and snout have 
broadened and flattened into a condition which recalls the early 
Permian zatracheids Acanihostoma and Zatrachys. The con- 
.striction. of the frontals into a narrow bar, together wilh the 



6 BREVIORA No. 157 

shortening- of the skull table, loss of the intertemporal bone 
and lateral displacement of the postorbital and supratemporal, 
are unexpected extremes of specialization in so early a fjeims. 
Such a condition did not recur, so far as we know, until the de- 
velopment of the plagiosaurs in late Triassic time (cf. Panchen, 
1959, %. 16). 

The specializations of Spathiccplialus evidently adapted it to 
life as a bottom-dwelling fish-eater. Amonp- labyrinthodonts of 
this ecotype — e.g. Erpeiosaurus, Zairacliys, Capitosanrus, Eu- 
pelor [Buettneria], GerrotJiorax — the lower jaw articulation 
tends to be aligned with the occipital condyle, so that the mouth 
could be opened by raising the snout while the thorax remained 
prone on the bottom (cf. Watson, 1951, pp. 67-70). A similar 
tendency is evident in ^paihiccplialus, for its occipital and quad- 
rate condyles are more nearly aligned than those of any other 
loxommid except the late Westphalian species of Blegalocephalus. 

Surely the extent of modification which Spathicephalns had 
achieved by late Mississippian time presupposes a long previous 
history — yet to be revealed — for tlie Loxommidae. 

ASSOCIATED FAUNA 

A Gyracauthus spine, a supposed Sagowdus quadrate and 
fragmentary fish remains were found in the flagstone layer with 
the mandible of PhoIiderpeton{ ?) hrcionensc in 1956. From this 
same layer the 1960 field party collected the type of Spathicepha- 
lus pereger and the proximal third of another Gyracanthiis spine 
(PU 17185). 

About 12 feet lower in the section, bones coated with red iron 
oxide occur in an 18-inch zone of highly calcareous mudstone, 
rich in ostracodes and Spirorhis, which is intercalated with layers 
of limey shale. A Gyracanfhus spine (PU 17186), a skull bone 
of the iungfish Sagenodus (the left "E" plate, PU 17187), and 
scales and scrap of the crossopterygians Megalichtliys and Strep- 
sodus (PU 17188, 17189) represent a typical assemblage of 
Carboniferous fresh-water fishes. Amphibian remains from this 
horizon include a left clavicle, a phalanx and a ventral scale (PU 
17190), all apparently embolomerous. 

STRATIGRAPHY 

The Point Edward formation is assigned to the lower part of 
the Canso group. While th(» Canso cannot yet be correlated pre- 
cisely with European and other American sequences, it appears 



1962 NEW SPATHICEPHALUS 7 

on the basis of fossil floras (Bell, 1944, pp. 23-24) and inver- 
tebrates (Copeland, 1957, pp. 6-8) to be more or less equivalent 
stratigraphically to the Lower Namurian of Europe. Since in 
current practice the lower boundary' of the Upper Carboniferous 
system coincides with that of the Namurian, the Point Edward 
formation is classified in European terms as basal Upper Car- 
boniferous. In North America, however, the Carboniferous is 
subdivided differently : its upper half constitutes the Pennsyl- 
vaniau system which includes equivalents of the Upper but not 
the Lower Namurian. Thus the Point Edward formation is clas- 
sified in American terms as Upper Mississippian (Weller et al., 
1948). 

The Scottish specimens of SimthicepJialus were found in the 
Loanhead No. 2 ironstone of the Midlothian coalfield, a bed which 
lies in the upper half of the Limestone Coal group (Tulloch and 
Walton, 1958). This group is believed to be approximately 
equivalent to Goniatite Zone Ei (characterized by Eumorpho- 
ceras) and the lower part of Brachiopod-Coral Zone D3 (charac- 
terized by DibunopJiyllum) ; these zones correspond to the Lower 
Namurian of continental Europe (Trueman, 1954). At the risk 
of confusion it should be added that although the Eumorphoceras 
Zone forms the base of the Upper Carboniferous, the flora and 
the fish fauna of this zone show Lower Carboniferous affinities 
and are separated from their Upper Carboniferous counterparts 
by a distinct biotic break. The sequence of amphibian-bearing 
beds in the Scottish Carboniferous has been reviewed by Westoll 
(1951) and most recently by Panehen and \Yalker (1961). 

In summary, although the correlations involved are still rather 
tentative on both sides of the Atlantic, current stratigraphic 
practice assigns an early Namurian age to both the Scottish and 
Nova Scotian species of Spathiceplialus. The epoch in which 
they lived was evidently one of biotic transition between the 
earlier and later phases of the Carboniferous. 

In view of the stratigraphers' uncertainty', the possible utility 
of Spaihiccphalus as an index fossil is worth investigating. For 
the most part the loxommid amphibians are too imperfectly 
known for their distribution to be stratigraphically significant. 
The best -known genus, Megalocephalus [Orthosaurus], ranges 
through early and middle Pennsylvauian time, i.e. from West- 
phalian substage A to late WestiDhalian D (Panehen and Walker, 
1961; Baird, 1957). But since ^paihkeplialus is the most ex- 
tremely specialized member of the family it may have existed 
over a comparatively short span of geologic time. 



8 BREVIORA No. 157 

SUMMARY 

A partial skull roof of the loxommid labyrintliodont Spathi- 
cephalus, previously known only from the Limestone Coal group 
of Scotland, has been found in the Point Edward formation of 
the Canso group near Sydney, Nova Seotia. The suture pattern 
(hitherto incompletely known) of this extremely specialized 
genus is described and derived from that of more typical loxom- 
mids. Spathicephahis-hea.rmg beds of Scotland and Nova Scotia 
are considered to be early Namurian in age, corresponding to 
the base of the Upper Carboniferous in Europe and the top of 
the Mississijipian system in North Ameriea. *S'. pcreger n. sp. 
is the first-described temnospondylous ami)hibian of Mississip- 
pian age from the Western Hemisphere. 

ACKNOWLEDGEMENTS 

This study owes much to the hospitality and cooperation of 
Director Donald K. Crowdis and staff members William F. and 
Jane McNeill Take of the Nova Scotia Museum of Science in 
Halifax. P. B. Van Houten analyzed the source rock ; A. L. 
Panchen provided most helpful advice on matters of British 
stratigraphy; Edward S. Belt has generously shared the results 
of his current research on Canso stratigraphy and sedimenta- 
tion. Financial support from the AVilliam Berryman Scott Re- 
search Fund of Princeton University (administered by Glenn L. 
Jepsen) and the National Science Foundation made possible the 
field work. 

REFERENCES 

Baird, Don^ald 

1957. Rhaehitomous vertebrae in tlif loxonmiid aiiiphiliian Megalo- 
ccphaJns [abstract]. Geol. Sot-. Ameriea Bull., vol. 68, p. 1698. 
Bell, W. A. 

1944. Carboniferous rocks and fossil floras of northern Nova Scotia. 
Canada Geol. Survey Mem. no. 238, 277 pp., 79 pis. 
COPELAND, M. J. 

1957. The arthropod fauna of the Upper Carboniferous rocks of the 
Maritime Provinces. Canada Geol. Survey Mem. no. 286, 110 
pp., 21 pis. 
Panchen, A. L. 

1959. A new armoured amphibian from the Upper Permian of East 
Africa. Roy. Soc. London Phil. Trans. (B), vol. 242, pp. 207- 
281, pi. 8. 



1962 NEW SPATHICEPHALUS 9 
AND A. D. Walker 



1961. British Coal Measure lahyriiithodont localities. Ann. Mag. Nat. 
Hist., ser. 1.3, vol. 3, pp. 321-832. 
ROMER, A. S. 

1941. Earliest land vertebrates of this continent. Science, vol. 94, 
p. 279. 

1945. A'ertebrate paleontology. Univ. Chicago Press, ix + 687 pp. 

1947. Review of the Lalnrinthodontia. I\Ius. Comparative Zool. Bull., 
vol. 99, pp. 1-368. 

19.j8. An embolomere jaw from the Mid-Carboniferous of Nova Sco- 
tia. Mus. Comparative Zool. Breviora, No. 87, 8 pp. 
AND R. Y. Witter 

1942. Edops, a primitive rhachitomous amphibian from the Texas Red 
Beds. Jour. Geol., vol. 50, pp. 925-960. 

Trueman, Arthur (Ed.) 

1954. The coalfields of Great Britain. London: Edward Arnold, xi + 
396 pp., 7 pis. 
TULLOCH, W., AND H. S. Walton 

1958. The geology of the Midlothian Coalfield. Geol. Survey Scotland 
Mem., vii + 157 pp., 5 pis. 
Watson, D. M. S. 

1926. The evolution and origin of the Amphibia. Roy. Soc. London 

Phil. Trans. (B), vol. 214, pp. 189-257. 
1929 ["1923"]. The Carboniferous Amphibia of Scotland. Palaeon- 

tologia Hungarica, vol. 1, pp. 219-252, pis. 1-3. 
1951. Paleontology and modern biology. New Haven: Yale Univ. 
Press, xii -f 216 pp. 
Weller, J. M., et ax. 

1948. Correlation of the Mississippian formations of North America. 
Geol. Soc. America Bull., vol. 59, pp. 91-196. 

Westoll, T. S. 

1951. The vertebrate-bearing strata of Scotland. Interuat. Geol. 
Cong., Rept. 18th Session (11), pp. 5-21. 




CX 

o 



o 
■f. 



p 



s o 



IB 






X S 



BREVIORA 

Meseiuinti of Comriparative Zoology 

('amhkii)gk, Mass. May lM), 1*)()2 Ximukk I")S 



A FOSSIL GERRIIOSAIiR FROM THE MIOCENE 
OF KENYA (REPTILIA: CORDYLIDAE) 

By Richard Estes^ 

Many thousands of fossils, both plant and animal, have been 
collected since 1947 in the Kavirondo Gulf area, Lake Victoria, 
Kenya. Most of these fossils come from sediments on Rusinp-a 
and Mfanwanu Islands, within the Gulf. The vertebrate fauna 
is diverse, though principally mammalian ; thus the discovery 
of the fossil lizard described here is of special interest. Much 
of the fossil mammalian fauna has been described in a British 
Museum of Natural History series (see e.g. LeGros Clark and 
Leakey, 1951; AVhitworth, 1958). Part of the extensive seed-nut 
flora has recently been described by Chesters (1957). Leakey 
(1952) has noted and figured some of the remarkably well-pre- 
served invertebrates. 

The age of the fossils in this sequence has been assigned en- 
tirely on faunal grounds, and is usually considered to be early 
Miocene (Burdigalian). The reasons for this age determination 
are given in the papers mentioned above. The presence of both 
archaic and modern elements in the mammalian fauna as well 
as our lack of knowledge of the Cenozoic sequence in tropical 
areas make it dif^cult to assign a firm date to these deposits. 
Suggestions ranging from Oligocene to "considerably younger 
than the lower Miocene" (LeGros Clark and Leakey, 1951) have 
prompted Whitworth (1958, p. 45) to state that '"it is clear 
that the question of geological age may have to be revised when 
the description of the East African fossils is completed." 



1 Department of Biolnjr.v, Boston rniversii.v, and Research Associate, Museum 
of Comparative Zoology, Harvard I'nixrrsit.v. 



2 BREVIORA No. 158 

Class Reptilia 

Order Sauria 

Family CORDYLIDAE 

Subfamily GERRHOSAURINAB 

Gerrhosaurus cf. G. imajor Dimieril 

Bef erred specimen. — Coryndon Museum, Nairobi, M. F. W. 
1955/1. 

Locality. — Mfaiifiranu Island, Lake Victoria, Kenya, Africa. 
Area B, of Red Earth Series as mapped by Wliitworth (1953). 
An extensive flora and invertebrate fauna have also been col- 
lected from these deposits, but have not yet been described. 

Age. — Early Miocene (Rurdip-alian ?) . 

Preservation and major feat ares of the specimen. — As pre- 
served, the total lencrth of tlie specimen is 55 millimeters. It 
consists of most of the head and the (greater part of the neck, 
though it is probable that much more if not all of the entire 
body was originally present. The external features, many of 
them from the soft anatomy, are in part completely replaced 
with calcite. Some bone is preserved internally, but selective 
replacement has occurred. The external calcite covering is about 

3 or 4 millimeters thick, and internal to this the neck aud pos- 
terior skull region are filled with a soft waxy black sediment 
which contains small caleite crystals. No trace of the jiosterior 
bones of the skull is visible. The head is slightly twisted to the 
left, and the half-open mouth is filled with calcite. A striking 
feature of this specimen is the replacement of the tongue by 
calcite, though no important details of structure are visible. 
Dorsally, especially in the neck and posterior skull region, the 
specimen is heavily crushed ; otherwise, there is little distortion. 
The shape of the depression in the top of the skull, the half-open 
mouth, and lack of crushing in the facial segment suggest that 
the animal may have been stepped on, perhaps by the sharp 
hoof of a grazing animal. This might have caused the animal's 
death, or could also have happened shortly after it was buried 
by sediment. 

The palatal, facial, and marginal bones of the snout are broken 
or missing. Most of the teeth are eroded, though their outlines 
are preserved as imprints in the calcite filling of the mouth, but 
in some places a few tooth crowns still remain, as well as frag- 
ments of tooth shafts. 



1962 



MIOCENE GERRHOSAUR FROM KENYA 



The eve sockets are eoniplctely filled with eah-ite, thoiiji^h the 
right one is badly eroded. On the h^ft side, the ealeitc hus taken 
an impression of the uncUn-siih' of the supraorbital plates, but 
their number cannot be deternuned. One of the most unusual 
features of this specimen is the preservation of the external 
shape of the left eye. in calcite. The lids are sharply delineated, 
and between them, a low, domed area reflects the outline of the 
cornea or perhajis the undei'lying lens. 




Fig. 1. Gcrrliosaiiru.s ef. G. major, T-owlm- Miocene, Kenya, right lati'ial 
view, X 1.5. 



The outline of the outer ear and tympanum is perfectly ])re- 
served on the right side, though on the left much of it is broken 
away. The anteroposterior diameter of the ear opening is less on 
the left as a result of twisting of the head. Many of the granular 
scales which surround the ear region and extend into the lateral 
fold are clearly visible. The tij) of the extracolumella is strongly 
imprinted on the tympanum. The lateral fold is prominent on 
both sides, but is especially clear on the right, where it has been 
stretched open by twisting of the body on the median axis be- 
fore deposition. 

Very little remains of the posterior skull bones. A thin film 
of prefrontal and palatine surrounds the left eye, and directly 
below, small portions of the vomers protrude. Maxillae and 
dentaries are badly broken, and little remains other than thin 
edges of their internal processes. Fnder both eyes parts of the 
internal faces of the jugals are visible as imprints. 

Dorsally, there are imprints of seven or eight postcranial scale 
rows. They end anteriorly near a smooth flat surface Avhich 



4 BREVIORA No. 158 

represents the skull roof, but whatever bone or scale imprints may 
have been preserved are no lonprer present. 

Description. — The larcre imbricate osteoscutes of the throat 
reo'ion are compound; each is formed of multiple polyo'onal or 
trapezoidal osteoderms. The anterior ones are small, smooth, and 
subequal. the jjosterior ones elongated and either smooth or 
faintlv wi'inkled. The posterior borders of the osteoscutes are 
roumled or sHu'litly squared. The throat osteoscutes are in seven 
loiiii-itudinal rows between the lateral folds, and they alternate 
ratlier than bein<i: aligned in straight transverse rows. Pos- 
teriorly, their arrangement is somewhat distorted as a result of 
post-mortem dislocation of some of the scales. Two pairs of large 
non-imbricate chin shields seem to have lieen present, formed of 
small subequal polygonal osteoderms. 

The lateral folds aiipear to have contained small or granular 
scales, bul this is uncertain. Both folds extend as far forward 
as the ear. 

The subtriangular outer ear opening is covered ventrally with 
small squarish or lenticular scales which grade into those of the 
lateral fold. On the posterior border of the ear opening, several 
marginal i-ows of small scales are followed by a row of slightly 
larger ones. The tympanic shield is strap-shaped, narrow, and 
not at all posteriorly expanded. 

The dorsal scales of the neck, of which imprints of seven or 
eight straight transverse i-ows ai-e preserved behind the occipital 
margin, are large and subrectangular. 

Abrasion has removed almost all imprints of scales from the 
cheek, but a few large ones are visible anteriorly. 

The pleurodont teeth are robust, tall, and columnar. Imprints 
of their closely-spaced shafts indicate that replacement teeth 
were formed in subcircular basal excavations. Preserved tooth 
crowns are faintly tricuspid. 

Discussion. — Union of the gerrhosaurs and cordylines as sub- 
families of the Cordylidae, as recently suggested by McDowell 
and Bogert (1954), is undoubtedly correct. The presence of 
compound osteoderms in cordylines {s€us}( .stricfo), a character 
not mentioned by these authors, further emphasizes the separa- 
tion of this group from the anguimorphs and allies them with 
the scincomorphs. 

Until now, th(^ cordylids iiavc liad no clear fossil record. 
Pseudolacerht inncronata (Filhol) has been tentatively placed in 
this family by Romer (1956, p. 552). De Stefano (1903, p. 413) 



1062 



MIOCENE GERRHOSAUR FROM KENYA 



and Filliol (1877. ]>. 489) do not mention any characters of 
taxononiii- value, anil Filhol's illustration (ibid., fig. 423) is 
vague and diagranimatie. Iloffstetter (1944, p. 553) considers it 
possibly a skink, but notes that vertebrae similar to those of 
Cordylus {sensu lain) occur in the same deposit. Later, he indi- 
cates (1955, p. 621) that these fossils "rappellent les pieces 
homolog'i(|ues de Tactucl Cordiihi^," but the assignment is still 
tentative. 

The fossil described here is referable to the subfamih^ Gerrho- 
saurinae on the basis of the large, broadly imbricate, rounded 
or slightly squared throat scales, contrasting with the much 
smaller, non-imbrieate, diamond-shaped throat scales of the cor- 
dylines. Chamaesaura has larger throat scales than other cordy- 
lines, but they are anteroposteriorly elongate and mucronate, 
rather than smooth and transversely widened as in gerrhosau- 
rines. 




Fig. 2. Gerrhosaurus cf. G. major, Lower Miocene, Kenya, left lateral 
view, X 1.5. 



Though the published generic characters of Gerrhosaurus are 
in soft anatomy and scale details not preserved here (Loveridge, 
1942, p. 488; FitzSimons, 1943, p. 268), similarity between this 
fossil and Recent members of the genus seems to indicate that it 
belongs here. The throat scales, what remains of the dorsal 
scales, shape of the ear and lateral fold all agree closely with 
these characters of the Recent genus. Moreover, a number of 
resemblances discussed below strongly suggest reference to the 
Recent species G. major. 



6 BREVIORA No. 158 

1. Tli(^ fossil has the sizo and o-oneral proportions of a large 
atlult individual of (i. major. The latter is the lar<>est of the 
species of (icrrhosaurus ; most other species are considerably 
smaller. 

2. The slia]ie of the opening' of the external eai- in 0. major 
may be rounded or sliohtly angular dorsally. Tn the available 
specimens, G. m. major most frequently shows the rounded con- 
dition, but a. m. (iraiidis usually has a more angular dorsal 
edge, as in the fossil. 6'. m. hottegoi resembles G. m. major in 
this character. 

3. Tn oerrhosaurs. the shape of the tympanic shield is con- 
sidered taxonomically significant. This scale lies on the anterior 
border of the outer ear, and in all species of Gcrrhomurns except 
(t. major, is thin, flattened, and often expanded to cover and 
])rotect the cavity of the outer ear (see e.g. FitzSimons, 1948, 
figs. 150, 156). Loveridge (1942, pp. 515, 518) states that tym- 
panic shields of G. fiavigularis are also narrow and band-like as 
in G. major. This is grossly true, but in detail the two can be 
distinguish(^d easily. G. flavujnlaris has a narrowly crescent- 
shaj)ed tympanic shield (see e.g. FitzSimons, ihid., fig. 154; cf. 
fig. 164 of (r. m. (jra)idh) which is thin and flattened, w4iile that 
of G. major is strap-shaped, and thickened. 

4. Another similarity to G. m. grandis is the presence, on the 
posterior border of the outer ear opening, of a small anterior 
row of scales, flanked by a larger posterior row. Tn G. m. major 
and G. m. holfcgoi these scales tend to be subequal. This charac- 
ter varies somewhat, and in any case the time separation as well 
as lack of further preserved characters precludes reference of 
this fossil 1o one of the living subspecies. However, this charac- 
ter and that given as number 2 above seem to suggest a closer 
relationship to (r. m. grandis and G. m. hotfegoi than to G. m. 
major. The other subspecies, G. m. zechi, is known from only a 
few specimens, none of which were available to me. Tt is very 
closely related to G. m. hotiegoi and its status is not clear at 
this time. 

Disti'ibntion of Tvecent GrrrJio.'iairnis major 

G. major occurs today in principally arid savanna along the 
eastern coast of Africa, north to Eritrea and south to Zululand. 
G. m. grandis, the most southern subspecies, is found from Zulu- 
land north to Morogoro, Tauganyika. G. m. f)WJor is a coastal 



1062 



MIOCENE GERRIIOSAUR FROM KENYA 



subspecies, principally in Taiifianyika, but reachinfr as far north 
as Kenya. G. m. botUijoi ranjies from central Tang-anyika noi'th 
throup-h central Kenya and reaches north to coastal Eritrea, far- 
tlici- noi-th than any otlicr gci'i'liosanr. and is the only subspecies 
of a. nutjor found today in tlic Ivaviroiido (iulf I'eg'ion of Lake 
Victoria, tlie same region as tiie occurrence of the fossil. The 
l)rol)lematical (}. m. zcchi has, so far as known, a disjunct distri- 
bution limited to the northern Belgian Congo and Togo. 

CONCLUSIONS 

The fossil described here is closely related to, and perhaps 
conspeciiie with, the Recent species GcrrJiosaurus major. Thus 
it is extremely probable that the habitat of the lizards repre- 
sented by the fossil was semi-arid or arid savanna, like that of 
the modern species. The presence of a mammalian fauna of 




Fig. 3. (ierrho.saurus cf. <i. ijnijar, Lower Miocene, Keny;i, ventral view, 
X 1.5. 



dominantly savanna aspect in these deposits supports this con- 
clusion. Whitworth (1953, p. 82) states that ephemeral lakes 
were probably present in this region in the late Miocene, allow- 
ing the savanna mammals access during the periods of desicca- 
tion indicated by the sediments. He also points out that similar 



8 BBEVIORA No. 158 

situations occur today in areas of the northwestern Sudan. Ches- 
ters (1957), on the basis of the flora, has concluded tliat a "o'al- 
h^'v-tvpe forest in which trees festooned with climliers overhuna' 
the watercourses" lived near to the site of deposition, and that 
many of the fossil nuts and seeds represent livino- tropical Af- 
rican weuera. 

Close relationship of the fossil with the Recent species indi- 
cates that at least part of the pattern of speciation seen within 
the genus today is of considerable anticpiity. Moreover, similari- 
ties of the fossil to some of the Recent subspecies of G. major 
perhaps indicate that some of the geographic variants seen today 
were beginning to apjiear, as far back as the Miocene. The two 
Recent subspecies which the fossil most closely resembles, G. m. 
hoiiegoi and G. m. grondh, are northern and southern popula- 
tions which intergrade in the area immediately south and east 
of Lake Victoria. It is interesting, but highly speculative, to 
suggest that the occurrence of the fossil near the present area of 
intergradation of these two Recent subspecies might either indi- 
cate a stage in the development of the Recent subspecific pat- 
terns, or an intergrade between the two which could possibly be 
duplicated today if sufficient specimens were available. 

Moreau (1951, esp. pp. 877, 881) has gathered evidence which 
suggests that at least from the mid-Cenozoic to the present, cen- 
tral and eastern Africa had a climate and broadly defined vege- 
tational types which differed relatively little from those occurring 
there today. If this is so, the above alternatives are quite pos- 
sible, yet additional fossil evidence, both biotie and climatic, is 
necessary to accept or reject either of them. 

SUMMARY 

The well-preserved head of a fossil lizard from the Lower 
Miocene (Burdigalian?) of Mfanganu Island, Lake Victoria, 
Kenya, is tentatively referred to the Recent species GcrrJiosaurus 
major (Reptilia: Cordylidae). Many external features of soft 
anatomy are preserved as casts in calcite, including the eye, 
tongue, and tympanic membrane. Close relation.ship to the Re- 
cent semi-arid or arid savanna species indicates a similar habitat 
for the fossil, a conclusion corroborated by the savanna aspect of 
the fossil mammals from contemporaneous deposits on nearby 
Rusinga Island. This .specimen is one of the oldest vertebrate 
fossils even tentatively referred to a Recent species and must in- 
dicate that at least part of the pattern of speciation seen in 
Recent gerrhosaurs is of relatively ancient origin. 



1962 MIOCENE GERRHOSAUR FROM KENYA 9 

LITERATURE CITED 

Chesters, K. I. M. 

1957. The Miocene floia of RiisiiiKn Islniul, L;ike Vietorin, Kenya. 
Palaeontogr., Abt. B, 101:;ji>-71, 4 figs., L'l pis. 
De Stefano, G. 

1903. I sauri del Queicy appartenenti alia collezione Rossignol. Atti 
fioc. Ital. Sci. Nat., 42:382-418, 2 jils. 

FiLlIOL, II. 

1877. Reclieiches sur les pliosplioriles du Quercy. Etude des fossiles 
qu'on y rencontre et specialement des maniiniferes. Ann. ScL 
Geol., 8:1-561, 55 pis. 
FlTZSlMONS, V. F. 

1943. The lizards of South Africa. Transvaal Mus., Mem. no. 1, xv -\- 
528 pp., 384 figs., 24 pis. 

IIOPFSTETTER, R. 

1944. Sur les Scineidae fossiles. I. Formes europeennes et nord- 
americaines. Bull. Mus. Nat. Hist. Nat. Paris, 16:547-553, 2 figs. 

1955. Squamates du type moderne. In: Piveteau, Traite de Paleon- 
tologie, 5:606-662, 26 figs. 

Leakey, L. S. B. 

1952. Lower Miocene invertebrates from Kenya. Nature, 169:624- 
625, 2 figs. 

LeGros Clark, W. E., and Leakey, L. S. B. 

1951. The Miocene Honiinoidea of East Africa. Brit. Mus. Nat. Hist., 
Fossil Mammals of Africa, no. 1:1-117, 28 figs., 9 pis. 
LOVERIDGE, A. 

1942. Revision of the African lizards of the family Gerrhosauridae. 
Bull. Mus. Conip. Zool., Harvard Univ., 89:484-543. 
McDowell, S. B., Jr., and Bogert, C. M. 

1954. The systematic position of Lanthaiiotu.s and the affinities of the 
anguinoniorphan lizards. Bull. Amer. Mus. Nat. Hist., 105:1- 
142, 43 figs., 16 pis. 
Moreau, R. E. 

1951. Africa since the Mesozoic, with particular reference to certain 
biological problems. Proc. Zool. Soc. London: 121:869-913, 4 
tables. 
ROMER, A. S. 

1956. Osteology of the Reptiles. Univ. Chicago Press, xxi + 772 pp., 
248 figs. 

Whitworth, T. 

1953. A contribution to the geology of Rusinga Island, Kenya. Quart. 
Jour. Geol. Soc. London, 109:75-96, 2 pis. 

19.")8. Miocene ruminants of East Africa. Brit. Mus. Nat. Hist., Fossil 
Mammals of Africa, no. 15:1 50, 18 figs., 12 tables. 



10 



BREVIORA 



No. 158 




- i) i^i 



S M s 



03 


Ul 


h^ 


a> 




S 


^ 


"Z 


O 






-^j 


o 


rt 


-/] 


'A 




o 




o 


^ 




.LJ 










F— 1 


-4—' 


!-• 


_ 


o 


73 


^ 


^. 


A 






>i 












*w 


-r 


V^ 


§ 


,__. 


;2 




— 


• -H 


^ 








^ 


-^- 


• ^ 


M 


^ 


i^ 








? 




— ' 


f-i 


^ 


•^ 


^ 


F^ 


^ 








^ 


o 


Ph 


2 


X 




^ 


'T 
O 


>» 


GO 


-a 




>■ 


^ 








-^ ^* 


K 








be 


C3 


»-M 


o 


S "5 


O-i 


o 










^ 


^ 


_cS 


p_^ 






^ 




-kJ 


, — • 




^4-^ 






o 


oT 




=1- 


be 




o 






^ 


3 




p< 






cS 


bl 




k< 


.s 




be 






o 






o 


M 






-4^ 




^^ 


o 




"~" 


•z 




^ 


>: 




o 






•f; — 


-4-" 




e 


^ 






M 





B R E V I O R A 

Miaseiunti of Compsirsitive Zoology 



Cambridge, Mass. May 31, U)(iL' Nimber 159 

AGE IN A SMALL SAMPLE OF BLUEFISH 
{P03IAT03IUS SALTATRIX (LINNAEUS) )i 

By Richard H. Backus 

Woods Hole Oceanographic Institution and 
Museum of Comparative Zoology 

Little is known about the life-history- of the bluefish (Poma- 
tonius saltafrix) even though this lono'-known species is widely 
distributed in warm and temperate seas and is valuable for food 
and sport. Not only have the eggs and larval young of this 
interesting fish not been positively identified, l)ut nothing exact 
is known of its rate of growth although the "annuli" on its 
scales prove to be quite easily read. Perhaps it is the great 
population fluctuations of this fish and its erratic a]:»pearance 
in many waters which have discouraged naturalists from study- 
ing it. Bigelow and Schroeder (1953) give a summary of what 
is known of bluefish life history. Little has been added since 
their account was written. 

To learn something of the growth of the species, advantage 
was taken of the abundance of bluefish around Woods Hole dur- 
ing the autumn of 1961 and of the angling expertness of Asa 
Wing, Henry Cain, and Carl Grant, Jr. These men saved 
heads and scale samples together with records of fork length 
from the 34 fish that they caught between October 9 and 19 in 
Great Harbor at Woods Hole, in Woods Hole passage, and 
along the east shore of Buzzards Bay north of AVoods Hole. 

The saccular otolith of the bluefish has been figured hj Le Gall 
(1934) and by Sanz Echeverria (1950). Le Gall {op. cit.) also 
figured the scales, and in another paper (1935) he said that his 
examination of the scales in North African specimens suggests 
that "young individuals attain their adult size and their first 
sexual maturity at the age of 4 or 5 years." On the other hand, 

1 Woods Hole Oceanographic Institution Contribution No. 1234 



2 BREViORA No. 159 

Boreea (1986) says of Black Sea bluefish : "During the second 
year they reach dimensions of 14-20 cm and can attain the first 
sexual maturity. ' ' 

Table 1 
Fork length and age in a small sample of hlucfish. 



Fork Length 


Age 




Pork Length 


Age 


(inches) 


(annull) 




(inches) 


(annuli) 


13.9 


1+ 




15.2 


1 + 


14.0 


1+ 




15.2 


1 + 


14.0 


1+ 




15.5 


1 + 


14.0 


1+ 




15.5 


1 + 


14.0 


? (1 + 


or 11+) 


15.5 


1 + 


14.1 


? (1 + 


or 11+) 


15.5 


11 + 


14.1 


1+ 




15.8 


1 + 


14.2 


1+ 




16.0 


1 + 


14.2 


1+ 




16.2 


11 + 


14.4 


1+ 




16.5 


1 + 


14.5 


1+ 




17.0 


11 + 


14.5 


1+ 




17.5 


11 + 


14.8 


1+ 




18.0 


TI+ 


15.0 


1+ 




19.0 


11+ 


15.0 


11+ 




19.0 


11+ 


15.0 


? (1 + 


or 11+) 


20.1 


11+ 


15.1 


1+ 




24.0 


III + 



An otolith extracted from a specimen in our sample 14.4 inches 
in fork length measured about 0.45 by 0.15 inches. The otolith is 
much sculptured and does not seem suitable for use in age de- 
termination by simple visual inspection though it might be so 
suited if x-ray methods were used. Since the scales of this fish 
Hi'c large, and those examined l)y us seem to show the annuli 
rather clearly (Figure 1), we have I'elied on the scah^s alone 
for the ages reported here. 

Fork length and age for the specimens in our sample are 
given in Table 1. Fork lengths were taken to the nearest eighth 
of an inch, were converted to decimals and rounded to the nearest 
tenth. Ages (expressed in annuli) were determined by examin- 
ing at least 10 legibh* scales from each specimen. In three cases 
consistent results could not be obtained. Thus, age has been deter- 
mined with some confidence in 31 instances. Of these, 21 speci- 
mens show an age of I+, nine an age of II+, and one an age of 



1962 AGE IN BLUEFISH 3 

III+. Specimens of age 1+ range from I'-iA) to 16.5 inches in fork 
length and have a mean length of 14.85 inches. Specimens of age 
11+ range from 15.0 to 20.1 inches and have a mean length of 
17.48 inches. The sole 111+ specimen measures 24.0 inches in fork 
length. Because of the small size of our sample these data can do 
little more than show that snapjiers (as small bluefish are called) 
of four to nine inches, seen in the autumn (Bigelow and Schroe- 
der, 1958), are indeed young of the year and that such fish ap- 
proximately double their length in the succeeding 12 months. 

Bluefish left the "Woods Hole area during the course of a five- 
day northeast storm which commenced on the evening of October 
19 and lasted until the early hours of October 25. The last fish 
in our sample was caught on October 19. None were caught after 
the passage of the storm although the fishing effort continued 
and produced good catches of striped bass (Roccus saxatiUs). 
Surface temperature records for Great Harbor, Woods Hole, 
show fluctuations between 64.9 and 64.5^F for the period 
October 6 to 12, a decline from 64.5 to 60.8°F from October 12 
to October 19, the day the storm began, a decline from 60.8 to 
55.0°F during the storm, and small fluctuations about 55°F for 
the remainder of the month. One may suppose that at about 
60°F the bluefish were near the minimum temperature that they 
can tolerate at this stage of the life cycle and that the catastrophe 
of the storm, with the accompanying further drop in tempera- 
ture, was enough to start them on the journey to their winter 
haunts. 

LITEEATURE CITED 

Bigelow, Hexry B. and William C. Schroeder 

1953. Fishes of the Gulf of Maine. U. S. Fish and Wildlife Service 
Fishery Bulletin, 74 (Volume 53) : 1-577. 
BORCEA, I. 

1936. Xotes sur la Biologie du Poniatome (Lufar; dc la Aler Noire. 
C. E. Aead. Sei. Eoumanie, 1 : 222-223. 
Le Gall, Jeax 

1934. Le Tassergall ou Bluefish (Pomatovius saltatrix Laeepede = 
Temnodon saltator Linne). Eev. Trav. Office Peches Maritimes, 
7: 27-85. 

1935. Le Tassergall ou Blue Fish. Bull. Soc. Sci. Xat. Maroc, 15: 
232-233. 

SaNZ EcHEVERRfA, JOSEFA 

1950. Notas sobre otolitos de peces procedentes dc las costas del 
Sahara. Bol. Inst. Espanol Ocean., 27: 1-14. 



BREVIORA 



No. 159 







m 



»S»f "^ ' '' 







ANNULUS I 



ANNULUS H 



Fig. 1. Scale from a bluefish {Pamatomits saltatrix) 20.1 inches in fork 
length taken in Woods Hole passage on October 17, 1961, showing two annuli. 



BREVIORA 

Museiimi of Comparative Zoology 

Cambridge, Mass. June 12, 1962 Number 160 



TWO NEW ARTHROPOD CARAPACES FROM THE 

BURGESS SHALE (MIDDLE CAMBRIAN) 

OF CANADA 

By W. D. Ian Rolfe 



Three carapaces of an undescribed artliropod were found by 
the writer when curating the large ^Museinn of Comparative 
Zoology collection of Burgess Shale arthropods, collected by 
P. E. Raymond, H. C. Stetson, W. E. Schevill and C. H. Burgess 
in August, 1930. Subsequent search through the material in 
the U. S. National Museum, Washington, D. C, yielded eleven 
further specimens and two specimens of another new form, 
which had been set aside for description Ijy C. E. Resser. The 
writer is indebted to Dr. G. A. Cooper for permission to borrow 
and describe the U.S.N.M. material and to Dr. II. B. Whitting- 
ton for the photographs and for criticism of the manuscript. 

The USN^M specimens came from Walcott's quarry at locality 
35k near Field, British Columbia (Walcott, IDll, pp. 51-52; 
Resser, 1929, p. 2 : = locality Sllf of Rasetti, 1951. pp. 37-38, 
103, 129). The MCZ specimens were also probably collected 
from this quarry, although it is impossible to be certain of this 
since the 1930 expedition also collected from "a second layer 
. . . very fossiliferous . . . some seventy feet further up the 
mountainside" (Raymond, 1930, p. 32;' 1935, p. 205). This 
second locality possibly corresponds with Rasetti 's Sllg (1951. 
pp. 38, 104, 130) and details of the stratigraphy of these two 
horizons are given in that work. Letters a and b following a 
specimen number indicate that part and counterpart are present. 

Carapace shape alone is insufficient to determine the affinities 
of any arthropod, as Roger has pointed out (1946, p. 59), and 
hence it seems better to group such isolated carapaces together 
as follows. 



2 BREVIORA No. 160 

TRILOBITOIDEA or CRUSTACEA incertae sedis 
PeOBOSCICARIS gen. nov. 

Type species. Prohoscicaris agnosia sp. nov. 
Diagnosis. Carapace valves only known, large, with antero- 
dorsal region produced into a spatulate beak. 

Proboscicaris agnosta sp. nov. 
Plate 1, figures 1, 2; Text-figure 1 

Diagnosis. Anterior beak prominent ; length of posterior mar- 
gin U.48 to 0.82 of greatest depth of carapace ; posterior margin 
indented at or near midpoint. 

Description. Valves elongate, ranging from shallow to deep. 
Since the orientation of the valves is unknown, the straight to 
slightly concave margin will be treated as dorsal and the pro- 
duced region as anterior. Postero-dorsal angle rounded and 
obtuse ; posterior margin slightly indented at or near its mid- 
point. Ventral margin moderately convex in posterior half to 
three-quarters of carapace length ; strongly concave in anterior 
region and separating off a tongue-shaped anterior beak. This 
beak has been lost from USNM 139866 (see Text-fig. 1) and 
it seems likely that the distinct outlines shown by USNM 
139869 and 139873 (Text-fig. 1) are only due to tlie loss of 
this region. The absence of a recognizable rim or doublure pre- 
vents certainty on this point. 

The valves were doubtless joined by a membranous hinge 
along the dorsal border as in Canadaspis. Only one specimen 
shows evidence of two valves preserved, and this is shown on 
Text-figure 1, USNM 139872. Only the anterior beak of the 
right valve is preserved and this is displaced anteriorly rela- 
tive to the almost complete left valve. 

A reticulate pattern may be seen in patchy areas on USNM 
139867 and 139873. The wrinkles subparallel to the ventral 
margin of USNM 139866 (Text-fig. 1) are clearly due to flat- 
tening of the originally convex test. Specimens USNM 139867, 
139870, 139871, 139873, MCZ 5979/2 and MCZ 5979/3 are 
blotched by the alga Morania parasitica Walcott, previously 
recorded on Canadaspis and figured by Walcott (1919, p. 232, 
pi. 50, fig. 1) on a carapace of Hurdia victoria AValcott. 



1962 



NEW BURGESS SHALE ARTHROPODS 




=5 

o 



=0 

o 



o "^ 






■-4-I a 

S o 
-1 o 












o 






4 BREVIORA No. 160 

Remarks. The form of the carapace is so distinct that little 
confusion is possible with previously described species. Such a 
small beaked specimen as USNM 139874 approaches Canadaspis 
perfccia (Walcott) in outline, but the posterior and antero- 
ventral embayments readily distinguish the new form. Some 
specimens of Hurdia victoria in the Museum of Comparative 
Zoology show an indentation of the ?posterior margin similar 
to that in Prohoscicaris, and in addition show an identical retic- 
ulation of the carapace surface, so that fragments of the pos- 
terior ends of the two forms might prove difficult to distinguish. 
Large-mesh reticulation also occurs in Tuzoia and Carnarvonia 
(Walcott, 1912, pp. 157-158, 165, 187, 189) so that this char- 
acter is of little value for suggesting relationships. Small-mesh 
reticulation is visible in sjiecies of CaryocariH, Dictijocaris and 
Conca vicar is (as well as in the olenellid trilobites: Raw, 1936; 
Moore, 1958, fig. 5.22), and in Ceratiocaris and Monfecaris such 
reticulation can be shown to arise from differential corrosion 
of the cutieular prisms (Rolfe, 1962a, pp. 45-47). The cuticle of 
the Burgess Shale specimens is too poorly preserved to ascertain 
whether this reticulation is sculptural or structural. 

Holotype. USNM 139871. Plate 1, figure 2 and Text-figure 1. 

Other material. The twelve specimens shown on Text-figure 
1: USNM 139866a/b. 139867a/b, 139868-139870. 139872-139875. 
MCZ 5979/1, 5979/2a/b. 5979/3a/b. Another specimen, USNM 
139876, is a fragment of the anterior end only. 

Dimensions of holotype. Maximum length parallel to hinge 
line : 98 mm. Maximum depth perpendicular to hinge line : 
52 mm. 

PrOBOSCICARIS IXGEXS sp. nov. 
Plate 1, figure 3 ; Text -figure 2 

Diagnosis. Carapace valves only known ; anterior beak rela- 
tively small ; length of posterior margin ca. 0.28 of greatest depth 
of carapace ; posterior margin sigmoidal. 

Description. The ventral margin is more of a simple skewed 
curve than the convex and concave outline of P. agnosta. The 
posterior margin is shorter and hence is situated more dorsally 
than in P. agnosta and in addition this margin is sigmoidal 
rather than indented. The carapace margin is curled under, 
except along the dorsal border, suggesting the marginal rim 
common in the later phyllocarids. 



1962 NEW BURGESS SHALE ARTHROPODS 5 

The surface of the carapace is smooth but little is preserved 
of the original test save blotches of filmy black material. Occa- 
sional circular areas of silver sheen on the holotype may repre- 
sent Morania parasitica. 



Fig. 2. Outline drawing of the holotype of Proboscicaris ingens sp. nov. 
— USX:\I 139865b. Postulated anterior to left. 

Remarks. Again the carapace shape is distinctive, though the 
asymmetrical outline recalls that of Isoxys, which, however, is 
smader and has the anterior and posterior dorsal extremities 
acuminate rather than truncate. 

This species is the largest of the known Burgess Shale arthro- 
pods in terms of surface area of the carapace and was doubtless 
the one which Walcott had in mind when he wrote, "there are 
also fragments of the carapace of a very large form that possibly 
may be related to Hurdia victoria" (1912, p. 183). Individuals 
of H. victoria may be longer but they are also slenderer. Such 
large carapaces are of particular interest in this fauna since 
it is among them that a suitable adult for the hypothetically 
larval Waptia jieldcnsis might be sought (Fedotov, 1925, pp. 
386, 389; Heldt, 1954, p. 180: Tiegs and Manton, 1958. pp. 292, 
314; cf. Henriksen, 1928. p. 14; Stormer, 1944. p. 100). In 
this connection is seems worth recalling the striking resemblance 
of Marria, from this same deposit, to a crustacean nauplius 
(Ruedemann, 1931, p. 8) or metanauplius. A comparable Upper 
Ordovician form, Paramarria, occurs in association with an arch- 
aeostracan carapace, Galenocaris (Wells, 1944). If Paramarria 
is a naupliar stage, and the larval stage of Galenocaris, and its 



6 BREVIORA No. 160 

aspect is not merely due to convergence for a planktonic exist- 
ence (as that of Minietastcr and Bostricliopus seems to be), it 
would contrast with Kecent Leptostraca. In the latter group, 
development is direct, the young hatching at a late stage. Simi- 
larly, Naraoia might l)e regarded as a larval merostomoid. 

It is possible that J', ingens is simply an older instar of P. 
agnosta. However, such radical changes in shape are not com- 
mon except in early ontogeny, and it seems better to distinguisli 
this form as a separate species. 

Holotype. USNM 139865a/l) — part and counterpart. 

Dimensions of holotype. Maximum length parallel to hinge 
line: 156 mm. Maximum depth ])er]^endicular to hinge line: 
75 mm. 

Other material: USXM 139890 — a fragment of the posterior 
of a carapace valve, which must have exceeded 150 mm. long 
by 95 nun. deep when complete. 

DISCUSSION 

The laek of limbs or body si'gments precludes any discussion 
of the affinities of this new genus and the problem of classifica- 
tion of these early crustacean-trilobitoid forms has been sum- 
marized elsewhere (Rolfe, 1962b). 

It seems worthless to classify such isolated carapaces above 
the generic level in view of the limited number of characters 
available. Many of the genera attributed to phj'llocarid families, 
or made the types of new families such as the Isoxyidae (junior 
synonym of Tuzoiidae Raymond, 1935) and Pseudoarctolepididae 
of Brooks and Caster (1956, p. 13), will need to be brought 
together under the inccrtae sedis category shown above. 

Some idea of the relative abundance of Proboscicaris agnosta 
in the Burgess Shale fauna may be gained from the following 
list of numbers of individuals of non-trilobite arthropods col- 
lected by the 1930 MCZ expedition, and recently curated by 
the writer : 

TRILOBITOIDEA 

Burgessia hella Walcott 54 

Emeraldella or fMolaria spp. indet. 3 

Leanchoilia superlata Walc. 12 

Marrclla splcndens Walc. 202 

Naraoia compacta Walc. 3 



1962 NEW BURGESS SHALE ARTHROPODS 7 

Opahinia regalis Wale. 1 

iSidneyia inexpectans Wale. 9 

fYohoia plena AValc. 1 

TRILOBITOIDEA or CRITSTACEA inceriae scdis 

Anomalocaris canadensis Whiteaves ca. 22 

Canadaspis ohliqua (Wale) 7 

C. ovalis (Wale.) 1 

C. perfect a (Wale.) 76 

C. sp. indet. 12 

Fieldia laticeolata Wale. 1 

Hurdia triangiilata Wale. 1 

H. victoria Wale. 16 

Isoxys acutangulus Wale. 10 

Prohoscicaris agnosta sp. iiov. 3 

Protocaris ef. pretiosa Resser 1 

Tuzoia retifera- Wale. 1 

T. sp. indet. 2 

EEFERENCES 

Brooks, H. K. and K. E. Caster 

1956. Pseudoarctolepis sharpi, n. gen., n. sp. (Phyllocarida) from the 
Wheeler Shale (Middle Cambrian) of Utah. Jour. Paleont., 
30:9-14. 
Fedotov, D. 

1925. On the relations between the Crustacea, Trilobita, Merostomata 
and Arachnida. Akad. Nauk S.S.S.R., Leningrad, Bull., YI 
Serie, 18:383-408. 
Heldt, J. H. 

1954. Waptia fiehienain Walcott et les stades larvaires des pen6ides. 
Bull. Soc. Sci. nat. Tunis.. 6: 177-180. 
Henriksen, K. L. 

1928. Critical notes upon some Cambrian arthropods described by 
Charles D. Walcott. Vidensk. Medd. fra Dansk naturhist. Fore- 
ning i Kjerbenhavn, 86:1-20. 
Moore, R. C. 

1958. Introduction to Historical Geology. 2nd ed. McGraw-Hill, New 
York, Toronto, London, 7 -|- 656 pp. 
Rasetti, Franco 

1951. Middle Cambrian stratigraphy and faunas of the Canadian 
Rocky Mountains. Smithsonian Misc. Coll., 116, No. 5: 5 + 
277 pp. 



8 BREVIORA No. 160 

Raw, Frank 

1936. Mesonacidae of Coniluy in iShrupsliiie, with a discussion of 

cdassification within the family. Quart. Jour. geol. Soc. London, 

92:236-293. 
Raymond, P. E. 

1930. Report on invertebrate paleontology. Ann. Rep. Mus. Conip. 
Zool. for 1929-1930:31-33. 

1935. Leandhoilia and other Mid-Cambrian Artluopoda. Bull. Miis. 

Comp. Zool., 76:20.5-230. 
Resser, C. E. 

1929. New Lower and Middle Cambrian ('iusta<u'a. I'roc U.S. Nat. 

Mus., 76, Art. 9, 18 pp. 
Roger, Jean 

1946. Les invertebres des eoiu-lies a poissons du Cretace Superieur du 

Liban. Mem. Soc. geol. France, Nouvelle Serie, 23, No. 51, 

92 pp. 
ROLFE, W. D. I. 

1962a. The cuticle of some Middle Silurian ccratiocaridid Crustacea 

from Lanarkshire, Scotland. Palaeontology, 5:30-51. 
1962b. Grosser morphology of the Scottish Silurian phyllocarid crus- 
tacean, Cerafinraris papilio Salter in Murchison. .Tour. Paleont. 

(ill press). 

RUEDEMANN, RUDOLF 

1931. Some new Middle Cambrian fossils from British Columbia. 
Proc. U.S. Nat. Mus., 79, Art. 27, 18 pp. 

ST0RMEE, LeIF 

1944. On the relationships and idiylogeny of fossil and recent Arachno- 
morpha. Skr. Norske VidenskapsAkad. Oslo, I, Math.-Kl., No. 
5, 158 pp. 
TiEGS, O. \V. and S. M. Maxton 

1958. The evolution of the Artluopoda. Biol. Rev., 33:255-337. 
Walcott, C. D. 

1911. Cambrian geology and paleontology, II. No. 3 — ^liddle Cam- 
brian holothuriaiis and medusae. Smithsonian Misc. Coll., 57:41- 
68. 

1912. Camljrian geology and paleontology, II. No. 6 — Middle Cam- 
Inian Branchiopoda, Malacostraca, Trilobita, and Merostomata. 
Sinith.sonian Misc. Coll., 57:145-228. 

1919. Cambrian geology and paleontology, W. Xo. 5 — Middle Cam- 
brian Algae, Smithsonian Misc. Coll.. 67:217-260. 
Wells, J. W. 

1944. Two new planktoiiic crustaceans from the Maquoketa Shale 
(Ordovician of Illinois). Amer. .lour. Sci., 242:436-441. 



1962 



NEW BURGESS SHALE ARTHROPODS 









* mf 



Plate 1 

Tlie scale beneath the Figures represents five centimeters. 
Figs. 1-2. Proboscicaris agnosia gen. et sp. nov. 1. Left valve and dis- 
placed anterior beak of right valve of carapace. USNM 139872. 2. Holo- 
type — USNM 139871 with anterior of valve at right. The silver blotches 
are the encrusting alga Morania parasitica Walcott. 3. Proboscicaris ingens 
sp. nov., holotype — U8XM 139865b. Postulated anterior to left. From an 



BREVIORA 

MiUseiuBi of Coimpajrative Zoology 



Cambridge, Mass. July 16, 1962 Number 161 



A COMPARATIVE STUDY OF THE 
RESPIRATORY MUSCLES IN CHELONIA 

By R. V. Shah 

Division of Comparative Anatomy and Embryology, 

Department of Zoology, M.S. University of Baroda 

Baroda, India 



INTRODUCTION 

The unique skeletal modifications in Chelonia have greatly 
influenced the morphological features of the soft parts in these 
animals. Of these modificatioiis the rigid shell formation is the 
most prominent characteristic feature of the Chelonia ; this has 
made their body wall immovable and hence the normal respira- 
tory movements characteristic of amniotes are lost. This loss 
of active body wall movements has given much reason for dis- 
cussion about the respiratory mechanism adapted by the chel- 
onians, and from the time of Malpighi and Cuvier many attempts 
have been made to explain the phenomenon. Some have sug- 
gested that the throat movements similar to those seen in frogs 
are responsible for bringing about the expirations and inspira- 
tions in Chelonia ; others have supposed that the movements 
of the limbs and the neck indirectly effect the expiration and 
inspiration. Recenth' McCutcheon (1943) has summarized the 
evidence that, on the contrary, the movements of certain 
abdominal muscles, the dmphragmaticus,^ the transversus ab- 
dominis, the serratus magnus and the ohliqnus abdominis, bring 



1 a matter of terminology must be mention^''! here. McCutcheon (1943) de- 
scribed the flank cavity muscles in Mnlaclemys cmtiafa where the diaphragmati- 
cns muscle as described b,^• him sofniis to be the ^anie as the muscularis striatum 
piilmonnlr and quite different from the dinplinigwnticus as described in this 
paper. Owen (lSf>6). describing the musculature of Emys europea, describes the 
diaphrngmnticus as formed of three parts originating' from the carapace; two 
parts insert on the wall of the Itmg while the third one inserts on the plastron. 
This description compares well witli the account given here, except that the two 
parts of the muscle described by Owen as inserting on the lung are here regarded 
as tne muscuhiria strUitiim ptihuonnle, and only the third part which inserts on 
the plastron is the triie diaphragmaticus. 



BREVIORA 



No. 161 



about expiration and inspiration. He interprets the throat move- 
ments in Chelonia as functioning in olfaction and not in respira- 
tion. George and Shah (1954) have studied the respiratory 
mechanism in Lisseniys and have confirmed McCutcheon's view 
that the abdominal muscles are effective for respiratory move- 
ments and that the throat movements are only for olfaction. 
In addition, they have also described the presence of an extra 
pair of muscles which cover the lungs completely in Lissemys. 
These muscles are composed of striated muscle fibres. On con- 
traction of these muscles the pulmonary air is pushed out of 
the lungs, and on their relaxation the atmospheric air is taken 
in. Thus these lung muscles, the muscularis striatum pulmonale, 
aid the action of the abdominal muscles in bringing about the 
expiration and inspiration. 



M.S.P. 




Fig. 1. Diagrammatic sketch of the disposition of the respiratory muscles 
in Cyclaxiorbinae where the lungs are completely covered by the muscularis 
striatum pulmonale. 



George and Shah (1955, 1958 and 1959) made a compara- 
tive study of the abdominal muscles and of the lung muscles 
in some additional chelonians: Lissemys punctata (all three 



1962 



RESPIRATORY MUSCLES IN CHELONIA 



subspecies), Geormyda trijuga, Trionyx gangeticus, Testudo ele- 
gans, Malacoclicrsus iorneri and Eretmochelys imhricata. Ac- 
cording to their observations the lung muscle, the muscularis 
striatum pulmonale, covers the lungs completely in Lissemys 
punctata, partially in Geoemyda, while the muscle is totally 
absent in the rest of the forms they studied. Of the flank cavity 
muscles the diaphragmaticus and the transversus: abdominis are 



O.A. 




Fig. 2. Diagrtuiimatie sketch of the disposition of the respiratory muscles 
iu Trionyehinae. 



well developed in Lissevnys and Trionyx where they join with 
each other to form a continuous muscle sheath covering the 
visceral organs including lungs. The diaphragmaticus in Geo- 
emyda trijuga and Eretmochelys imhricata does not join with 
the transversus ahdominis to form a continuous muscle sheath, 
but there is a bridge of connective tissue between them. In 
Testudo elegans and Malacochersus torneri the diaphragmaticus 
muscle is totally absent leaving only a thin membranous sheath 
of connective tissue in its place. The other flank cavity muscles, 
Adz. the serratus magnus and the ohliquiis abdominis are present 
with slight variation in all the animals they studied. 

In the light of these observations on the respiratory muscles 
in a very few chelonians, it was thought desirable to examine 



BREVIORA 



No. 161 




n.s.p.L. 



T^.S.P.M. 



Fig. 3. Diagrammatic sketch of the disposition of the respiratory muscles 
in Malacleviys terrainn terrapin. 




Fig. 4. Diagrammatic sketch of the disposition of the respiratory muscles 
ill Pseudemys floridana and Fscudemys texana. 



1962 



RESPIRATORY MUSCLES IN CHELONIA 



more forms representing, as far as possible, almost all the major 
groups of the order Chelonia, and make a comprehensive com- 
parative study of these muscles to get an overall idea of the 
morphological features of the respiratory mechanism adapted 
by the animals of this order. 

For this study some fifty different cryptodiran and nine 
pleurodiran forms were selected. A list of the animals chosen 
is given below. 

This work was carried out at the Museum of Comparative 
Zoology at Harvard University, Cambridge, Massachusetts, 
U.S.A. I am thankful to Dr. A. S. Romer, then Director of 
the Museum, and Dr. E. E. Williams, Curator of Herpetology, 
for all the facilities given and for their constant help and en- 
couragement during the course of the study. 



tvA.D. 



M.S.P.L.- 



C.T 




r\.s.T.T^- 



Fig. 5. Diagrammatic sketch of the disposition of the respiratory muscles 
in Emydinae forms in which the diapliragmaticus muscle is absent. 



MATERIAL STITDIED 

All the animals selected for the study were alcohol preserved 
and were found in excellent state of preservation. Careful dis- 
sections of the flank cavity muscles, the diaphragmaticus, the 
f)-ansv(rsus abdominis, the serrafus magmis and the obliqnus 



BREVIOBA 



No. 161 



abdominis and the lung muscle, the miiscularis striatum pulmon- 
ale, were done on these animals and followino- is the report of 
the comparative study. 

List of chelonians selected for the present study: 
CRYPTODIRA 

TESTUDINOIDEA 
Testudinidae 



Eniydinae 

Chineniys reevesii 
Chrysem.ys picta dorsalis 
Chrysemys picta marginata 
Chrj'semys picta picta 
Clemniys caspica caspica 
Clemmys caspica lepiosa 
Clemmys guttata 
Cleininys mutica 
Cuora aniboiuensis 
Deiroclielys reticularia 
Emydoidea blandingii 
Em^'s orbicularis 
Geoemyda luanni 
Geoemyda puuctularia funerea 
Geoemyda spinosa 
Geoemyda trijuga 
Graptemys kohni 
Graptemys pseudogeograpliica 
Kachuga tectum tectum 
Malaclemys terrapin terrapin 
Malayemys subtrijuga 
Oeadia sinensis 
Pseudemys floridana 
Pseudemys texana 
Terrapene yucatana 

TesUidininae 
CJeoclielone pardalis 
Pyxis arachnoides 
Testudo graeea 
Testudo hermaiini 
Testudo horsfieldii 
Testudo kleinmanni 

Chelydridae 

Kinosterninae 
Sternotherus carinatus minor 
Sternotherus odoratus 

Chelydrinae 
Chelydra serpentina 



TRIONYCHOIDEA 

Trionychidae 
C yclanorMnae 

Cyclanorbis sp. 
Cycloderma frenatuin 
Lissemys punctata 
(all three subspecies) 

Trionychinae 

Dogania subplana 
Trionyx gangeticus 
Trionyx sinensis 
Trionyx triunguis 

CHELONOIDEA 
Chelonidae 

Caretta caretta 
Chelonia mydas 
Eretraochelys imbricata 
Lepidochelys olivacea 

DERMOCHELYOIDEA 

Dermochelyidae 
Dermochelys coriaeea 

PLEURODIRA 

Pelomedusidae 

Pelomedusa subrufa subrufa 
Pelusios subniger 
Podocnemis expansa 
Podocnemis lewyana 
Podocnemis unifilis 

Chelidae 

Chelodina longicollis 
Emydura krefti 
Hydromedusa niaximiliani 
Plnynops geoffroana 
Platemys platycephala 



1962 



RESPIRATORY MUSCLES IN CHELONIA 



DESCRIPTION OP THE MUSCLES 

Of all the respiratory muscles mentioned above, the lung 
muscle, the muscularis striatum pulmonale, shows the most ex- 
treme variation in Chelonia. It is so well developed in the forms 
belonging to the subfamily Cyclanorbinae that it covers the lung 
completely while in the other subfamily, Trionj'chinae, the muscle 
is totally absent. In Cyclanorhis sp. and Cyclodcrma frenaium 
(Fig. 1) the muscle arises from the carapace in the vicinity of 
the second and third thoracic vertebrae and also from the lateral 
side of these vertebrae. The fibres arising from the carapace 
run over the entire dorsal surface of the lung and when they 



M.T). 




M.5.P.M- 



Fig. 6. Diagrammatic sketch of the disposition of the respiratory muscles 
in Kachnga tectum. 



reach the outer, anterior, and the posterior limits of the lung 
they turn onto the ventral side and continue to run towards 
the entrance of the bronchus. The fibres arising from the lateral 
side of the thoracic vertebrae run over the medial side of the 
lung and then come onto the ventral side and reach the entrance 
of the bronchus. All the fibres of the muscle closely adhere to 
the wall of the lung. From the place of the origin of the fibres, 
the muscle could be arbitrarily divided into two parts, a lateral 



8 



BREVIOBA 



No. 161 



part which arises from the carapace and a medial part which 
arises from the lateral side of the thoracic vertebrae. The nuis- 
cularis striatum pulmo'nale in Lisseinys punctata (Fig. 1) 
(George and Shah, 1954) differs slightly from the one in Cycla- 
noi'Ms and Cycloderma; the muscle is otherwise very similar in 
its course and insertion in all the three genera of Cyclanorbinae. 
In Lissemys the muscle arises entirely from the carapace and 
does not have its lateral part arising from the side of the verte- 
brae. The muscle in all the forms of the group Cyclanorbinae is 



C T.B 




Fig. 7. Diagrammatic sketch of the disposition of the respiratory muscles 
in marine turtles. 



innervated by the branches of the intercostal nerves. As said 
before, the muscle on its contraction pushes out the pulmonary 
air of the lungs, and on its relaxation the atmospheric air rushes 
in. 

In Trionyx gangeticus, Trionyx sinensis, Trionyx triunguis 
and Dogania suhplana (Fig. 2) belonging to Trionychinae, the 
niuscnlaris striatum pulmonale muscles is totally absent. 

All the forms belonging to the Emydinae have a niuscnlaris 
striatum pulmonale which partly covers the lung. In this group 
the muscle shows great variation in different species. 



1962 



RESPIRATORY MUSCLES IN CHELONIA 



n.D. 




Fig. 8. Diagrammatic sketch of the disposition of the respiratory muscles 
in Testudininae, except Pyxis arachnoides. 




XA 



H.SV.L. 



Fig. 9. Diagrammatic sketch of tlie disposition of the respiratory muscles 
in Pyxis arachnoides. 



10 



BREVIORA 



No. 161 



In Malaclemys terrapin terrapin, Clemmys caspica caspica, 
Clemmys guttata, Grapteniys pseudogeographica, Graptemys 
kohni, Ciiora amhoinensis, Pscudemys fioridana and Pseudemys 
texana, the muscidaris striatum pidmonule, though only partly 
covering the lung, is well developed compared to other Emydinae. 
In these emydines the medial part of the muscle arising from 
the side of the thoracic vertebrae is well developed and quite 
extensive, while the lateral part of the muscle with its origin 
from the carapace, though well developed, is comparatively small 



M.D. 




M.S. P.M. 



T.A 



Fig. 10. Diagrammatic sketch of the disposition of the respiratory muscles 
in Pleurodira forms in which the muscuJaris siriatum pulmonale is partial 
and the diaphragmaticus muscle is absent. 



in extent. However, in Malaclemys terrapin terrapin, Graptemys 
pseudogeographica and Graptemys kohni the lateral part of the 
muscle is comparatively more developed than in the other emy- 
dines listed above. The place of oriain of the lateral part of the 
muscle in these three forms is parallel to that of the medial 
part of the muscle (Fig. 3). In Pscudemys fioridana and Pseu- 
demys texana the place of origin of the lateral part of the 
muscle is perpendicular to that of the medial part of the 
muscle (Fig. 4). 



1962 



RESPIRATORY MUSCLES IN CHELONIA 



11 



111 Emys orhicularis, Kachuga tectum tectum, Ocadia sinensis, 
Vhrijsemys picta picta, Chnjsemys picta dorsalis, Chrysemys 
picta marginata, Deirochelys rcticularia, Emydoidca hlandingii, 
Clenimys mutica, Chinemys rcevesii, Malaycuiys siibtrijuga, Geo- 
emyda punctularia funerea (Figs. 5 and 6), Geoeinyda manni, 
Geoemyda spinosa, and Geoemyda trijuga (Fig. 12) the medial 
part of the muscular is striatum pulmonale is very poorly devel- 
oped and only covers a very small portion of the anterior 
medial side of the lung. The lateral part of the muscle is also 
less developed compared to that of Malaclemys and others and 
shows variations in its extent, never covering more than a small 
portion of the anterior and anterolateral side of the lung. 



M.D 



M. S. P. 



C.T. 



T.A 



O.A; 




Fig. 11. Diagrammatic sketch of the di-spositiou of the respiratory muscles 
in Podocnemis. 



Six species belonging to the Testudiniuae, Testudo hcrmamii, 
Testudo graeca, Geochelone pardalis, Testudo horsficldii, Testudo 
Ideinnianni (Fig. 8) and Pyxis arachnoides (Fig. 9) have been 
examined. In all these except Pyxis arachnoides the muscularis 
striatum pulmonale is totally absent and a thin sheet of con- 
nective tissue is present in its place. In Pyxis arachnoides (Fig. 
9) there is a poorly developed lateral part of the muscularis 
striatum pulmonale present, covering only a very small part 
of the anterior region of the lung. The presence of part of the 



12 



BREVIORA 



No. 161 



muscle in Pyxis appears to be a case of an intermediate stage 
between the typical condition of the Testudininae, on one hand, 
where the muscle is absent, and that of Bmydinae, on the other, 
in which it is better developed. 

In Stcrnotheriis odoratus and Sternotherus carinatus yyiinor, 
belonging to Kinosterninae, the muscularis striatum pulmonale 
is similar in its origin, course and insertion to that described 
for the Malaclemys terrapin terrapin (Fig. 3). In Chelydra ser- 
pentina of the Chelydrinae the muscle is completely absent and 
instead a thin layer of connective tissue is present in its place. 

In Chelonia mydas, Caretta caretta, Lepidochelys olivacea and 
Eretmochelys imhricata (Fig. 7), which all belong to the family 
Chelonidae, the muscularis striatum pulmonale is totally absent. 
Even in Dermochelys coriacea (Fig. 7), of the Dermochelyoidea, 



OA. 




C.T.BH— ^ 



T, A. 



Fig. 12. Diagrammatic sketch of the disposition of the respiratory muscles 
in sucli Emydinae as Geoemyda spinosa, etc. 



the muscle is absent. Thus, it appears that none of the marine 
forms possess any lung muscle, and that unlike the land forms 
or the fresh water ones there is no variation in this regard. 

All the members belonging to the suborder Pleurodira possess 
a partial muscularis striatum pulmonale rather similar to that 
seen in the Emydinae (Fig. 5). The members of the genus 



1962 RESPIRATORY MUSCLES IN CHELONIA 13 

Podocneniis show an unusual extension of the muscularis stria- 
tum pulmonale. In Poclocnemis nnifilis and the other two species 
of Poclocnemis, the medial part of the muscle, after its usual 
origin, runs forward adhering to the medial wall of the lung 
and (unlike the normal condition where the fibres terminate 
on the lung near the entrance of the bronchus) after reaching 
the anterior limit of the lung continues forward and closely 
adheres to the dorsolateral side of the pericardial membrane. 
Finally, these fibres insert on the membrane at the level of 
the anterior side of the auricles (Shah, in press). Such a 
pericardial extension of the m,uscularis striatum pulmonale is 
not found in any of the other pleurodirans that were studied. 
No trace of such an extension is present in any of the Crypto- 
dira, nor does there appear to be any previous record of the 
presence of a striated muscle layer on the pericardium in any 
vertebrate. 

FLANK CAVITY MUSCLES 

The diaphragmaticus and the transversus abdominis form the 
expiratory set of the flauk cavity muscles in Chelonia, while 
the serratus magnus and the ohliquus ahdominis form the in- 
spiratory set (McCutcheon, 1943 ; George and Shah, 1954, 1958 
and 1959). 

In all Chelonia the transversus abdominis muscle is well de- 
veloped. In the Cryptodira studied, the transversus ahdominis 
muscle is the most highly developed in the Trionychoidea, and 
in this group it joins with the anteriorly placed diaph ragmaticus 
muscle of the same side to form a continuous muscular sheath 
which envelops the visceral organs including the lungs. The 
muscle arises in all the chelonians from the posterior half of 
the carapace, but the place of origin is not constant in all forms 
since great variations in its extent are seen in different individ- 
uals. In Pseudemys floridana and Pseudemys tcxana the trans- 
versus abdominis muscle extends almost up to the level of the 
apex of the heart on the ventral side (Fig. 4). In no Emydinae, 
whether the diaphragmaticus muscle is present or not, does the 
transversus abdominis muscle have the extensive spread seen 
in all the Trionychoidea. In those Emydinae where the dia- 
phragmaticus muscle is present, there is a bridge of connective 
tissue between it and the transversus abdominis of the same 
side. Such a bridge of connective tissue between the diaphrag- 
maticus and the transversus abdominis muscle is present in all 
the species of the genus Geoemyda and all the marine chelonians. 



14 BBEVIORA No. 161 

The diaphragmaticus muscle, in all the cheloiiians in which 
it is present, arises from the nndersurface of the second or 
third costal plates of the carapace. Its place of origin is oriented 
transversely with respect to the vertebral column. In all the 
forms of Trionychoidea (Figs. 1 and 2) the muscle is very 
highly developed and, as mentioned above, it joins the trans- 
versiis ahdo)iiinis muscle of its side to form a continuous mus- 
cular sheath to envelop the viscera. In some Emydinae the 
muscle is present (Fig. 12) ; in others it is absent (Figs. 3, 4, 
5, and 6). No members of the Testudiuae have the diaphrag- 
maiicus (Fig. 8) ; there is a thin layer of connective tissue in 
its place. 

In all the chelonian studies the inspiratory muscles, the .s'^r- 
ratus magnus and the ohliquus ahdominis (Figs. 1 to 11), are 
present with such slight variation that these are not worth 
detailed discussion. On contraction of these muscles the volume 
of the body cavity is increased and thus a negative pressure 
is created in this cavity and so the lungs expand. On expan- 
sion of the lungs the atmospheric air rushes in and in this way 
inspiration is brought about. 

DISCUSSION 

From the present stud}^ of the respiratory muscles in Chelonia 
it is evident that there is a great deal of noticeable variation in 
two muscles, the diaphragmaticus and the muscularis striatum 
pulmonale. The variations in these two muscles range from a 
highly developed condition to a total absence, with all inter- 
mediate stages. The other respiratory muscles are always pres- 
ent, and although slight variations in different forms are seen, 
these are very minor ones. 

The presence of the muscularis striatum pulmonale in all the 
Cyclanorbinae, where the muscle covers the lung completely, is 
regarded as a primitive character which is retained in these 
forms. The early ancestral chelonians presumably developed 
these muscles as a substitute for the intercostal muscles lost 
when their body w-all was covered by the rigid shell and could 
not have the normal movements which are the main component 
of the respiratory mechanism in all other amniotes. The muscu- 
laris striatum pulmonale must thus have been of survival value 
to the early ancestral chelonians, and it is retained fully in all 
the Cyclanorbinae but shows a gradual trend toward total dis- 
appearance in other chelonians. Some chelonians, the Triony- 
chinae, Testudiuae and all the marine forms, have totally lost 



1962 RESPIRATORY MUSCLES IK CHELONIA 15 

this muscle. It is quite obvious that the presence of the muscle 
covering the lung is a hindrance to full expansion of the lungs; 
it must therefore have developed as a stop gap arrangement to 
tide over the loss of the body wall movements until some better 
physiological adaptation for respiration was achieved. Unpub- 
lished Avork by the author on the blood of some cheloniaus shoAvs 
some interesting results. The oxiphoric capacity of the blood of 
Lissemys punctata Avhere the muscularis striatum pulmonale 
covers the lungs completely is much less than that of tlie blood 
of Trionyx or Tcstudo elegans where the muscle is totally absent. 
The oxiphoric capacity of the blood of Gcoemyda trijnga, where 
the muscularis striatum pulmonale muscle is incompletely de- 
veloped (Fig. 12), shows intermediate values. Thus from the 
study of blood some light is thrown on the new physiological 
adaptations which have taken place, substituting for some of 
the morphological adaptations of the primitive forms. More 
work on the physiology of respiration in different chelonians 
will be necessary for a better understanding of the problem of 
respiratory mechanism in this order. Some aspects of this are 
being worked on at present in my laboratory at the University 
of Baroda. 

ABBEEVIATIONS USED IX FIGUEES 

Br. Bronchus 

CT. Connective tissue 

O.T.B. Bridge of connective tissues between diaphragmnticus and the 

iransrersus abdoviinis muscles. 
D. Diaphragmatwus muscle 

H. Heart 

Lu. Lung 

M.D. Connective tissue in place of the diaphra gmuticus muscle. 

M.S. P. Muscularis striatum pulmonale muscle covering the left lung 

M.S.P.i Muscularis striatum pulmonale muscle cut horizontally and the 

right lung removed so as to show the place of origin of the 

muscle 
M.S.P.L. Lateral part of the muscularis striatum pulmonale 
M.S.P.M. Medial part of the muscularis striatum pulmonale 
O.A. Obliquiis abdomiiiis muscle 

P.M. Pericardial membrane 

P. Ext. Pericardial extension of muscularis striatum pulmonale. 

S.M. Serratus magnus muscle 

T.A. Transversus abdominis muscle 



16 BREVIORA No. 161 

EEFERENCES 

George, J. C. and R. V. Shah 

1954. The occurrence of n striated outer muscular sheath in the lungs 
of Lissemys punciata granosa Schoepff. J. Anim. Morph. Phys- 
iol., 1: 13-16. 

1955. Respiratory mechanism in Chelonia. J. Anim. Morph. Physiol., 
1: 30-32. 

1958. The structural basis of the evolution of the respiratory mech- 
anism in Chelonia. Proc. XVth Int. Congress of Zool. London. 
Papers read by title: 24: 1-2. 

1959. The structural basis of the evolution of the respiratory mech- 
anism in Chelonia. J. Anim. Morph. Physiol., 1: 1-9. 

McCUTCHEON, F. H. 

1943. The respiratory mechanism in turtles. Physiol. Zool., 16: 255- 
269. 
Owen, R. 

1866. Anatomy of vertebrates. Vol. I. London, xxxvii + 650 pp. 
Williams, E. E. and S. B. McDowell 

1952. The plastron of the soft shelled turtles (Testudinata, Triony- 
chidae). A new interpretation. J. Morph., 90: 263-275. 



BREVIORA 

Museum of Comparative Zoology 



Cambridge, Mass. Ji^ly 25, 1962 Number 162 



AUSTRALIAN CARABID BEETLES X. BEMBIDION 

By p. J. Darlington, Jr. 

Museum of Comparative Zoology, Cambridge, Mass. 

This is the tenth in a series of papers on Australian Carabidae. 
Some earlier parts, including a list of localities at which I col- 
lected in 1956-1958 and a discussion of transition of wet forest 
carabid faunas from New Guinea to Tasmania, are given under 
References. 

The present paper deals with the Australian species of Bem- 
hidion {sensu lato). This is a zoogeographically important genus, 
which tends to be bi-zonal in distribution, occurring mainly in 
the north and south temperate zones of the world (Darlington 
1959, pp. 332-333). The distribution, ecology, relationships, and 
possible history of the Australian forms are therefore note- 
worthy and will be summarized after discussion of the separate 
species. 

I am indebted to Prof. Carl H. Lindroth for dissecting males 
of all the Australian species and telling me how he thinks they 
are related to European and North American forms. I could 
have made the dissections myself, but I am not familiar with the 
genitalic characters of northern Bemhidion and could not have 
interpreted the characters of the Australian species. However, 
neither Prof. Lindroth nor. I have investigated most of the 
Asiatic species or those of New Zealand or southern South 
America. This paper is therefore only a limited contribution 
to the zoogeography of Bemhidion. 

At the time of Sloane's last study of Australian Bembidiini 
(1921), he knew five Australian species of Bemhidion and two 
o/ Cillenus. Two supposed "Bemhidion" of Blackburn's that 
Sloane did not know {hoharti and watt sense) are in fact not 
Bemhidion but Tachys. I plan to treat them in mj' next paper. 
I have series of all five real Bemhidion known to Sloane and 



2 BREVIORA No. 162 

have seen no other native species, and it may be that these five 
species of the gemis (excluding Cillenns) are all that are 
native in Australia, although it is too soon to be sure about this. 
References and synonymy of the species are given by Sloane 
(op. cit.) and will usually not be repeated here. The species 
should be identifiable by the following key, which is based partly 
on Sloane 's key (1921, p. 193). All the species are winged and 
presumably able to fly, except that the wings are dimorphic in 
proprhim (q.v.). 

Key to Australian Species of Bembidion 

1. Large (c. 5.2-6.5 mm.) ; dull bronze, elytra with 2 incomplete transverse 
pale fasciae ; clypeus with several fine converging grooves on each side 
(introduced from South America) brullei 

— Smaller; not marked as above; (native") 2 

2. Frontal sulci long, impressed and converging on clypeus ; upper surface 
of insect dull or shining ; elytron with 6 or 7 dorsal striae 3 

— Frontal sulci shallow and rather short, not crossing clypeus; upper 
surface dull ; elytron with 7 dorsal striae 5 

3. Dull brown with vague paler elytral markings; whole upper surface 
microreticulate ; elytron with (5 dorsal striae (stria 7 absent or faint); 
length c. 4.2-4.8 mm errans 

— Shining, upper surface not microreticulate; elytron 7-striate; size 
smaller 4 

4. Elytron with 2 seta-bearing punctures on 3rd interval, none on yth ; 
color black, elytral apices and sometimes lateral subapical spots slightly 
paler; length c. 3.3-3.9 mm hlackburni 

— Elytral intervals 3 and 5 each with several inconspicuous seta-bearing 
punctures; color irregular dark brown; length c. 2.9-3.4 mm. proprium 

5. Prothorax transverse, sides not sinuate posteriorly; color much like 
following species except apical testaceous area of each elytron broken 
into subapical and apical marks which are often narrowly connected 
along outer elytral margin and sometimes connected near suture too ; 
length c. 3.0-4.0 mm. jacksoniense 

— • Prothorax cordate, sides sinuate posteriorly ; color greenish or bronzed 
with elytral apices conspicuously testaceous, the testaceous areas broadly 
lunate; length c. 4.0-4.7 mm opulentum 

Bembidion (Notaphus) brullei (Gemminger and Harold) 

variegatum Brulle 1843, p. 44 (not Say). 
Bemhieidium hrvllei Gemminger and Harold 1868, p. 409. 

Form as figured, broader and less convex than usual in 
Australian species of genus; bronze, appendages irregularly 
testaceous and fuscous, elytra with complex but variable pale 



1962 



AUSTRALIAN CARABID BEETLES 



marks which usually involve epipleuri, parts of elytral bases 
inside humeri, marginal channels, apices (rather vaguely), an 
irregular interrupted fascia before middle often including very 
elongate pale areas on intervals 7 and 8 and isolated spots on 3rd 
intervals, and an irregular incomplete post-median fascia; sides 
of abdomen also pale; upper surface dull, with close reticulate 
microsculpture, isodiametric on head and pronotum, vaguely 




Betnbidion hrullei (Gemminger and Harold), from between Murray Bridge 
and Meningie, South Australia. 



transverse (but nearly isodiametric) on elytra. Head .76 and 
.76 width prothorax (in S 9 measured) ; eyes large and prom- 
inent; antennae of moderate length, middle segments c. 2I/2X 
long as wide ; front slightly convex ; frontal grooves subparallel 
(slightly curved) and poorly defined on disc, converging and 
more sharply impressed across clypeus, and latter with addi- 
tional fine longitudinal (converging) grooves or wrinkles; men- 
turn with entire tooth. Prothorax transversely subcordate with 
rather broad base; width/length 1.45 and 1.44 (in measured 
6 9); base/apex 1.19 and 1.17; base/head 1.07 and 1.06; sides 
broadly arcuate for much of length, moderately sinuate before 
basal angles ; latter well defined and nearly right or slightly 



4 BREVIORA No. 162 

obtuse; lateral margins moderate, each with usual 2 setae; disc 
convex, with anterior transverse impression poorly defined, mid- 
dle line distinct but slightly abbreviated at both ends, basal 
transverse impression poorly defined; baso-lateral impressions 
deepest inwardly, bottoms irregular or slightly convex, slightly 
wrinkled but not much punctate, limited externally by longi- 
tudinal carinae distinct from prothoracic margins. Elytra c. 
Ys wider than prothorax (E/P 1.36 and 1.35) ; humeri prom- 
inent but rounded ; sides subparallel for much of length ; mar- 
gins moderate (wider than in opulentum) , ending anteriorly 
opposite ends 6th striae (opposite 5th in opidentum) , forming 
translucent shelves before subapical sinuations ; striation entire, 
striae slightly impressed and rather strongly punctate especially 
on disc ; intervals slightly convex, 3rd with 2 seta-bearing punc- 
tures about % from base and less than % from apex. Inner 
wings fully developed. Lower surface microreticulate but mostly 
not distinctly punctate ; anterior process of raetasternum strongly 
margined between middle coxae, the margin strongly rounded- 
angulate at middle. Legs without obvious distinctive characters. 
Secondary sexual characters: $ with 2 segments each front 
tarsus moderately dilated and squamulose below; S with 1, 
$ 2 setae each side last ventral segment. Length c. 5.2-6.5 ; 
width c. 2.0-2.6 mm. 

Known in Australia only from six specimens taken by myself 
beside the road between Murray Bridge and Meningie, South 
Australia, September 1957, probably beside one of the series 
of more or less saline lakes near the mouth of the Murray River. 
However, my collecting there was done under difficulty, in the 
face of light rain driven by strong wind, and I did not dis- 
tinguish the species in the field. Specimens of Benil)idio7i errans 
and proprium were taken on the same occasion. 

Superficially, this species looks rather like Bemhidion (Nota- 
pkus) dentellum Thunberg and related species of the Northern 
Hemisphere, and Prof. Lindroth says its genitalic characters are 
those of a true Notaphus: "The armature of the internal sac, 
even in details, comes very close to that of approximatum Lee. 
and coloradense Hayw." of North America. The sculpture of 
the clypeus of brullei is distinctive but, I think, not of sub- 
generic value. 

I was on the point of describing this as a new Australian 
species when I discovered specimens of it in the Museum of 



1962 AUSTRALIAN CARABID BEETLES 5 

Comparative Zoology from South America, from several locali- 
ties in northern Argentina. It was described more than a hun- 
dred years ago from specimens taken on the seashore at Monte- 
video, and it is apparently common in the La Plata region and 
at Cordoba, Argentina. It has presumably been introduced into 
southern Australia by shipping. 

Bembidion (Ananotaphus) errans Blackburn 

Netolitzky (1931b, pp. 181-182) made this the type of sub- 
genus Ananotaphus, which may be considered to include also the 
two following species: propriur)i Blackburn and hlackburni 
Csiki. The three species in question are somewhat alike in form 
and agree in having deep frontal sulci, and the male genitalia 
"show a certain, though not very evident, agreement in the 
internal sac" (Lindroth). However, the three species differ con- 
siderably among themselves, and their relationship to other sub- 
genera is doubtful, so far as the genitalic characters are con- 
cerned. 

Blackburn and also Sloane (1921) thought that errans oc- 
curred only near the coast, and I agree, although it is not con- 
fined to obviously saline habitats. It has been previously recorded 
from southern "Western Australia, South Australia, and Victoria. 
I have a series from southern Western Australia, from several 
localities including the vicinity of Perth and Pemberton, collected 
by H. Demarz, and I took two specimens between Murray Bridge 
and Meningie, South Australia, and four near Hobart, Tasmania. 
In Western Australia Sloane found it "on the muddy margin 
of the Vasse River within the tidal influence," and some of 
Demarz 's specimens were taken at Mandura salt lake. My South 
Australian specimens were probably taken by slightly saline 
ponds near (east of) the mouth of the Murray River (see under 
preceding species). My Tasmanian specimens were on the 
grassy-muddy bank of a small pond in flat, lowland country north 
of Hobart. This was essentially a fresh-water habitat, but it was 
only a few miles from the coast and there may have been a 
trace of salt there. A series of B. proprium and one specimen 
of hlackhurni were taken at the same place. 

Bembidion (Ananotaphus) proprium Blackburn 

Sloane (1921) knew this species too only from localities on 
or near the coast of southern Australia, including South 



6 BREVIORA No. 162 

Australia, Victoria, and southern New South Wales (Wollon- 
gong). One of his specimens was "beside a little rivulet near 
where it entered the sea" (presumably at Wollongong). My 
mainland specimens too are from coastal localities, from between 
Murray Bridge and Meningie (see under B. Irullei) and between 
Meningie and Kingston, South Australia (taken by myself), and 
from Seaford, Victoria (taken "under beach drift" by W. L. 
Brown). However, I took a series in Tasmania, north of Hobart, 
beside fresh water, though still near the coast (see under 
errans). The species thus seems to have about the same ecological 
distribution as errans. 

My seven mainland specimens are all fully winged. Most of 
the sixteen from Tasmania have the inner wings somewhat re- 
duced, about as long as the elytra, slightly folded or crumpled 
at apex, and evidently unfit for flight, but one of the Tasmanian 
specimens is fully winged or nearly so. 

Bembidion (Ananotaphus) blackburni Csiki 

Csiki 1928, p. 159. 

(Jubiinn Blackburn 1888 (not Heer, not Wollaston). 

Sloane (1921) records this species from South Australia, 
Victoria, and southern New South Wales, and notes that it 
occurs beside fresh water, in some cases much farther from the 
coast than the preceding species. I took it at Winchelsea, Vic- 
toria, and found it common in Tasmania. It occurred in wet 
places at low altitudes near Hobart (see under errans) and near 
Ellendale, but it was commoner on the mountains, near Lake St. 
Clair (over 2400 ft. altitude) and Great Lake (c. 3400 ft.), 
for example. On the mountains in Tasmania hhckhurni occurs 
not only beside standing water and in other wet places but 
sometimes also on wet, open heaths near and above tree line. 
Here it runs in sunlight in and on the dense mats of moss and 
other vegetation that cover much of the ground in open places. 
This is a true subantarctic habitat. Benihiclion hlackhurni is the 
only Australian species of the genus that reaches a subantarctic 

habitat. 

On the wet mountain heaths, B. blackburni is a member of a 
small association of superficially similar species of small black 
Carabidae including Cyphotrechocles gibbipennis (Blackburn), 
Amblystomus nigrinus Csiki {niger Blackburn), and Euthcnarus 
nigelhis Sloane. These species look so much alike, superficially, 
that it is not easy to distinguish them in the field with the naked 



1962 AUSTRALIAN CARABID BEETLES 7 

eye. They occur together on warm days on wet moss pads, etc., 
on the mountain heaths as well as in other wet places. This is a 
striking example of convergence of species of unrelated carabid 
genera. I do not know its ecological significance. I should add 
that, although these species occur together under some circum- 
stances, they do not have identical ecological limits elsewhere. 

Bembidion (Philochthus) jacksoniense Guerin 

[Subgenus SloanephUa Netolitzky is here declared a synonym of Philochthus, 
for reasons given below (new synonymj')-] 

See Sloane (1921, p. 193) for sj'-nonyms and references, and 
see Netolitzky (1931b, p. 182) for subgenus SloanephUa, based on 
this species. However, Netolitzky himself notes the great simi- 
larity in most characters of jacksoniense and species of the 
northern (Europe, western Asia, North Africa, etc.) subgenus 
Philochthus, and Prof. Lindroth finds ' ' general agreement in the 
arrangement of the internal sac of the male genitalia in jack- 
soniense and subg. Philochthus . . . ," and it seems to me there 
is more to gain by recognizing the relationship and putting 
jacksoniense in Philochthus than by stressing the differences. 
Netolitzky says the principal difference is that the metasternal 
process between the middle coxae is margined in the northern 
Philochthus, not in jacksoniense. Netolitzky adds that in this 
character and in frontal sculpture jacksoniense closely resembles 
B. " Notapliojuimus" {=Notaphocampa) opulentum (below). I 
think these facts may indicate (1) a close relationship between 
jacksoniense and the northern Philochthus and (2) a less close 
one between Philochthus and Notaphocampa, with opule7itum 
perhaps in some ways a connecting link. 

Sloane says jacksoniense is found over the whole continent of 
Australia, beside fresh water, and this is probably essentially 
true. However, it may be absent in small areas in the east and 
perhaps in larger ones in the north (I am not sure about this) 
and it has not yet been found in Tasmania, and it certainly 
occurs in brackish and perhaps alkaline habitats (inland) as well 
as by strictly fresh water. It is veiy common in Western Au- 
stralia but less so in the east, although I have specimens from 
several eastern localities ranging from Townsville in tropical 
Queensland south to the Blue Mountains of New South Wales 
and Mt. Kosciusko. In some localities it occurs with the follow- 
ing species, opulentum. In light-trap material from Cooper 
Crossing, Lake Eyre, it is represented by a few specimens among 
many opulentum. 



8 BREVIORA No. 162 

Bembidion (Notaphocampa) opulentum Nietner 

[Subgenus Notaphomimus Netolitzky is here declared a synonym of Nota- 
phocampa, for reasons given helow (new synonymy).] 
sobrinum auet. (not Boheman). 

Sloane (1921, p. 193) gives Australian synonyms and earlier 
references to this species. Netolitzky (1931b, pp. 175-178) makes 
an (I think) unnecessary special subgenus, Notaphomimus, for 
it. Except for the difference in frontal sculpture, which I think 
is over-stressed by Netolitzky, opulentum seems very close to 
Bemhidion (Notaphocampa) niloticum Dejean, and the two 
species are closely allied in geuitalic characters (Lindroth). 

I have discussed the distribution of opulentum {'"sohrinum") 
recently (1959, p. 339), but my statement that the species ranges 
from Africa through tropical Asia, etc., was apparently wrong, 
although made on good authority (Netolitzky, Andrewes). There 
seem to be two slightly different species: foveolatum Dejean 
{sohrinum Boheman) of Africa, Madagascar, etc., and opulentum 
Nietner of the Oriental-Australian region (see Jeannel 1946, 
pp. 370-371, and Basilewsky 1952, p. 177). B. opulentum occurs 
through tropical Asia, part (but not all) of the Malay Archi- 
pelago, part of tropical and south temporate eastern Australia, 
and apparently also in New Caledonia. It exhibits some 
geographical variation (Netolitzky, loc. cit.) : the Australian 
form can be called subspecies riverinae Sloane. In Australia it 
is widely distributed in the eastern part of the continent at least 
from the latitude of Townsville south to Tasmania and west to 
South Australia, but I do not think it has yet been found in 
Western Australia. (Sloane knew it only from Queensland, 
New South Wales, and Victoria.) It usually occurs near the 
margins of standing or slowly moving fresh water. Six specimens 
that I took behind the bank of the Burdekin River near Charters 
Towers, Queensland, in March 1958, were running on wet mud 
and beside a stagnant pool of flood water. A specimen from near 
Waratah, Tasmania, was taken on a log floating in water at the 
side of a small pond. And three from west of Renmark, South 
Australia, September 1957 were, I think, taken on mud behind 
the bank of the Murray River. However, the species apparently 
occurs also in inland saline areas, for I have a long series taken 
at light at Cooper Crossing, Lake Eyre, South Australia, in 
February 1956 (collected by Dr. G. F. Gross). That this species 
is so widely distributed in Asia but has not yet spread through 
the whole of Australia suggests that it has reached Australia 



1962 AUSTRALIAN CARABID BEETLES 9 

comparatively recently, possibly by way of the Lesser Sunda 
Islands and Timor, for it does not seem to occur in New Guinea 
or Cape York. 

Bembidion, subgenus Cillenus 

Cillcnus is commonly considered a subgenus of Bcmhidion. 
Its gross range is from Europe and China to Australia, Tasmania, 
and New Zealand, but it is strictly coastal. Different species of 
it occur by running water near the coast, or on ocean beaches, 
or actually between the tide lines. I have discussed its distribu- 
tion and possible history elsewhere (1953; 1959, pp. 333-334). 

Two species of Cillenus are known from (eastern) Australia: 
a larger, duller one (albovirens Sloane) from tropical Queensland 
(from Townsville, not Cairns as incorrectly stated by Sloane in 
1921), and a smaller, more slender, more shining one (mastersi 
Sloane) from Sydney and the north coast of Tasmania. Both 
species probably live on the ocean beach. Both are (irregularly) 
testaceous, as beach-living Carabidae often are. Both Australian 
species are fully winged and one or both probably fly. My single 
specimen of albovirens probably flew to light at Townsville, al- 
though I did not actually see it do so. Some other Cillenus in 
other regions including New Zealand are flightless. 

SUMMARY AND DISCUSSION 

The five native Australian species of Bembidion (excluding 
Cillenus, which has evidently had an independent history) are 
distributed as follows. The three species of subgenus Ananota- 
jyhus occur in the southern part of Australia, and all of them 
extend to Tasmania too. Of these three, errans and proprium 
apparently occur only at low altitudes near the coast, sometimes 
in saline but also sometimes in fresh or nearly fresh habitats 
beside water, while blackburni is more widely distributed. It 
sometimes occurs at low altitudes with the two other species, but 
it has also entered the subantarctic zone on Tasmanian moun- 
tains. B. (Philochthus) jacJcsoniense occurs almost throughout 
Australia (but not Tasmania) in both fresh and saline (interior) 
habitats, usually beside standing or slowly moving water. And 
B. (Notaphocampa) opulentum is widely distributed in the 
Orient and in tropical and south temperate eastern Australia. 
It reaches Tasmania but apparently not Western Australia. It 
occurs in about the same variety of habitats as jacksoniense. 



10 BREVIORA No. 162 

These five native Australian species of BemhkUon are appar- 
ently descended from tln-ee successive invaders. Tlie first was 
the ancestor of the endemic subgenus Ananoiaphus, which has 
difl'erentiated in Australia and formed three verj^ distinct species. 
All three are now confined to the southern edge of Australia plus 
Tasmania; one of the three (hlackhurni) extends into sub- 
antarctic habitats on mountains in Tasmania; and one {prop- 
rium) is undergoing- wing atrophy. This looks like an early stage 
in penetration of the southern cold temperate zone by an orig- 
inally tropical or northern winged group of Carabidae, and 
evolution of a derivative flightless stock adapted to southern 
cold temperate habitats. The ancestor (s) of the Australian and 
Tasmanian "Trechns" may have gone through a stage like this. 

The second invader was the ancestor of Bemhid/ion (Philoch- 
thus) jacksoniense. The latter has spread throughout Australia 
(but not Tasmania) and is now a thoroughly distinct species, 
although its relationship to the northern species of the sub- 
genus is still clear. In southern Australia it overlaps the edge 
of the range of the species of Ananoiaphus but (in my experi- 
ence) it does not often actually occur Avith them and perhaps 
does not compete with them very much noAV, although it may 
limit their distribution northward. 

The third invader is Bemhidion (Notaphocampa) opulentum. 
The fact that this species is now widely distributed in southern 
Asia (with a very close relative in Africa) but has apparently 
not yet spread through the whole of Australia, and the fact that 
the Australian population is only subspecifically diiiferentiated, 
suggest that the species has reached Australia rather recently. 
In eastern Australia it occurs with jacksoniense, in the same 
habitats. The tAvo species may compete, and opulentum may be 
replacing jacksoniense, which seems to be much less common in 
eastern Australia, where opulentum occurs, than in the west, 
where opulentum does not occur. 

This history of three successive invasions, the later invaders 
perhaps competing with and modifying the distributions of 
earlier ones, accounts very well for the present distribution of 
Bemhidion in Australia. The question then is, what has been 
the history of the ancestral forms outside Australia? 

Bemhidion is now dominant in the north temperate zone but 
is very poorly represented in the tropics, where the genus is 
replaced by swarms of Tachys. The obvious, but not necessarily 
the correct, guess, therefore, is that the three Bemhidion that 
have invaded Australia have somehow crossed the tropics from 



1962 AUSTRALIAN CARABID BEETLES 11 

the north temperate zone. This would be consistent with the 
distribution of Philochthus, which is now Avholly north temperate 
except for jacksonicnse isolated in Australia. 

However, another, perhaps more probable history can be sug- 
gested. The endemic Australian subgenus Ananotaphus vaguely 
resembles Notaphocampa in form. Ananotaphns has deep frontal 
sulci and Notaphocampa opulentum has shallow ones, but opu- 
lentum is very close to Notaphocampa niloticmn Dejean in most 
characters including male genitalia (Lindroth), and niloticum 
has deep frontal sulci. Ananotaphus and Notaphocampa are in 
this way linked by niloticum, which, incidentally, ranges from 
North Africa across tropical Asia north to temperate Japan. 
Similarities between Notaphocampa opulentum and Philochthus 
jacksoniense are noted under the latter in the preceding pages, and 
iacksoniense is clearly related to northern species of Philochthus. 
These are hints that Ananotaphus, Notaphocampa, and Philoch- 
thus, which include all the Bcmhiclion (except Cillenus) native 
in Australia, may belong to one group which has had a rather 
complex evolutionary and geographical history, and the group 
may have been primarily tropical. B. opulentum and niloticum 
are widespread in the tropics now ; all the Australian Bemhidion 
are salt-tolerant, which is characteristic of tropical Bemhidion 
(Darlington 1953, p. 14), and at least one European Philochthus 
{aeneum Germar) is salt-tolerant too (Lindroth) ; and evolution 
in the tropics would facilitate the successive invasions of Au- 
stralia that have occurred. 

I suggest, then, that subgenera Ananotaphus, Philochthus, 
and Notaphocampa may all be derived from one salt-tolerant 
group of Bemhidion that has evolved primarily in the Old World 
tropics, has invaded Australia three times, and has invaded the 
north temperate zone at least twice, once as Philochthus in Eu- 
rope, etc., and once as Notaphocampa niloticum in Japan. This 
hypothetical history cannot be critically tested until the phylo- 
geny of the whole genus Bemhidion is better understood. Geni- 
talic characters do not clearly confirm it. If the forms in ques- 
tion have evolved as suggested, their history has probably been 
complex and may have involved additional groups of Bemhidion 
that have not reached Australia. 

Cillenus is another, independent subgenus of Bemhidion (per- 
haps it should be considered a related genus) that has crossed 
or evolved in the tropics, in saline habitats, and reached Australia 
(Darlington 1953: 1959, p. 333). 



12 BREVIORA No. 162 

It is noteworthy that, although several stocks of Bemhidion 
have reached the western part of the Malay Archipelago from 
Asia apparently by "mountain hopping," by somehow dispersing 
from mountain to mountain and from island to island at con- 
siderable altitudes (Darlington 1959), none has reached Aus- 
tralia or New Guinea in this way. No Bemhidion has been found 
on mountains in New Guinea, and the Australian species are 
primarily lowland forms and seem to be descended from salt- 
tolerant ancestors that crossed or evolved in the tropics at low- 
altitudes. 

EEFERENOES 

Blackbuun, T. 

1888. [Australian Benibidiini.] Trans. R. Soc. South Australia, 10: 

38-45. 
1901. [Australian Benibidiini.] Trans. R. Soc. South Australia, 25: 
120-124. 
Basilewsky, p. 

1952. [Types of African Bemhidion, etc.] Bull. Ann. Soc-. Ent. 
Belgique, 88: 173-194. 

Brulle, a. 

1843. (In) d'Orbigny, Voyage dans I'Amerique meridionale, 6, Part 
2. Insectes. 
CsiKi, E. 

1928. Junk-Sehenkling Coleopterorum Catalogus, Carabidae, Harpa- 
linae I. 
Darlington, P. J., Jr. 

1953. [Cniemis.] Coleopterists ' Bull., 7: 12-16. 

1959. Bemhidion and Trechus of the Malay Archipelago. Pacific 

Insects, 1: 331-345. 
1961a. Australian carabid beetles IV. List of localities, 1956-1958. 

Psyche, 87: 111-126. 
1961b. Australian carabid beetles V. Transition of wet forest faunas 
from New Guinea to Tasmania. Psyche, 68: 1-24. 
Gemminger, M. and B. de Harold 

1868. Catalogus Coleopterorum, 1. 
Jeannel, R. 

1946. Coleopteres carabiques de la region malgache, part 1. 
Netolitzky, F. 

1931a. [Palearctic Bembidion.] Koleopterologisehe Rundschau, 28: 

28/1-124/96, D/97-70/166. 
1931b. Uberpriifungen afrikanischer und australischer Benibidiini. 
Wiener Ent. Zeitung, 47: 169-183. 
Sloane, T. G. 

1921. [Australian Bembidiini.] Proc. Linn. Soc. New South Wales, 
46: 192-208. 



BREVIORA 

MiuseiLim of Compairative Zoology 



Cambridge, Mass. Jilv 26. 1962 Number 163 

NEW W0RM-LIZAKD8 

(ANCYLOCRAXirM AND A3IPHISBAEi\A) 

FROM SOUTHEASTERN TANGANYIKA TERRITORY 

By Arthur Loveridge 

1. ANCYLOCRAxn':\r 

On the 27tli May. lf).")0. Mr. C. J. P. lonides collected at 
Newala an Anciilocrauium whicli, later that same year, he do- 
nated to the British Museum (Nat. Hist.). AYhile expressing 
the liopc that more material would be forthcoming'. I tentatively 
identified it with A. harkeri Loveridge (1946, Proe. Biol. Soc. 
\Yashington, vol. 59. p. 78, figs.), a species then, as now. known 
only from the 6 holotype (M.C.Z. 48950). 

Last year Mr. lonides was successful in securing six addi- 
tional Ancylocranium from NcAvala, the most southerly record 
known for any member of this ]ieculiar genus. I find that all 
six differ from ^1. harleri of Lindi District in precisely the same 
characters as did the first, as recorded in my manuscript notes. 
Consequently, I propose to designate these Newala Anci/locrn})- 
iiim as a southerly form of harkeri . viz. 

Ancylocranium barkeri newalae subsp. nov. 

Holotype. Museum of Comparative Zoology No. 67001 (lonides 
No. 9356), a presumed 6, from Newala, Southern Province, 
Tanganyika Territory. Collected by C. J. P. lonides, Esq., on 
12th June, 1961. 

Paratypcs. lonides Nos. 8844, 9351-3, 9355. 9359, of which 
three are now in the Museum of Comparative Zoology (M.C.Z. 
67002-4) and three in the British Museum Nat. Hist.) (BM 
1959.1.5.18. 1962.177 and 1962.178). 



2 BREVIORA No. 163 

Diagnosis. Head shields substantially the same as figured for 
.4. harkrri from which. howeA-er, it may be distiiifriiished as 
follows : 

Median ventrals about six times as broad as their fellows; 

complete caudal annuli five or less. 

27 CIS + 9) or 29 flS + 11) sej^ments in a midbody annulus; 
205-215 annuli on body, normally 3 on underside of tail from 
post-anal rin<r to conical tip; known only from 7 specimens 
from Newala _ _ h. newalae 

31 (20 -f 11) sep-ments in a midbody annulus; 222 annuli on 
body, 4 on underside of tail from post-anal rinpr to conical 
tip ; known only from ,j liolotype ex Mbemkuru River, Lindi 

district h.harkeri 

Median ventrals scarcely broader than their fellows ; com- 
plete caudal annuli nineteen or more. 

32 (16 + 16) or 34 (18 + 16) segments in a midbody annulus; 
301-327 annuli on body, 19-23 on underside of tail; known 
from 7 specimens ex Kilwa and also Mtene, Rondo Plateau 
(lonides writes me that he has since obtained 20 more from 

the Rondo Plateau, but these have not been studied) ionidesi 

Description. Rostral enormous, compressed, arched, with sharp 
cutting edge ; remaining head shields also as in the typical form 
except that the median upper labial is higher and relatively 
larger: also there is only a single anterior chin-shield (i.e. post- 
mental) in all paratypes though that of the type (which is the 
youngest) bears a faint trace of a longitudinal groove. 

Segments in a midbody annulus 27 (18 + 9) (in two para- 
types 29 (18 + 11) ) : annuli on body 205 (208-215 in paratypes) ; 
annuli on underside of tail from post-anal ring to conical tip 3 
(4 in I 9353 only), above tail 6 (7 in all paratypes) ; anals 5 
(as a result of the fusion of the median pair seen in &. harkeri, 
shown by traces of a longitudinal suture in h. newalae para- 
types I 9351-9352). 

Color. In alcohol. Rostral horn-colored, rest of head and 
body white ( ! flesh-pink in life), tip of tail purplish brown, the 
pigmentation extending backwards on the upper surface to be- 
yond its conical end, and undoubtedly serving as a pseudo-head 
for the amphisbaenid while burrowing. 

Size. Total length of holotype $ , 228 (218 + 10) mm., which 
is the smallest of the series. Largest, apparently a 9 , 255 (245 
+ 10) mm. is now in the British Museum. 



1962 NEW TANGANYIKA WORM-LIZARDS 3 

Discussion. Sexiiis'. None of the spven specimens appears to 
he breediii». As indicated above, h. newaJae is the most special- 
ized of the three AnciiJocranium occnrrino; in Tanpranyika Terri- 
tory. Its very shoi-t tail renders sexino; difficnlt witliont damage, 
but the type of h. harkcri was a verified S in which the tail 
length was included 24.1 times in its total length. Quite prob- 
ably tail lengtli is no indication of sex in this species, but if it 
is one might expect S S to have longer tails. If this is the case 
then four of the Newala lizards are likely to be S S and three 
( I 9852, 9355, 9359) are 9 9. In that event the largest example, 
whose measurements are given above, would be a 5 , whereas 
when I examined it two years ago I thought it a 9 . 

In 7 h. ncwalae, "l i i : tail 22.8 to 25. G times in total length. 
? 9 9 : tail 26.1 to 29.3 times in total length. 

In 1 b. harkeri, 6 : tail 24.1 times in total length. 

In 7 ionidesi, 1 S $ : tail 10.5 to 11.7 times in total length. 
9 : tail 12.7 times in total length. 

For figures and description of the strikingly different Ionidesi 
see Loveridge, 1955, Journ. East African Nat. Hist. Soc. vol. 22. 
p. 177. figs. 

2. Amphisbaena 

The genus Amphisbaeua furnishes us with an interesting 
series of stages in the specialization of worm lizards by reduc- 
tion of head shields, annuli. etc. From the same district in 
which the Ancylocranimn was discovered, two more undescribed 
amphisbaenids have been obtained by Mr. lonides. Were it not 
for the fact that his name already figures in both genera as a 
result of earlier discoveries made by him, one would like to 
show appreciation for his generosity in enriching the collec- 
tions of the Museum of Comiiarative Zoology and British Mu- 
seum (Nat. Hist.) by naming one of these novelties after him. 
Instead, and as they may eventually be assigned subspecific 
rank, it is as well to name each one after its type locality. 

Newala, situated on the edge of the Makonde Plateau at 
2,600 feet, is headquarters for the district of the same name and 
some sixty air miles from the Eondo Plateau where occurs an- 
other member of the genus with a quite distinctive arrangement 
of head shields. The new species may be known as : 



a 



4 BREVIORA No. Id'.i 

Ampittsbaena newalaensis, sp. nov. 

Ilohifjlix. Mnspum of Comparative Zoolopry No. 6700.') (Ion- 
ides No. *»n7). a i from NeAvala, Newala District. Southern 
Province, Taniianyika Territory. T'ollected by C. J. P. Tonides, 
on 19 January, 1061. 

Varahipes. lonides Nos. 9112. 9114-6. 9118-22. of wliich fonr 
(■ now in tlie Mnspuni of Comparative Zoolooy (M.C.Z. 67006-9). 

and five in tlie British Museum (Nat. Hist.). Collected by lonides 

from 17 to 20 January, 1961. 

Diof/nosis. Referable to the subp-enus Cynisca, in which it is 
intermediate between .1. cirvrhecl-i (Werner) with whose head 
squamation vciraJaoisis ay-rees. and .1. rondooisis Loveridpe. 
witli which it concurs in tlie numl)ci- of segments in a midl)ody 
annulus. Ti-cnds ina^' most readily lie seen from the followinjr 
key embracinu' variational I'anpe — approximately 20 in eacli 
of the other thr(M' species, of each of which 1 have seen ;")() or 
more specimens. 

Posterior temporal lacking (beinir fused with ]>arietal). 
20 (10 + 10) segments in a midbody annulus; 227-247 annuli 
Oil liody. 28-28 on tail: Kondo Plateau jviulocnsis 

rostciioi- leiiij)oi'al i)resent immediately below parietal. 
20 (10 + 10) seiiiitents in a midbody annulus; 239-255 annuli 
on body. 22-24 on tail: ^lakonde Plateau ik iroloensis 

22 ! 10 + 12) sepmenis in a midliody ainiulus; 252-277 ainiuli 
on l)ody. 22-27 on tail: Nangiu-nwe, Newala )iangun(Wfnsis 
24 (12 + 12). rery rarely 22 (10 + 12) segments in midbody 
annulus; 264-280 annuli on body, 25-28 on tail: Mbanja and 
Lindi, Lindi District ewerhecl-i 

Df scfiption. Due to fusing of the surviving head shields in 
the subgenus Ciinlsca. their complicated nomenclature is some- 
what difficuli to follow. The student is therefore referred to 
figures 22 (( ini-hccki) and 23 ifoiidocnsls) on pages 393-394 
of my Pevision oF the Afi-ican Lizards of the Family Amphis- 
baenidae (1941. P.nll. Mus. Comp. Zool.. vol. 87, no. 5). The 
lalei'al view of civcrhrcki there shown, will be found to corre- 
spond with the head of )icic(!lacnsis in displaying behind the 
temporal a scale (]iosl('rior temjioran immediately below the 
large ])arietal. This a])idies to all ten Newala lizai-ds, as it did 
to all fifty fv'crbvcki that 1 collected at Mbanja on the Lindi 
coast. \'iewevl from above, the ])ari('tal suture of both n<tral(nn- 
sls and H<iii(iiintir( iiKis is markedly siiorter than as figured for 



1962 NEW TANGANYIKA WOBM-LIZARDS 5 

ewerhecki. Also in both new species, immediately behind the 
parietals there are frequently traces of occipitals resnltine from 
a marked tendency to enlar^'ement of the median pair of scales 
in the first annulns. 

Even so 1 have inchided this annulus as the first body annnlus 
and counted backwards to. and inclndino-, the one beai'inp- pores 
ill males, i.e. tUv one immediately precedino- the anals. 

Seo-ments in a midhody annulus 20 (10 + 10), as is also the 
case in every paratype ; aiinuli on body 241 (239-255 in para- 
types, viz. 239-243 in 6 6 , 246-255 in 9 9 ) of which the median 
pair of ventrals are only about liv, times as broad as the adja- 
cent ones; annuli on underside of tail from post-anal rino- to 
conieal tip 24 (as in all paratype $ S, 22-23 in three 9 9); 
anals 6 (5 in one paratype. due to fusion), the median pair 
much enlar<>-ed. the immediate flanking' pair of laterals trans- 
versely divided (or entire in some paratyjies). 

Color. In alcohol. Grayish white ( ' flesh-]unk ui life), uni- 
form except for the e.rfrrme tip of tail which is purplish. 

Sizr. Total length of holotype 6 . 144 (127 + 17) mm., largest 
6 in the series; largest 9 (I 9114), 152 (13(i + 16) mm. 

In the six i 6 the tail is included in tlie total length 8.4 to 
M.3 times: in the three undamaged 9 9 from 9.5 to 10.4 times. 

AmPHISBAEXA XAXGI'Rl'WENt^Is; sp. UOV. 

Holotype. Museum of Comparative Zoology Xo. 67010 (lonides 
Xo. 9249). a 3 from X'^anguruwe. ca. 1600 feet, 8 miles south 
of Xewala. Xewala District, Southern Province. Tanganyika 
Territory. Collected by C. J. P. lonides, on 16 May. 1961. 

Paratypes. :\LC.Z. 67011-19 (lonides Xos. 9250-9355). being 
37 ci o [aU used for statistics) and 69 9 9 (of which 62 were 
measured but only 42 had annuli counted i. AVith same data 
as type but collected 12-16 May. 1961. 

Diagnosis. ISee Diagnosis for ^4. niwalaensi^, where the pre- 
liminary jiaragrajih and key are e(|ually applicable to nanguru- 
irt nsis. 

Description. xVgain, see remarks under Description of ihe 
preceding species, for the head shields of nangurnu'ensis arc 
substantially similar to those of ewerhecki. 

Segments in a midbody annulus 22 (10 + 12), but counts 
made on only a score of paratypes: annuli on body 263 (252-277 
in paratypes. viz. 252-2G!-* in 6 6. 261-277 in 99) of which 



6 BREVIORA No. 163 

the median pair of ventrals are only about l^/o times as broad as 
the adjacent ones; annnli on underside of tail from post-anal 
ring to conical tip 27 (22-26 in 37 paratype $ i , 22-25 in 42 
9 9); anals 6 (4 in four paratypes, due to fusion), the median 
pair much enlarged, the immediate flanking pair of laterals 
entire, transversely divided, or reduced to a tiny wedge 

Color. In alcohol. Grayish white (? flesh-pink in life), uni- 
form except that much of the tail, both above and below, may 
be purplish. 

Size. Total length of holotype $ , 160 (143 + 17) mm., largest 
of the 38 S $ ; largest 9 (T 9250), 170 (153 + 17) mm. 

Total length of smallest 6 , 124 (111 + 13) mm., of smallest 
9 , 100 (90 + 10) mm. 

In the 38 (J (5 the tail is included in the total length from 
8.7 to 10.1 times, average 9.4 times ; in the 62 9 9 from 9.4 to 
10.9 times, average 10.1 times. 



/^^ 



BREVIORA 



MniseiiirTti of Coimparsitive Zoology 

Cambridge, Mass. Aigust 22, 1962 Number 164 



NOTES ON THE HERPETOLOGY OF HISPANIOLA. 

7. NEW MATERIAL OF TWO POORLY KNOWN ANOLES 
ANOLIS MONTICOLA SHREVE AND 
AX LIS CHRISTOPHEI WILLIAMS. 

By Ernest E. Williams 



Recent expeditions in Haiti have obtained material of two 
ancles, one of which {A. inonficula Shreve) had been previously 
recorded on the basis of the unique type specimen, and the other 
(A. christophei Williams) was known only from type and para- 
type. 

Examination of the new material sugg-ests that the two species, 
though in a number of respects strikingly different, may yet be 
related. It is, therefore, appropriate to discuss these two species 
jointly. Comparison is made also with Anolis darlingtoni Coch- 
ran = .4. ctheridgei new name,^ regarded by Cochran (1939, 
1941) as allied to monticola. 

Anolis monticola 

A. monticola Shreve 1936 was described from a single male 
(lacking most of the tail and darkened by formaldehyde) col- 
lected by P. J. Darlington in "tlie northern and eastern foothills. 
Massif de La Hotte, 1000-4000 feet, Haiti." 

No other specimens have been reported since the original 
description. However, specimens had been collected for the 
American Museum of Natural History by W. G. Hassler in 
1935 in the vicinity of Aux Cayes and Camp Perrin. These, 
like the type, are in a dark phase and show only the faintest 
trace of pattern. In Hassler 's notebook, on the other hand, there 



1 Etheridge (unpublished thesis, University of Michigan) has shown that the 
genus Xiphocercus cannot be retained. Thus. Xiphocercux diirlingtoni Cochran 
l'.i.';."i joiiKs the ijcnus AjioHk. am; Aiwli-y (liirliiuitoni Cdchran liC'.it iiuiHt in conse- 
tiuenee be renamed. I propose, with Miss ('(ichran's kind consent, tliat tlie later 
named species be called Anolis etheridgci. 



2 BREVIORA No. 164 

is an excellent description of colors in life. The American Mu- 
seum specimens (as well as Hassler's notes) were examined by 
Max Hecht some years ago and the material tentatively referred 
to monticola Shreve. 

In 1960, A. S. Rand and J. Lazell (collecting in Haiti with the 
aid of a grant from the American Philosophical Society) re- 
turned with the first well-preserved specimens of this very beauti- 
ful small lizard. 

A. S. Rand reports the circumstances under which this mate- 
rial was collected as follows : 

"The monticola were found along the trail one-half day's ride 
above Camp Perrin. The trail there ran along a hillside mostly 
cut over and grown up to dense bushes and grass. Along this 
trail were many hendersoni subsp. in the bushes, particularly 
where they were shaded by occasional trees. I remember also 
seeing cyhotes and one ricordii. 

' ' At one point we came to a small patch of rocks . . . boulders 
of various sizes, some very large, heaped one on top of another 
a bit like but less extreme than the boulder heaps in the 
Panduras Mts. in Puerto Rico. This area had not been cleared 
and growing up from among these boulders were sizeable trees 
and much smaller woody vegetation. The result was a dense, 
heavily shaded patch of bush isolated in an open sunny area. 
It was in this spot on the trees, bushes, fallen branches among 
these large boulders that we found the monticola." 

The available sample of Anolis monticola now consists of the 
following specimens:^ 

Haiti. Department du Sud. Norther n and eastern foothills, 
Massif de La Hotte, 1000-4000 ft.: MCZ 38296 (type) . Mountains 
25 miles north of Aiix Cayes on Jeremie Road: AMNH 49818, 
49845, 50108-9, MCZ 56139 (formerly AMNH 50110). In moun- 
tains on road to Jeremie about 8 miles from Camp Perrin, 2000- 
3000 ft.: MCZ 56140 (formerly AMNH 50097). Tomheau Cheval 
between Camp Perri^i and Beaumont : MCZ 62998-6300'. Grande 
Cayemite: MCZ 58026. 

The last specimen requires special discussion. It is the 
Eyerdam specimen (formerly MCZ 25483B) discussed by Miss 
Cochran (1941, p. 179) as a juvenile coelestinus with keeled 
ventral scales. Though like many Eyerdam specimens poorly 



1 Museums from which specimens are cited in this paper are alilireviateil as 
follows : AMNH, American Museum of Natural History : MCZ, Museum of Com- 
parative Zoology ; UMMZ, University of Michigan Museum of Zoology. 



1962 TWO POORLY KNOWN ANOLES 3 

preserved and dried, it seems a typical ynonticola with the char- 
acteristic swollen middorsals as well as keeled ventrals. It is 
important in extending the range of monticola to this small 
island off the north coast of Haiti's southwestern peninsula. 
Hassler's and the more recent collection were all made in a 
relatively small area in the southeastern foothills of the IMassif 
de La Hotte. However, the type, like the Grande Cayemite speci- 
men, came from north of the Massif de La Hotte. (Recent evi- 
dence, still unpublished, indicates that there may be striking 
differences between the forms to the north and to the south of 
this ridge.) 

The new specimens show that the species reaches at least 
46 mm snout -vent length. 

Color notes and sketches are available for A. monficola both 
from W. G. Hassler and from the Rand-Lazell expedition. 

Hassler's color notes are for specific specimens: 

1. 3Iale (Field no. 74, now AMNH 49845). General dorsal 
color Hooker's Green. Saddles brown green, three in number, 
narrowest middorsally, one across shoulder, two between fore 
and hind legs. A light crescent in the temporal region. Throat 
and belly dark olive green. Legs barred. Tail barred. Eyes 
Antwerp Blue, sometimes changing to greenish. Edge of orbit 
yellowish brown. Skin of fan (which is relatively small) blue, 
scales light and dark green. Occurring also in a dark phase 
almost without pattern. 

2. Female (Field no. 70, AMNH 50097 = MCZ 56140). Back 
brownish, bounded laterally by a wavy reddish line edged with a 
dark line. Sides below the dark line olive shading to very light 
yellowish green. Sides of neck yellowish green. Tail nearly 
plain brown. Legs reddish brown posteriorly. Belly nearly white. 
Upper lips bluish. Throat yellowish with some green. 

I have examined also by courtesy of Mr. Hassler (now of the 
Fort Worth Children's Museum, Fort Worth 7, Texas) a color 
sketch, made by Melville P. Cummin, of the head and neck of a 
live male from "mountains on Jeremie road about 30 miles 
from Cayes, 2000-3000 ft. alt., August 28-29, 1935." This shows 
very well the blue of the chin and the yellow green ground color 
of occiput and nape as well as the black white-centered ocelli 
which occur laterally on the occiput and nape of some male 
specimens. 

Hassler's photographs of a live male and a female contrast the 
ocelli and the strong banding of the body in the male with the 
almost patternless female. 



4 BREVIORA No. 164 

The observations of Eand and Lazell are in excellent agree- 
ment with those of Ilassler. A pencil and crayon sketch from 
life is included in their field book and Lazell 's description of the 
same specimen, now MCZ 63004, may be paraphrased as follows : 

Ground color yellow green, top of head darker. Body banded 
with velvety black. Upper and lower jaws blue, this color con- 
tinued as a stripe fading to whitish above shoulder. An area 
behind the eye and a V-shaped band across the neck ochre yellow. 
In the black mark on the neck behind the yellow is a long, thin 
sky-blue spot. Another such blue spot in the black area on the 
occiput and anterior to it, still within the black area, a larger 
white spot with a pale pink center. Legs banded, with a brown- 
ish wash. Venter and fan bright yellowish green, but base of 
fan with a bluish wash on both scales and skin. 

Lazell comments (in agreement with Hassler) that the males 
can turn very dark after capture, obscuring the markings to 
an extreme degree, such that only the markings on head and 
neck are still visible, but the jaws always retain their bluish 
color and the fan its bright yellow green with a blue wash. (The 
type was apparently preserved in this phase ; no markings at 
all are visible, but the blue of the chin remains evident.) 

All the males in the Rand and Lazell collection have two 
prominent pairs of ocelli, one on the occiput, one on the nape. 
In the figured specimen the light spots in the interior ocelli are 
divided. In the preserved specimens the light zone between the 
two pairs of ocelli may be lighter than the ground color or not. 
(This is one of the areas described by Lazell as ochre yellow in 
life.) Banding on body and limbs is always conspicuous; on the 
flanks each dark band is divided by a lighter, narrow, vertical 
streak. 

The two females obtained by Rand and Lazell show the lighter 
mid-dorsal zone bounded by a wavy dark line mentioned by 
Hassler. Rand, who has described these specimens, cites the 
color of the middorsal zone as greenish brown. The inward curves 
of the dark line are gentle, the convex portions of the line are 
produced into points by larger or smaller spots of color even 
lighter than the broad middorsal zone. There is blue on the 
sides of the chin, and the throat is green and the belly whitish 
in life. 

Anolis christophei 

The species A. christophei Williams 1960 was based on two 
female specimens obtained in the vicinity of La Citadelle of 



1962 TWO POORLY KNOWN AXOLES 5 

Kino- Christophe, one collected by W. J. Eyerdam for the Mu- 
seum of Comi:)aratiye Zoology in 1927, and one collected by 
W. G. Hassler for the American Museum of Natural History in 
1935. 

A series of this species, including the previously unknown 
males, has now been secured again in the vicinity of the Cita- 
delle by Luc and George Whiteman collecting for the Museum 
of Comparative Zoology as part of a collecting program in Haiti 
partly supported by N.S.F. grant G 16066. Unfortunately these 
are without notes on color in life or any record of dewlap color. 
They are recorded as occurring on walls and on the ground. 

The type (I\ICZ 25485) and paratype (AMNH 49736) are now 
supplemented by the following specimens (all from La Cita- 
delle) : 

MCZ 66900-19, UMMZ 122818(5). 

The color as ascertained from preserved specimens is not very 
different from that described by Cochran (1941) for the type, 
as quoted in the original description. 

Male. Snout mottled. Two darlv supraorbital stripes, neither 
sharply defined, the posterior stripe coalescent with an arc of 
dark pigment surrounding a light area which encloses the inter- 
parietal. A scalloped, crescent-shaped transverse dark mark 
on the occiput. A butterfly-shaped mark middorsally in front of 
shoulder, light-edged behind. Three similar marks between the 
fore and hind limbs and above the sacrum, but in each of these 
the anterior margins are ill defined. Tail indistinctly banded. 

On each side a light line from the ear arching downward to 
join a dark-edged light line just above root of arm, continuing 
halfway along the body. Flanks above and below light line 
strongly mottled with darker. Limbs mottled light and darlc. 

Labials above and below darkly pigmented. Venter washed 
and powdered, Avith throat and undersides of limbs and tail 
darker, with some obscure light spotting, least evident on the 
tliroat. Dewlap scales white, skin grey. 

Female. Pattern as far as discernible the same but much more 
obscure. Throat and limbs more distinctly spotted with white. 

Even in the male the pattern, while complex and distinct, is 
not prominent. Near hatchlings show the same pattern especially 
well-defined, but even in these the contrast of dark and light 
areas is not really bold or striking. In all cases specimens must 
be fully immersed in liquid before the pattern becomes at all 
evident. 



BREVIORA 



No. 164 



Comparison op A. christophei, A. monticola and A. ETHERrooEi 

A. monticola, A. christophei and A. etheridgei are all small 
anoles (44-48 mm snout-vent length) of, as far as known, quite 
local distribution. (See map.) 

Only A. etheridgei is known to be montane, reaching at least 
6000 feet elevation. A. monticola, despite its name, is not known 
above 3000 feet, and A. christophei is known only from the Cita- 
delle which is not more than 2000 feet high. 

All three belong to the set of Hispaniolan anoles that have the 
ventral scales arranged in transverse rows. They differ from 
all other Hispaniolan anoles (except the very different giant 
anole A. ricordii) in having the subocular scales separated from 
the supralabials by a row of intervening scales. In the other 
Hispaniolan anoles the suboculars and supralabials are in 
contact. 

The three species share other minor details of squamation : 
10-14 scales across the snout ; 6-7 loreal rows ; 6-7 supralabials to 
center of eye ; temporals and supratemporals finely granular ; 
interparietal smaller than ear : only one sublabial in contact 
with the inf ralabials ; lamellae under phalanges ii and iii of 
fourth toe 14-19. 

They are somewhat similar also in basic color pattern. All 
have the dorsum transversely marked. In monticola the dorsal 




 Anolis monticola 



AAnotis chrislophe 



Anolis etheridgei 



Figure 1. Map of the distribution of Anolis monticola, A. christophei 
and A. etheridgei on Hispaniola. 



1962 TWO POORLY KNOWN ANGLES 7 

marks extend on to the flanks as transverse bands ; in christopliei 
and etheridgci they are restricted to the dorsum as saddle-like 
marking's and in etheridgci rather broken up into mottling. ^ 
Monticola is strongly sexually dimorphic in pattern; etheridgei 
weakly so; chrisfophci hardly so at all. 

In spite of all resemblances, however, we deal with three 
strongly marked species, almost as distinct from each other 
as from other Hispaniolan anoles. 

Table I compares the differences in squamation between the 
three taxa. 

As Cochran suggested in the description of darlingtoni ( = 
etheridgei), this species seems the closest relative of monticola. 
Christ ophei in its lack of the specializations of the other two 
forms — in particular, in its possession of a well-developed dew- 
lap — • seems the most primitive of the trio. 

The bright bold patern of male A. monticola would seem prima 
facie an obvious compensation for the extreme reduction of the 
dewlap, which is in most species a striking species-recognition 
mark. We do not have a description of colors in life of A. 
etheridgei; it would be reasonable to expect a more vivid pattern 
than appears on the available specimens. We also lack ade- 
quate information on the color in life of christophei. In regard 
to the latter species I can only state that the material recently re- 
ceived and in a state of preservation that normally retains well 
any vivid pattern seemed nondescript and obscurely patterned 
until very closely examined. 

Whether there are any more members of this small sub-group 
of Hispaniolan anoles will have to be determined by more thor- 
ough search of the island. A. etheridgei is still known only from 
four localities in the Cordillera Central of the Dominican Repub- 
lic (Loma Vieja, Loma Rucilla, Valle Nuevo, Constanza), A. 
monticola from four localities in the foothills of the Massif de 
La Hotte and on the island of Grand Cayemite, and A. christophei 
only from the single locality, the vicinity of La Citadelle. These 
are widely separated areas. If related species are equally local 
in distribution, they may well have been missed. Of course, we 
lack at the moment any information which would permit sig- 
nitieant hypotheses on the history of these forms. There is indeed 
evidence that suggests that these three forms are one branch of 
an extremely interesting intra-Hispaniolan radiation ; this con- 
ception, however, will be documented in another paper. 

1 1 rely on Miss Cochran's description of A. darlingtoni = etheridgei for evi- 
dence of transverse, dorsal, saddle-like markings in this form. At the time of 
writing twenty years later they are not at all evident on the specimens. 



BREVIORA 



No. 164 








■t.3 




s 




o 


Ci 


X 


t 


•e* 


m 


5^ 


M 


<i) 


p 










^-i 


o 


5^ 


« 




in 




^ 




l-H 




cS 




« 



■* 





O 








-M 








■«^ 








ce 








g 








c^ 




1^ 




J3 


••?^ 


w^ 


1— 1 




Si. 


o 

33 


J 


CM 


o 


33 


n 

< 


O 
1^ 




05 

o 


M 


s 


'1 


C5 




u 




m 




eS 




OJ 




P< 




Si 




o 




K> 




O 




(M 



e 


o 






1 


« 


«»• 


tc 


s 


o 


Q 


h 


S 






w 




o 




-a 




« 




SQ 



o 






rr* 


'« 


1> 


n 




Ci 


'S 


h 


«i 


n 


r« 


QJ 




'3 


•\ 




"3 






C3 




ti 


bi 


p 


Q 




^ 


=w 


<u 








f^ 




rS 


• rH 






5i 


OS 


00 


^ 




or. 


xn 
o 



QJ 




£ 


^ 


§ 


O 

o 


ri 


s 


o 


(n 


.(H 






1 


^ 


S 


a 


6 


OJ 


s~ 


■73 


ti 


,_^ 


o 


R 




-M 


■2 


fl 


« 


O 


^ 


u. 




"VI 


<ai 


c; 


§ 


• r-l 


^ 


m 


t;^ 


<» 








m 


o 



0) 



s 

o 
a: 



o 

0; ^ 



CO O 



tS 




<v 




-*^ 




cS 




>-> 




e« 




ft 




o 




00 




05 
-2 


 iH 


'3 


m 


.^ 


1:3 


.a 


.rH 






.-• 




c 


03 


O) 


13 


CO 


a" 




0) 


<~— 




« 


02 


-*-i 


a; 


'rS 


i-H 

CCI 


o 


OS 


03 




^^ 


CO 


ft 




3 


>i 


to 




T3 




<U 




-t^ 




c: 




t-. 




c3 




ft 




QJ 




CO 




oa 




C/ 








"o 




tH 





.1= « 



,0 


XI 





Ol 


03 




^ 


rH 






#-H 








'O 




V 




->J 




ri 




h 


b 


CO 


f 


ft 


^ 


<» 


03 


en 


a 




C 




05 


M 













Zj 


03 


^ 


•rH 


• r- 


'« 





<v 



s s 



-2 i 

'■9 rt 

O M 

t. CO 

ft . 



eS 




S 




0) 




>. 




t— 1 




-!.:> 




ft 




g 




X2 




C« 




•40 









s 


ut 


00 









oj 


;h 






;3 

o3 


ft 


t< 


fi 





C3 


t4 


J= 


w 


+- 


•+^ 


^ 


p 


_o 


03 




^1 




<o 




F— 1 




»-H 




o3 








CO 




>> 








f^ 




2' 












f~ 




-c> 




a 




Oj 








'2 






tH 


-*•- 





G 


•rH 


CO 


Ih 





O) 




■^r 




co 






ft 


b 




aj 


c 


-^ 


CS 


g 


,C 


c« 


-(•^ 



ft 



t; 


O; 


CO 






CO 






ft 


■►^ 


B 


c3 


03 





■^ 





cu 


Lh 


^ 


« 


c3 


+H 

(5 


s 



a 






^ CO 



o 

»H 

o 



o 

cS 

Ih 

ft 



o e! 

ft CO 








cS 




«H 









aj 




N 




• l-< 




05 




;^ 




e 







l-i 


,_H 


r5 


cS 




-♦•^ 


"3 


c 






^ 


03 


<4H 


Vr 




ft 


tH 


3 


_o 


Ol 








.2 


CO 


'{•M 





O) 


ft 


+H 



CO ''I 



1^ 




CO 




« 




N 




• fH 




00 




?f^ 




!;' 




c 






t^ 






03 


03 














c3 




ft 


Ih 


S 





CO 






fH 


tH 








05 


';h 





« 


ft 


+H 



1962 



TWO POORLY KNOWN ANOLES 



9 






be 


3 




VI 






o 


,^ 








ai 


^ 


0) 


§ 


&; 




-*j 


-i-* 


OJ 




o 






U> 




o 


m 


r^ 


o 






> 


CB 


01 


[» 


"« 


CO 


-M 


^_, 




•1 



« 

M 



o 



D 


' — 


3^ 


c 


M 






f^ 






L^ 


3 


O 


[» 


r- 


sT 


a 


p 


3 




o 


o 


^ 


CO 




b 


•* 


V 


^ 


> 


C8 


X 


ri 


3 




'4-' 


_2 




13 


tf 



c; 


;I3 




eS 


>» 

^ 


1 


r^ 




SJ 


tn 


-*- 


<V 




1— « 




t) 


1—5 


oo 









a 3 -3 

^ .rH O 

? ca 3 





3 

o 




r3 




bo 




o 


1 








aj 






f— J 


3 




.s 


3 


X 






OJ 


•r^ 






•4^ 


Tt* 








3 
O 


X 

o 


C 


■? 


<o 


W 




-M 




" 


o 


pfi 






'*-' 






aj 


u 


3 












^^ 


X 




cS 


X 
0) 


>^ 




"x 




S 






^ 














1— 1 


-^ 


cS 


o 


X 


X 

> 




X 


X 


n 
< 


o 


X 


O 




o 

It 


o 


> 




H 


X 


;^ 


^ 


-^- 


X 

o 


§ 


X 


S 






■? 


X 


S 


T3 


15 


•^ 


rj: 




"^ 




<a 


t3 


3 


:■" 


'C 



.2 ^ 

5 § ^ " 



S 2, o 



4^ 


.2 


X 

3 




^ 








z> 




O 




'$■ 




o3 


s 


> 




o 

ft 


1^ 
X 

X 
X 


3 

o3 
& 

X 


-1-5 


a 
£ 

3 


0) 
3 

'fcH 


a7 
> 

X 


+3 
O 

o 

a 




p 
3 


S 


r^ 


X 


X 


)5 




3 


3 


fO 

o 

X 


13 

5 
> 


X 

3 
3 


X 

13 

X 

p 


_7r 


■^ 


s: 


3 




s 


15 




p 


+-• 


1-1 

O 


■^ 


■^- 



-w 


3 




u 


v 




3 


O) 




3 
o 






^ 












■f^ 


>» 




bj3 


l-< 






o 




o 






^ 


+J 




i 


X 

o 






p< 




k> 


X 




4J 


Ol 




TS 


13 




■? 


X 


_x 


^ 




« 




■*! 


'1 


13 




_3 


3 


• r-t 


^ 


a 


fe 





»— 1 

I— < 
X 






c« 

X 

o 



<;] 



a> 




X 




M 




» 




P» 




X 








M 




3 




;h 


aj 


-M 


-^ 




3 


3 


3 


• fh 


O 




t^ 


TJ 


a 


-3 

01 


3 


^ 


aT 




.4^ 


^ 


3 


3 


o 




>-4 


3 


^ 


m 


J2 


X 


e 




■PH 


Is 






X 


3J 


o 



3 5 



t>l 


3 


-2 


X 


^^ 








i 


;2 


3 


'o 


t^ 


«H 



3 


TJ 




'.^ 


3 




X 


be 

3 




aj 




> 


^ 


'P 






a 
M 








»-*i 






3 


-*^ 








13 


nd 


00 


X 



10 BEEVIORA No. 164 

LITERATTJEE CITED 

Cochran, D. M. 

1935. New reptiles and amphibians collected in Haiti by P. J. Darling- 
ton. Proe. Boston Soc. Nat. Hist., 40: 367-376. 

1939. Diagnoses of three new lizards and a frog from the Dominican 

Republic. Proc. New England Zool. Club, 18: 1-3. 
1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus., 177: 
1-398. 
Shreve, B. 

1936. A new Anolis and new Amphibia from Haiti. Proc. New Eng- 
land Zool. Club, 15: 93-99. 

Williams, E. E. 

1960. Notes on Hispaniolan herpetology. 1. Anolis christophei, new 
species, from the Citadel of King Christophe, Haiti. Breviora, 
No. 117: 1-7. 



1962 



TWO POORLY KNOWN ANOLES 



11 




o 
o 

CO 

to 

N 
Q 






Si 

o 









BREVIORA 

MiaseiLiiiri of Comparative Zoology 



Cambridge, Mass. August 22, 1962 Number 165 

AN EXTINCT SOLENODONTID 
INSECTIVORE FROM HISPANIOLA 

By Bryan Patterson 

During the summer of 1958, Drs. Clayton E. Ray and A. 
Stanley Rand carried on field work^ for this museum in Puerto 
Rico and the Dominican Republic. Particular attention was 
paid to cave deposits, and a number of previously unexplored 
caves were examined. The cave containing the material re- 
ported on here is in the Sierra de Neiba near Rancho La 
Guardia in the Province of San Rafael, Dominican Republic; 
it has no local name. Situated in a limestone cliff and accessible 
only by a nearly vertical climb of some 30 feet, the cave con- 
tains a number of chambers connected by narrow^ passages. 

Bones were encountered in the deposits on the floor of the 
antechamber and of the first chamber beyond it. In the ante- 
chamber two layers are present, an upper, dark grey to black 
one some six inches thick, and a lower, reddish-brown one eight 
to ten inches in depth. The upper layer is either lacking or 
negligible in depth in the first chamber, the lower there attaining 
a thickness of six to twelve inches. Scanty remains from the 
upper dark layer include Rattvs, which clearly indicates post- 
Columbian age. No introduced forms were encountered in the 
reddish-brown layer, which contains indigenous rodents and 
the three species of Nesopliontcs- known to have inhabited the 
island, together with scanty remains of bats, birds, lizards, and 
solenodontids. In addition to the bones of the extinct form de- 
scribed below, one specimen of Solenodon paradoxus was ob- 
tained. This is the left horizontal ramus of a young individual 



1 Partly supported by a Sigma Xi-RESA grant-in-aid. 

-N. paramicnis, luipomicrus and zamicrus of Miller (1929). A', '•panimicru.s," 
the largest of the three, agrees in size with the Cuban 2f. micrus, from which 
Miller separated it on molar characters. I have examined several hundred speci- 
mens of the Hisiianiolan form and compared them with Cuban material. The 
supposed differences are not constant and I have so far been unable to find others 
that might validate Miller's species. 



2 BREVIORA No. 165 

with lo in process of eruption and alveoli of the other teeth 
(M.C.Z. no. 7260) ; as far as I am aware this find constitutes the 
first fossil (or subfossil) record of the species. Judging from the 
good preservation and completeness of most of the bones from 
the reddish-broAvn layer, Dr. Ray (MS field notes) is inclined to 
doubt that their occurrence is a result of owl-roost accumulation. 
Most of the material, including all the solenodontid remains, 
was found in the first chamber of the cave. As regards the age 
of the layer, all that can presently be said is that it is almost 
surely pre-Columbian. 

During their stay in the Dominican Republic, Drs. Ray and 
Rand were accompanied by Professor Eugenio de Jesus Marcano 
F., Universidad de Santo Domingo, whose aid was invaluable 
in all phases of the work. It is a pleasure to name the extinct 
solenodontid in his honor. Dr. Ray will discuss the rodents from 
this and other localities in the course of his comprehensive 
studies on the Antillean Rodentia. I am indebted to Miss Linda 
Loring for the cleaning, sorting and cataloguing of the other 
groups represented in the collection. The drawings are by Mrs. 
Dorothy Marsh. T have been fortunate indeed as regards com- 
parative material, thanks chiefly to the excellent series of 
Solenodon in the Mammal Department of the museum. Thirty- 
six mandibles, fifteen humeri and six ulnae of S. paradoxus and 
three mandibles of ^. cuhanus have been available. I have not 
seen limb bones of S. cuhanus, but published figures (e.g. Peters 
1863) indicate that these do not differ appreciably, either in 
structure or in proportions, from those of the larger species. 

INSECTIVOEA 

SOLENODONTIDAE 

Antillogale^ gen. nov. 

Type: — A. marcanoi sp. nov. 

Distribution: — Quarternary, Hispaniola. 

Diagnosis: — Differing from Solenodon as follows: P4 and 
lower M smaller relative to size of jaw, lower M with lingual cleft 
between paraconids and metaconids deeper, paraconid wings 
directed more anteriorly, paraconids consequently farther from 
metaconids, low ridge between bases of paraconids and metaconids 
isolating slight valleys at bases of trigonid basins, anterior 

] The Antilles, plus 70X77 , -weasel. 



1962 EXTINCT SOLENODONTID INSECTIVORE 3 

ciispules below bases of paraconids very weakly developed, heels 
less broadly shelf-like labially. Post-dental portion of ramus 
larger relative to anterior portion. Ilnmeriis and ulna shorter, 
mueh wider relative to length, ulna with pit proximo-medial to 
sigmoid notch. 



^e' 



Antillogale marcanoi sp. nov. 

Type: — M.C.Z. no. 7261, incomplete right ramus of mandible 
with P3-M0 and alveoli of other teeth. 

Hypodigm: — Type and the following specimens: M.C.Z. nos. 

7262, incomplete left ramus with P4-M2 and alveoli of Po, M3 ; 
7266, posterior portion of left ramus with alveoli of Mo, juvenile ; 

7263, right humerus lacking ectepicondyle ; 7264 ; left humerus 
lacking proximal epiphysis; 7265, right ulna. 

Horizon and locality: — Late Pleistocene or Recent; unnamed 
cave 2 kilometers SE of Rancho La Guardia, Municipio de 
Hondo Yalle, Provineia de San Rafael, Republica Dominieana. 

Diagnosis: — As for the genus. 

Description: — Knowledge of the lower incisors, canine and 
anterior premolar derives solely from alveoli. On this evidence, 
the relative sizes of the incisors and the degree of enlargement 
of I2 were essentially as in Solenodon, as was the relative size of 
the single-rooted canine. The anterior premolar would appear 
to have been larger than P^, to about the degree seen in S. para- 
doxus. P3 is small relative to P4, nearly as much so as in ^. 
paradoxus, and much smaller, both actually and relatively, than 
in S. cnhanus. In structure it is very similar to the corresponding 
tooth of S. paradoxus, and rather different from the larger, more 
globular one of S. cuhanus. The paraconid of P4 is a larger cusp 
than in the majority of specimens of S. paradoxus and much 
larger than in *S^. cuhanus. The essential characters of the molars 
have been given in the diagnosis and can be seen in the figures : 
I may add that, as far as these teeth are concerned, Antillogale 
could only with difficulty be distinguished generically from 
Apternodus. 

The horizontal ramus is shorter and more slightly built than 
in Solcnodon and, instead of the two, or even three, mental 
foramina almost invariably present in that form, there is only 
a single large one, situated beneath P3. The postdental portion 
of the mandible is relatively large and robust. "Whereas the 
horizontal ramus is rather shorter than in S. cuhanus the pos- 
terior part is rather larger and longer, being intermediate in 



BREVIORA 



No. 165 



1 



rfl 










a 

Figurfi 1. a) AntiUogale marcanoi, type, M.C.Z. no. 7261, incomplete right 
ramus, dorsal view, b) Solenodon paradoxus, M.CZ. no. 12384, crown view 
of right P3-M2. X 3. 



these I'espects between cuhanus and paradoxus. The coronoid 
procesis, so far as can be judged from the juvenile M.C.Z. 7266 
and the incomplete M.C.Z. no. 7262, appears to have been less 
tapering than in the living species. The base of the anterior 
border of the process is convex, as in S. cuhanus, and not ex- 
cavated as it is in S. paradoxus. The masseteric fossa is small, 



1962 



EXTINCT SOLENODONTID INSECTIVORE 



relatively and actually, oval in outline, and not very sharply 
defined. The marginal process, for the insertion of M. digastricus, 
is comparable to those of the living forms. The angle is not 
complete in any of the specimens but enough is preserved in 
M.C.Z. no. 7262 to demonstrate that, in contrast to S. pa7'adoxiLs 
and in agreement with S. cuhanus, the ventral border is not 




a 




Figure 2. AntillogaJe marcanoi. a) M.C.Z. no. 7261, type, incomplete 
right ramus, lateral view, b) M.C.Z. no 7266, incomplete left ramus, lateral 
view, X 2. 



downcurved and that, consequently, there is no lunate notch 
between angle and marginal process. 

The limb bones differ markedly in proportions from those of 
Solenodon. The humerus of Antillogale may be summed up as 
a bone having the width but not the length of that of S. para- 
doxus, and hence massive. The relative lengths of humerus and 
ulna appear to be essentially the same in both. The ulna, like 



6 



BREVIORA 



No. 165 



the humerus, is shorter than but equally as wide as that of the 
living species; other differences of note between the two are 
the presence in the fossil of a very distinct pit proximo-medial 
to the sigmoid notch and of a rugose interosseous border. 





Figure 3. Lateral views of left rami of a) Solenodon panidoxus, M.C.Z. 
no. 34828, L) Antillogale marcanoi, M.C.Z. no. 7262, and e) Solcnodon 
cubanus, Y.P.M. no. 1203. In b) areas in solid outline are restored from 
M.C.Z. nos. 7261 and 7266. X 3/2. 



1962 



EXTINCT SOLENODONTID INSECTIVORE 






^•^f% 



Figure -4. Anterior vie^vs of humeri (a-e) and ulnae (d, e). Antillogale 
marcanoi, a) M.C.Z. no. 7264, b) M.C.Z. no. 7263, (!) ]\[.C.Z. no. 7265. 
Sohnodon pdmdoxus, c) and e) ^M.C.Z. no. 12416. X 4/3. 



8 BREVIORA No. 165 

Measurements in mm. 

M.C.Z. nos. 

7261 7262 

Ps, length 2.7 — 

width 2.0 — 

P4, length 3.1 3.2 

width 2.3 2.5 

Ml, length 3.0 3.4 

width 2.9 — 

M2, length 3.3 3.4 

width 3.0 3.3 

Depth of ramus beneath Mi 6.5 — 

7263 7264 

Humerus, total length 40.0 — 

a.-p. diameter of proximal end 9.9 — 

tr. diameter of proximal end 10.9 — 

a.-p. diameter at center of shaft 7.1 6.6 

tr. diameter at center of shaft 7.3 6.5 

a.-p. diameter of distal end 5.1 4.9 

tr. diameter of distal end — 17.7 

tr. diameter of trochlea — 9.2 

7265 

Ulna, total length 45.6 

a.-p. diameter of olecranon 4.8 

tr. diameter of olecranon 7.4 

tr. diameter of proximal articular surface 6.4 

a. p. diameter at center of shaft 4.3 

tr. diameter at center of shaft 2.3 

a.-p. diameter at distal end 3.9 

tr. diameter at distal end 2.8 

Discussion: — Tlie Solenodontidae have thus far been known 
from one genus with two species. Splitting of these taxa has of 
course been attempted. Soletwdon cubamis has been made the 
type of Aiopogale by Cabrera (1925), and Barbour (1944) has 
described a second Cuban species, S. poeyanus, based on pelage 
characters. Barbour explicitly stated that he could detect no 
cranial or dental distinctions. Aguayo (1950) and Koopman and 
Ruibal (1955) have regarded this form as a subspecies of 
cuhanns. The latter authors noted that some fragmentary remains 
(M.C.Z. no. 7054) from a cave in Camaguey, which they referred 
to the living species, were somewhat larger than Recent specimens 
of cubanus examined by them. In recent years, members of the 
Sociedad Espeleologica de Cuba have obtained further material, 
including some skulls, from the Province of Havana. Thanks 



1962 EXTINCT SOLENODONTID INSECTIVORE 9 

to Sr. Oscar Arredondo, I have seen most of these specimens ; they 
also represent individuals rather larger than, although not other- 
wise different from, any in the small sample (3) of cuhanus 
available to me. It is of some interest that the type of poeyamcs 
(M.C.Z. no. 6957) is also rather larger than this sample, a point 
not mentioned by Barbour; in fact there is agreement in this 
respect between the type and these fossils (or subfossils). To 
conclude from this that poeyanus is distinguishable from cuhanus 
on the basis of size and that the fossils so far found are referable 
to the former would, I think, be premature. The available series 
are too small to rule out accidents of sampling. The size differ- 
ence is slight and if the size range of cul)mius is comparable 
to that shown for paradoxus by the adequate series at hand we 
could be dealing simply with segments of a normal distribution. 
Reinforcing caution is the fact that one of the two mandibles 
from Maisi, Oriente (M.C.Z. no. 10065), mentioned by Allen 
(1918) is smaller than the other fossils. Whatever the solution 
of this minor problem may prove to be, I agree entirely with 
Aguayo and with Koopman and Ruibal that subspecies, at most, 
are involved. There is at present no good evidence of more than 
one species of Solenodon in Cuba. Atopogale is recognized, either 
as a genus or as a subgenus, by some authors. There can be no 
question that paradoxus and cuhanus are clear-cut taxa but they 
seem to me, as to others, to merit no more than specific rank. 
AntiUogale marcanoi helps to clarify matters here ; it is sharply 
distinct from either of the living species and the latter share 
most of the characters that differentiate them from it. 

AntiUogale is quite evidently a member of the Solenodontidae. 
The structure, although not the proportions, of the known limb 
bones ; the general structure of the mandible, and especially the 
presence of a marginal process; the small Ij and the degree of 
enlargement and mode of implantation of I2 (as revealed by 
the alveoli) — all these characters combine to place this con- 
elusion beyond reasonable doubt. Only in lower molar structure 
is there a closer resemblance to members of another family, the 
Apternodontidae. The difference here I believe to be the result 
of specialization in the Solenodon phylum. The molars of the 
two living species give the impression of having undergone 
anteroposterior compression of the trigonids. A majority of the 
characters in which they differ structurally from those of 
Aniillogale could be directly correlated with such a change. 
Aside from molar structure, the differences between the extinct 
and the living species are chiefly of a proportional nature, as 



10 BREVIORA No. 165 

pointed out above. AntillogaJe evidently had a somewhat shorter 
facial region and much shorter, more heavily built fore limbs 
than Solcnodon. It may well have been more fossorial in habits. 
Solowdon with its tAvo species, one on each of the larger West 
Indian islands, has always been one of the most isolated of mam- 
mals, zoogeographically — and even taxonomically — speaking. 
As knowledge of the past history of mammals has improved, this 
isolation has become, if anything, even more apparent. In earlier 
days it was at least possible to assume a common ancestry with 
if not membership in the Tenrecidae. There is now no real 
evidence for such an assumption — tenrecids, as far as the record 
goes, appear always to have been African and Madagascan in 
distribution —  and we must look elsewhere for close relatives and 
possible ancestors. The only group that seems to me to come close 
to fulfilling the requirements is the Apternodontidae, a family 
which at least had the merit of inhabiting North America, the 
only probable source area.^ The described forms, the Oligocene 
Aptcrnodus and Oligorycfes, were certainly not ancestral to the 
solenodontids. but apternodontids are now knoAvn in North Amer- 
ica at least as far back as the Bridgerian Eocene." The possibility, 
I would go so far as to say probability, exists that solenodontids 
were derived from relatively unspecialized apternodontids that 
inhabited the Central American peninsula during the earlier 
Tertiary. Rafting of the ancestral stock to the Antilles, for 
rafting was certainly involved, may have taken place in the later 
part of the Eocene, at roughly the same time as the rafting of 
the ancestors of caviomorph rodents and platyrrhine primates to 
South America. This event, if it occurred at the time suggested, 
would have insured for the solenodontids a very long residence 
in the West Indies. There has, I feel, been some reluctance to 
accept such a possibility, a reluctance based, consciously or un- 
consciously, on the fact that Solenodon alone has hitherto repre- 
sented the family and on the assumption that we now have an 
adequate idea of the Pleistocene, or at least pre-human, faunas 
of the archipelago. I strongly doubt if we do have a good 



1 Allen's belief (1918) that solenodontids were derived from nesophontids. a 
vifw recently supported liy McDowell (]ft.")S). is not at all convincing'. McDowell's 
attemnt to ri^id Aiitenifiiiiiit out fit the Insectivorji is ntti'rlv unrealistic, being 
in large part based on misinterpretation of the fossils he examined. 

2 I am indel)ted to Dr. Craig' C. lUaclv for the opportunit.v of examining a speci- 
men from Tabernacle Bntte. Wyoming. This is tlic earliest kr.own entberiaii 
with zalambdodont molars. Contrary to earlier statements (including one of 
mine made on the b;isis of the literature), the molars of the I'aleocene I'dliirn' 
riirtvft are not zalandidodont in structure, nor are they even pre-zalaiubdo<lont 
in the way those of Potamoyalc are. 



1962 EXTINCT SOLENODONTID INSECTIVORE 11 

knowledge of the faunas (cf. Koopman and AVilliams 1951) ; as 
regards birds at any rate Sr. Arredondo and I will demonstrate 
the contrary in a forthcoming paper. And now, in Antillogale, 
we have a hint that there may have been a radiation of solenodon- 
tids on the Antilles in some degree comparable to that of the 
tenrecids on Madagascar. 

REFEKENCES 

Aguayo, C. G. 

1950. Observaeiones sobre algunos niamiferos cubanos extinguidos. 
Bol. Hist. Nat. Soc. Felipe Poey, 1 : 121-134. 

Allen, G. M. 

1918. Fossil mammals from Cuba. Bull. Mus. Comp. Zool., 62: 131- 
148. 
Barbour, T. 

1944. The solenodons of Cuba. Proe. New England Zool. Club, 23: 1-8. 
Cabrera, A. 

1925. Genera Mammalium. Inseetivora, Galeopithecia. Madrid: 1-232. 
Koopman, K. F. and R. Ruibal 

1955. Cave-fossil vertebrates from Camaguey, Cuba. Breviora, Mus. 
Comp. Zool., no. 46: 1-8. 
Koopman, K. F. and E. E. Williams 

1951. Fossil Chiroptera collected by H. E. Anthony in .Jamaica, 1919- 
1920. Amer. Mus. Novitates. no. 1519: 1-29. 

McDOWEHLiL, S. B. 

1958. The greater Antillean insectivores. Bull. Amer. Mus. Nat. Hist., 
115: 113-214. 
Miller, G. S. 

1929. A second collection of mammals from caves near St. Michel, 
Haiti. Smithson. Misc. Coll., 81, no. 9 : 1-30. 
Peters, W. 

1863. Uber die Saugethier-Gattung Solenodon. Abhl. K. Akad. Wiss. 
Berlin, 1863: 1-22. 



BREVIORA 

MiuiseimTi of Comparative Zoology 



Cambridge, Mass. September 5, 1962 Number 166 



LECTOTYPES OF SPECIES OF OGCOCEPHALIDAE 

SELECTED FROM SYNTYPES IN THE 

MUSEUM OF COMPARATIVE ZOOLOGY 

By Margaret G. Bradbury^ 

Hopkins Marine Station 
Pacific Grove, Calif. 

During the 1891 expedition of the U. S. steamer "Albatross" 
to the eastern tropical Pacific, about 80 specimens of fishes of 
the family Ogcocephalidae were taken. Garman, who studied 
the collection, concluded that it represented twelve species, ten 
of which were described by him as new (Garman, 1899). The 
remarkable thing- is that subsequent collecting in the same area 
has produced additional examples of only about half of these 
species; the rest remain known only from specimens taken on 
the original expedition. 

It became necessary for me to examine this valuable "Albatross" 
collection in the course of work on the family Ogcocephalidae. 
Then it was discovered that most of the specimens upon which 
Garman based his original descriptions had been designated 
as syntypes, a common practice at the time. In the interest of 
stabilizing the nomenclature of this group, I have selected 
lectotypes for the species heretofore represented by syntypes. 
The list of species below includes only those for which Garman 
wrote original descriptions. They are arranged in the order 
in which they appear in his paper. 

Ogcocephalus porrectus 

The original lot contained four specimens from "Albatross" 
Sta. 3368A, and of these Hubbs (1958) selected the largest speci- 
men as lectotype; it bears the number MCZ 28733. The other 
three specimens are now removed to MCZ 41594. 

^ This investigation Avas supported by Public Health Service Fellowship 
No. 8543 from the Division of General Medical Sciences, Public Health 
Service. 



2 BREVIORA No. 166 

Halieutopsis tumifrons 

The only two specimens known of this species are those 
originally taken by the "Albatross." It was not possible for 
me to determine which of the two had been figured by Garman, 
so the choice of a lectotype was arbitrarily made, the two ex- 
amples being closely similar. The lectotype, MCZ 28729, is from 
"Albatross" Sta. 3400A. The second specimen, from "Albatross" 
Sta. 3413A, bears the number MCZ 28730. 

DiBRANCHUS HYSTRIX 

Garman assigned three examples to this species, and from 
these I have chosen the specimen that appears to be the one 
illustrated in the original description ; therefore the lectotype 
is MCZ 28726, "Albatross" Sta. 3375A. The other two specimens 
are: MCZ 28727, "Albatross" Sta. 3392A, and MCZ 28728, 
"Albatross" Sta. 3362A. 

DiBRANCHUS SCABER 

Two specimens represented this species originally, the larger 
of which is apparently that figured in the original description. 
It has the number MCZ 28724, "Albatross" Sta. 3431 A, and is 
hereby designated the lectotype. The second specimen, MCZ 
28725, "Albatross" Sta. 3364A, is a specimen of Dihranchus 
hystrix, not Dihranchus scaher. The lectotype of Dihranchus 
scaher is therefore the only known specimen of the species. 

DiBRANCHUS ASPER 

This species was described from a single specimen, MCZ 
28723, "Albatross" Sta. 3418A ; accordingly this specimen is 
the holotype. My own studies show Dihranchus asper to be the 
same as Dihranchus hystrix; of the two nominal species, I hereby 
place Dihranchus asper in the synonymy of Dihranchus hystrix 
for the reason that no figure accompanies the description of 
Dihranchus asper. 

DiBRANCHICHTHYS NUDIVOMER 

Altogether, two lots totalling seven specimens represent this 
species. A specimen in relatively good condition from MCZ 
28719, "Albatross" Sta. 3353 A, was selected as the lectotype; 
there was one other specimen in the lot with the lectotype, and 



1962 LECTOTYPES OF OGCOCEPHALIDAE 3 

it is now removed to MCZ 41595. The second lot of five speci- 
mens bears the number MCZ 28718, "Albatross" Sta. 3395A. 
These, alonis: with an eighth example from "Albatross" Sta. 
3395A which had been sent to the U. S. National Museum 
(USKM 57867), are the only known specimens of this species. 

Malthopsis sparsa 

Garman originally assigned three lots totalling 19 specimens 
to this species. Of these it is impossible to tell exactly which 
example was used for the illustration in the original descrip- 
tion, and the lectotype was chosen more or less arbitrarily from 
a series considered to be in good condition and which most 
resembled the original illustration. The lectotype, MCZ 28717, 
bears a tag indicating that it came from "Albatross" Sta. 3386A, 
but the situation is not so clear with respect to most of the 
specimens that were originally in the same lot with the lectotype. 
Of these there are eight specimens as follows : two specimens 
with the same station number as the lectotype, i.e., "Albatross" 
Sta. 3386 A; one specimen tagged "Albatross" Sta. 3354A ; five 
specimens with no tags and no indication in the original de- 
scription as to whether they came from station 3386A or 3354A. 
These untagged specimens together with those from station 
3386A are now removed to MCZ 41596. The specimen from 
"Albatross" Sta. 3354A is certainly an example of MaltJwp- 
sis spinosa, not Malthopsis sparsa, and is now removed to MCZ 
41597. 

The remaining two lots of Malthopsis sparsa stand as fol- 
lows: MCZ 28715, "Albatross" Sta. 3396A, six specimens; MCZ 
28716, "Albatross" Sta. 3385A, four specimens. 

Malthopsis erinacea 

From the original series of eight, the one specimen that ap- 
pears to have been used for the illustration accompanying the 
original description is hereby designated the lectotype ; it bears 
the number MCZ 28712, "Albatross" Sta. 3402A. A second 
specimen originally in the same lot with the lectotype is now 
removed to MCZ 41598. The other six specimens bear museum 
numbers as follows: MCZ 28711, "Albatross" Sta. 3358A, one 
specimen; MCZ 28713, "Albatross" Sta. 3425A, one specimen 
(this is probably not a specimen of Malthopsis erinacea, but it 



4 BREVIORA No. 166 

is small and difficult to place at this time without more com- 
parative material) ; MCZ 28714, ''Albatross" Sta. 3418 A, four 
specimens (these four are actually examples of Malthopsis 
spinosa, not Malthopsis erinacea). 

Malthopsis spinosa 

Two rather similar specimens were assigned to this species by 
Garman, but I am unable to tell which was used for the illus- 
tration accompanying the original description. I have selected 
MCZ 28710, "Albatross" Sta. 3392A, as the lectotype. The re- 
maining specimen from "Albatross" Sta. 3393 A, originally with 
the same museum number as the lectotype, is now removed to 
MCZ 41599. 

Malthopsis spinulosa 

From a series of 21 specimens from "Albatross" Sta. 3394A, 
I have selected a specimen on the basis of its similarity to the 
illustration used with the original description and the fact of 
its relatively good condition. This specimen bears the number 
MCZ 28709, and the remaining 20 specimens of the lot are 
removed to MCZ 41600. Prom my own examination of this 
material in conjunction with newer collections, I must conclude 
that Malthopsis spinulosa is the same as Malthopsis spinosa. 
Inasmuch as the original descriptions of both nominal species 
are accompanied by illustrations of equal quality, I give prefer- 
ence to Malthopsis spinosa as the name to represent this taxon 
by virtue of its page priority. Thus, Malthopsis spinulosa falls 
into the synonymy of Malthopsis spinosa. 

I am extremely grateful to Mrs. Myvanwy Dick and to Dr. 
Giles AV. Mead, who both took great trouble to assist me in the 
loan of material. 

LITEEATUEE CITED 

Garman, S. 

1899. Eeports on an exploration off the west coasts of Mexico, Central 
and South America, and off the Galapagos Islands, in charge of 
Alexander Agassiz, by the U.S. Fish Commission Steamer 
"All>atross" during 1891, Lieut. -Commander Z. L. Tanner, 
U.S.N., commanding. XXVI. The Fishes. Mem. Mus. Comp. 
Zool. Harvard Coll., 24: 1-431, pis. 1-88, A-N. 
HuBBs, Carl L. 

1958. Ogcocephalus darwini, a new batfish endemic at the Galapagos 
Islands. Copeia, 1945 (3) : 161-170, pis. 1-5. 



BREVIORA 



Mnasemm of Cooipsirative Zoology 



Cambridge, Mass. September 5, 19B2 Number 167 

Bathyclupea schrocderi, a New Bathyclupeid Fish 
from the Weslerii Tropical Atlantic 

By 
Myvanwy M. Dick 

Museum of Comparative Zoology, Harvard University 

The species described below is based on specimens collected by 
the Atlantis during the Harvard-Havana Expedition to Cuba, 
and by the Oregon (exploratory vessel of the U.S. Bureau of Com- 
mercial Fisheries) during her work in the Gulf of Mexico. To 
Mr. Harvey R. Bullis, Jr., director of the Oregon's program, go 
my thanks for representatives of both western Atlantic species 
of Bathyclupea. I am also indebted to Mr. Loren P. AVoods of 
the Chicago Natural History Museum, and to Dr. Leonard P. 
Schultz, U.S. National Museum, for the loan of comparative 
specimens of related species and for additional representation 
of the new form. 

Bathyclupea schroederi, new species . . 

Holotype: A specimen 140 mm. in standard length, taken at 
Atlantis Station 2987, 23°23'N., 79°39'W., 280 fathoms. MCZ 
41498. 

Paratypes: MCZ 39416, 4 specimens, Atlantis Station 2987, 
23°23'N., 79°39'W., 280 fms. ; MCZ 39380, 1 specimen, Atlantis 
St. 2987a, 23°22'N. ; 79°39'W., 260 fms. ; MCZ 40600, 1 specimen, 
Oregon St. 2635, 17°37'N., 63°28'W., 220-235 fms.; CNHM 
65145, 1 specimen, Oregon St. 1872, 16°41'N., 82°20'W., 300 
fms.; CNHM 65147, 1 specimen, Oregon St. 1888, 16°41'N., 
81°02'W., 250 fms.; CNHM 65146, 1 specimen, Oregon St. 1886, 
16°55'N., 81°12'W., 275 fms. 

Diagnosis: The following species of Bathyclupea have been 
described : Bathyclupea hoskynii Alcock, 1891, from the Andaman 
Sea (the type species) ; Bathyclupea argentea Goode and Bean, 



2 BREVIORA No. 167 

1895, from Nevis, West Indies; Bathychipra ynnylayana Weber, 
1934, from the Flores Sea, Indonesia ; Bathyclupea megaceps 
Fowler, 1937, from off Mindanao, Philippines ; and Bathyclupea 
gracilis Fowler, 1937, from off Makyan Island, Moluccas. 

Bathyclupea schroederi differs from all of these by havino- a 
spine and 37 to 39 rays in the anal fin (cf. 33 or fewer). It can 
also be distinguished from B. hoskynii by its less deep body (3.9 
to 4.4 in standard length; cf. 3.3), and from B. argent ea and B. 
maylayana by the placement of the anal fin, which originates in 
the anterior half rather than in the posterior half. B. schroederi 
can be distinguished from B. megaceps by the relative length of 
the base of its anal fin, which is longer than the head in the new 
form but shorter in B. megaceps. The length of the head in B. 
schroederi, 3.3 to 3.4 in standard length, is shorter than that 
of B. gracilis (3.0 in length). 

Description: The description which follows is based on the 
holotype and paratypes, 120 to 144 mm. in standard length, 
listed above. 

Counts and proportional dimensions are provided in Table 1. 

Body laterally compressed, its greatest depth at origin of anal 
fin, 3.9 to 4.4 in standard length. The dorsal profile is nearly 
straight. There is a well defined, rather straight lateral line bear- 
ing 38 pored scales which are somewhat more adherent than the 
scales elsewhere on the body. Scales cycloid, deciduous, thin and 
relatively large. Head naked with large mucous cavities. Verte- 
brae 10-20-1, total 31. 

Head 3.3 to 3.4 in standard length, interorbital flat, 16.3 to 
17.1 in standard length. Eye mainly in the posterior half of the 
head. Nostrils small, contiguous, almost superior. The mouth is 
nearly vertical. Minute conical teeth on the lower jaw, a single 
row of minute teeth on the vomer and premaxillary. The anterior 
end of the vomer protrudes into the mouth, frequently extend- 
ing forward to beneath the tip of its snout. 

There are four gill arches. Moderately long gill rakers, with 
.slight protuberences, on the first and second arches ; rakers much 
reduced and stublike on the third and fourth arches. Gill rakers 
on the first arch 3 + 16. The gill opening is wide, the branchi- 
ostegal membranes free from one another and from the isthmus. 
Pseudobranchiae present. 

The pectoral fin is large, pointed, the upper rays the longest, 
about 3.5 in standard length. The ventral fins are anterior to 
the pectorals, short, very slightly separated. The rays are rather 



1962 



NEW BATHYCLUPEID FISH 



like the ribs of a fan, fitting into a shallow groove when con- 
tracted. The dorsal fin originates in the postmedian half of the 
body, the anal fin in the anterior half. 

Color in alcohol: Opercle and abdominal wall dusky. In life 
the general color was silvery. 

It is a pleasure to name this species for William C. Sehroeder 
of the Department of Fishes, Museum of Comparative Zoology, 
who collected the holotype and several of the paratypes. His 
many publications, the collecting he has done, and his work in 
the department have greatly enriched the field of ichthyology. 



TABLE 1 



Counts, and proportional dimensions expressed in percent of standard length, 
of the fiolotype and paratypes of Batfiyclupea schroederi. 





M 5 
1 




rvj-o 

CO 




^ CT- 
en 




1^ 
S in 




§2 


Standard lengtfi (mm.) 


144 


144 


140 


139 


137 


137 


130 


120 


116 


Greatest deptfi of body (percent) 


27.8 


26.4 


25.0 


26.4 


25.5 


27.0 


24.6 


25.0 


23.3 


Greatest depth of body at midpoint 


26.4 


23.6 


25.0 


25.0 


24.1 


24.1 


23.1 


25.0 


23.3 


Least depth of caudal peduncle 


10.4 


9.7 


9.3 


9.5 


9.5 


8.7 


9.2 


9.2 


8.6 


Greatest width of body 


10.4 


9.7 


7.9 


9.5 


9.5 


8.7 


7.7 


9.2 


7.8 


Greatest width of head 


12.5 


11.8 


12.2 


12.2 


12.4 


11.7 


11.5 


12.5 


11.2 


Snout to origin of dorsal fin 


54.2 


55.6 


55.0 


53.6 


54.7 


53.2 


54.6 


50.8 


51.7 


Snout to origin of anal fin 


43.8 


43.1 


42.8 


42.8 


45.2 


43.8 


43.1 


47.5 


44.8 


Snout to insertion of ventral fin 


27.8 


27.8 


28.6 


28.6 


31.4 




26.9 


29.2 


29.3 


Snout to insertion of pectoral fin, 
ventral angle 


28.5 


27.8 


27.8 


29.3 


29.9 


29.9 


26.9 


28.0 


29.3 


Length of base of dorsal fin 


10.4 


10.4 


10.7 


10.6 


10.9 


10.9 


10.8 


10.0 


10.3 


Length of base of anal fin 


51.4 


49.4 


47.1 


49.3 


48.2 


51.1 


48.4 


48.3 


50.9 


Distance between insertion of ventral 
fin and origin of anal fin 


20.7 


20.2 


17.8 


17.8 


20.4 




17.7 


16.7 


17.2 


Length of head 


28.5 


28.5 


30.0 


29.3 


29.2 


30.6 


27.7 


30.0 


32.0 


Length of snout 


8.3 


8.3 


8.6 


8.6 


8.7 


8.7 


8.5 


9.2 


7.8 


Greatest diameter of eye 


11.8 


11.1 


11.4 


11.4 


11.7 


11.7 


10.8 


11.7 


11.2 


Width of interorbitai space 


5.6 


5.6 


5.0 


5.0 


5.1 


5.1 


5.4 


5.8 


6.4 


Length of upper jaw 


13.2 


13.2 


12.2 


12.2 


12.4 


13.1 


12.3 


12.5 


12.9 


Counts: 




















Dorsal fin 


1-9 


1-9 


1-8 


1-9 


1-8 


1-9 


1-9 


1-8 


1-9 


Anal fin 


1-37 


1-38 


1-39 


1-37 


1-38 


1-39 


1-38 


1-39 


1-37 


Pectoral fin 


1-28 


1-28 


1-29 


1-28 


1-28 


1-28 


1-29 


1-28 


1-29 



BREVIORA 



No. 167 




X:: 



o 

M 
en 

00 

Ci 

T-l 

N 



O 



> 
o 






Hi 



K5 



D 
O 



BREVIORA 

Museium of Coimpsirative Zoology 

Cambridge, Mass. Sp:i'temher 7, 19()2 Number KiS 



TWO NEW SPECIES OF FOSSIL TALPID 
IXSECTIVORES 

By Katherine M. Reed 



During work on the talpid siilifamily Proscalopiiiae, two new 
sealopine moles came to my attention and are described in this 
note. 

I am grateful to ^Ir. Richard Tedford, University of Cali- 
fornia Museum of Paleontology, and Dr. C. W. Hibliard, Uni- 
versity of Michigan Museum of Paleontology, for the loan of the 
material. The work was carried out at the ]\Iuseum of Compara- 
tive Zoology. I am obliged to Professor Bryan Patterson for 
critical comment and to ^liss Barbara Lawrence for access to 
the collections of Recent inseetivores. I also appreciate the com- 
ments and assistance of Dr. J. R. Macdonald and Dr. Mary 
Dawson. The illustrations were made by Mr. Richard Stafford. 

The following abl)reviations are used : 

UCMP — University of California ^luseum of Paleontology 
UMMP — University of IMichigan ^Museum of Paleontology 
1., w., trig., tal. — length, width, trigonid, talonid 

TALPIDAE 

Subfamily Scalopinae 
DOMXIXOIDES Green 1956 

DOMNINOIDES VALENTINENSIS n. Sp. 

Type: UCMP 33152, right ramus with Pj, P;.-4. Mo-. 

Hrjpodigm: Type and UCMP nos. 36150-36157, including iso- 
lated teeth, partial rami and some liml) bones. UCMP 29215 and 
29215-A refer to limb material marked "float." 

Horizon and locality: Late ^Miocene, Valentine formation, from 
the quarries near the middle of the exposed Valentine formation 



2 BREVIORA No. 168 

at the Port Niol)rara locality on the quarter section line between 
the NW and SW (|iiarters of Sec. 24, T 34 N, R 26 W, Cherry 
County, Nebraska, UCMP locality Y3218 (See IMacdonald 1947). 
The specimens marked "float" are, according to 'Sir. Tedford, 
"nndonl^tedly Barstovian, but come from an undetermined 
horizon within the Valentine." 

Diagnosis: No diastema between P^ and P4 ; slightly larger 
and more robust tlian DonniiuoirJes ripareiisis; metastylid defi- 
nite on Mo.' 

Description : The only uj)per tootli in tlie material, and tlie first 
known to belong to this genus, is a broken, isolated P^. This 
tooth has a blade-shaped paracone. A very narrow anterior 
cingulum widens to a shelf-like lingual cusp and extends up the 
jiosterior side of the tooth to join the paracone crest posteriorly. 
The lingual cusp is Avidest opposite the stoutest jiortion of the 
paracone. The tooth has at least two, jiossibly three roots, two 
labial, one lingual. 

In the lower dentition, P^ is small, conical and single rooted. 
P2 is not represented in the material but is double rooted. P.^ is 
consideraldy larger than P^, double rooted as in D. rijxirnisis, 
and with a slight heel. It is situated very close to P.,, with no 
diastema between these teeth. P4 is double rooted with a very 
small anterior cuspule. It has a larger heel than Po., the heel 
sloping downwards labially to a small cingulum which connects 
the heel to the anterior cuspule. The main cusp is conical. 

Ml is not represented in the material studied; thus no com- 
jiarison with D. riparcnsis is ])ossible in this resj^ect. Relative 
root sizes are in accord with this species. In ^2 the paraconid 
is smaller than the metaconid, which is the highest lingual cusp. 
The entoconid is the stoutest lingual cusp. A crest, the crista 
obliqua. runs from the hypoconid to the metastylid, as in D. 
ripa)'e)isis. The protoconid is higher than the hypoconid and is 
slightly labial to the metaconid. There is a large anterior cin- 
gulum, wide labially and with an irregular border, that extends 
around the paraconid to the opening of the trigonid valley. The 
posterior cingulum is small and does not reach to the lingual' 
face of the tooth. The talonid has a narrow opening. In M.-., tbe 
size relations of the cusps are as in M^. All exami)les of this tooth 



iDr. J. R. Macdoiiald has kiiullj- sent iiip a eopv of a forthcoming nianuscript 
in which lie clpseril)ps a new species of D':i)uiinoi(l' x. D. rnlriitiiirtisis is ilistiii- 
guished from this species by lacking any trace of cingnla on the lingiinl face of the 
molars and l>y the crista obli(|iia nnining to tlie metastylid rather than to the 
metaconid. 



10()2 NEW FOSSILS TALPIDS 3 

are worn, hut there are stroiifr supgestioiis of a metastylid and 
of a relatively wide anterior ein<»:uhnn. Xo posterior einfrulmn is 
present. The talonid is relatively longer than in ^Iv. 

There are usually two mental foramina which vary in posi- 
tion. In UCMP 36151, they are below the anterior roots of P4 
and between P4 and Mi ; in Ur^IP 33152, between P2-.S and the 
roots of P4. In IK'^IP 36152. there is only one foramen, l)elow 
the anterior root of P4. In D. riparensis, the foramina ai-e below 
P;{ and between the roots of P4. The ramus is mueh like that of 
D. riparc7isis and is in general similar to that of Macdonald's 
species, although slightly stouter than in either. 

Some comparison of the limb material of the fossil has been 
made with modern species. It must be noted that the majority 
of tile limb material is "float" and association with the teeth 
could ])erhaps be doul)ted: the extremely talpid-like nature of 
both does, however, strongly suggest association. The humeri 
of the fossil are very similar to but slightly smaller than those 
of Talpa europaea. They are larger than in Parascalops hreweri 
and wider than in Condylura. 

Discussion: Macdonald has suggested (j^ers. comm.) that the 
jaws of his new species may represent the lower dentition of 
Proscalops sccundiis. This now seems extremely unlikely, first 
because of the discovery of P^ of Domninoides valentinensis. 
which is very unlike that of the Proscalopinae, and second, the 
greater development of the anterior cingula on lower molars 
than I would expect on the basis of the known trends in the 
Proscalopinae (See Reed 1961). Wilson (1960) has recently de- 
scribed lower molars which he assigns to Proscalops sp. cf. P. 
secundus; these are much nearer to the proscalopine type of 
molar than to that of Domninoides. 

It is evident that Doiiini)ioides is a talpid, not a soricid as 
Green (1956) originally described it. 

]\Ieasurements 

[^CMP 33152 M. UCMP 36152 .AIo UCMP 33152 M3 

1. 3.1mm 1. 3.2 mm 1. (approx.) 2.4 mm 

w. trig. 2.5 w. trig. 2.2 w. trig. 1.9 

w. tal. 2.7 w. tal. 2.7 w. tal. 1.6 

depth of jaw below :\Ii, UCMP 33152 : 3.7 mm. 



4 BREVIORA No. 168 

HesPEROSCALOPS Hibbard 1941 
Hesperoscalops sewardensis n. sp. 

Hesperoscalops rexroadi Hibbard 1953, p. 23, fig. ID. 

Type: UMMP 27276, partial right ramus with M2-3, fragment 
of a left ramus, anterior part of a left ramus with worn Mj, ulna, 
part of a scapula, and a femur. 

Iliipodigm : Type only. 

Horizon and localitij: Late Pliocene or early Pleistocene, from 
area of Saw Rock local fauna, NE 1/4 Sec. 35, T 34 S, R 31 W, 
Seward County, Kansas. 

Diagnosis: Distinctly larger than H. rcrroadi and with greater 
develo])ment of basal accessory cusi)ules. 

Description and discussion: The difference in size between the 
new species and specimens of H. rexroadi in which the teeth are 
in nearly the same state of wear indicates that these cannot be 
wear differences in the same species and that the two are distinct. 
Although the cusp pattern is very similar, as Hibbard states 
(1953, p. 23), the anterior cingular cuspule on Mo is larger and 
better developed than in H. rexroadi, as is the anterior basal ac- 
cessory cuspule on ^lo. If Hesperoscalops rexroadi is ancestral 
to Scalopus, this new, but closely related species must represent 
an extinct side line, for //. seicardensis is considerably larger 
than Sealojjus aquaticiis and has much better developed basal 
cuspules and cingula. 

^Measurements 

H. sewardensis 

UMMP 27276 M. UMMP 27276 M3 

1. 3.0 mm 

w. trig. 2.5 

w. tal. 2.6 

depth of jaw below Mo on lingual face : 4.5 mm. 
length of Mo -3 : 5.8 mm. 

H. rexroadi 

UMMP 27278 Mo UMMP 27278 M3 

1. 2.3 mm 1. 2.2 mm 

w. trig. 1.9 w. trig. 1.6 

w. tal. 2.1 w. tal. 1.5 

depth of jaw below jMo on lingual face : 3.2 nun. 

length of M2-3 : 5.0 mm. 



1. 




2.8 mm 


w 


trig. 


2.4 


w 


tal. 


1.7 



1962 NEW FOSSILS TALPIDS 5 

REFERENCES 

Green, M. 

1956. The Lower Pliocene Ogallala-Wolf Creek vertebrate fauna, 
South Dakota. Jour. Paleontology, vol. 30, no. 1, pp. 146-169. 
HiBBARD, C. W. 

1953. The insectivores of the Rexroad fauna, Upper Pliocene of Kan- 
sas. Jour. Paleontology, vol. 27, no. 1, pp. 21-32. 
Macdonald, J. E. 

194:7. A new shrew from the Niobrara River, Upper Miocene of Ne- 
braska. Am. Jour. Sci., vol. 245, pp. 123-126. 
1962. The Miocene fauna from the Wounded Knee area of western 
South Dakota. Bull. Am. Mus. Nat. Hist, (in press). 
Reed, K. M. 

1961. The Prosealopinae, new sul)family of talpid insectivores. Bull. 
Mus. Comp. Zool., vol. 125, no. 14, pp. 473-494. 
Wilson, R. W. 

1960. Early Miocene rodents and insectivores from northeastern Colo- 
rado. Vertebrata, Univ. of Kansas Paleontologieal Contribu- 
tions, article 7, pp. 1-92. 



BREVIOBA No. 168 





, 


X 






'A 




, 






CO 


^< 




CO 


. 


lO 


X 




ft 


CI 

co' 


>n 


a 


o 


. 


t>-_ 


o 


• r-i 


CO 


•rH 

a 


o 






o 


o 




& 


03 


o 


• r— 1 


Qj 


o 


cu 


P( 


> 

C3 


9 

05 




o 
o 

o 






ai 




M 


■r. 




<1^ 



k; 


>o 


lO 




l- 




1^ 

Cl 


I-H 


1— 1 


co' 







' « " > « 





^ 


-M 


^ 




be 


rq 


CO 



& 




c^ 




u) 


> 


.s 


« 








;_i 


f—^ 


1— 1 


^ 






-4— 


'^-J 


^4h 


? 


•/: 




■■K 






O 


O 


p 






p 




_bc 


^ 


^ 


o 


c 





s 




rH 


Cj 


o 


.. 


.*" 


r^ 


2 










-t- 






P 




v£ 


K* 


>■ 




-J—' 


•"^ 


■^ 


)-^ 


t~ 








^-A 


-f- 




w 




o 


9 


PLI 




-a 


"3 


< 






to" 




i-H 
CO 


10 


10 

?— 1 


^ 


g 




oi 


tH 


CO 


rH 




t^ 


CO 


Cl 


CO 


Ph 


CO 


TO 
CO 




■2" 


O 


Oh 




o 




Ph 






Hi 






CO 










'^O 


^ 


s 


•S 


■5 




to 








"32 

4j 








to 




o 


^ 


SC 




C) 


'^ 


?^ 




^ 




=0 


•^ 












^ 
■^ 


. 


cj 


ci 




=c 


"TS 


o 


••!* 


10 








la 


•^ 


1 


o 




^ 






h^ 




s 


bil 


^ 


i--' 








p 


s 

o 


s 


bi 


bi 




T— 1 


S 








s 




bi 


P^ 
















. ■■'■' be 



Ph 



BREVIORA 

Miiseiuim of Coeiparsitive Zoology 



Cambridge, Mass. October 15, 1962 Number 169 

NEW RECORDS OF INSHORE FISHES FROM THE 
ATLANTIC COAST OF PANAMA 

By Ira Rubinoff^ and Roberta W. Rubinoff 

During February, March, and April, 1961, a collection of in- 
shore fishes from both coasts of the Isthmus of Panama was made 
in connection with a studj' of the effects of geographical isola- 
tion on fish speciation. This collection consisted of 2095 speci- 
mens of 136 species. A total of 907 specimens of 47 species was 
taken from the Pacific coast and 1188 specimens of 89 species 
were collected on the Atlantic coast. Fourteen species in the 
latter collection Avere found to represent extensions of known 
ranges and some of these are sufficiently abundant to be consid- 
ered resident members of the Panamanian fauna. The purpose 
of this paper is to report the new records from this area. Notes 
are also included on three species represented in previous collec- 
tions from Panama by only one or two specimens each. All the 
specimens discussed herein are in the collections of the Museum 
of Comparative Zoology. 

Rotenone poisoning of coral pools and lagoons was the princi- 
pal method of collecting, although occasionally sandy beaches 
were seined. All of the new records are from coral reef areas, 
particularly from the reefs bordering the Galeta Point Naval 
Station near Coco Solo in the Canal Zone. 

Most of the smaller eels were collected when the incoming tide 
overflowed poisoned coral pools onto the exposed surfaces of the 
reefs and into small crevices and burrows in the coral. When 
the poison reaches these burrows the inhabitants are driven onto 
the flat of the reef and are easily collected. Many young speci- 
mens of Muraenidae as well as most of the specimens of Morin- 
guidae and Echelidae were collected in this manner. 

iThis research was supported in part by a Public Health Service Training 
Grant to the Department of BioloRy at Harvard University. Cambridge, Massa- 
chusetts, and in part by a fellowship from the Woods Hole Oceauographic Insti- 
tution during the summer of 1061. 

Contribution No. 12.'5,'? of the Woods Hole Oceauographic Institution 



2 ]!REVIORA No. 169 

In their classic work on the marine fishes of Panama, Meek 
and Hildebrand (1923, 1925, 1928) collected 23G species from 
the Atlantic coast, of which only five were eels. In other collec- 
tions from this area, the apodes are one of the poorest repre- 
sented groups. Numerous species are known to range along the 
Atlantic coast north and south of Panama but have yet to be 
recorded there. Of the nine species of apodes in our collection 
seven represent new records for this area. 

Specimen lengths are standard lengths to the nearest milli- 
meter in all cases except the eels, for which total lengths are 
given. 

The authors wish to thank the following people for making 
this part of our study possible : Dr. Sydney Galler and Mrs. 
Helen Hayes of the Biology Branch of the Office of Naval Re- 
search, Dr. L. P. Schultz of the United States National Museum, 
and Dr. Martin Moynihan and Mrs. Adela Gomez of the Canal 
Zone Biological Area. 

Dr. James E. Bohlke of the Philadelphia Academy of Sciences 
most kindly identified some of the eels. 

SPECIES NOT PREVIOUSLY RECORDED FROM THE 
ATLANTIC COAST OF PANAMA 

XENOCONGRIDAE 

Kaupichthys atlanticus Bohlke 

MCZ 41455-58 

Five specimens, 50-205 mm. long, were taken from coral reefs 

at Galeta Point. These specimens agree with the description of 

Bohlke (1956). 

Range : Tropical western Atlantic from Bermuda to south of 

Jamaica. 

MORINGUIDAE 

Aphthalmichthys mayeri (Silvester) 
MCZ 41459-61 

One specimen, 275 mm. in length, was taken from the edge of a 
reef southwest of Las Palmas mountain about two -thirds of the 
distance from ]\Iaria Chifpiita to Porto Bello. This fish had a dis- 
tinctly pink head and pink-orange body. This color faded to a 
dull yellow two days after preservation. Three specimens, 150- 
330 mm. long, were taken from the exposed surface of Galeta 
Point Reef. 
Range : Bermuda, Florida, Puerto Rico. 



1962 NEW RECORDS OF PANAMANIAN FISHES 3 

ECHELIDAE 

Myrophis egmontis Jordan 
MCZ 41462-64 

Twelve specimens, 81-231 mm. in leno-th, were obtained from 
Galeta Point. These specimens may be differentiated from Eche- 
lidae previously recorded from Panama by the following char- 
acteristics : origin of dorsal fin behind the vent, and absence of 
teeth on the vomer. These eels were found in situations similar 
to those of Aphthalmichthys mayeri. 
Range: Florida, Bahamas, West Indies. 

MURAENIDAE 

Enchelycore nigricans (Bonnaterre) 

MCZ 41465-67 

Seventeen specimens of this eel. 81-545 mm. in lenp'th, were 

taken from the reefs at Galeta Point. 

The slitlike posterior nostril diagnostic of Enchelycore is an 
unreliable character in separating young specimens from other 
Muraenidae. Separation of the young of E. nigricans from 

Gymnothorax moringa which it closely resembles, was facilitated 
b}' two characters. The anterior nostrils of Enchelycore are 
shorter than those of G. moringa and the upper jaw of Enchely- 
core has a series of 4-6 long canine teeth medial to the outer row 
of teeth on both sides. All Gymnothorax moringa which we ex- 
amined have 1-3 teeth in this series. 
Range : Bermuda, West Indies. 

Gymnothorax viciniis (Castelnan) 

MCZ 41468,69 

Four specimens, 145-330 mm. in length, were taken at the Galeta 

Point reefs. 

Range: Bermuda, West Indies to Brazil, Cape Verde Islands and 

Africa. 

Gymnoth&rax moringa (Cuvier) 
MCZ 41470-73 

The collection contains nine specimens, 64-185 mm. in total 
length, taken from the reefs at Galeta Point. 
Range: Atlantic coast of America from Florida to Brazil, Ber- 
muda, Bahamas, West Indies, St. Helena. 

Uropterygius Bohlke (n. sp.. in manuscript) 
MCZ 41475 



4 BREVIORA No. 169 

One specimen 190 mm. long was collected at Galeta Point. Our 
specimen was identified by Dr. James Bohlke as a species he is 
currently describing and it will be designated as a paratype. 

AULOSTOMIDAE 

Aulostomus maculatiis Valenciennes 

MCZ 41474 

One specimen, 185 mm. long, was taken at Galeta Point. 

Range : Bermuda, Florida, Gulf of Mexico, Central American 

coast of Caribbean, Bahamas, West Indies. 

HOLOCENTRIDAE 

Holocentrus coruscus Poey 
MCZ 41476 

One specimen, 45 mm. long, from Galeta Point, has the folloAv- 
ing meristics: D-XI, 12; A-IV, 8; gill rakers 9. Color of fresh 
specimen : dorsal surface of head red ; body red and white lateral 
stripes ; caudal fin red ; anal and second dorsal dark red at distal 
portions ; interspinous membranes of dorsal peppermint striped, 
black spot distally between first three dorsal spines. With the 
exception of the number of gill rakers this specimen fits the de- 
scription of H. coruscus by AVoods (1955) in his revision of the 
Western Atlantic species of IIoloce7itrus. 
Range : Bermuda, Florida, Bahamas, West Indies. 



r 



fVPOGONIDAE 



Apogonichthys stellatus Cope 
MCZ 41477 

Two specimens, 12 mm. and 29 mm. in length, were taken from 
the edge of a reef southwest of Las Palmas mountain about two- 
thirds of the distance from Maria Chiquita to Porto Bello. 
Range : Bermuda, Florida, Bahamas, West Indies. 

LABRIDAE 

Thalassoma hifasciatum (Bloch) 

MCZ 41478-83 

Fifty-eight specimens 19-83 mm. in length, were taken at Galeta 

Point. Although this species has not been previousl^v recorded 

from Panama it is one of the most abundant representatives of 

the reef pool fauna. Many more examples were seen than were 

collected. 

Range : Bermuda, Florida, Bahamas. West Indies. Honduras. 



1962 NEW RECORDS OF PANAMANIAN FISHES 5 

CANTHIGASTERIDAE 

Canthigaster rostratus (Bloch) 
MCZ 41484, 85 

We collected eleven specimens, 19-42 mm. in length, at Galeta 
Point. Ten of these specimens have the fin formula D-10, A-9 ; 
one specimen has D-9, A-9. These specimens agree with the de- 
scription of Breder (1927). Jordan and Evermann (1898), 
Evermann and Marsh (1900), and Nichols (1930) report speci- 
mens with the fin formula D-6, A-8. For a partial explanation 
of this discrepancy see Breder (1927). 
Range : Bermuda, Florida, West Indies, Venezuela and Madeira. 

CLINIDAE 

Lahrisomus I'alislierae (Jordan) 

MCZ 41486, 87 

Six specimens, 25-68 mm. in length, were found in Galeta Point 

reef pools. One 56 mm. specimen taken in the first week of April 

possessed enlarged ovaries from which ova 0.5 mm. in diameter 

were obtained. A 54 mm. male also taken at this locality had 

enlarged testes. 

Range : Florida to Brazil. 



^tr-*^ 



Ldhrisomus nigricinctns Rivero 
MCZ 41488 

One male specimen 45 mm. long was taken at Galeta Point. It 
was compared with the holotype, MCZ 34150, and with the de- 
scription given by Springer (1958). 

Springer (1959) reports the range extension of L. bucciferus 
and L. guppyi to the Atlantic coast of Panama. L. nuchipinms 
reported by Meek and Hildebrand (1928) and our specimens of 
L. kalisherae and L. nigricinctns bring the total recorded number 
of Atlantic Panamanian Lahrisomus species to five. 

SPECIES RARELY RECORDED FROM THE 
ATLANTIC COAST OF PANAMA 

With the exception of Dinematichthys cayorum these species 
were not collected by Meek and Hildebrand (1923, 1925, 1928). 

BLENXIIDAE 

Rupiscartes atlanticus (Cuvier and Valenciennes) 
MCZ 41489-91 



6 BREVIORA No. 169 

Eight specimens, 39-74 mm. in length, from the coral reefs at 
Galeta Point are in the collection. The previous record of this 
species from Panama was a single specimen taken by Fowler 
(1916) at Toro Point. 

Range : Bermuda, Atlantic and Pacific coasts of tropical Amer- 
ica, West Indies. 

Salarichthys textalis (Quoy and Gaimard) 
MCZ 41492 

Three specimens, 24-42 mm. in length, were taken at Galeta 
Point. One 31 mm. specimen was previously collected at Cale- 
donia Bay, Panama, by Breder (1925). 
Range : Bermuda, Florida, AVest Indies, Brazil. 

BROTULIDAE 

Dinematichthys cay or urn (Evermann and Kendall) 
MCZ 41493-96 

Twelve specimens of this species (Ogilbia cayorum of Meek and 
Hildebrand), 26-51 mm. in length, were collected at Galeta Point. 
On April 6 some adults were found to contain well developed 
embryos which could be seen through the body wall. These were 
extruded in gelatinous strings when a slight pressure was exerted 
on the abdomens of the females. They were about 5-7 mm. in 
length and had small yolk sacs. When placed in a bucket of sea- 
water the embryos were free swimming although apparently pre- 
mature. Fowler (1916) and Meek and Hildebrand (1928) each 
found only one example of this species. 
Range : Bermuda, Florida, Bahamas. 

LITEEATUEE CITED 

BOHLKE, J. E. 

1956. A synopsis of the eels of the family Xenocongridae (including 
the Clilopsidae and Chilorhinidae). Proc. Acad. Nat. Sci. Phila- 
delphia, 108: 61-95. 
Breder, C. M. 

1925. Notes on fishes from three Panama localities: Gatun spillway, 
Eio Tapia and Caledonia Bay. Zoologica, 4 (4) : 137-158. 

1927. Scientific results of the first oceanographic expedition of the 
"Pawnee" 1925, Fishes. Bull. Bingham Ocean. Coll., 1 (1): 
1-90. 
Evermann, B. W., and M. C. Marsh 

1900. The fishes of Porto Eico. U. S. Fish Conim. Bull, for 1900: 
49-350. 



1962 NEW RECORDS OF PANAMANIAN FISHES 7 

Fowler, H. W. 

1916. Cold-blooded vertebrates from Costa Eiea and the Canal Zone. 
Proc. Acad. Nat. Sci. Philadelphia, 68: 389-414. 
JoRDAK, D. S., and B. W. Evermann 

189(5, 1898, 1900. The fishes of North and Middle America. Bull. U. S. 
Natl. Mus., 47 (1-4): 1-3313. 
Meek, S. E., and S. F. Hildebrand 

1923, 1925, 1928. The marine fishes of Panama. Field Mus. Publ. 
Zool. Ser. Chicago, 15 (1-3): 1-1045. 
Nichols, J. T. 

1929, 1930. The fishes of Porto Eico and the Virgin Islands. (New 
York Academy of Sciences) Scientific survey of Porto Eico and 
the Virgin Islands, 10 (2, 3): 161-399. 
Springer, V. G. 

1958. Systematics and zoogeography of the clinid fishes of the sub- 
tribe Labrisomini Hubbs. Publ. Inst. Mar. Sci., 5: 417-492. 

1959. A new species of Labrisomus from the Caribbean Sea, with notes 
on other fishes of the subtribe Labrisomini. Copeia, 1959 (4) : 
289-292. 

Woods, L. P. 

1955. Western Atlantic species of the genus Eolocentrus. Fieldiana : 
Zool., 37: 91-119. 



BREVIORA 

Mmseiuim of Comparative Zoology 

Cambridge, Mass. Xovkmhkk lo, \\)ii'2 Number 170 



THE BRAIN OF THE EMU (DROMAEUS 
NOVAEHOLLANDIAE, LATH )' 

1. Gross Anatomy op^ the Braix and Pineal Body' 

B>- 

Stanley Cobb 

and 
Tii.LY Edinger 

The histology of the cerebral hemisphere of the einii has been 
extensively studied by Craigie (1935a, 1935b, 1940) and three 
diagrams of the hemisphere have been published. Drawings of the 
whole brain have also appeared in the literature (Strong, 1911; 
Kiienzi, 1918), and a photograph was published hy Anthony 
(1928). No description of the whole brain, however, is to be 
found. Since the emu is, next to the ostrich, our largest living 
bird, and since it belongs to a taxonomically controversial group, 
it seem>s of value to describe the brain and compare it with the 
brains of other birds. Moreover, the emu is considered, by Py- 
craft (1900) and many others, to be one of the most primitive of 
birds. The concept of "primitiveness" will be considered in 
the discussion at the end of this paper. 

MATERIAL 

Three specimens of Droiiiacus novachollandiae were collected 
by S. J. J. Davies in November 1960 in Western Australia for 
Professor Ernst Mayr, Director of the Harvard Museum of Com- 
parative Zoology. Two of them were kindly given to us by Pro- 
fessor Mayr for neurological study. The lieads had been cut off 



1 'I'liis spellirifT of Dronidciis is not the (iiic iU'CPiitpd by soJiii' newer cliecklists. 
but beciUise Dromiccius (an alternative si)elling) is the jierpetuation of a grapho- 
logical error (Newton, 1896) and because Dromaius is a less proper Latinization, 
it seems better to vise the older form. 

2 From the Museum of Comparative Zoology and the Department of Neurology 
and Psychiatry, Harvard University, and the Laboratory for Psychiatric Research, 
Massachusetts General Hospital. 

This investigation has been aided by grants from the Foundations Fund for 
Psychiatry and the National Institute of Neurological Disease and Blindness, 
grant #03429-02. 



2 BREVIORA No. 170 

and skinned, the eyes liad been removed, and the speeiniens had 
been fixed in 10 per cent formalin solution in the field. After 
about two months the heads were packed in moist condition and 
shipped in cellophane bags to the Fnited States. Here the brains 
were removed from the skulls after making photographs of vari- 
ous stages of the dissections. The brains were then fixed in fresh 
neutral formalin solution (10 per cent) for a month. One brain 
(that of Emu #85) was divided into its com])onent ])arts for 
weighing and special histological studies. The other brain (Emu 
4^104) was removed, photographed, fixed for a month in 10 per 
cent formalin as above, and embedded whole in celloidin for 
serial sectioning. Both are brains of adult males. The first (#85) 
appears somewhat larger and weighed 27.7 grams ; the second 
(#104) weighed 25.1 grams. 

DESCRIPTION 

The position of an avian brain within the skull is determined 
by many developmental factors. The most obvious are the shape 
of the bill, the size and position of the eyes, the habitual posture 
of the bird, and the size and shajie of the brain itself. Starck 
(1955) has given an excellent discussion of these relationships 
and emphasizes the importance of the size of the eye and the 
position of the orbit. One way of describing the position of the 
brain is to measure the angle between the cerebral axis and the 
axis of the bill (Cobb, 1959). In tlie emu this angle is about 27° 
(see Fig. 1), an angle somewhat smallei- than that of the gull 
{Larui< ar(/( itfafus: 84°) and the grouse {Boiiasa kdiIxHus: 36°), 
but distinctly greater than that of the cormorant {Phalacrocorax 
ourifu^: 15°) which has the sti-aightest (most extended) type of 
skull and an exceptionally small hraiii-bill angle. 

Besides showing the relation of the bi-ain to the skull. Figure 
1 shows the olfactory bulb and memln-anous sac of the olfactory 
nasal chamber ; the bulb s(>ems to be in direct contact with the 
chamber, but closer scrutiny shows that there is a space bridged 
by the short olfactory nerves. When the light, dijiloic bone of 
the bill is removed, the sac which forms the lining of the olfactory 
chamber is revealed. It is a fairly tough structure containing 
blood vessels and many nerve fibers. It is crossed anteriorly by a 
branch of the first division of the trigeminal nerve. The main 
nerve trunk of this division is seen passing through the orbit, close 
to the ()])tic nerve and u]) to a ])oint just below the olfactory bulb. 



1^62 THE BRAIN OK THE EMU 3 

This is the main sensory nerve from the bill, innervating' tiie skin 
and vibrissas Its large size suggests tliat taetile sense in the bill 
is acute and important. 

On opening the olfactory chamber, the most posterior of the 
three nasal chambers, a well-developed turbinal mound (superior 
or olfactory concha) is seen on the lateral Avail. It is covered with 
a soft, yellowish epithelium, which becomes thinner and less 
yellow as it spreads out over tlie dorsal and mesial aspects of the 
chamber. A vertical section through the nasal chambers of the 
bill at this level (Fig. 3) reveals that the concha is raised to, a 
height of about 5 mm. and is slightly constricted at its base, but 
is not folded into a spiral like the conchae of some vultures and 
albatrosses (Bang, I960). A specimen for microscopic examina- 
tion was taken from the dorsal surface of the olfactory chamber ; 
it shows cells and cilia typical of olfactory epithelium. About 
5 cm. anterior to the olfactory concha there is a large nostril 
(Fig. 1), which is the external opening of the anterior nasal 
chamber. 

The emu has large eyes and the orbits are spacious. As one 
sees in Figure 1 the brain lies mostly behind the orbit with the 
olfactory chambers in the bony structures just in front. The 
optic nerve enters the chiasm and passes directly to the optic 
lobe of the opposite side of the midbrain (Fig. 2C). The large 
fascicles of nerve fibers can be seen as they cross. The optic lobe 
is a large and conspicuous structure (Figs. 1. 2B and 2C'). Tn the 
lateral view only about one-fifth of it is covered by the overlying 
hemisphere. In Figure 2B (in which the parts of the brain are 
slightly separated) the relation of the optic lobe to hindbrain 
and forebrain is emphasized. It is clearly a part of the midl)rain. 
In fact, the optic lobes are homologues of the corpora bigemina 
of reptiles, and of the anterior corpora quadrigemina of mam- 
mals. They have taken a A-entrolateral position in birds, perhaps 
because it was easier there to make room for the extraordinary 
tectal development in this class of vertebrates. 

The emu brain when viewed from above (Figs. lA and 2A) 
impresses one by its triangular shape, with cerebral hemispheres 
broad posteriorly and narrow anteriorly. The olfactory bulbs 
protrude, forming the anterior pole of the hemisphere. On the 
vertex the two sagittal elevations of hyperstriatum stand out con- 
spicuously and are separated from the lateral parts of the hemis- 
pheres by a distinct sulcus, tlie vallc^cula (Portmann and Stinge- 
lin, 1961). 



4 BREVIORA No. 170 

The cerebellum is larger in comparison to the forebrain than 
in passerine birds. It has a greater diameter dorsoventrally than 
laterally (Figs. 1 and 2) although the auricles ]n-otrude laterally 
on each side. These lobes, composed of flocculus and nodule, are 
the only ones that complicate the simple conformation of the 
cerebellum, the corpus cerebelli being largely a mid-line organ 
corresponding to the vermis of mammals. Between the anterior 
surface of the cerebellum (culmen and declive) and the posterior 
poles of the cerebral hemispheres there is ample space for the 
pineal stalk and gland. 

The lateral view of the brain (Fig. 2B ) shows the relative sizes 
of the main subdivisions. For this photograph the forebrain, 
midbrain, and hindbrain were slightly pulled a])art. The hemi- 
spheres of the forebrain are well developed and extend backwards 
covering parts of the optic lobe and of the cerebellum. The great- 
est diameter of the hemisphere is 36 mm. and the greatest diam- 
eter of the olfactory bulb is !) nun., giving a ratio of 4 to 1 or 
25 per cent. This places the emu among those birds that have 
large olfactory bulbs (the Gruiformes, Caprinudgiformes, Procel- 
lariiformes, Podicepidiformes, and Apterygiformes). In a list 
of 47 different species of birds, arranged according to the relative 
size of the olfactory bulb, the largest at the top. the kiwi would 
come first and the emu seventh (Cobb, 1960). The anterior end 
of the hyperstriatum accessorium (sagittal elevation or AVulst) 
is close to the olfactory bulb, and the posterior end shades off into 
the neostriatum before reaching the occipital pole of the hem- 
isphere. Thus the emu has a large Wulst that reaches well back 
towards the occipital pole (Figs. 1 and 2) and well forward to a 
point close to the olfactory- bulb. 

A comparison of the external configuration of the brain of the 
emu with that of other birds shows that it resembles most sonu' 
herons and ducks. Tn comparing it with Stingelin's (1958) 
photographs, it is seen to be strikingly similar to the brain of 
Ixobrychus minutus (see his fig. 21 "Zwergreiher"). 

Seen from below (Fig. 2C ) the conspieuous characteristics of 
the emu brain are ; ( 1) the large, separated olfactory bulbs, form- 
ing the anterior pole; (2) the flatness of the ventral aspects of 
the lateral parts of the two cerebral hemispheres; and (3) the 
pair of big optic lobes shaped like flasks with their necks joined 
in the optic chiasm. The cerebellum is so narrow that it is almost 
hidden by the medulla oblongata, only the flocculi .showing on 
each side. The roots of the third, seventh, eighth, ninth, and 
tenth cranial nerves show in this view. 



1962 THE BRAIN OF THE EMU 5 

The brain of emu ^85 (after formalin fixation) weighed 27.7 
grams; liis body weight was 34 kg. The brain of emn #104 
weiglied 25.1 grams (also after formalin fixation ) ; body weight 
31 kg. This gives a ratio of brain weight to body weight in emu 
#85 of 1/1227 and in emu #104 a ratio of 1/1235. Little sig- 
nificance. h()\v('^•el■, should be given to these ratios because it is 
known that a living emu may vary 30 to 40 per cent in weight 
during a year due to conditions of food, climate and water supply. 
The first brain was separated into 8 pieces, for weighing, as 
follows : 

Olfactory bulb (right) (injured) 

Olfactory bulb (left) 0.12 grams 

Cerebral hemisphere (right) 8.85 grams 

Cerebral hemisphere (left) 8.8 grams 

Optic lobe (right) 7.3 grams 

Optic lobe (left) 7.3 grams 

Cerebellum 4.6 grams 

Brainstem 3.9 grams 



p?' 



The brainstem (defined by Portmann. 1I)4(). and named 
"Stammrest") is the basal mass of nerve tissue made i\p of 
thalamus, midbrain (with optic lobes removed) and hindbrain 
(with cerebellum removed). Portmann 's purpose was to choose 
as his common denominator that part of the brain \vhich varies 
least in its size relative to the size of the whole bird. That part is 
obviously the brainstem. He then compares its size to other parts 
of the brain and, by dividing the weight or volume of the stem 
into the eoresponding value for another part, he obtains his index. 
This ''index of cerebralization" he finds for an emu to be 4.18, 
obtained by dividing the weight of the "Stammrest" into the 
combined weight of the two hemispheres. In our emu #85 this 
index is 17.6/3.9 = 4.5. According to Portmann 's list the figures 
4.18 and 4.5 both place the emu far below parrots and ravens, 
but above loons, grebes, and quail. He believes that this quotient 
gives an expression of the "level of integration" of the brain 
for each species. 

The Pineal Body 

In the dissection of enni #104, a large })art of the post-central 
area of the calvarium was left intact and carefully lifted off the 
brain. The pineal stalk was thus torn away at its attachment to 
the diencephalon. It is 10 imn. in length and remained attached 



6 BREVIORA No. 170 

to the pineal body (Fig. 4). The body itself is embedded in the 
dura and lies in a dciiression of the cranial roof between the an- 
terior and posterior fossae. The dorsal position of the epiphysis 
is thus clearly demonstrated ; it lies between cerebrum and cere- 
bellum at the level of their dorsal surfaces. The stalk leaves the 
brain at a point just rostral to where forebrain joins midbrain. 
The pineal body is round and firm, slightly flattened dorso- 
ventrally. It is yellowish in contrast to the white skull. The 
fibrous envelope is continuous with dura which has strong bands 
spreading laterally and anteriorly. Removed from the mem- 
branes, the pineal body is roughly triangular, 7 mm. long on each 
side. With stalk attached, it Aveighs 0.1 gm. after formalin fix- 
ation. 

DISCUSSION 

The description of the gross anatomy of the brain of the emu 
brings up five points for discussion : 1 ) the size of the brain, 
2) the question of primitiveness, 3) the general shape of the 
brain in relation to the base of the skull, 4) the size and position 
of the Wulst, and 5) the topographic relations of the pineal body. 

The size of the brain in relation to body size and "intelligence" 
has been the subject of much study and many pronouncements. 
Suffice it to say here that in our opinion the relation of brain 
weight to body weight (so called cephalization) is a ratio too 
simple to give information of much significance. Portmann's 
(1952) pioneer work in describing an index of encephalization 
is an advance in the right direction. Body weight in birds is too 
grossly variable to be used in com])arison to the much more stable 
brain weight. Small birds may show rapid and marked change 
in weight. There is good evidence that some birds may lose from 
30 to 50 per cent of their body weight in 24 honi-s during a mi- 
gratory flight (Odum et al., 1961 ; Helms and Drury, 1960). The 
emu, being flightless, lives in a fairly uniform environment and 
does not go through the prolonged exertion of migratory flights. 
Its ratio of brain weight to body weight might, therefore, be 
relatively stable. Actually, in Dromaeus novaeJiollandiae this 
ratio is approximately 1/1230 (see p. 5). From the weights s'iven 
by Crile and Quiring (1940) we deduce that the ratio for an 
ostrich (Strnthio canielus massaicKs) is 1/2929; for a sparrow 
(Passer domesUcus) it is 1/23 ; and for a hummingbird {AmazUia 
tzacatl) it is 1/24. This does not mean that the hummingbird 
has a "better" brain than the emu. It merely indicates that the 



1962 THE BRAIN OF THE EMU 7 

body controlled by the brain of the hummingbird is just as com- 
plex as the body of the emu, thouo'h much smaller. The question 
as to which brain is "better," or more highly evolved, is mean- 
ingless unless one asks. ' ' Better for what ? ' ' Obviously, the hum- 
mino;bird's brain is better for Hij>ht and the emu's better for 
running. 

Another factor relative to brain size must be considered. It has 
been pointed out by Sholl (1956) that small brains are in general 
more closely packed with nerve cell bodies than large brains 
which have more glial structures between neurons. Man has 10.5 
nerve cell bodies per cubic micron ; a mouse has 142.5. 

In short, the need is to learn what parts of the brain, control- 
ling what organs, are larger or smaller in each family of birds. 
With more investigation into the quantitative anatomj' of the 
brain, some of these questions may be answered. Fritz (1949) 
has estimated the volume of four parts of the striatum in four 
different species of birds ; he found significant differences, but no 
correlation with Portmann 's cerebral index. 

Many authors have spoken of the emu, and in fact all ratite 
birds, as primitive, but their concept of primitiveness is not clear. 
Some seem to call these birds primitive because they are flightless 
and have no keel on the sternum (Leach, 1923), others because 
they have a straight type of skull base (Streckschadel) (Mari- 
nelli, 1928, p. 156). Stingelin (1958) considers those birds, with 
a small ^\^ulst which lies neither far forward nor far back, to be 
the less evolved type. The point would seem to be that one must 
not apply the term primitive in a general way to the emu (or 
probably any other bird). One should specify in what respect a 
given type or family is less evolved ("primitive") and in what 
respect it is more evolved (specialized). Even then, the gaps in 
our phylogenetic knowledge do not allow us to say whether the 
ratite sternum is due to a devolution from carinate ancestors or 
an evolution from cursorial reptiles. The presence of feathers 
and the avian type of brain suggest strongly a descent from 
flying ancestors. In respect to running and adaptation to life in 
open plains one feels confident in saying that the emu is highly 
evolved. 

Much work has been done on the development of the avian 
skull. Pertinent to an understanding of the shape of the emu's 
brain are three recent lines of investigation. Duym (1951) 
described the bending of the base of the skull in different birds 
and specified four types — the stretched or extended type of 



8 BREVIORA No. 170 

skull and three de<>-rees of bendiiifj'. Dullemeijer (1960) has re- 
lated the shape and size of the pi-ineipal parts of the brain to the 
amount of bendin<i' of the cerebral axis and has described four 
classes on this basis: 1) stretched skulls with little bending, 2) 
bending of 20 degrees, 3) bending of about 70 degrees, 4) bending 
of about 120 degrees. Starck (1955) has shown the great im- 
portance of the size of the eye and oi-bit in d(4cnnining the shape 
of the brain and skull. 

In the enui one finds a rather extended type of skull and a very 
large eye and orbit. Our measurements show that there is an 
angle of about 27 degrees between cerebral axis and bill axis, 
and a bending of the cerebral axis of about 13 degrees. Thus the 
emu falls into class "1" in Dullemeijer 's grouping. We agree 
with him in emphasizing that "the position and shape of the 
brain parts is influenced by the position of the bill and the posi- 
tion and size of the eye." The development of the bill in birds 
has been remarkably variable, and with these special develop- 
ments come variations in the bones of the skull and in the con- 
formation of the brain. 

As mentioned on page 4, the general shape of the brain of the 
emu resembles that of Lrohrychus but shape in itself is not very 
significant phylogenetically or physiologically in comparing 
birds' brains. One feature, however, may be of interest: the size 
and position of the Wulst, because this ganglion is conspicuous 
on the surface of the cerebral hemisphere and because its size in 
relation to that of the hemisphere appears to vary. Unfortunately, 
however, there are not enough quantitative data concerning the 
size of the Wulst in various types of birds to mak(^ any statements 
about its significance. 

Ill the lateral and dorsal views of the emu's brain the AVulst 
is conspicuous (Figs. 1 and 2). Its anterior end is almost in 
contact with the olfactory bull). Tiie posterior end reaches back 
to within 4 iinn. of the occipital i)ole of the hemisphere. This 
rounded ridge is long and lies parallel to the interhemispheric 
fissure — hence, the name used by L. Edinger et al. (1903) 
"Sagittalwulst," translated as "sagittal elevation" bj^ Portmann 
and Stingelin (1961). Its position in the emu is like that in the 
pigeon, a bird whose brain resembles Stingelin 's (1958) "Grund- 
typus." But the Wulst of the emu in relation to the rest of the 
hemisphere is both longer and broader than that of the pigeon. 

Stingelin emphasizes the importance of the position of the 
Wulst. In his chapter entitled "Comparison and extent of stri- 
atal fields" there is a comparative description of the striatal 



1962 THE BRAIN OF THE EMU 9 

ganglia in 18 species, with clear diagrams of each. He believes 
that in the "highly evolved" ("hochevolutiert," p. 38) species 
there is a tendency to marked frontal enlargement of the hem- 
isphere. This is achieved in two ways. In developmental line A 
the frontal pole is largely Wulst, the rostral end of which is in 
contact with the olfactory bulb. In developmental line B the 
frontal pole is made from the neostriatum and ventral hyper- 
striatum, the Wulst having receded to a position on the vertex 
by successive caudal shifts. From these observations Stingelin 
deduces a morphological rank ("Formwert") in relation to the 
basic type ("Grundtypus"). In developmental line A, crows and 
owls are considered the more highly developed groups ; in line B, 
the higher ones are snipe, spoonbill (PlataUa) and parrot, with 
a plover considered as "lower" and the lapwing as "middle." 
This rank order seems to be entirely based on cerebral anatomy. 
Reference is made neither to fossils nor to other characteristics 
sucli as brain axis or anatomy of skull. 

As descriptions of the different relationships of one ganglion 
of the brain to another, the figures and exposition of Stingelin 
have great value, but taken as indicating evolutionary levels they 
may be misleading. Until one knows the lines of descent from 
reptilian and avian ancestors, descriptions of such "Entwick- 
lungsgerichtungen" and levels of evolution with "hoher" and 
"niederer Formwert" are not justified because evolution is the 
process of phylogenetic transformation, a phenomenon that can- 
not be observed except in consecutive phases of an ancestral line. 

The "Horizontalmodifikationen" ( Stingelin 's fig. 32) are cer- 
tainly of interest as showing differences between the brains of 
living families of birds, but these modifications are not a basis 
for conclusions concerning evolutionary ancestry. In short, we 
doubt if any living bird has a conformation of the brain that we 
are justified in calling primitive. The data for making such a 
judgment are inadequate. On the other hand, research into the 
relative size of various parts of the brain, such as Fritz (1949) 
has done in Portmann's la])oratory. may give us important leads 
as to the degree of cerebral developments. Such investigations 
would be especially useful if correlated with behavior. 

We wish to emphasize the possibility of drawing erroneous con- 
clusions when the anatomy of living forms is used as evidence 
for describing an evolutionary process, disregarding the evidence 
from fossils. We welcome the opportunity to point to one feature 
of the emu brain as a gra])liic argument against a persistent, but 
erroneous theorv. This feature is the position of the pineal body. 



10 BREVIORA No. 170 

SO obvious -vvhen one looks at the dorsal surfaee of the brain. The 
theory is that absence in birds and mammals of a second epi- 
thalamie appendage in the pineal area, the "parietal eye" of 
reptiles, is due to enlargement of cerebrum and cerebellum in the 
two descendant classes. That concept supposes that the covering 
over of the diencephalon and mesencephalon by the cerebrum and 
cerebellum in Aves and Mammalia ol)structed the access of epi- 
thalamic appendages to the brain surface. This process is be- 
lieved to have caused devolution and loss of the predominantly 
sensory, stalked, second organ in the pineal complex which sur- 
vives only in lizards and the tuatara, the parietal eye. 

Actually, among birds, a pineal organ reaching to the level of 
the cerebral and cerebellar vaults, attached to the skull roof, is 
not an exceptional occurrence. This is found in our Drotnaeus 
and has been previously reported in brains of other ratites 
{Struthio, Rhea, Apteryx) and, as a button-shaped convexity, on 
the endocranial cast of the extinct Dinornis (Starck, 1955). The 
difference between these recently and carefully prepared speci- 
mens and those figured in the literature is not a real difference, 
but a matter of preparation. 

Kiienzi's (1918) diagrammatic figures of the brains of 36 dif- 
ferent kinds of birds give the impression that no bird possesses an 
externally visil)le pineal organ, as do almost all macroscopic fig- 
ures of avian brains in the literature, including Strong's (1911) 
figures which show Dromaeus. Kiienzi, however, mentions in 
several places (pp. 28, 52, 89) that the pineal body is too firmly 
embedded in the meninges to be removed with the brain ; he re- 
ports (pp. 70-71) that the pineal body in all birds studied occu- 
pies a median space between the posterior borders of the hemi- 
spheres and the front end of the cerebellum, its distal end reach- 
ing approximateh" to their dorsal level. Our observations on 
Buho, Corvus, Gallus, Columha, and Larus agree witli those of 
Kiienzi ; all have pineal bodies extending into the dura. A recent 
study on ten embryonic and three later developmental ])hases of 
Larus (Wetzig. 1961) also clearly testifies against the theory of 
mechanical suppression in l)irds of the second (the stalked) organ 
in the pineal complex of reptiles. The epiphysis extends to the 
level of the prospective skull roof in an early transitory phase, 
and again in the last phases of embryogeny. It is then, and re- 
mains in the adult, fitted into the space between cerebrum and 
cerebellum. It is club shaped, its apex coalesced with the dura in 
contact with the roof of the skull. 



]i)62 THE BRAIX OF THE EMU 11 

Thus, there is more space available than was assumed. The 
club shape (witli the largest circumference distal) suggests that 
the form is governed by the space available. While we are well 
aware that the shape of a predominantly glandular organ (Stam- 
mer, 1961) has no great significance, we do wish to draw atten- 
tion to the shape of the epiphysis of the cinu. When, as in our 
emus, the distal expansion is abrnjit, its connection with the 
corpus diencephali a mere stalk (Fig. 4), the avian pineal body 
strikingly resembles not the reptilian pineal organ, which is more 
or less sessile, but the reptilian parapineal vesicle with its nervous 
and vascular stalk — that is, the parietal eye. Krabbe (1961) 
observed this similarity in an embryo Cygnus. Many species of 
birds similarly demonstrate that there is no obstruction to the 
development of a parietal eye. 

The reason for the absence of a parietal sense organ in birds 
is obviously not mechanical suppression ; it is to be found in the 
fossil record, namely in pre-avian phylogeny. The presence of a 
parietal eye is reflected in a corresponding foramen of the roof 
of the skull. The fossil record of skulls plainly shows that the 
organ was first present and then lost in innumerable phyletic 
lines within the classes Pisces, Amphibia, and Kept ilia. Birds are 
an offshoot of the great reptilian subclass Archosauria. Among 
the many hundreds of known skulls from its various orders, only 
two have the parietal foramen. Significantly, both the specimens 
showing that heritage from Palaeozoic ancestors belong to the 
earliest forms identifiable as archosaurian, each representing the 
beginning of a suborder of the order Thecodontia, "stem archo- 
saurs" (Mesorhinus: Jaekel, 1910, and Erythrosuchus: Huene, 
1911). These openings in the parietal bones of Archosauria oc- 
curred for the last time at the beginning of the Mesozoic era in 
the earliest Triassic times. It follows that parietal eyes were 
lost, not within the evolution of birds, but in remote reptilian 
ancestors some 80 million years befoi-e the first, late Jurassic, 
appearance of feathered animals, and ])resumbaly more than 
]00 million years before the modern type of avian brain was 
evolved. 

In the Mammalia, likewise, both recent and fossil conditions 
plainly contradict the assumption that possession of a parietal 
photoreceptor became impossible because of progressive brain 
evolution. In various groups of mammals now living, much or all 
of the midbrain is dorsally exposed in a gap between cerebrum 
and cerebellum. Actually, it has long been known that in some 
bats, lagomorphs, and rodents the pineal gland extends into the 



12 BREVIORA No. 170 

dura niat(M- at tlic caudal end of the interhemisplierie fissure, and 
this condition has now been found prevalent in studies of a lar^ie 
material from a great number of genera (Pilleri, 1960). Tlie 
usual absence of a pineal organ in specimens of the rabbit brain 
is the result of its having been torn off:' with the tentorium during 
preparation. Furthermore, endoeranial easts show that in most 
early Tertiary mammals there w^as a considerable gap between 
cerebrum and cerebellum. The parietal foramen was obliterated, 
i.e. the parietal eye had been lost in a pre-mammalian phase of 
evolution, in this case, in mammal-like Reptilia shortly before 
the emergence of the new class. 

SUMMARY 

A description of the gross anatomy of the brain of Dromaeus 
novaehollandiae is presented on the basis of two specimens from 
Western Australia. The brain is of the extended type. The 
olfactory bulbs and sagittal elevations of the forebrain, and the 
optic lobes of the midbrain are comparatively large. The index 
of encephalization is 4.5. The brains weighed 27.7 and 25.1 
grams, respectively. The pineal body lies in a shallow fossa in the 
roof of the skull and weighed 0.1 gram. It is pointed out in 
discussion that there is no good reason for considering this brain 
to be primitive, and that phylogenetic relationships cannot jus- 
tifiablv be deduced from the anatomv of the brains of living birds. 

LITEEATUBE CITED 

Anthoxy, R. 

191^8. Lei;oiis sur k' i-civeau. (i. Doiu et Cio (Paris). 
Austin, O. L. 

1961. Birds of the woild. Ooldon Press (New York). 
Baxg, B. G. 

1960. An;itoiiiii-al evidence for olfactory function in some species of 
birds. Nature (London) 188: 547-.549. 
Cobb, S. 

19.59. (Jn the aiigk' of tlie cerebral axis in the American woodcock. 

Auk 78: 55-59. 
1960. A note on the size of the avian olfactory liulli. Epilepsia 1: 
395-402. 
C'raigie, E. II. 

1935a. The cerebral lieniisiiheres of the kiwi and of the emu (Aiitcri/x 
and Dromireius). .1. Anat. London 69: 380-393. 



1962 THE BRAIN OF THE EMU 13 

1935b. Tlie hipijocamiial mid |i;ii;iIii|)pocamp;il coitcx of flic emu 
(Droniiccius) . ,1. coiiip. Xciirol. 61: -463-591. 

19iU. The cerebral cortex in palaeugiuithine and neognathine bird.s. 
J. comp. Neurol. 73: 179-234. 
Crile, G. and D. P. Qiirixg 

1940. A record of the body weight and certain organ and gland weights 
in 3(>9U animals. Ohio J. Sci. 40: 219-259. 

DULLEMEI.JER, P. 

1960. Shape and size of the biain ])aits as architectonic factors in tlie 
skull of birds. Acta nioiphol. neerl.-scandin. 4: 96. 

Dl'YM, M. 

1951. On the head posture in birds and its lelation to some anatomical 
features. Proc. Kon. Xed. Akad. Wetensch. (C) 54: 202-211, 
260-271. 
Edinger, L., A. Wallenberg and G. Holmes 

1903. Untersuehungen iiber das Vorderhirn der Viigel. Abh. sencken- 
berg. naturf. Gis. 20: 341-426. 
Edinger, T. 

1961. Fossil brains reflect specialized behavior. World Xeurol. 2: 
934-941. 

Fritz, Walter 

1949. Vergleichende Studien iilier den Anteil von Striatumteilen am 

llemisphiirenvolunien des Vogelhirns. Eev. suisse Zool. 56: 

461-491. 
Helms, C. W., and W. H. Drury, Jr. 

1960. Winter and migratory weight and fat field studies on some 
North American buntings. Bird Banding 31: 1-40. 

HuENE, F. v. 

1911. Ulier Erijtlirosueliii.'f. Yertreter der neuen Eeptil-Ordnung Pely- 
cosimia. Geol. Pal. Abh. 14: 1-6(1. 
Jaekel, O. 

1910. Ulier einen neuen Belodonten aus dem Buntsandstein von Bern- 
burg. Sber. Ges. naturf. Freunde Berlin: 197-229. 
Krabbe, K. H. 

1961. La glande pineale. Worl<l Neurol. 2: 94-102. 
Kr'ENZl, W. 

191S. Versuch einer systematischen Morpliologie des (iehirns der Vogel. 
Rev. Suisse Zool. 26: 17-111. 
Leach, J. A. 

1923. An Australian Bird Book. Whitconie and Tombs f Melbourne V 
Marinelli, W. 

1928. fiber den Schadel der Schnepfe. Palaeobiologica 1: 135-160. 
Newton, A. et al. 

1896. Dictionary of Birds. Adam and Black (London). 
Odum, E. p., C. E. Connell, and H. L. Stoddard 

1961. Flight energy and estimated flight ranges of some migratory 
birds. Auk 78: 515-527. 



14 BREVIORA No. 170 

PlLLERI, G. 

1960. Beitiage zur vergleichenden ^loipliologie des Nagetiergehirns. 
Acta Aiiat, 42: 1-88. 

PORTMANN, A. 

1946. Etudes sur la eerebralisation ehez les oiseaux. Alauda 14: 1-20. 
1952. Die allgemeine l)iologi.sche Bedeutuiig der Cerebralisations- 
Studien. Bull, .sc-hweiz. Akad. mod. Wissoiiscli. 8: 253. 
PoRTMANN, A. and W. Stingelin 

1961. Biology and Comparative Physiology of Birds. Marshall, A. J. 
(ed. ). Academic Press (New York), volume 2: 1-36. 

Pycraft, W. p. 

1900. On the inorphology and phylogeny of the Palaeognathae (Rati- 
tae and Crypturi) and Neognathae (Carinata). Trans. Zool. 
See. London 15: 149-290. 
Sholl, D. a. 

1956. The Organization of the Cerebral Cortex. Methuen and Co. 
(London). 
Stammek, a. 

1961. Untersuchungen iilier die Struktur und die Innervation der 
Epiphyse bei Yogeln. Acta Univ. Szeged., Acta Biol. N.S. 7: 
65-75. 
Starck, D. 

1955. Die endokraniale Alorpholo'gie der Ratiten, besonders der Aptery- 
gidae und Dinornitludae. Morph. Jalirb. 96 : 14-72. 
Stingelin, W. 

1958. Vergleichend-morphologische Untersuchungen am Vorderhirn der 
Vogel auf cytologischer und cytoarcliitektonischer Grundlage. 
Helliing and Lichtenliahn (Basel). 
Strong, R. ^Nf. 

1911. On the olfactoi'v organs and smell in l)irds. J. Morphol. 22: 
619-658. 
Wetzig, H. 

1961. Die Entwicklung der Organe des Zwischenhirndaches (Epiphyse 
und Plexus choroideus anterior ) bei der Sturnimove, Larus canus, 
L. Morph. Jahrb. 101: 406-431. 



1962 



THE BRAIN OF THE EMU 



15 




in 



00 



to 









Si 

pq 



E z£ 



o 

CO 



-01 ^ 



O 



-iH 



UJ ii 



bo 



OS 



ID 

be 



16 



BREVIORA 



No. 170 




Figuri.' 1I>. riiotogiaph, life size, of dorsal view of same dissert ion 



1962 



THE BRAIX OF THE EMU 



17 



FIG 2^^ DORSAL 



FIG 2^ LATERAL 



FIG 2^ VENTRAL 




Figure 2. Three views of the brain of the emu ( Drnmaeus novaehollan- 
diae) #104. Life size. F, forebrain hemisphere, E, hindbrain, il, mid- 
brain, showing optic lobe (T) and optic- chiasm (C), V. vallecula, TT, Wulst 
or hyperstriatum aceessorium. 



18 



BREVIORA 



No. 170 



FIG 3 



FIG 4 



FIG 



B 




Figure 3. Section tlirough the nasal chambers cut in frontal vertical 
showing the olfactory (or posterior) chamber (P) into which protrudes the 
olfactory condha (C) covered with yellow olfactory epithelium. Below is 
seen part of the middle nasal chamber (M). The two chambers are di^aded 
by the sepfiiin. They connect anteriorly witli the anterior chamber and the 
external nostril. Life size. 

Figure 4A. Photograph, life size, of pineal body (PB) lying in shallow 
cavity of the ealvarium, posteroanterior view. The stalk (S) protrudes 
downward. 

Figure 4B. Ventral view, looking upwards at under surface of ealvarium. 
The stalk (S) is bent backwards. 



BREVIORA 

Mmseiiioi of Comparative Zoology 



Cambkidgk, Mass. 1)kcembp:r 14, 11H)2 ^'^MB^:R 171 

NOTES OX AMPHLSBAEXIDS ( AMPHI8BAEXIA : REP- 

TILIA). 6. REDESCRTPTIOX AXD RANGE EXTEXSION 

OF AMPHISBAENA SFURRELLI BOULENGER. 

By Carl Gans 

Department of Biology, The University of Buffalo, 
Buffalo 14, Xew York 



111 1915 Boulenofer (p. 659) described the new species of 
Amphishaeua spurrdJi, characterized primarily by the presence 
of tubercular or subcoiiical segments on the dorsal surface of 
the tail. The two syiitypes were collected at "Anda Goya 
[Colombia], at the junction of the R. Condoto and San Juan." 
The only subsequent record of the species (Burt and Burt, 1931. 
p. 40) is the citation of a single specimen (A.M.X'.H. 18261) 
from the neighborino: locality of ''Boca de la Raspadura, " 
Colombia, without su])p](MU(Mitary description. 

The present note is based upon a re-examination of these 
three and of two additional specimens, one of which extends the 
range of A. spnrrcUi into Panama. The original description has 
been amended and rewritten according to the standard pattern 
(Gans and Alexander. 1962). Simple, non-idealized illustrations 
are included in the present paper. 

It is a pleasure to acknowledge the support of the X'ational 
Science Foundation (XSF G-9054. G-21819). Examination of 
the types was made possible by assistance from the estate of 
Leo Leeser. Specimens Avere examined through courtesy of 
C. M. Bogert. The American Museum of N^'atural History 
(A.M.X.H.), Miss Alice G. C. Grandison. British Museum 
(Xatural History) (B.M. ). R. F. Inger and H. Marx. Chicago 
Natural History Museum (C.X.H.M.), and E. E. Williams, 
Museum of Comparative Zoology (M.C.Z.). I am particularly 
grateful to Dr. Federico Medem who made the C.X.H.M. speci- 
men available, and to Miss C. Rhodes for technical assistance. 



BREVIORA 



No. 171 



Amphisbaena spurrelli Boulenger, 1915. 

Amphubaena spHrrelli Boulenger, 191.5, p. (359. Terra typit-a : ' * Anda Goya, 
at the jiiiK'tion of the R. Coiidoto and San .luaii," Colonihia. LECTO- 
TYPE: B.M. 19ir3.10.lM.9 (liy present designation). J'ARATYPE: 
B.M. 1915.10.21.8. 

Diagnosis: A form of Ampliisbacna without fusions of head 
scah^s; with 4 oval [not round] prech)aea] pores; and witli the 
dorsal and lateral surfaces of the eaudal tip covered with eonieal 
or tubercular segments. Specimens have 218 to 222 body annuli ; 
18 to 20 caudal annuli ; 16 to 18 dorsal and 16 to 18 ventral seg- 
ments per midbody annulus; and one row of postgenial and one 
row of postmalar chin shields. There is no visible autotomy con- 
stricton of the tail. Autotomy takes place after the seventh 
annulus. 

Notes 0)1 tin types: Boulenger (191."), p. (i59) illustrated the 
smaller of his syntypes, which has here been chosen as a lecto- 
type. The types, still extant and in good condition, suggest that 
his illustrations were idealized, and s(>veral of his counts [shown 

78 




Fig. 1. Amphisbarna .^purrrlli. Map showing lo'-alities mentioned in 
te.xt. Anda Goya and Boea de la Rasi>adura are actually cdoser together 
than can be indicated on a map ilrawii to this scale. 



1962 AMPHISBAENA SPURRELLI 3 

ill brackets in the table] erroneous. Tbe erroi-s do not affect the 
validity of the species. 

I)( scvipiiou : ^leristic charactcM's are listed in the table. Fiynre 
."] shows the head scalation, l-''ij:ure 4 the sef):nientatioii of cloaca 
and tail, P^igures 5 to 8 ])hotographs of head, niidbody pattern 
and tail. 

Presei'A'ed specimens are various [faded] shades of brown 
dorsall}', somewhat lighter vent rally. The darker dorsal color 
is in part produced by a darkening of the rectangular center of 
each segment, the contrast with the lighter segmental margins 
giving the impression of dark spots. The fully dark dorsal spots 
descend the sides to approximately the third ventral below the 
lateral groove on each side. Ventrad from this the dark center 
shrinks drastically or may fade out entirely. The anterior fifth 
(M.C'.Z. 8f)784), or tlir head alone, lacks the dark colored seg- 
ments. 

The head scalation shows some variability and no major fu- 
sions. An azygous rostral tiarely visible in dorsal view is fol- 
lowed by three pairs of enlarged cephalic shields in contact 
along the dorsal midline. The nostrils pierce the first pair 
(nasals). The second pair (prefrontals) are the largest segments 
of the head. There are three supra- and two and a half infra- 
labials, as the third infralabial extends considerably beyond the 
angulus oris. The suj^ralabials are large, the second much the 
largest. The CX.H.^I. s))ecimen has this segment subdivided 
differently on both sides. The second infralabials are the largest 
segments on the lower .jaw. Small segments lie beyond 
the angulus oris in the position of fourth supralabials. The 
mental is shaped like a truncated wedge with a posteriorly con- 
vex tip. The postmental is hexagonal and elongate. It lies in 
lateral contact with the medial edges of the second infralabials. 
as well as the anterior poi-tion of the medial edges of the rela- 
tively short, wedge-shaped malars. There are one to two rows 
of postgenial segments, follo\\ed by a single postmalar row, the 
lateral segments of which are slightly enlarged. 

The head is relatively blunt, flattened slightly dorsoventrally 
and oval in cross-section. The lower jaw is but slightly shorter 
than the upper. The sides of the head would, if extended, inter- 
sect some distance anterior to the rostral tip, even in adults 



BREVIORA 



No. 171 



E 
E 



c 

0) 



27 



25 



23 



21 



19 



17 



o» 



20 



22 



24 



26 



28 



30 



Snout - Vent Length - cm 



Fig. '2. Amphishama spiirrclli. Scatter diagram showing plot of tail 
length versus snout-vent length for the available specimens. The lectotype 
is shown as a hollow circle. 



in which the bulge of the temporal musculature changes the out- 
line. The attachment of the skin to the crest of the skull pro- 
duces a concave dishing of the interfrontal suture, particularly 
in adult specimens. 

The first body annulus curves forward to contact the frontals. 
Its dorsalmost segments may be somewhat enlarged, and one of 
the specimens has an intercalated dorsal half-annulus. The 
second through fifth annuli are narrowed, and the fourth marks 
the level of the head joint or the point at which the bulge of the 
temporal musculature returns to normal. 



1962 



AMI'HISBAENA SPURRELLI 






Fig. 3. AmpJiisbaena .fpurreUi. Dorsal, lateral and ventral views of the 
head of A.M.X.H. 18261 from Boca de la Raspadura, Colombia. The line 
equals 1 mm to scale. (V. Cummings, del.). 



BREVIORA 



No. 171 






Fig. 4. AmpliLsbacua spiimlli. noisnl, lateral ami ventral virws of tlu' 
cloaea aiul tail of A.M.X.H. ]8l^(il from Boi-a de la Raspaduia, Cdlomliia. 
Note the extent of cone formation on the tail ami the Hattened, .slit-sha]itMl 
precloacal pores. Tlie line equals 1 mm to scale. (V. C'umniinf»H, del.). 



The dorsal gTOOve is only indicated on the head. The ventral 
groove is indicated mainly as a gap between aligned segments. 
The lateral grooves start about one and a half head lengths be- 
hind the head joint, and are well defined by a double row of 
triangular segmental fragments. The middorsal segments are 
almost twice as long as wide, the midventral segments are almost 
twice as wide as long. 



1962 A.Ml'lilSHAEXA SPURRELLI 7 

Tlip oval pi'ccloacal pores lie in a single uninterrupted row 
ol" normal sized seg-ments antcrioi- to the predoaeal shield. The 
preeloaeals ai-c cliaraeterized by a eentral group of four some- 
what elongate segments. The posteloaeals, slightly gi-eater in 
number, eliaraeteristieally have a set of two to four midventral 
and enlarged segments and, tiaidving these, several split segments 
entering the doaeal sides. The cloaca may be entii'ely prolapsed. 

The tail becomes gradually wider posterior to the eloacal slit 
and somewhat higher as well. The ventral surface appears plane 
and an ext(Misi())i of the pi-ecloacal region. The terminal third 
of the tail shows reduction, with the tip about twice as high as 
wide. The segments of the dorsal and lateral surfaces are 
strongly t nbcrculate or cone-shaped. This character facilitates 
diagnosis of specimens with intact tails. Caudal autotomy takes 
place behind the seventh ])ostcloacal annulus (cf. Vanzolini, 
1951, p. 2.3). 

Range: Lowland ri\er valleys of northwestei'n South America, 
from extreme northern Colombia (Choco) to southern Panama. 

Distrihulioit rcconh: COLOMBIA: Choco Province: Anda 
Goya, mouth of Rio Condoto ( l)()ulenger, 1915); B.M. 
19i5.10.21.8 (PAIJATYPE). 1915.10.21.9 (LECTOTYPE ) ; 
C.N.H.M. 18098S [E. K. Dunn.' leg. per F. Medem|. Boca de la 
Raspadura (Burt and Burt, VX^\ ) ; A.M.X.H. 18261. PANAMA: 
Tucuti branch. Tuira River [H. C. Clark, leg.] M.C.Z. 39784. 

LiTi:K.\'r["'Ri': cited 

Boi'LENGER, George Alhert 

19in. Descriptions of a new AmpJiishaiiui nnd a new snake discovered 
l^y Dr. 11. Or. F. Spuriell in soutliern ("olninl)i;i. I'roc. Zool. Soc. 
London, (1915), i)p 659-61. 
BiRT, Charles E. and .\[ay Daxiieim Dirt 

1931. Sonth American lizards in tlie collection of the American Mu- 
senm of Natural History. Bull. Amer. Mus. Xat. Hist., vol. 61, 
art. 7, PI). 227-395. 
Gans, Carl Axn Alexander Allan Alexander 

1962a. Studies on amphisbaenids (Amphisbaenia, Reptilia), 2. On 
the amphisl)aenids of the Antilles. Bull. Mus. Comp. Zool., 
vol. 128, no. 3, pp. 65-158. 
N'anzolixi, Paulo Emilu) 

1951. Anipliisbaena fiiliginnm. Contribution to the knowledge of the 
Brasilian lizards of the family Amphisbaenidae Gray, 1825. 
6. On the geographical distribution and differentiation of 
.1 iiiiiliishaena fuliginosa Linne. Bull. Mus. Comp. Zool., vol. 
lUti, no. 1 , i)p. l-()7. 



8 



BREVIORA 



No. 171 




Fig. f). Amphisboena fipiin-fUi. Lateral view of the liead of the topotype, 
C.N.H.M. 180988, from Aiida (ioya, rohiiiiliia. Note tlie irresuhir sulj- 
division of the second suprahibial. 



1962 



AMPHISBAENA SPURRELLI 



9 




Fig. (i. .1 iiiiihislKii'iKi siHirnlli. Dorsal ( k'ft ) and ventral (riuiit) viuw.s 
of A.M.X.H. spec-inien at midbody. Note the darkening of the dorsal seg- 
mental centers. 




Fig. 7. Amphisbaena spurreUi. View of caudal tip erossdighted to empha- 
size the knolilied nature of the tenninal segments. 



10 



BREVIOKA 



No. 171 




Fig. 8. Amphishaena spurrelli. Ventral view of the cloaea and tail of 
the A.M.N. H. .specimen. Note the onset of autotoniy at annulus .seven, also 
the contrast between the plane ventral and eonieal laterodor.sal i-andal 
segments. 



1962 



AMPHISBAENA SPURRELLI 



11 



Si. 

=0 



■5 



^-1 
o 















^ ^i' 
H > 



x — 






W 



"& 


"71 

+ 


+ 


11 

+ 


11 

+ 


o 


•t 


t^ 


o 


or. 


^ 


ci 


o; 


11 


11 




11 


CI 


n 


11 






11 M 



CC 



I'l 



Z: 3C 



00 
QC 



m 






+ + 
n o 

11 (M 



oc -^ 



CO 



11 



•^ tC OC ■£ -^ 



in 



or 


-t 


11 


f-H 






11 


HI 


11 


11 


11 


11 







L-^ 




Lt 








X 


^^ 


— ^ 


r— < 








Cl 


'"'' 


o; 


X 




"~ 


-~ 


^— 


_« 


_- ^ 


OO 
























. 1 








• 1 


—  


~ 


^' 


~ 


o 


'jH 


^- 


' — 


ii 


-— 


S: 


^ 


X 














t^ 


— ' 


_ 


i; 


i.» 


r" 


* 


OS 












«rt 


CO 


















~, 


3 


S 


M 




N3 

d 


<;' 


;^ 




M 




d 


S 



BREVIORA 

Mmseiinii of Comparative Zoology 



Cambridge, Mass. December 14, 196 2 Number 172 

A NEW SPECIES OF THE RODENT PIPE.STONEOMYS 
FROM THE OLIGOCENE OF NEBRASKA 

By Raymond Alf 
Webb School of California, Claremont 

In 1956 John C. Donohoe described from the Pipestone Springs 
formation of Chadronian age in the Montana Oligocene two jaw 
fragments of a new rodent, Fipestoneomys bisulcatus, that he 
placed in the family Aplodontidae. 

The purpose of this paper is to record the occurrence of Pipe- 
sioneomys in the Chadron formation of Nebraska, to suggest that 
it be placed tentatively with the castorids rather than with the 
aplodontids, and to propose a new species of the genus. 

North of Crawford, Nebraska, well known exposures of Oligo- 
cene sediments occur along the Pine Ridge escarpment. For 
several years, through the kindness of Frank Arner of Crawford, 
Nebraska, the Webb School of California has had the privilege of 
collecting fossils on the Arner ranch. In Sec. 26, T. 33N., R. 
53W., Sioux County, Nebraska, are several harvester ant mounds 
in the Chadron formation, and from these have been collected a 
rich microfauna including Pipestoncomys. 

I wish to thank W. D. Turnbull of the Chicago Natural History 
Museum for the loan of the type specimen of P. hisnlcatua fur 
study, and Bryan Patterson of Harvard University, and Dr. 
Mary Dawson of the National Science Foundation for advice. The 
new species is named in honor of Professor Bryan Patterson. Tlic 
drawings are by Nick Strekalovsky. The abbreviations refer to 
the following institutions or collections: C.N.H.M., Chicago Na- 
tural History Museum; M.C.Z., Museum of Comparative Zoology; 
R.A.M., Raymond Alf Museum, Webb School, Claremont, Cali- 
fornia. 



2 BREVIORA No. 172 

Family CASTORIDAE ? 

PiPESTOXEOMYS PATTERSOXI 11. Sp. 

Type: M.C.Z. no. 7113, fragment of left maxilla with Mi-2. 

mjpodigm: The type and M.C.Z. nos. 7110, RP* ; 7106, LP-*; 
7108, LP4 ; 7102, LMi ; 7111, RM2 ; 7104, LP4 ; 7107, RP4 ; 7103, 
RMi ; 7109, RMj ; 7112, RMi ; 7101, LM^ ; and R.A.M. nos. 3072, 
RM2 ; 1683, LM2 ; 671, LM2 ; 1283, RP4 ; 2105, RP4 ; 2104, RMj. 

Horizon : The base of the Chadron formation along Pine Ridge 
is marked by a basal conglomerate resting on the weathered sur- 
face of the Pierre shale. The top is characterized by the upper 
purple-white layer, a continuous purple-tinted white limestone 
designated by Schultz and Stout (1938) as the boundary be- 
tween the Chadron and the Brule formations of the White River 
group in this area. About half way between this and the basal 
conglomerate there is a second limestone lens, which is referred 
to as the lower purple-white layer. The specimens come from well 
below the lower purple-white layer. 

Localtiy: Sec. 26, T. 33N., R. 53W., Sioux County, Nebraska. 

Diagnosis: Differing from P..hisulcatus as follows: paracone 
and metacone of Mi"2 not rounded, paracone and anteroloph not 
sharply separated, anteroexternal and anterointernal lakes sepa- 
rate, relatively small ; Mi smaller, more rectangular in outline. 

Description : The paracone and metacone of M^'^^ though dis- 
tinct and higher than the remainder of the crown, are not the 
rounded cusps that they are in P. hisulcatus. In P. hisulcatus the 
paracone is separated from the anteroloph by a deep groove com- 
municating with the anteroexternal lake, whereas in P. pattersoni 
the separation is very faint. In P. hisulcatus the anteroexternal 
and anterointernal lakes communicate by a slight groove. In 
P. pattersoni the corresponding lakes are completely separate and 
are also relatively smaller and are more delicately outlined. There 
is no trace of a mesostyle. P^ is larger than either M^ or M^. A 
deep groove separates the anteroloph from the rest of the tooth 
and communicates openly with a deep valley that separates the 
paracone-metacone loph from the hypocone. On the anteroloph 
there is a lake below the anterocone and an embayment projecting 
inward to the protocone. As on the upper molars, a mesostyle is 
lacking. 

The lower first molar of P. pattersoni differs in shape from the 
corresponding tooth in P. hisulcatus, being much more rectangu- 
lar. It is also smaller and more delicate. P4 is larger than Mj. 



19G2 



NEW OLIGOCENE RODENT 






Fig. 1. Pipestone&mys pattersani. A, M.C.Z. no. 7113, left maxillary frag- 
ment bearing Mi-2, type. B, M.C.Z. no. 7110, RP^. C, M.C.Z. no. 7103, RMi. 
D, M.C.Z. no. 7107, RP4. X 25. 



BREVIORA 



No. 172 



The anterolophid, metalophid, and posteroloi)hid are well devel- 
oped and separated by deep grooves that terminate at the meta- 
conid and entoconid. There is a lake posterior to the metaconid 
and one on the posterolophid adjacent to the hypocones. 

Discussion: Pipestoncomys, especially in the light of the new 
evidence reported here, does not fit well into the Aplodontidae. 
The upper Eocene Eohaplomys, the earliest known aplodontid, 

1 '^ 3 
has the dental formula (Stock, 1935; McGrew, 1941). 

_L U J. o 

This formula is retained to the present. The presence or absence 
of P^ is indeterminable in the holotype of P. hisulcatus, but a 




2. mm 



» 

Fig. 2. Pipestoiieomys bisulcatus. Right M^"^, crown view, C.N.H.M. no. 
UM 409, type. 



small fragment of the maxilla anterointernal to a P"^ of P. pattcr- 
soni (M.C.Z. no. 7106) shows no sign of the presence of P^. P| 
are large relative to the molars. The anterior margin of the mas- 
seteric fossa in P. bisidcatus extends to beneath the middle part 
of P4, and the ramus is relatively thick and deep. These charac- 
ters are not typical of the early aplodontids, but are quite like 
those occurring in castorids. The cheek tooth structure seems to 
me to accord better with that of castorids than with that of aplo- 
dontids. A lower molar of P. pattersoni (M.C.Z. no. 7112) was 
ground down to show the pattern of a well worn tooth. The median 
external groove (hypoflexid) persists but its medial portion be- 
comes transformed into a lake, forming a pattern similar in gen- 
eral to that found in, e.g., Paleocastor (cf. Fig. 3 and Stirton 1935, 
Fig. 30). The incisor enamel of Recent Castor, Miocene Paleo- 
castor, and Oligocene Agnotocastor is characterized by an orange 



1962 



NEW OLIGOCENE RODENT 



color (Wilson 1949), that of aplodontids is not pigmented. 
Among the many incisors picked out of the ant mounds are a 
few with orange color, and those of appropriate size could well 
pattersoni. 



belong to P 




Fig. 3. Pipestoneomys pattersoni. M.C.Z. no. 7112, EMi ground down to 
show pattern near base of crown. X 25. 



The past and present distribution of aplodontids is also of some 
interest in this connection. Not only is the range of Recent Aplo- 
dontia limited to a strip along the west coast of North America, 
but no unquestioned fossil form has been found east of the Great 
Basin. 1 If this picture of geographic distribution is adequate, we 
may have another item of evidence that argues against aplodontid 
affinities for Pipestoneomys. 

Pipestoneomys seems to me to fit more reasonably into the Cas- 
toridae than into the Aplodontidae, and I know of no other 
family of appropriate range to which it might be referred. If this 
tentative assignment is correct, the genus is the earliest beaver so 
far known, although its position in castorid phylogeny is un- 
certain, 

EEFERENCES 

DONOHOE, J. C. 

1956. New aplodontid rodent from Montana Oligocene. Jour. Mam- 
mal., 37: 264-268. 
McGrew, p. O. 

1941. The Aplodontoidea. Geo!. Ser., Field Mus. Xat. Hist., 9: 1-30. 



iShotwell. in his interesting studj of the aplodontids (I'J-jS), mentions Pipes- 
toneomys only in passing, noting that P. bisulaatits occurred well to the east of 
thf other known fossihs. 



6 BREVIORA No. 172 

ScHULTZ, C. B. and T. M. Stout 

1938. Classification of Oligocene sediments in Nebraska. Bull. Univ. 
Nebr. State Mus., 4: 15-52. 
Shotwell, J. A. 

1958. Evolution and biogeography of the aplodontid and mylagaulid 
rodents. Evolution, 12: 451-484. 
Stibton, E. a. 

1935. A review of the Tertiary beavers. Univ. Calif. Publ. Bull. Dept. 
Geol. Sci., 23: 391-458. 
Stock, C. 

1935. New genus of rodent from the Sespe Eocene. Bull. Geol. Soc. 
Amer., 46: 61-68. 
Wilson-, E. W. 

1949. Early Tertiary rodents of North America. Carnegie Tnst. Wash- 
ington, Publ. 584: 71-164. 



Table 1 

Pipestoneomys. Upper Dentition 

Measurements in millimeters 

P. pattersoni . P. bisuleatus 

C.N.H.M. UM 409, type 
Crown length 

Ml M.C.Z. 7102 1.62 1.78 

M.C.Z. 7113 1.47 

M2 M.C.Z. 7111 1.48 1.6 

M.C.Z. 7113 1.23 

E.A.M. 671 1.48 

E.A.M. 1683 1.62 

E.A.M. 3072 1.44 

P4 M.C.Z. 7106 2.16 

M.C.Z. 7108 2.07 

M.C.Z. 7110 1.98 

Greatest width 

Ml M.C.Z. 7102 1.71 1.95 

M.C.Z. 7113 1.67 

M2 E.A.M. 1683 1.62 1.81 

E.A.M. 671 1.44 

M.C.Z. 7111 1.35 

M.C.Z. 7113 1.67 

E.A.M. 3072 1.35 

P-t M.C.Z. 7106 1.98 

M.C.Z. 7108 2.09 

M.C.Z. 7110 1.94 



1962 NEW OLIGOCENE RODENT 

Table 2 

Pipestoneomys. Lower Dentition 

Measurements in millimeters 





P. pan 


ersoni 




Crown length 






Ml 


M.C.Z. 


7101 


1.44 




M.C.Z. 


7103 


1.48 




M.C.Z. 


7109 


1.74 




M.C.Z. 


7112 


1.89 




E.A.M. 


2104 


1.62 


P4 


M.C.Z. 


7104 


1.62 




M.C.Z. 


7107 


1.51 




E.A.M. 


1283 


1.80 




E.A.M. 


2105 


1.71 


Greatest width 






Ml 


M.C.Z. 


7101 


1.51 




M.C.Z. 


7109 


1.62 




M.C.Z. 


7112 


1.62 




M.C.Z. 


7103 


1.35 




E.A.M. 


2104 


1.53 


P4 


M.C.Z. 


7104 


1.35 




M.C.Z. 


7107 


1.42 




E.A.M. 


1283 


1.48 




E.A.M. 


2105 


1.62 



P. bis III cut US 
C.N.H.M. UM 408 



dP4 1.78 



1.69 



dP4 1.18 



BREVIORA 



MiuiseiLim of Comparative Zoology 

Cambridge, Ma>s. I)k( f.mhkh 24, 19(iL' Xu.mbei; l?,'! 

NEW (SPECIES UF LAND MOJ.HSKS VUijM THE 
REPUBLICA DOMINICANA 

By 

"WiLLiA.M J. Clench 

We are deeply indebted to Drs. Staidey Rand, Clayton Kay 
and Juan Rivero for two important collections of land moUusks 
collected in the Republica Dominicana. Both collections were 
made in areas hitherto unknown as far as the land inollusks were 
concerned. Of the foni- species herein described, two represent 
a subfamily and a <i'enus not previously known from IIispa)iiola. 
Probal)ly one-third of the species are known only from their 
type localities, the remaining two-thirds from a very few locali- 
ties. For this reason it is impossible to give significant distribu- 
tional patterns of any one species, and but few generic ])atterns. 
Our present knowledge of the land and freshwater mollusks of 
Hispaniola is [irobably only about ten j)ercen-t, compared with 
that of Cuba. 31uch of th.is is due, of course, to the fact that 
there are no local interested persons in IIisj)aniola. Cuba has 
had a host of highly trained naturalists, such as .Juan (Jundlach, 
Charles Wriglit, Filip Poey, Rafael Arango. Carlos de la Torre, 
P. J. Bermudez and C. (A. Aguayo. All have made extensive 
collecting trips throughout most of Cuba and their interest 
stimulated many others to collect. As a conse(juence, there are 
comparatively few areas in Cuba where the fauna still remains 
unknown. Conversely. Hispaniola has had none of the local 
stimulation, and what is known of the area has been gathered by 
outsiders on a few expeditions such as those made by Augnste 
Salle in 1847-18.31, D. V. AVeinland in IS.")?. Justus Iljalmarson 
in 1858, Paul Bartsdi m 11)17, 1!)2(). and l!)2!), \V. J. Eyei-dam 
in 1927, C. R. Orcutt in 1929-1980, D. C. Pease in 1932 and 
W. J. Clench. R. A. McLean and TI. D. Russell in 1987. Many 
others have contributed matei-ial. imicli of it secondary to other 
zoological or botanical pursuits (see Crosse 1891. and Bartsch 



2 BREVIORA No. 173 

1946). There remains much territory on this island about whicli 
nothing' is known, iiarticnlai'ly tli(^ central mountain system, the 
Cordillera Central. 

IlKLICIXIDAE 

Helicina .ji'LIAK, new species 

Phlte 1. figure 2 

Description. Shell reaeliing 8 nnn. in greater diameter, im- 
perforate, subo-lobose, smooth and shining. Color a bright ])ea- 
green to yellow with the first IV2 post-nuelear whorls with a 
broad band of brownish purple, as well as a peripheral band of 
the same color. Columellar area white aiul occasionally mar- 
gined witli blue. There may Ix' one or more axial bands which 
occur only from the peripheral band to the suture above. AVhorls 
4 and convex. Spire depressed and obtus(\ Aperture subtri- 
angular in shape. Outer lip reflected, wiiite. and with a mod- 
erate dejiression behind. Parietal area thinly glazed. Columella 
very sliort. Sculpture consisting of very fine gi-owth lines. 
Height Widtli 

5.5 mm. 8.U mm. lIoh)type 

5.1mm. 7.0 mm. I'aratype 

Types. The holotype is in the Mu-eum of Comparative Zool- 
ogy, no. 168267. from Colonia Ramfis. 20 km. W of San Cristobal. 
Republica Dominicana. Di-. Juan Rivero collector. March 19"7. 
Paratypes from the same locality in the Museum of Com]iarative 
Zoology and the Academy of Natural Sciences of Philadelphia. 

Remarks. This species differs from Helicina ririelis Lamarck 
to which it is j:)robably related, by being much smalhn- and l\v 
having several color j)hases in addition to tlie axial brown line 
or lines which extend from the peripheral band to the suture 
above. //. viridis reaches 14 mm. in width and 0.5 nun. in height 
and is a pure pea-green in color. The two species are the same 
in shell outline. 

Proserpina marcanoi, new species 

Plate 1, figure 8 

Description. Shell reaching 4.7 nnn. in width, depressed, im- 
perforate, smooth and shining. Color a liglit whitish green. 
Whorls 5 and convex. Spii-c depressed. Aperture semicircular. 



1962 LAND MOLLUSKS IliOAI IIISPANIOLA 3 

Outer lip simple. Parietal wall glazed and supporting a high 
and thin lamella. Columella short and snppoi-ting a second 
lamella whieli is about one-third as high as the parietal lamella. 
Umbilical area slightly depressed. Suture defined but not im- 
pressed. Sculpture consisting of exceedingly fine growth lines. 

Height Width 

2.4 mm. 4.7 mm. Ilohjtype 

2.2 mm. 4.4 nnu. Pai-atype 

2.1 nnn. 4.8 mm. 

Typts. The holotype is in the Museum of Comparative Zool- 
ogy, no. 1881)11. from Colonia Ramfis. 20 km. W of San 
Cristobal, Kepublica Dommicana. Two paraivpes are in the 
Museum of Comparative Zooh)gy from the same locality. Dr. 
Juan Rivero collector, IMarch 1!)57. 

Remarks. This is the first si)ecies of the Proserpininae to be 
recorded from Hispaniola. The snbfamily lias been known 
elsewhere only from Jamaica and Cuba. In relationship, it 
appears to be nearest to P. dcjtrcssa d'Orbigny, a species Miiich 
is widely distributed in Cuba. It differs from dcpressa l)y being 
smaller in size, proportionally higher, and in having a columclhir 
himella nearly twice as high as the Cuban species. 

This speeies is named for Eugenio ^larcano of the Instituto 
de Investigaciones Botanieas y Zoologieas de la Cniversidad de 
Santo Domingo, who eollected with Dr. Juan Rivero when he 
was at Colonia Ramfis in IMarch 1957. 

TRCXCATELLIDAE 

GeOMELAXIA { ]\1eRRILLL\XA ) RIVEKOI. new SI)ecies 

Plate 1, figure 4 

Dcscrlpiion. Shell reaching 7.(5 mm. in heiglit (truncated), 
extended, iin])erlV)rate and strongly sculptured. Color a dull 
white. Whorls 8 and moderately convex. Aperture holostoma- 
tous and alta<'hed to the body whorl. Parietal and palatal lips 
reflected. Columella short and slightly ohliiiue. Suture im- 
pressed. Sculjiture consisting of numerous axial costae with 
exceedingly fine spiral threads Ix'tween the costae. Operculum 
unknown. 

Height Width 

7.6 nnn. 2.0 mm. Holotype 



4 BREVIORA No. 173 

Types. The holotype is in the Museum of Comparative Zool- 
ojiy. IK). 2.'}()505, from Coloni;) Ramfis, 20 km. W of San Cristo- 
bal, Kepubiiea Dominieana, eolleeted by Dr. Juan Kivero in 
March 1957. 

Remarks. This is the second known species of (ieomelania 
fr(»m Hispaniola. It differs from Cioiiu lania haitiensis Wein- 
land in being twice as large and in having the axial sculpture 
coarser and by having 8 whorls, (i. haitiensis having only 5. 
Jn relationship, it appears nearest to Geomelania elonyata 
Pfeiffer from (Ji'iente, Cuba, and differs from it by being a 
little larger, having one more whoi'l and in being a little more 
coars(dy sculptured. 

SACDIDAE 

Zapiivsema randf, new species 
Plate 1, figure 1 

Description. Shell large, reaching ()7 unn. in height, smooth, 
rather thin and globose. Color i)robal)ly a yellowish brown. 
Whorls -1' •> and strongly convex. Outei- ]i]i simple, jmrietal 
wall with a thin glaze. Suture slightly indented. Columella 
long, sinuous and somewhat oblique. Sculpture consisting of very 
fine axial growth lines. 

Height Width 

65.0 mm. 67.0 nun. Holotype 

67.0 mm. 67.5 nnn. Paratype 

Typ( s. The holotype is in the Museum of Com])arative Zool- 
ogy, no. 230508, from Cueva de San Fi-ancisco, Cerros de San 
Francisco, Mun. Pedro Santana, San Rafael, Republica Domini- 
eana. Collected by Stanley Rand and Clayton Ray, Augast 1958. 
There is single paratype from the same locality. 

Renunl.s. This species is placed pi-ovisionally in the genus 
ZapJiyseind, a genus heretofore known only from Jamaica and 
Xavassa islands. It is about three times as large as the lai-gest 
Jamaican species, Zaptnisrnid niacniurrayi C. B. Adams. Both 
specimens were dead when found in tlu' cave and appear to be 
((uite old as they are white and completely devoid of perios- 
tracum. 

This s])ecies is as large as the largest PoUidontes r/if/antea 
Scapoli from llisjiainola and Zaelu'i/sia petitiana d'Orbigny 
from the Triindad Mountains of Cuba. Its large size and thinner 
shell makes it moi'e capacious tlian eithei- of these two species. 



1962 LAND MOLU'SKS FROM HISPANIOLA 5 

This is by far the largest species in the family Sagdidae, a 
family which occurs in the West Indies, southern Ignited States,^ 
and south through Central America and northern South Amer- 
ica. Its greatest generic development is centered in the island 
of Jamaica. 

REFERENCES 

Bartsch, Paul 

1946. The operculate Iniul luolliisks of the family Aiinulaiiidae of 
the Ishxiul of Hispaniohi and the Bahama Archipelago. Bull. 
U.S. Nat. Mus. 192: 1-3. 
Crosse, H. 

1891. Fauiie nialac-ologi(|iu' tenestre et fluviatile de I'lle de Saint 
Dominge. Jour. Coneiiyl. 39: (59-95. 
Turner, R. D. 

1960. Land shells of Xavassa Island. Bull. Mus. Comj). Zool., 122: 
233-2-14. 



lA single jrenns. Microphusiila. with two spcci^-s. iiigersolli lihuid and cookci 
Pilsbry extends mirth Ihrougli tlic K(iel<y Mountain states to southern British 
("ohiinliia. 



PLATE 

Fig. 1. Zapln/soiut randi Clench. Cuc-va de Saii Fiaiu'isco, Mun. Pedro 
Santana, Republica Doiiiinicana (aliout natural ,si/.e). 

Fig. 12. IlcUcina juliae Clench. Colonia Ranifis, '20 km. W of 8an 
Cristobal, Republica Dominicana (7.4X). 

Fig. 3. Proserpi}ia marcanoi Clench. Colonia Hanilis, 20 km. W of San 
Cristobal, Repnl)lica Dominicana (13.8X). 

Fig. 4. (icomclania (Merrilliana } rircroi Clencli. Colonia Ranifis, 
20 km. W of San Cristolial, Republica Dominicana (about 9X). 



BREVIORA 

Miaseiiaoi of Co;mparative Zoology 

CAMHKllMiE, ;M.\SS. I)K( KMHKK 24, 1 9G2 XUMBKR 174 

A NEW AUCTOCVOXll) FKO.M TilE J'A1.P]()('EXE 

OF WYOMING 

By Bryan rattci-son and Pan! <). MeOrew ^ 

Since 1959. the Department of (xeoloo-y of* the University of 
Wyoming and the ]\Inseum of Comparative Zoology, Harvard 
University, have been carrying on a joint program of field and 
laboratory work in the later iMesozoic and earlier Cenozoic verte- 
brate-bearing formations of Wyoming, paying special attention 
to those deposits amenable to washing techniques. Much of the 
work accomplished thns far has been done in the Shotgun member 
(Keefer, 19(il) of the Fort Union formation in the northern part 
of the AVind River Basin. 

Within this member is a bone Ix'd. some twelve to twenty-four 
inches in thickness, that is extraordinarily rich in vertebrate re- 
mains, mostly single teeth of mammals, crocodiles and small 
sharks. The bed occurs at the base of a sandstone (unit 21) of 
Reefer's section at Twin Buttes ) 217 feet above the base of the 
Shotgun member, which is 1265 feet in thickness at this locality. 
Keefer has shown tliat ov(M-lying the lower Fort I'nion in this 
region are two essentially contemporaneous, interfingering rock 
units, the lacustrine (or marine!) AValtman shale and the mai-- 
ginal Shotgun member. The mammalian fauna from the bone l)ed 
in the Shotgun is a rich one. Fi-oni Keefer 's sample collection, 
Gazin ( //( Keefer. 1961) has recoi-ded : J'tilodiis. Miiiu todoit, Ec- 
typodus. A)tc(ni')(Ii)ii. I-J iicosiiiadoii .\ ('fit()ps(dts, P( y(id( vii s .\ 
Gelastops, Biacodon^, ])0ssibly Aphro)ion(.s, I'cntacodon, jiossibly 
Zfnuicfn-is. cf. I'ronoiho'h cii s. CUu iiodoti. Trier uffs. J'rriptii- 
chus. Anisoiichits. Pro}ni(Jcl(i( iiKs, Liiomxihis?, (ridl( ifiiKi and 
PantoJanibda. Our extensive collections will add a number of 
forms as the work of identification pi-oceeds. The Shotgun mem- 
bei- at Twin Buttes has vielded very little in the way of vertebrate 
material outside of the l)one bed.- Thus fai- we have found only a 

1 l)cp;irliiiri]i of (it'olofry. riiivcrsiiy <>( \V.\ uniiiii;. 

2 During the 1{»61 season a concentration similar to Imt less ricli lliaii tliat 
of the bone befl was found some 00 feet low<'r in the member. 



2 BREVIORA No. 174 

few frafinients of a medium-sized pantodont. We propose that the 
assemblage be known as the Shotgun local fauna, from the name 
of the member in which it occurs. The age, so far as can be deter- 
mined at present, appears to be early Titfanian. At Shotgun 
Butte, the type locality, where the member has a thickness of 
2,830 feet (Keefer, 1961), l'li< nacodHs s]). and Plesiadapis sp. cf. 
P. cookci, have been found in the uj)per i)art of the sequence 
(Keefer and Troyer, 19r)6). 

We are greatly obliged to l)v. William R. Keefer for his help 
in the field. For the opportunity to examine comparative material 
we are indebted to Dr. C. Lewis Gazin, U. S. National Museum, 
Dr. Malcolm ('. McKenna, The American Museum of Natural His- 
tory, and Dr. Craig C. Black, Carnegie Museum. During the field 
season of 1960 we were assisted by James A. Jensen, Lee A. 
Wooder.son, John Zamecnik, Floyd Andrews, Clyde T. Williams. 
Richard P. Timmerme.yer, Charles P. Lyman, Jr., and Robert F. 
Wallin. The ]ihotographs arc the work of Miss Linda Loring. The 
following aI)l)reviations are used: A. M.N. II., American ^luseum 
of Natural History; M.C.Z., Museum of Comparative Zoology: 
I'.AV.. Fniversity of Wyoming. 

AHCTOCVONIDAE 1 .Muiiay 

OXYCLAENINAE Matthew 

OOLPOCLAEX rs -gen. nov. 

Ttjpc sijccies: C. keeferi sp. nov. 

Known distribution : Paleocene, vavW Titfanian, Wyoming. 

I)i(i(jnosis: Enamel of molars strongly wrinkled, all crests cren- 
ulated, accessory cuspules numerous. Prine-ijjal cusjis of upper 
.\[ high, massive, tightly grouped; central basin small; protocone 
nearly central in position with long lingual slojx' ; conules large, 
blunt, separated by grooves from principal cus]is; hypocone small 
on ^P-, i-udimentary on 'SI'': cingnla sti'ong, not continnoiis 
around jtrotocones ; external cingulum intcrruptetl by labial con- 
linuation of sharp cleft between paracone and metaconc; ni)per 
M wide relative to length. Trigonids of lowei- M high, short ; i)ai-a- 
conid internal in position, closely ap])ressed to metaconid, i)ai-a- 
conid crest well dcx'cloped : postci'ior ci'cst connecting protoconid 

1 Out- <(1' us (li.I'.l proiPdscs to liarislCr tlic A rcl myiniiil.-ii' Irniii the ( 'rciKlonlM 
to the Coiid.vliirthni. 'I'liis traiist'cr .iiid tiic \:jiiois (|iicsli(iiis tliiit it raises wiU l(c 
discussed in ;i later jiaiier. 

-Kidpcis, a fold and cluviius; in alliisinii to I lie \vrinl<le(i en.iinej of tlie molars 
anil, in particular, to the curiousl.v folded trij,'onids. 



i''>>- XKW i'Ai,i;()(i:\i-: AKc'rocvnxii) ;; 

and metaeonid present: protoeonids and metaeonids with slioii, 
centrally sitnated erests rnnning lino-nally and labially. respee- 
tively. forming a third, transverse trigonid crest : deejjest jxjrtion 
of talonid basin near lingnal side : hypoconnlid of ^I.-j long, broad, 
higli. cuspidate. 

COLPOCLAKXrS KEEFERI ' s]l. nov. 

Type: M.G.Z. no. 8355, LM.,. 

Hypodigm: Type and F.W. nos. li)31. LM... ; ll»3l2. KM.-., ; 1933, 
LMi; 1934. RM3; 1935. L.M^. : :\I.r.Z. nos.' 8356. KM,, "(much 
worm ; 8357. LM2 ; 8358. ini.. 

Horizon and locality: Shotgun local fauna. Shotgun member of 
the Fort Union formation; XEI.4 SEf^., sec. 31. T. (i X.. R. 3 E.. 
% mile SW of the more iiortlici-ly of the Twin Rnttes. Fremont 
Comity. AVyoming. 

J)in(/)iosis : Sole known species of th(^ genns diagnosed above. 

1) F SCRIPT lOX 

Colpoclaenus kcrferi is notable for the massive, high, princij^al 
cusps of the upper molars, the elevated trigonids and the extreme- 
ly rugose enamel. The latter feature disappears after very little 
wear, as is demonstrated by a moderately worn M2 (M.C.Z. no. 
8358). There can be. we believe, no doubt that the various lower 
molars represent the same species, and their association with the 
n]i])ers would appear to be demonstrated by excellent occlusion 
between .AI.C.Z. no. 8357, LM^, and U.W. no. 1935, L.M.. as well 
as by the general structure. 

The upper molars are considerably wider than long. ^V - being 
e.ssentially quadrangular, ^l^ suboval in outline. The i)rotocone 
is the largest of the three principal cusps; its apex is a little lin- 
gual to the center of each tooth of the series, and its lingual face 
is very long and gently sloping, rather shorter in ]\F" than in 
M^"-. The paracone is more labial in position than the metaeone 
and is slightly larger and higher, being approximately the same 
height as the protocone. Paracone and metaeone are grooved on 
their basinward faces and are separated by a well marked cleft. 
The large blunt proto- and metaconules are separated from the 
primary cusps by grooves and are united by crests to the anterior 
and posterior cingulum respectively; the protoconule is smallei- 



1 For Dr. W. R. Keefer who discovered the Shotgun bone bed in the (■llur^se 
(if his worl; on the Fort Union of the Winn River Basin. 



BREVIORA 



No. 174 




Figure 1. Colpoclaenus keeferi gen. et. sp. nov. Upper molars: a, LM-, 
M.C.Z. no. 8357; fe, Li\[l, U.W. no. 1933; c, EM^, U.W. no. 1934. X3. 
Stereoscopic views. 



1962 NEW PALEOCENE ARCTOCYONID .") 

than the metaconnle on ^l^-, al)ont eqnal to it in size on "Sl'^. The 
ratiier shallow basin enclosed by these cusi)s and conules is situ- 
ated in the labial half of the tooth and bears various i)Oorly de- 
fined cuspules on its floor. A hypoeone is differentiated on Mi"-, 
on which it is considerably smaller than the coiiulcs. I)iit is bai-ely 
distinguishable on M^. Apart from an interruption on the exter- 
nal face, where the cleft Ix'tween paraeone and metacone con- 
tinues on to the labial margin, a strong', crenulated cinguluiii runs 
from the hypoeone around the tooth to the antero-internal corner, 
at Avhicli point the cingulum is broken in AP. M- shows no j)i'()to- 
stylar enlargement here and M"^ a very slight one. With the ex- 
ception of a slight rise in the cingulum at the parastylar site on 
^r"^, there is no indication of external styles. Anterior and pos- 
terior cingula extend farther lingually on M^ than on the jjreced- 
ing teeth and the two are almost united by a median lingual 
cuspule. Several cuspiiles are present on the posterior face of 
M^ above the cingulum. 

^Fo is slightly constricted at the junction of trigonid and talo- 
nid but is otherwise nearly quadrangular in outline ; the trigonid 
and talonid are approximately e(iual in length and width. The 
trigonid is somewhat higher than the talonid and narrows apically. 
The protoeonid is a large massive cusp that forms nearly half of 
the trigonid. The heavy, cuspidate paraconid crest runs from the 
apex of the protoeonid to the lingual side of the tooth, where the 
paraconid is poorly ditferentiated. The blunt metaconid is but 
little smaller than the protoeonid and equal to it in height ; the 
apices of the two cusps are connected by a papillate crest that is 
more lightly built than the paraconid crest. Blunt, Avrinkled 
crests run directly lingually and labially down the facing slopes 
of the protoeonid and metaconid. These al)ut (U.W. no. l!):^.')) or 
fuse (M.C.Z. no. 8358 ) to form a third, central transverse crest. 
A sinuous, antero-posterior cleft partially separates the trigonid 
apex into labial and lingual halves. All this complicated structure 
is shallow and would rapidly be ol)literated by weai-. The pos- 
terior face of the trigonid is sloping and bears various small, 
papillate crests; that behind the metaconid is the most prominent 
but there is no trace on it of a metastylid. The very large, blunt 
hypoconid nearly equals the protoeonid in size and makes up 
nearly half of the talonid. The cuspidate crista obliqua is low 
and fades away toward tlie base of tlie talonid. Two minor crests 
run forward from it to the trigonid base, isolating a vei-y sitiall 
fossette. The hypoconulid is low but relatively large, and in the 



BREVIORA 



No. 174 




Figin'o 1". Colpocliit mis l-crfrri gen. et .sp. iiov. ^li 
h. M.C.Z. no. S3;")8. X'^. Stt'rt'oscu]ji(' views. 



I. r.W. 710. 193; 



unw()i-ii r.W. no. 19.'^.") is ti-iciispidatc and set off fi-oiii tlic .'ulja- 
(M'lit (Mis|)s by sliallow <iTO()V('s. As is shoAvn by ^I.C'.Z. no. )^'.\~)H. 
it is ()l)literat?d as a distinct ensp after only a little wear. Tlif 
ciitoconid is hioher hut scarceh' larg'er than the hypocoiiulid and. 
like it, at least in T.W. no. If).')."), is ti'iensjiidate. From it a ens])i- 
date ridge runs down to tlie base of tlie trio-onid. The talonid 
l)asin is dee])est in its anterolin^'nal ])ortion, and tlie floor beai's 
a nunilier of poorly defined euspnles. A strong-. ])apillate eiugu- 
hiiii i-nns from a nnint beneath the pai-aeonid around the labial 
side of the tootli to the hypoeonulid ; in I'.W. no. If):^."), but not in 



19G2 



NE\V l'Al,i;()(i:XE ARCTOCYONID 




Figure 3. Colpoelaenu^ keeferi gen. et sp. nov. M;? : a, V.W. iki. 1931; 
b, M.r.Z. no. 8355, type; c, U.W. no. 1932. X3. Stereoscopic views. 



8 BREVIORA Xo. 174 

M.C.Z. no. 8358, tliis rises to a euspule in the slight notch between 
trigonid and talonid. 

The third molar is slightly narrower and rather longer than the 
second; it tapers bluntly toward the rear and, as in Mo, trigonid 
and talonid widths are approximately equal. The trigonid is 
shorter and conspicuousl.v higher than the talonid, notably more 
so than in Mo, tapers ajiically as on that tooth and is slightly 
oblique to the long axis, being longer on the lingual than on the 
labial side. The crown of the trigonid resembles that of Mo in 
general, differing in the lesser development of the central trans- 
verse crest, which results in the fonnation of a shallow basin. The 
posterior face is nearly vertical, with wrinkled enamel. The hypo- 
conid is the largest cusp of the tooth ; it is low, blunt, elongate and 
bears a crest that is bowed laterally. The crista obliqua is lacking 
in U.W. no. 1931 and is represented only l)y vestigial, incomplete 
crestlets in the type and T.W. no. 1932; the hypoconid crest con- 
tinues on to the base of the protoconid. The broad, flat-crowned 
hypoconulid is the highest of the talonid cusps and is almost as 
large as the hypoconid. It is set off by shallow grooves and is itself 
varyingly grooved and cuspidate. The entoconid resembles that 
of M2 both in size and in structure. As on M2, the talonid basin 
is deepest antero-lingually and its floor is vaguely cuspidate ; it is 
open anterior to the entoconid due to the absence of a crest on the 
posterior slope of the metaeonid, which on the preceding tooth, 
together with the entoconid crest, forms a slight dam at this point. 
The external cingulum runs from the anterior face of the trigonid 
to the hypoconulid, showing some tendency toward formation of a 
cuspule between hj'poconid and hypoconulid. The tooth is very 
slightly constricted at the junctions of the trigonid and talonid 
and of the hypoconulid with the adjacent cusps. 

DISCUSSION 

We have been able to compare the material referred to Cul- 
poclaenus keeferi with specimens representing most of the de- 
scribed arctocyonids. Colpoclaenus is a very distinct genus; the 
combination of characters it presents — summed up in the diag- 
nosis — separates it sharply from all other forms, although cer- 
tain of them do approach it in one character or another. Anacodon 
resembles it in having accessory cuspules and strongly wrinkled 
enamel, and also in the jiossession of three transverse crests on the 
trigonid. as sliowu hv an uiiwoi'u Mo of .1. ursidens (A.M.N.H. 



1962 NEW PALEOCENE ARCTOCYONin 9 

no. 92). Clacnodon has moderately wrinkled eiuiinel and com- 
parable crests on the molar tri<i-oiiids (Gaziii, lll.ltj, pi. 7, fig. 5). 
Other forms — Thrypfacodoii, Triccntcs, Mimofriccnt< s, Para- 
doxodon — have the enamel wrinkled to varying degrees but do 
not otherwise resemble Colpoclaotus. In Protogonodon the proto- 
cone has a fairly long lingual slope. The trigonid is high in Oxij- 
claenus, CJfricwus, Spannxiiodon, Profhri/ptacodoH, and Metachri- 
acHs. There is a tendency in some of these, especially in Prothryp- 
tacodon, for the trigonid height to decrease from ^l^ to M^.. ; in 
Colpochu tins the trigonid is higher on Mo than on Mo. Colpo- 
claenus thus ])resents a curious combination of primitive charac- 
ters, such as high principal cusps in the upper molars, high trigo- 
nids in the lowers, and advanced ones, such as a degree of wrink- 
ling of the enamel and proliferation of accessory cuspules that is 
exceeded only in Anacodon. As the family is currently sub- 
divided, Colpoclaeniis must be placed in the Oxyclaeninae, l)ut its 
acquisition of arctoeyonine-like wrinkling perhaps gives added 
point to Simpson's suggestion (1937, p. 172) that the Arctocyo- 
ninae may be a partially artificial assemblage composed of several 
lines that were independently following similar adaptive trends. 

The last low'er molars of Colpoclaenus are quite primate-like in 
general appearance and in some structural details. When our 
knowledge of the animal was confined to these teeth and to the 
slightly worn ^Mo we were persuaded that we were dealing with a 
primate astonishingly large for the Paleocene"^. Recognition of 
the upper molars rendered such an identification less likely and 
the finding of the unworn Mo seems to us to have ended all pos- 
sibility of it. We wryly sympathize with Cope and with Osborn 
and Earle who also at times believed an oxyclaenine (Chriacus) 
to be a jiriniate. 

Measurements in mm. 







I" 


.W. no. 19:j.3 
Ml 


M.C 


.Z. no. s:;."ii 
M2 


r.w 


. no. 
M3 


1934 




L 




7.1 




8.0 




6.4 






W 




11. i; 




— 




8.8 






U.W. 11(1. 
193.-I 




.M.C.Z. 
no. 83.58 


Type, M.C.Z. 
no. 83.55 


V.W. no. 
1931 


L'.W. no. 
1932 


M.C.Z. 
no. 8356 




M-j 




M2 


M:; 


M.-i 


Ms 




M:-. 


L 


8.8 




9.1 


11.2 


10.5 


lO.-l 




10.9 


W 


7.0 




7.(') 


6.4 


6..3 


6.2 




6.3 



1 UnfortunatPlv this enthnsiasm fonnd Hfctinjr expression on page 9 of the 
Society of Vertebrate Paleontology News Bulletin No. 61, February, 1961. 



10 BREVIORA X(». 174 

BEFERENCES 

Gazix, C. L. 

1956. Paleoc-eiK' luiinuualian faunas ui the Bison Basin in soutli-eentral 
Wyoming. Smithson. Misc. Coll., 131, No. 6: i-iv, 1-57. 
Keefkk, W. B. 

lOiil. The Waltman shale and Shotgun members of the Fort Union 
formation (Paleocene) in the Wind Eiver Basin, Wyoming. Bull. 
Amer. Assoc. Pet. Geol., 45: 1310-1323. 
Keefee, W. R. and M. L. Troyer 

195r). 8tratigra])hy of the Upper Cretaceous and Lower Tertiary rocks 
of the Shotgun Butte area, Fremont County, Wyoming. U.S. 
Geol. Surv., Chart OC 56, Oil and Gas Inv. Ser. 
Simpson, G. G. 

1937. The Fort Union of the Crazy Mountain field, ^Fontana, and its 
mammalian faunas. Bull. U.S. Nat. Mns., 169: i-x, 1-287. 



BREVIORA 

Mmseuiinii of Coimparative Zoology 

Camhkidgp:, Mass. 1)k( kmhkk l24, 19(i2 Xt\mhkr 1 7o 

A PICRODOXTID IXSErTlVOKE( .') FROM TTTE 
PALEOCEXE OF WYOIMTXa 

By Paul O. McGrew ^ 

and 

Bryan Patterson 



Among: the interesting- materials fonnd in the course of recent 
field work in early Tertiary deposits in Wyoming, conducted 
jointly by the Fuiversity of Wyoming- and the ^luseum of Com- 
parative Zoology, are several fragments of maxillae and man- 
dibles and a number of isolated teeth of a picrodontid. Only two 
other records of these problematical little mammals are known: 
Picrodiis silherliugl Douglass 1908 from the Fort Union (Lebo) 
of Montana and Zanijcteris paleocena Matthew 1917 from the 
Tiffany of Colorado, the former based on lower, the latter on 
upper teeth. We confidently assume that our upper and lower 
molars are referable to the same species. On the evidence of the 
upper molars this species cannot be ])laced in Za)iycteris and 
on that of the lowers it cannot be excluded from Picrodiis. We 
suspect it to be new. but the differences from, and the parts in 
common with, P. silbcrlingi are so few tliat we refrain for the 
present from any attempt at diagnosis. Gazin's tentative record 
of Zanijcteris from the Shotgun member of the Fort Union for- 
mation in the Wind River Basin (19(il, p. .11. and //; Keefer, 
1961) is almost surely based on this form. 

The acknowledgments made in a previous paper (Patterson 
and McGrew. 1962) apply equally to this one, except that the 
photographs printed here were taken by Mr. John F. Cutler. 
Abbreviations are as follows: U.W., University of Wyoming; 
M.C.Z., Museum of Comparative Zoology. 

1 Department of Geology. University of Wyoming. 



2 BREVIOBA No. 17.") 

INSECTIA^()RA( ?) 

PICRODOXTIDAE Simpson 
PiCRODUS Drnio-lass 

i'lCRODUS SI). C'l". P. SlLiSEKLlNGI DoU^lass 

Material: U.W. no. 1780, fragment of left maxilla with M^ -. 
incomplete alveolns of M-'^; M.C.Z. no. 8363, fragment of rig-ht 
maxilla with M^ incomplete alveolns of M- ; r.AV. no. 1781. 
fonr complete and three partial ]\Ii ; M.C.Z. no. S422, tlirec M, : 
M.C.Z. no. 8423, ^U- 

Horizon and localities: Paleocene, early Tiffanian. U.W. no. 
1780 is from the Bison Basin Saddle locality. Fort T^nion for- 
mation, El o, sec. 28, T. 27 X., R. 95 \V., Fremont Co., Wyoming 
(Bell, MS.; Gazin, 195(3). All the other specimens are from the 
Shotgnn local fanna, Shotgnn member. Fort Union formation, 
% mile SW of the more northerly of the Twin Bnttes in the 
northern part of the Wind River Basin, NE I4, SE i^., sec. 31, 
T. 6 N., R. 3 E., Fremont Co.. Wyoming (Keefer, 1961"; Patter- 
son and McGrew, 1963). 

All specimens were enconnteretl in the course of washing 
operations. 

DESCRIPTION 

The ni)i)ei- molars of the Wyoming foi-m agi'cc witli those of 
Zanyctcris paleocena (Matthew, 1917; Simi)son, 1935) in basic 
structure yet diffei- considerably in detail, especially as regards 
M^. This tooth is nearly ((uadrangnlar, rather than triangular, 
in outline, an outpocketing of the posterior cingnJum forming 
a postero-internal angle. The protocone is more antei-ioi- in ])osi- 
tion and more directly internal to the paracone. The protoloph, 
within which the protocone is completely subordinated, is a 
heavy, transversely aligned crest that extends almost to the in- 
ternal base of the paracone, from which it is separated l)y a 
shalloAv notch. The anterior face of the tooth slopes gently n|)- 
ward; the anterior eingulum (double in l^.W. no. 1780) is 
strong compared with that of Zanijcteris. The antero-external 
corner of the tooth is rounded and bears no stylar pi-ojection. 
A faint, irregular crest extends, with interruptions, from the 
base of the protocone to the metacone. It is situated almost 
wholly within the central basin, of which the lingual and postero- 
lingual margins are formed ])y the eingulum. The enamel within 



196: 



A PALEOCENE PICRODONTID 







Fig. 1. Picmdu.s .sp. cf. P. silherlinf/i Douglass, a, U.W. no. 1780, frag- 
ment of left maxilla with M' -. h, M.C.Z. no. 8363, fragment of right max- 
illa with M^. Approximately X 8. Stereoscopic views. 



4 BREVIORA Xo. 175 

tlic (iroa pneiosod by the ciii^ulnni and 1li(» aforemontioned low 
e-rest is fully <\s wrinkled as that over the main iiortion of the 
central basin. Picrodns sp. is rather nioi'e advanced than Z. 
paleocena in the essentially coni])lete incorporation of this cin- 
gular area into the basin and in the presence of the small postero- 
internal outpocketing already mentioned/ Paracone and nieta- 
cone are mere elevations at tlie ends of the lono:, nearly straight 
ectoloph. An external cingulnm is ])resent posterior to the iiai-a- 
cone, where it forms a stout, rather wide and featureless shelf 
that passes back into the postero-externally projecting meta- 
stylar area; a crest connects metastyle and nictacone. 

M- is (juite similar to the correspon(lin<>' tooth of Z . paleocena, 
differino- only in a few points. The tooth is slightly smaller rela- 
tive to M^ the postero-internal angle is sonunvhat more scinared, 
and the antero-external style is i-athei- more ])i-ominent and con- 
nected by a low inconspicuous crest to the tip of the paracone. 
The adjacent styles of AP- abut to form an apex fi-oiii which 
the outer margin of the cheek tooth series falls aw;iy anteriorly 
and posteriorly. 

The tirst lower molar diffei's only in detail from Ihat of /'. 
sill)erli)\(ji. The ])araconid is usually higher and slightly more 
independent and is invariably larger than the metaconid. Tn 
several teeth the latter cusp has almost or (piite lost its identity 
in the oblique ti-igoiud crest. Ti-igonid and talonid are con- 
sistently better ch'mareated externally. The crest that forms 
the external border of the talonid cui-ves inward at its anterior 
extremity to reach the center of the [jostcrior face of the trigonid 
a short distance below the protoconid rathei- than abutting 
against the base of the trigonid externally, as is the rule in P. 
silberlin(/i. A narrow, shallow gi-oove lies between the trigonid 
face and the incurving position of the crest. The main iX)rtion 
of the talonid crest, that forming the lateral and posterior 
borders of the basin, appears to be more cuspidate than in the 
Lebo sample; the cuspules number from four to idne rather than 
from two to three. The two cuspules on the lingual side of the 
talonid are extremely variable in the Shotgun sample, amount- 
ing in some specimens to little more than irregularities on the 
talonid margin. An interradicular crest is present. Mo resembles 
the corresponding tooth of P. silb( rlliu/i more closely than does 
-M, : Simpson's description (1937, ]). 138) applies practically 
verbatim to our single specimen. 

1 The iiostHi-ior ciii;^!!]!!!)! of >[i of Zaiii/ctiri.s iialeoccna is not as distinct from 
llif ilcntriil Icisiii lis Siin|is<iirs liiiiii-c (I'.i;;."). lin. (i I wmild siiu'^X'st. 



1962 



A PALEOCENK [MCKODOX-ril) 




Fig. L'. Picroduf: sp. cf. P. .silhcrlini/i Douglass. </. T'.W. mi. 1 7^1, "RAfi, 
approximately X 20. h, M.V.7.. no. 8422, mfi, apinoxiiiiatcly X 11. c, 
M.C.Z. no. 842;?, RM2, approximately X 11. Stereoscopic- views. 



6 BRBVIORA No. 175 

111 both upper and lower molars, the enamel at the base of the 
crown extends out far beyond the necks of the teeth, the external 
and posterior eiiigula of AD and the postero-external portion of 
the talonid of Mj being* especially projecting. The term exoedae- 
nodont has been applied to this condition, which, in the Insec- 
tivora, is encountered especially among the Dimylidae (Hlirzeler, 
194-1:). Iliirzeler and also Raban (1958, j). 896 u) believe that 
exoedaenodonty is an indication of a malacophagous diet, Saban 
citing in sup])ort the luimerous shells of Anodu)ita that occur 
in deposits that yield remains of Dimylinae. However this may 
be for diniylids witli their strongly ciisped molars, we do not 
believe that it indicates any such habit for the picrodontids.^ 
As Matthew recognized in liis description of Zanycteris, low 
crowned more or less flattened molars with finely wrinkled 
enamel of the sort that occur in the family almost certainl}' in- 
dicate a frugivorous diet. 

The small fragments of maxillae and mandibles add nothing 
to knowledge excei)t to suggest that the anterior root of the 
zvgoiiui m;iv not have been as stout as in Zanjirffris. 



MEASUREMENTS IN MM. 



Length M^ 
Width M' 
Lenoth M- 
Width M~ 



Oblique niaxiimim 

dianu'tor- M] 7 

Width talonid Mi in 



U. W. 17 


so 


M.C.z. 8;?6:i 


2.85 




2.95 


2.35 




2.4U 


1.35 







1.63 







OK >I 



Jjength Ml 
Width Mi; 



2. -10  


-2.85 2.6G ±.051 


.135 ±.03(5 


5.07 ±1.35 


1.00 


-1.40 I.IG ±.038(3 

M.C.Z. 8423 

1.90 
i.lo 


.122 ±.027 


10.5 ±2.34 



1 Nor. a fortiori, for such forms a.s soriculs and various inicrochiropterans that 
are exoedafuoiloiit to varyin.^ d»';rre('S. 
- As measured liy .Simpsou ( l!»,'i7. p. 13S). 



1962 A PALEOCENE PICRODONTID 



DISCUSSION 



JMatthew, in his description of Zanycteris paleocena, took no 
account of Picrodus silh€)-Ii)tyi. and indeed there was nothing- in 
Douo'lass' description and figures to invite comparison between 
tlie two forms. It remained for Simpson, with new material of 
P. silhcrlingi and tir.st-kaml knowh^dge of Z. paleocena, to show 
that they were closely related. As he pointed out (1937, p. 136), 
the species were distinct, l)ut in the absence of comparable parts 
it could not certainly l)e stated that this was also true of the 
genera. We now have lower molars indistinguishable generically 
from Picrodus found with upper molars clearly distinct from 
those of Zanycteris. On the face of it, then, the two previously 
described species would appear to belong to distinct genera. 
Howevei' i)robable, this cannot yet be considered as certain. The 
highly specialized lower molars may i)erhaps prove to be rather 
stereotyped within the family and hence unreliable for generic 
discrimination. Although quite unlikely, it is thus still conceiv- 
able that P. .siJberlingi and Z. paleocena may represent one 
genus, and P. sp. another. Finds of associated upper and lower 
teeth of the previously described forms will l)e required before 
all doubt on this score can be set at rest. 

As regards the broader question of picrodontid affinites, we 
are in complete agreement with Simpson (1937) that the family 
cannot be referred to the Chiroptera. The enlarged lower incisor, 
small upper canine, small i)remolars, long and slender muzzle, 
and origin of the anterior root of the zygoma opposite M^'^ are 
definitely non-ehiropteran characters. Since ^latthew's work, 
it has always been stated that the molars resembled those of the 
specialized Phyllostomatidae, members of the subfamilies Stur- 
nirinae, Phyllonycterinae and Stenoderminae. Thanks to the 
excellent collection of bats in the ^luseum of Comparative Zool- 
ogy, we have been able to make comparisons with nearly all 
members of these three groups. AVe are quite unimpressed by 
the resemblances between their molars and those of the picro- 
dontids. Essentially these are limited to wrinkled enamel and 
pointed trigonids, and are more than offset by numerous dif- 
ferences. 

Sturnira and the Phyllonycterinae have smooth enamel. In 
Sturnira, the upper molars have deep, antero-posteriorly run- 
ning central valleys and no traces of lophs; the lowers have the 
trigonid and talonid basins continent, the metaconid widely sep- 
arated from the protoconid, and the trigonid of Mj neither com- 
pressed nor elevated. In the phyllonycterines, the upper molars 



8 BREVIORA No. 175 

are triangular and lack lophs and a stylar area ; the lowers have 
the trig'onids scarcely elevated above the talonids, and all cusps, 
save the prt)toeonid of M^, incorporated in the rims of tlie talo- 
nid basins. The Stenoderniinae have wrinkled, often strongly 
wrinkled, enamel, l)ut the molar structure is very different from 
that of the picrodontids. The upper molars ai'c short in com- 
parison with their widths, 'SI- is large (except in I'ygodcrma), 
frequently larger tluiii AP, and the i)aracones and metaeones 
are high and trenchant (in Pygoderma the paracones only), 
standing well above the flattened lingual portions of the teeth. 
The trigonids are elevated and compressed to points, hut the 
compression does not involve the whole trigonid as it does in the 
picrodontids. The metaconid, when not subordinated in the 
crest running to the apex of the ])r()t()coni(l, is a distinct element 
situated low on the crown. The jiointed ])rot()c()nitls of both 
molars and premolars give every appearance of being involved 
in the same morphogenetic gradient. 

The few resemblances in molar structure between picrodontids 
and specialized j^hyllostonuitids scmmu clearly to ])e of the sort 
brought about by convergenee. The Alicrochiroptera can be elim- 
inated as possible relatives of the Picrodontidae, and we can see 
in the latter nothing suggestive of the JMegachiroptera. On the 
positive side, we can offer no really useful suggestion as to the 
ordinal affinities of the ])icrodontids. Simpson has reniarked 
that "reference to the l*rimates is merely a possibility, with no 
positive evidence to c(unmend it." The possibility, of course, 
exists, and such ])icrodontid characters as the small size of the 
trigonid comjjared with that of the talonid and the tight group- 
ing of the trigonid cusps could be cited as resemblances, how- 
ever vague, to early members of that order. Kesemblances of 
this sort scarcely constitute positive evidence, however. Kefer- 
ence to the Insectivora is e(|ually unsatisfactory. Our (|ueried 
placement of them there is strictly faute de mieux, due largely 
to reluctance to use the Primates as a scrap-basket order. 

REFERENCES 

Bell, W. G. 

MS. The geology of the southeastern flank of tlie Wind River Moim- 
[19.15]. tains, Fremont County, Wyoming. Pp. 1-204. On deposit in 
the library of tlie Deiiartment of Geology, The University of 
Wyoming. 



1962 A PALEOCEXE I'K IK iD' )X'1II) 9 

(;\zix, C. T,. 

1SI')I'. I'ak'ocoiU' ii!;iii:iii;ili;iii f';iiii:;is of tlio Bison Basin in south t-putral 

Wyoming. Sniitlison. Misc. Coll., 131, no. 6: i-iv, 1 ;")". 
lIKil. Occiiirences of Palcocene Maniiiialia in Tertiary basins of Wy- 

(imiiit;. Wyo. Geol. Assoc. KUli Ann. Field Conf. 47-52. 

1! i'liZKLKK, .1. 

11144. lieitriige ziir Kcnntnis dcr Oimylidae. Alili. S<di\vei/'.. I'al. Ges., 

65: 1-44. 
Kekfek, W. R. 

19()1. The Waltnian shale aiiil Shotgun inenibers of the Fort Union 

formation (Paleocene) in the Wind River Ba.sin, Wyoming. 

Bull. Anier. Assoc. Pet. Geol., 45: 1310-1323. 

M.VTTHKW, W. I). 

1917. A Paleocene hat. Bull. Amer. Mu.s. Nat. Hist., 37: .569-571. 
Patterson, B. and P. (). McCtRew 

19(il2. A new aretoeyonid from the Paleocene of Wyoming. Breviora, 
Mus. ("omp. Zool., Xo. 174: 1-10. 
Saban-, H. 

1958. Insectivora /7( Traite de I'aleontologie, edited by J. Piveteau, 6, 
vol. 2: 821-909. Paris: Masson et Cie. 

i~^IMPSON'. (i. G. 

19;!."). The Tiffany fauna, uppci- Paleocene. I. — Multituberculata, 
Marsupialia, Insectivora, and ?('hiroptera. Amer. Mus. Novit., 
no. 795: 1-19. 

19;!7. The Fort Union of the Crazy Mountain field, Montana, and its 
manjmalian faunas. Bull. U. S. Xat. Mus., 169: i-x, 1-287. 



BREVIORA 

Museminni of Comparative Zoology 

Cambridge, Mass. December 27, 1962 Number 176 



ON THE RACES OF KINIXYS BELLIANA GRAY 

By R. F. Laurent 
Museum of Comparative Zoology 

When I noticed the peculiarities of Kinixys beUiana from 
northeastern Congo, I suspected that these populations might 
represent a race already described by Riippell under the name 
of sclioensis, but I did not have Riippell's description. Dr. Mer- 
tens kindly gave me the necessary information and it became 
evident that these populations were different from sclioensis, and 
I therefore described the race meriensi (1956). Dr. Mertens {in 
litt.) declared that sclioensis was in his opinion a valid race, but 
he apparently changed his mind, since the rehabilitation of sclio- 
ensis has never appeared (Wermuth and Mertens, 1961). In 
preparation for a checklist of the turtles of Africa, I have now 
reinvestigated the validity of sclioensis and also the range of 
mcrtensi toward the east and northeast. 

Acknowledgments. I wish to thank Miss A. G. C. Grandison 
(British Museum of Natural History) and Mr. A. Opdenbosch 
(Technician in the Musee Royal de I'Afrique Centrale, Tervuren) 
who very kindly gave me invaluable information on specimens in 
their charge. My thanks are due also to Prof. G. G. Simpson for 
advice on statistical matters and to Dr. E. E. Williams for read- 
ing the manuscript and helpful criticisms. This study has been 
supported by National Science Foundation grant NSF G 17144. 

Kinixys belliana Gray 

This paper limits itself to the examination, with the aid of the 
ratios already used in the description of mertensi, of two ques- 
tions : (1) Is sclioensis Riippell recognizable as a northeastern 
race, and (2) does mertensi connect with the typical form by a 
smooth cline in which no objective limits could be formulated 
for the eastern distribution of the race? 



2 BREVIORA No. 176 

Measurements, additional to those taken for the description of 
mertensi (1956) (i.e. on other specimens), were taken on the 
MCZ specimens; Miss A. G. C. Grandison, with her customary 
kindness, sent the pertinent measurements for specimens from 
northeastern Africa and Uganda. Mr. A. Opdenbosch furnished 
also data on a large series of K. helliana from the Congo. 

Admittedly, the method of relying on other persons to take 
measurements has serious drawbacks since it can introduce bias 
in the data. For amphibians the discrepancies are very great 
indeed, but as far as turtles are concerned it has been hoped that 
the measurements are generally so easily definable that these dis- 
crepancies would be almost negligible. However, the comparison 
of the ratios already published (Laurent 1956) with those cal- 
culated on Opdenbosch 's measurements disclosed a considerable 
bias in the breadth of the carapace (maximum breadth, and 
breadth on level of lateral ends of humero-pectoral sutures). 
Even disregarding these data, the variation of 'bclliana is such 
that the validity of schoensis appears untenable on present evi- 
dence. [Of course, other characters may prove some day that 
the northeastern populations or even some others from east or 
southern Africa are subspecifically differentiated.] Thus, in re- 
gard to the four ratios used in my previous work (Laurent 1956) 
the situation is as follows. 

1. From the data at hand in 1956, the general shape of the 
carapace seemed definitely different in nicrtcnsi and "schoensis" 
and helliana. The carapace breadth (at the level of the lateral 
ends of the humero-pectoral suture) in per cent of the maximum 
breadth (back part of the carapace) was 87.2 to 97.8 (m = 92 for 
nine specimens) for helliana, 78.7 to 90.3 (r» = 85.67 for fifteen 
specimens) for mertensi and 83.8 for the type of schoensis. The 
overlap was slight enough to make unnecessary any test for sig- 
nificance of the difference. However, the data now at hand are 
definitely less conclusive. The difference seems to be still statis- 
tically significant between mertensi and the complex of popula- 
tions from eastern and southern Africa which we call helliana 
but which surely cannot be considered as homogeneous. Also, 
because bias is obvious in these rather difficult measurements and 
because other differences are more clearcut, it has not been found 
worthwhile to make any statistical calculations. The specimens 
from Sudan, Abyssinia and Somaliland do not appear to be 
significantly distinct from those from East Africa, though more 



1962 KINIXYS BELLIANA GRAY 3 

consistent measurements taken on more numerous specimens 
could some day disclose a valid difference. 

2. In my 1956 work the ratio between the median gular suture 
plus the median humeral suture and the breadth of the plastron 
at the level of the lateral ends of the humero-pectoral sutures 
was seen to show a very eleareut difference between hclliana and 
mertimsi — the figures being 56.7 to 73.3 (m = 66.87) for helli- 
ana (iV=9), 72.9 to 91 for mertensi (iV=15). The new data are 
ana {N=9), 72.9 to 91 (m = 84.92) for mertensi (iV=15). The 
new data are still reliable as the measurements are not likely to 
suffer from individual bias. The ratios now obtained are as fol- 
lows (expressed in percentage) : 

a) bclliana {N = 87) 49. 0-S2.S jh = 66.45 i; = 12.51 

b) mertensi (N = 22) (57)^ 75-94 »(= 84.18 f = 12.47 

c) "schoensis" (iV = 8) 55.7-71.7 m=Q9.V?, r = 10.80 

d) nogucyi {N = 3) 65.4-73-83.1 

The variation coefficients are high because the samples are 
heterogeneous from the point of view of size and consequently 
of age. In the mertensi-helliana and helliayia-" schoensis" com- 
parison, Student's t is respectively 8.45716 and 8.25106, which 
shows highly significant differences in view of the small inimber 
of specimens. 

3. The ratio between the pectoral suture and the sum of the 
gular and humeral sutures gives the following figures : 

a) helliana (N = 81) 25.6-69 m = 42.01 r = 18.30 

b) mertensi {N = 21) 0-31 in =19 v = 41.51 

e) "schoensis" {N = 8) 31.4-42.3 /;;= 34.50 v= 9.86 
d) nogueyi (N = 3) 27.8-27.8-39.1 

Notwithstanding an enormous variation coefficient in inert ensi, 
t shows a significant difference between mertensi and " schoen- 
sis" {t = 5.32) and mertensi and helliana (t = 12.71) but not 
between helliana and "schoensis." 

4. The ratio between the pectoral and the abdominal sutures 
gives the following figures : 

a) helliana (N = 86) 21.3-60 m = 39.89 i; = 25.81 

b) mertensi {N = 21) 0-31.2 m = 18.71 f = 42.98 

c) "schoensis" (N = 8) 29.4-37.4 tn = 33.5 v= 8.62 

d) nogueyi {N = 3) 22.4-36 

1 The low figure 57 comes from a juvenile. 



4 BREVIORA No. 176 

Only the difference between " scJioensis" and mertensi proves 
to be significant with a ^ of 5.03. 

Two other ratios not previously used but combining the same 
measurements have proven useful. 

1. The ratio between pectoral suture and the maximum width 
of the carapace : 

a) Ulliana {N = 81) 11.3-24 m = 16.51 t> = 18.50 

b) mertensi (N = 21) 0-13.8 m= 8.95 v = 41.87 

c) "schoensis" (N = 7) 11.6-15.3 m = 12.86 v = 13.83 

Student's t is significant between helliana and mertensi (9.48), 
and between "schoensis" and mertensi (6.65), dubiously signifi- 
cant between helliana and "schoensis" (3.11). 

The same relation between 'belliana and "schoensis" becomes, 
however, significant if analyzed by regression lines (t = 4.26). 

2. The ratio between the width of the plastron at level of the 
lateral ends of the humero-peetoral suture and at the abdominal 
suture : 

a) helliana (iV = 86) 110-200% )u = 145.45 i; = 11.30 

b) mertensi {N = 21) 103-147% m = 114.52 t; = 31.54 
Student's t (8.05) is highly significant. 

As a result of these computations, mertensi appears definitely 
valid, but "schoensis" seems to be so only for two ratios: 1. the 
ratio of the sum of humeral and gular sutures to the breadth of 
the plastron at the level of the humero-peetoral suture and 2. the 
regression line for the correlation between the length of the pec- 
toral suture and the maximum breadth of the carapace. 

From a taxonomic point of view, it does not seem advisable 
to revive "schoensis" on such slender evidence — -more espe- 
cially as the helliana sample is geographically highly heteroge- 
neous. A complex elinal variation from Rhodesia to Ethiopia is 
possible, since we have no data from the populations between 
Kenj-a and Ethiopia or Eritrea. Some records from Eritrea 
(Sordelli 1901, Calabresi 1927, Scortecci 1928) disclose a short- 
ening of the pectoral suture, as in mertensi. 

On the other hand, no east-we.st cline exists between mertensi 
and helliana: the British Museum specimens from Entebbe and 
Mount Elgon are clearly mcHensi. 

The two (luestions propounded have thus the following an- 
swers : 

(1) A race "schoensis" cannot now be revived for the north- 
eastern populations of Kinixys helliana. 



1962 KINIXYS BELLIANA GRAY 5 

(2) The distribution of Kinixys helliana mcrtensi includes 
Uganda, but its northern limit remains unknown. 

These conclusions are still provisional. More material could 
prove not only that "schoensis" is valid but that other subspe- 
cies can be recognized, or on the contrary that even mertensi 
merges in helliana through Sudanese and Abyssinian popula- 
tions. 

Material examined 

Kinixys hclliaria helliana Gray 

MUSEUM OF COMPARATIVE ZOOLOGY: Sudan: Torit (1). 
Kenya: Golbanti (1), Ithanga Hills (1), Kibwezi (1), Voi (5). 
Tanganyika: Amboni (1), Kilosa (1), Kiponda to Mitungu (1), 
Kitaya (1), Mikindani (4), Morogoro (1), Simo near Tabora 
(2), Turiani (1), Ujiji (2). Zanzihar Island: Zanzibar (1). 
Nyasaland: Cholo Mtn. (1), Mtimbuka (5). Northern Rhodesia: 
Isoka (1). Southern Rhodesia: Birchenough Bridge (1), Bula- 
wayo (1), Hot Springs (2), Lumani (1), Selinda Mtn. (3), 
Umtali (1). Trawsraai; Naauwpoort (1). Katanga: Kaipiri (1), 
Lukafu (1). 

BRITISH MUSEUM (NATURAL HISTORY) : Sudan: Ka- 
dugli (1). Ethiopia (1), Anseba (2). Somaliland: Berbera (1). 
near Berbera (1). 

MUSEE ROYAL de l'AFRIQUE CENTRALE (Tervuren, 
Belgium). Kenya: Kibwezi (1), Voi (1). Northern Rhodesia: 
Abercorn (3). Katanga: Kahamhaie (l),Kabinda (1), Kakanda 
(3), Kansenia (12), Kapiri (8), Lofoi sources (1), Lukafu (2), 
Lukonzolwa (4), Mwera (1), Ste. Walburge (2). Kivu: Ma- 
kunga (2). 

Kinixys helliana mertensi Laurent 



MUSEUM OF COMPARATIVE ZOOLOGY: Ituri: Mahagi- 
Port (1). 

BRITISH MUSEUM (NATURAL HISTORY) : Ucjanda: En- > 

tebbe (1), Mt. Elgon (2). 

MUSEE ROYAL de l'AFRIQUE CENTRALE (Tervuren, Bel- 
gium). Cow(/o, without locality (4). Uele: Dika, (3),Mauda (1), 
Niangara (1). Ituri (1) : Abimva (1), Gangala na Bodio (3), 
Mahagi-Port (3). Stanleyville District: Avakubi (1). 



6 BREVIORA No. 176 

Kinixys helliana nogueyi Lataste. 

MUSEUM OF COMPARATIVE ZOOLOGY: Dahomey: Bassila 
(1). Togo: Tohoun (1). Sierra Leone: Kabala (1). 

Two specimens from Lukolela (western Congo: M.A.C. 4648)^ 
and from the Kwango District (southwestern Congo : M.A.C. 
10736) suggest that a differentiated population exists in the 
lower Congo region, which somewhat resembles mertcnsi in hav- 
ing a short pectoral suture and a long abdominal suture. Other 
specimens are, of course, needed. 

Key to the races of Kinixys belliana 

A. Forelimb with 4 claws — Range : Western and northern Cameroon, west 
to Senegal K. b. nogueyi Lataste 

B. Forelimb with 5 claws (occasional specimens have 4 claws) 

1 — Eatio between the median gular suture + the median humeral suture 
and the breadth of the plastron at the level of the lateral ends of the 
humero-pectoral sutures: 75 to 94% (less than 75 in juveniles). Ea- 
tio between the pectoral suture and the sum of the gular and hu- 
meral sutures: 0-31% 

Range: Northeastern Congo and Uganda (presumably also Eepub- 
lique Centre Africaine and eastern Cameroon). 
K. b. mertens^i Laurent 

2— These ratios, respectively, 49 to 83% and 25 to 69%. 

Range: Sudan east to Eritrea and Somaliland, south to Natal, north- 
west to Angola and southern Congo. 
K. b. belliana Gray 

BIBLIOGRAPHY 

Calabresi, E. 

1927. Anfibi e rettili raccolti nella Somalia dai Proff. G. Stefanini N. 
Puccioni. Atti. Soc. Ital. Sci. Nat. Milano, 66: 14-60, pi. I. 

Laurent, R. F. 

1956. Contribution a 1 'herpetologie de la region des Grands Lacs de 
I'Afrique Centrale. I. Generalites. II. Cheloniens. III. Ophi- 
diens. Ann. Mus. Congo, Zool., 48: 1-390, pis. I-XXXI. 
SCORTECCI, G. 

1928. Rettili dell 'Eritrea esistenti nelle collezioni del 'Museo Civico 
di Milano. Atti Soc. Ital. Sci. Nat. Milano, 67: 290-339, figs. 
1-8, pis. VII-IX. 

SORDELLI, F. 

1901. Anomalia in una testuggine (Cinixys belliana) del Sudan Orien- 
tale. Atti Soc. Ital. Sci. Nat. Milano, 39: 111-114, figs. 1-2. 
Wermuth H. and R. Mertens 

1961. Schildkroten. Krokodile. Briickenechsen. G. Fischer, Jena. 

1 M.A.C. : Mus6e Royal de I'Afrique Centrale (Tervuren). 



BREVIORA 

Mmseium of Contiparsitive Zoology 

Cambridge, Mass. December 27, 1962 Number 177 



RHIPIDISTIAX CLASSTFICATTOX TX RELATION 
TO THE ORIGIX OF THE TETRAPODS 

By 

Keith S. Thomson 

Department of Biology and Museum of Comparative Zoology, 
Harvard University 

IXTRODUCTIOX 

111 an extensive study of the nasal region of the lower piiatho- 
stomes, Jarvik (19-1:2) eoneluded that the four families of rhipi- 
distiau Crossopterygii represent two distinct stocks (superorders 
according to Lehman, 1959) — the 'Porolepiformes' (Porolep- 
idae and Iloloptyehidae ) and the 'Osteolepiformes' (Osteolep- 
idae and Rhizodontidae). Of major interest to students of 
vertebrate evolution was his conclusion that the structures seen 
in the snout of 'Porolepiformes' were closely comparable to 
those seen in the recent LTrodela, and those of 'Osteolepiformes' 
were comparable to those of the recent Anura, and that here 
was evidence of a biphyletic origin of the tetrapods. His work 
was based upon detailed studies of two genera, PoroJcpis and 
Eu.sthcnoptcron (a rhizodontid). Jarvik 's material was unusual 
in that it showed details of the endocranial part of the snout 
which it is not usually possible to study in fossil crossopterygi- 
ans ; in recent years, however, more such material has been 
described (Vorobjeva, 1959, 1960a, 1960b; Kulcyzcki, 1960) 
and 0rvig (1957) has published a full treatment of the inter- 
relations of the Rhipidistia on the basis of the structure of the 
scales. For the last year or so I have been engaged in a study 
of the ethmoid region of the osteolepids Megalichthys and 
Ectosteorhachis, continuing the work started by Romer (1937, 
1941). 

Briefly, the newly available evidence does not support all of 
Jarvik's original conclusions. Because of the relation of this 



2 BREVIUKA No. 177 

\\(»i'k to the problem of tlic oi-ijiin of tlic tctrapods and particu- 
larlv to the currently disinitcd tlieories of the ancestry of the 
recent Amphibia, I haye decided to publish this short reyiew of 
the more ]iertinent points in adyance of a more thorough treat- 
ment. This may also be useful since the work of Vorobjeya, be- 
ing in Russian, may not be readily ayailable to eyeryone. 

This paper forms })art of the work to be submitted to the De- 
partment of Biology, Haryard Uniyersity, in fulfillment of the 
requirements for the degree of Doctor of Philosojihy ; it is my 
pleasure to acknowledge the constant help and guidance of Pro- 
fessor A. S. Romer during all this study; he and Dr. E. E. Wil- 
liams and Professor B. Patterson read and criticized the manu- 
script. I am also grateful for the friendly encouragement of 
Drs. Jaryik and ^ryig in Stockholm ; the former has allowed 
me to see an as yet unjiublished manuscript on this subject 
(Jaryik, in press). My friend ]\Ir. Simon Karlinsky provide*! 
the translations from the Russian. My studies haye been sup- 
ported by the awai-d of X.A.T.O. Science Studentship 3/60/955 
by H. M. Department of Scientific and Industrial Research. 
London, and by tlie .T(^ffries Wyman Scholarship at Harvard 
Uniyersity. 

MATERIAL 

It is necessary to establish the taxonomir status of certain of 
tlie rhipidistians concerned in this study; these are: 

McgalicIifJii/s Agassiz 1841- (including Ecio^fi orhavhis Cope 
1880). Carboniferous and Lower Permian of Europe and 
North America. 
PlatycephaIicJit]iys Vorobjeva 1959. Upper Devonian of 

U.S.S.R. 
Panderichthys Gross 194U Upper Devonian of U.S.S.R. 
Porolepis {P. ex grege posiKntiensis [Kade 1858]). Lower 

Devonian of Poland. 
Mfgalichthys-Ectosirorhach is. Ecfosfcorhochis was originally 
described by Cope (1880) from material from the Lower Per- 
mian of Texas; it w-as later (1891) referred by him to the Car- 
boniferous genus Megalichihiis. In recent years it has been sug- 
gested several times (see, for example, Romer, 1941) that the 
two genera are, in fact, distinct and that the Lower Permian 
material will have to be referred back to EctostcorJwchis. No 
formal diagnosis of this has been given ; however, in advance of 



1962 RIIIPIDISTIAX KELATI0X8I Ill's 3 

siR-h a diagnosis T shall follow precedent and refci- to tlic Per- 
mian fish, for {•onvenience, as ' EctostcorJiachis.' 

Mcgalichfhys and Eciostrorhachis are generally acknowledged 
to be members of the Osteolepidae. Berg (lf)58) niiites them 
with Owen's Parahatmchn.'i to form the separate family Para- 
batrachidae; however, available evidence, e.g. Bystrow (1950), 
shows that Megalichfhys is closely related to Osicoh pis itself. 

PoroJepis. Knlcyzcki's (1960) material of ForoJi pis is with- 
out doubt correctly referred to that genus. 

Platijcephalichihys and Pandcrichthys. These genera are in- 
volved in a long series of taxonomic shufflings concerning the 
genera Cricodus, Dendrodus and Polycodus; they may be con- 
sidered most conveniently together. The best way to review the 
situation seems to be to start with the description by Rohoii 
(1889) of several specimens from the Upper Devonian of Ru.ssia, 
some of Avhich he assigned to Dendrodus hiporcafus Owen (Ro- 
hon, 1889, plate 1, figures 1, 5, 9, a skull, and 2, 7 8, a tooth and 
two scales), while others he named Cricodus wenjucovi (Rohon, 
1889, plate 1. figures 4, 6, a skull from the River Ojatj, and 3, 
11. a skull and lower jaw from the River Sjass). Since these 
specimens are not given numbers by Rohon, I shall refer to 
them, for convenience, by the number of the figure by which 
they are illustrated. 

Gross (1933) placed tho.se specimens figured in plate 1, figures 
1, 3, 4, 5. 6, 9, together with new material from the Baltic Old 
Red Sandstone, in the genus Pclyplocodus Pander 1860, as the 
new combination Polyplocodus wenjucovi. Jarvik (1937), how- 
ever, put "figures 1, 3, 4, 5, 6. 9, 11" in Eusfhenopteron as £". 
wenjucovi. He also concluded that the names Cricodus, Den- 
drodus, and Prjlyplocodus should not be assigned to any fresh 
material because of the fragmentary and enigmatic nature of 
the originals. Hence Gross (1941) named a new genus Pan- 
derichthys in order to reassign material named by him (1930) 
Polyplocodus (Cricodus) rJwmholepis (see also Gross 1933. 
1936). Now there enters a possible source of confusion because 
Gross (1941) named a second new species — PanderichfJiys 
bysfrovi — for the material from the Baltic Old Red which he 
had named Polyplocodus wenjucovi in 1933 (see above), while 
leaving the Rohon material in Eusfhenopteron tvenjucovi. Xov- 
objeva (1960a) named a third species — Panderichthys stol- 
bovi. 



4 BREVIORA No. 177 

Til 1959 Vorobjeva described a new genus Platycephalichthys, 
with the type riaf)/C( pJialichfhifn hischoffi, based on recently 
collected material from the Tjiper Devonian of Russia, and in- 
cluded in this genus the Rohon specimen from River Ojatj ("fig- 
ures 4 and 6") as a new species Plafyccphalichthys rohoni. 
Later (1960b) she referred the remaining Rohon material to 
Eusthenodon Jarvik 1955 as Eusihenodon wenjucovi. 

Thus the two genera now comprise : 
Panderichfhys 

P. rhomholepis Gross 1941 

P. hystrovi Gross 1941 

P. stolhovi Vorobjeva 1960 
Plafycephalich th ys 

P. hischoffi Vorobjeva 1959 

P. rohoni Vorobjeva 1959 

From the structure of the scales and teeth, Plafyccphalichthys 
is a member of the Rhizodontidae. Panderichfhys was assigned 
by Gross (1941) to the Rhizodontidae but there seems little 
doubt (Vorobjeva 1960a, and Orvig 1957) that it should be 
placed with the Osteolepidae. 

SUBDIVISION OF THE RIIIPIDISTIA 

Jarvik (1942, p. 489) lists a series of dift'erences between 
Porolepis and Eusfhcnoptcron which he considers to be repre- 
sentative of a basic split within the Rhipidistia. He also (1942, 
pp. 417, 495) discusses the connections between these fishes and 
the recent Ami^hibia. In general, the same characters are in- 
volved in the two arguments. In the next few pages I shall 
review recent findings which indicate that certain of these char- 
acters no longer support Jarvik 's distinctions. These are all 
characters which may readily be determined in the fossils; this 
is in contrast with some of Jarvik 's points involving the passage 
of nerves and vessels, the refutation of which could be as diffi- 
cult as their interpretation. Some of Kulcyzcki's (1960) state- 
ments may be disputed, as I shall mention later, on the grounds 
that his material was insufficiently well preserved. This criti- 
cism cannot apply to Vorobjeva 's work, particularly since the 
most important of the characters she has described are, as already 
mentioned, easily ascertained. 

The recognition, in certain anatomical characters, of resem- 
blances between specific rhipidistians and recent amphibians, 
without the supporting evidence of a fossil lineage (lacking in 



1962 RHIPIDISTIAN RELATIONSHIPS 5 

the ease of the Urodela and Anura) becomes, to a certain ex- 
tent, a matter of subjective judgment. In the following pages 
I shall consider primarily the evidence for a basic distinction 
between 'Osteolepiformes' and 'Porolepiformes, ' for it is upon 
the validity of this supposed dichotomy that all subsequent 
phylogenetic hypothesis must rest. 

The nature of the anterior palatal fenestrae. According to 
Jarvik (1942, p. 489, etc.), an important difference between the 
"Osteolepiformes' and the 'Porolepiformes' lies in the nature 
of the paired palatal recesses present between the anterior edges 
of the vomers and the posterior rim of the premaxillae. In 
Eusthenopteron this region has the form of a shallow, partially 
sub-divided groove, the 'prenasal groove,' limited posteriorly by 
the edges of the vomers; in Porolepis there is a pair of '])its' 
extending backwards between and separating the vomers. These 
palatal recesses which are described by three names — 'fossae 
apicales, ' 'anterior palatal fenestrae,' and 'pre-nasal pits' — 
were assumed by Jarvik to have contained glandular organs 
homologous with the various intermaxillary glands of recent 
Amphibia. It was further proposed that the condition in Eus- 
thenopteron foreshadowed that of recent Anura, and the condi- 
tion in Porolepis tliat of Urodela. 

It had already been suggested, however, that these recesses 
served solely to receive the points of large tusk-like teeth set 
in the tips of the lower jaws (Holmgren and Stensio, 1936; 
Romer, 1937; and now also Kulcyzcki. 1960). The material at 
my disposal shows quite clearly that this latter explanation is 
the true one for the Osteolepidae. Plate I shows the tusk fitting 
into the palatal recess, leaving no room for any glandular struc- 
ture. That this is also the case in other 'Osteolepiformes' may 
be deduced from the similar large tusks of the lower jaws — for 
example, in Panderiehthys (Gross, 1941). In 'Porolepiformes' 
the situation is very similar, but in tliis case, as Jarvik has 
.shown, the teeth concerned are a pair of tooth whorls. This 
extremely interesting discovery provides, incidentally, positive 
indication that the familiar Onychodiis, long known from such 
tooth whorls, is, in fact, a 'porolepiform' rhipidistian (Jarvik, 
in press). 

The presence of the paired tooth wiiorls speaks, indeed, for 
tile unity of the 'Porolepiformes,' but the morphological rela- 
tionship between the type of palatal fenestra and the lower jaw 
dentition completely precludes any phylogenetic relationship 



6 BREVIOBA No. 177 

between the fenestrae in Rhipidistia and the jilandnlar organs 
of recent Amphibia. 

Evidence from, the cnniiaJ cavity. A point of resemblance, 
apparently possible of interin-etation as evidence of relationship, 
between Porolepis and the Hrodela is that in both "... the in- 
ternasal wall is broad and does not form any nasal septnm. It 
lodges the ethmoid part of the cranial cavity" (Jarvik, 1942, 
p. 417). This, moreover, is supposed to be in direct contrast 
with the 'Osteolepiformes' (and Anura) — by extrapolation 
from the situation in Eusthenopteron in Avhicli the internasal 
wall is relatively narrow and also solid. 

Kulcyzcki (1960) noted that his material of Porolepis did not 
show any forward extension of the cranial cavity between the 
nasal sacs; his material was perliaps not as good as might be 
desired fully to substantiate this view, since it consisted entirely 
of natural casts and not true bony remains. At the same time 
that Kulcyzcki 's work was published there appeared the work 
of Vorobjeva. Her material consisted of several portions of the 
skulls of Plotycephalichthys and Panderichthys, both of which 
are undoubtedly 'osteolepiform' and both of which, she states, 
have an ethmoid ext(»nsion of the cranial cavity. Her descrip- 
tion of PlatyeephalicJttJiys is interesting; ". . . a wide inter- 
nasal portion (of the braincase) with a cavity stretching forward 
almost to the front edge of the skull" (Vorobjeva, 1960a, trans.). 

In the face of evidence that a rhizodontid and an osteolepid 
have a 'pars ethmoidalis cranialis' and at least one species of 
Porolepis may not, one is forced to conclude that this character 
is of no significance in any attempt to distinguish supra-familial 
groupings within the Rhipidistia. One may further ])ear in 
mind that the extent of the cranial cavity need have no relation 
to the extent of the brain contained therein. A dramatic demon- 
stration of this is afforded by the coelacanth Latimcria (Mi]U)t 
and Anthony, 1958), in which a large cranial cavity contains 
but a small brain. Presence or absence of the pars ethmoidalis 
cranialis is thus a rather labile character in the Rhipidistia and. 
having no great anatomical or functional basis, no phylogenetic 
speculation may reasonably be drawn from it. 

Nasal apertures aud the )iasal cavity. All known 'Porolepi- 
formes' are characterized by the presence of two external nares 
and a choana. Further, the endocranial opening for the poste- 
rior naris is confluent with that for the choana. In Evsthenop- 
teron, on the other hand, there is but one external naris and this 



1962 RHIPIDISTIAX RELATIUXSHIPS 7 

is separated from the choana by the lamina iiario-choanalis of 
the endoskeletal nasal capsule. T'ntil the work of X'orobjeva 
this was believed to hold true for all 'Osteolepiformes" but 7^^//;- 
dcrichihijs is described fVorobjeva, 1960a) as liavin<r both an 
anterior and a posterior external naris. From her ti<>ures it is 
not possible to determine Avhether the described contiuenee of 
the posterior naris with tiie fenestra endochoanalis is (liic iiici'dy 
to a defect in the preservation oi- not. This is indeed a strano'e 
situation for a fish which is in all major respects to be consid- 
ei'ed 'osteolepiform.' 

Jarvik has other speculations based on the detailed configura- 
tion of the nasal cavity itself, but these seem to be more open to 
dispute in connection with phylogenies spanning 300 million 
years. Ectosteorhachis certainly lacks most of the ridges, grooves 
and depressions described in Eusth< nopfi ran and supposed to be 
typical of all 'Osteolepiformes.' 

Jarvik described in Eustlienopteron a foramen in the post- 
nasal wall which he states is the posterior endonarinal fenestra 
(Jarvik 's terminology). This is also present in Ectosteorhachis, 
but the presence of an anterior naris in Panderichthys may pos- 
sibly indicate that the single external naris of 'Osteolepiformes' 
is homologous with the posterior naris of Porolepis. In this case 
the foramen in the post-nasal wall, which Jarvik considers to be 
the forerunner of the tetrapod naso-lachrymal duct, may ])oh- 
sibly have to be interpreted in some other fashion, but consider- 
ably more evidence is needed to settle this point. 

Other structures. There are several other points, noted by 
Jarvik as indicative of a division within the Rhiiudistia, which 
are contradicted by the anatomy of Ectosteorhachis: 

1) Jarvik (1942, p. -t92) states that a difference between 
'Osteolepiformes' and 'Porolepiformes' is that in the former the 
vomers are in mesial contact, while in the latter they are separate 
from each other. Although in tiu^ sectioned material at my dis- 
posal the vomers are slightly displaced, it seems that these 
bones are not in mesial contact. Further, they lack the pos- 
terior extension passing back on either side of the tooth-bearing 
part of the parasphenoid, which has also been stated to be typi- 
cal of 'Osteolepiformes.' The vomers are thus much more simi- 
lar to those of Porolepis than to tbose of Eusthenopteran : they 
lie entirely anterior to the tooth-bearing part of the parasphe- 
noid. 



8 BREVIORA Xo. 177 

2) Jarvik states (1942, p. 492) that in 'Osteolepiformes' the 
parasphenoid is narrow and in 'Porolepiformes' it is broad. Bnt 
in Ectosteorhachis (an osteolepid) only the tooth-bearing part 
is narrow — the parasphenoid is continued forward and later- 
ally as a broad, if thin, film of bone fused to the ventral surface 
of the endocranium (cf. Romer, 1937, p. 19). 

3) Jarvik states that lack of the external parietal foramen is 
tyi)ieal only of 'Porolepiformes'; both MegalichfJnis and Ecto- 
steorliachis lack it, however. 

Kuleyzeki further criticizes Jarvik 's description of the snout 
anatomy of Porolepis, especially the detailed description of the 
nasal capsule and the canals for nerves and vessels (see espe- 
cially Kuleyzeki, 1960, pp. 81-94). On the other hand, Jarvik 
states (in press) that in most details the new material he has 
of Glyptolepis fully bears out his description of Porolrjns. 

DISCUSSION AND CONCLUSIONS 

The nature of the fossil remains, by which the Ehipidistia 
are known, is extremely variable, ranoiny from the excellent 
and fairly plentiful material preserved *in the round' of Eus- 
thenopteron, Ectosteorhachis and Glyptolepis (Jarvik, in press, 
and in preparation), to the fragments of skulls, isolated teeth, 
and scales typical of most genera. It is not surprising, there- 
fore, that most phylogenies and taxonomic studies have been 
based on the histology of teeth and scales. The latter approach 
is, of course, open to some doubts and reservations, but the re- 
cent work of 0rvig seems particularly important (see 0rvig, 
1957, p. 409 for phylogeny). 

It is especially interesting to note the positions assigned by 
0rvig to Pandcriclithys and Platycephalichthys. According to 
Jarvik 's interpretations of crossopterygian anatomy, the pres- 
ence of the pars ethmoidalis cranialis in both genera and the two 
external nares in Panderichthys would ally them with the Poro- 
lepis lineage. All other evidence, however (J^rvig and Vorob- 
jeva), opposes this view and maintains their 'osteolepiform' 
status. One is forced to conclude that these two characters are 
more labile than was formerly supposed ; they fail to show cor- 
relation with other, diagnostic, characters. 

Schmalhausen (1959) rejected Jarvik 's theories on the grounds 
that the two 'types' of skull merely reflected the relative pro- 
portions of the skull, Porolepis being broad-snouted and Eus- 
tJienopteron narrow-snouted. Schmalhausen was referring par- 
ticularly to the difference between the two types of anterior 



1962 



RHIPIDISTIAN RELATIONSHIPS 



palatal fenestra, the nature of whieh we have already seen to 
depend on the type of lower jaw dentition. Although Sehmal- 
hausen's idea is an attraetive one, it does not explain the dis- 
crepancy in the oeenrrenee of the pars ethnioidalis ci'aiiialis. 
The 'Osteolepiformes' contain both broad and narrow snouted 
forms, but the presence of the pars ethnioidalis cranialis is in- 
dependent of this factor : 

Pars ethnioidalis 



Osteolepidae 



Rhizodontidae 



Snout 


cranialis 


Broad 


Absent 


Narrow 


Present 


Narrow 


Absent 


Broad 


Present 



j Ectostf orliachis 
I Pandcrichfln/s 

j Eusfhenopfcron 
[ Platyccphaiichthys 

This character is not even related to the relative width of the 
internasal wall, which seems to be dependent on the relative size 
of the nasal capsules rather than on the external proportions or 
intrinsic 'osteolepiform'/'porolepiform' nature of the snout. 

I can think of no explanation of the apparently random oc- 
currence of these characters except that the Rhipidistia form a 
fairly close-knit group within which comparable variations can 
occur in all families. 

There is no doubt that a distinction of some sort can he ili-awii 
between a porolepiform and an osteolepiform assemblage (al- 
though the former is a much more compact group than the lat- 
ter). The controversy lies in the status to be assigned to eacli. 
The two groups have been interpreted by Jarvik (1942. I!).!."). 
1960) to represent a fundamental split within the Rhipidistia 
and, by extrapolation, within the recent Amphibia also. There 
are two major objections to the arguments which are presented 
to substantiate this hypothesis. Firstly, only one genus of 'Osteo- 
lepiformes' (Ei(sthenopfcrou) was available for consideration in 
any detail, and secondly, all the characters dividing the two 
groups are chosen with reference to the proposed relation to the 
Urodela or Aiiura and not at all with reference to the Rhipidis- 
tia in general. Taking into consideration the evidence presented 
above and bearing in mind other more detailed jioints such as 
are dis]nited by Kulcyzcki and to which 1 hope to return in a 
future work, it seems that the two groui)s of Rhipidistia arc 
more closely related than has been stated. Furthermore, those 
characters which might reflect a relationship with particular 



10 BREVIORA No. 177 

recent Amphibia are present in both croups of rhipidistians and 
are not, therefore, justifiably so considered. 

From my own studies of the Rhipidistia, I personally favour 
the view propounded by Berg (1958) that there are three <>roups 
of Klii])idistia : the Porolepiformes, the Osteolepiformes and the 
Rhizodontiformes. I would allow each group no more than 
super-familial rank. 

Literature cited 

Berg, L. S. 

1958. System der rezenten iind fossileii Fisehartigeii uiid Fisehe. 

31(t pji. Deutscher Verlag, Berlin. 
Bystrow, a. p. 

1939. Zahnstniktur der Crossopterygier. Acta. Zool., 20: 283-338. 
1950. Mieioscopic structure of tlie bones and teeth of the Carbonifer- 
ous crossopterygian tish MepaUclitliys (family Osteolepidae). 

Dokl. Akad. Xauk. U.S.S.B., 34: 119-121. (In Russian.) 
Cope, E. D. 

1880. Second contrilnition to the histuiy of the Vertel)rata of the 

Permian formation of Texas. Proc. Am. Phil. Soc, 19: 38-58. 
1891. On the character of some Palaeozoic fishes. Proc. U.S. Nat. 

Mus., 14: -447-463. 
Gross, W. 

193(1. Die Fische des mittleren Ohl Red Sud-Livhmds. Geol. Paliiont. 

Abh., N.F., 18: 121-ir)(i. 
1933. Die Fische des Intltischen Devons. Palaeontographica, 79, Alit. 

A: 1-72. 
1936. Beitrage zur Osteologie lialtis(diei- und rlieinischer Devon-Cros- 

sopterygier. Palaont. Z., 18: 129-155. 

1941. Uber den Unterkiefei ciniger devonistdier Crossopterygier. 
Abh. Preuss. Akad. Wiss., umth.-nat. Kl., no. 7: 1-51. 

Holmgren, N. and E. A. Stensio 

1936, Kranium und Visceralskelett der Akranier, Cyclostomen und 
Fisclie. In L. Bolk ei alio, Handbucli der Vergleichenden Ana- 
tomie der Wirbeltiere, vol. 4: 345-353. 

Jarvik, E. 

1937. On the species of Eusthenopteron found in Russia and the Baltic 
States. Bull. Uppsala Geol. Inst., 27: 63-127. 

1942. On the structure of the snout of crossopterygians and lower 
gnathostomcs in general. Zool. Bidrag, 21: 237-675. 

1955. The oldi>st tetrapods and their forerunners. Sci. Monthly, 30: 

141-154. 
1960. Theories de 1 'evolution des vertebres. 104 pp. Masson et Cie, 

Paris. 



1!)62 RHIPIDISTIAN RELATIONSHIPS 11 

(ill l>i't'ss I. On tlu' Porolepiformes and the origin of tlic iiidiji'les. (Unpub- 
lished, read before tbe Intei-national ('olloiniiutn oil Kvobition, 
Paris, May, l!t(jl.j 

KlLCYZCKI, J. 

1 !•<)(». Pnrolcpis (Crossopterygii) from the Lower Devonian of tlie 
Holy Cross Mountains. Aeta Palaeont. Polonica, 5 (1) : 65-106. 
Lehman, J. P. 

1959. L 'evolution des vertebres inferieures. 188 pp. Dunod, Paris. 
MiLi>()T, J. and J. Anthony 

1958. Anatomie de Latimeria ehalumnae. Vol. T. Squelette, Museles 
et Formations de soutien. 122 pp. Centre National de la Rech- 
erche Scientifique. Paris. 

Orvig, T. 

1957. Remarks on the vertebrate fauna of the lower Upper Devonian 
of Escuminae Bay, P.Q., Canada, with especial reference to the 
porolepiform crossopterygians. Arkiv. Zool., 10 (6): 367-126. 
ROHON, J. V. 

1889. Die Dendrodonten des devonischen Systems in Russland. Mem. 
Acad. Imp. 8ci. St. Peterslnirg, (7) 36 (14): 1-52. 
ROMER, A. S. 

1937. The braincase of the Carlioniferous crossopterygian Megalich- 

ihys nitidus. Bull. Mus. Conip. Zool., 82 (1): 1-67. 
l9-tL Notes on the crossopterygian hyoniandilnilar and braincase. J. 
Morph., 69 (1,): 141-160. 

SCHMALHAUSEN, I. I. 

1959. Concerning monophyletism and i)olyphyletism in relation to the 
problem of the origin of the land vertebrates. Bull. Mosk. 
Obshch. Isp. Pri. Biol., 64 (4): 15-33. (In Russian.) 

VOROB.JEVA, E. I. 

1959. A new genus of crossopterygian fish — Plaii/cephalichihjis from 

the Upper Devonian of Lovat. Palaont. J. Akad. Nauk. U.S.S.R., 

1959, no. 3: 95-106. (In Russian.) 
1960a. New facts aljout the genus Paiiderichthi/s from tlic Devonian 

of the U.S.S.R. Palaont. J. Akad. Nauk. U.S.S.R., 1960, no. 

1: 87-96. (In Russian.) 
1960b. Concerning the .systematic position of Eusfhenopteron wenju- 

covi (Rohon). Palaont. J. Akad. Nauk. U.S.S.R., 1960, no. 2: 

121-129. (In Russian.) 



Xo. 177 




Plate 1. 
Megalichtlni.s fidiu tlie Scottish Carboniferous. M.C.Z. 8941. Tip of the 
?noi:t in anterior view, showing iiremaxillary teetli, dentary tusks, and the 
anterior pahital fenestrae exposed by a natural break in the specimen: 

A. Photographed immersed in water. 

B. Semi-diagrammatie sketch of A, emphasizing the pertinent features. 



V / V-t-.^ 



BREVIORA 



Museium of Comparsitive Zoology 

Cambridge, Mass. December 27, 1962 Number 178 

ON A NEW SPECIES OP THE EARTHWORM CENTTS 
TBIGASTEE BENIIAM 1886 (OCTOCHAETIDAE)i 

By G. E. Gates 

University of Maine, Orono, Me. 

The presence, in Cuba, of an apparently endemic Trigaster is 
of such zoogeographical importance as to contra-indicate further 
and perhaps indefinite delay in publishing' avail;i1<le data. 

Tbigastei; (javernicola n. s]). 

"Cueva de Colon," Punta Caguanes, Yaguajay, Las Villas 
Province, Cuba, 0-0-1. G. S. Ta!)()ada. (Type in the Museum of 
Comparative Zoology. ) 

External characteristics. Size, 14:} by 5 mm. Segments, 276. 
Prostomium, epilobous, tongue open posteriorly but crossed by a 
deep transverse furrow near anterior margin of i. Setae, AB < 
CD < AA which is ca. = or slightly < EC DD ca. = ^l.C. First 
dorsal pore, at 8/9. 

Clitellum, faintly indicated, possibly saddle-shaped and reach- 
ing down only to A, on (xiii ? ) xiv-xix (xx ? ) . Spermathecal pores, 
minute, superficial, midway between A and B, two pairs, equa- 
torial in viii-ix. Female pore, at A and midway between 13/14 
and eq/xiv, on left side. Male and prostatic pores, as well as 
apertures of setal follicles in the male area, unrecognizable. 
Seminal grooves, deep and wide, about at B, between eq/xvii and 
xix/eq. Genital markings, unpaired, transversely placed, in AA, 
each with a deep transverse, central groove, primarily postsetal 
on xvi, primarily presetal on xx and xxi but perhaps crossing 
sites of 19/20 and 20/21. 

Internal anatomy. Septa, 6/7-10/11 muscular and opaque, 
11/12 membranous but with discrete muscular strands. Pigment, 



1 From research financed by the National Science Foundation. 



2 BBEVIORA No. 178 

red, recognizable only in region of mD, in or associated with the 
eirenlar musculature. 

Gizzards, large, in v and vi. Esophagus, deeply constricted at 
insertions of 11/12-13/14, in xi-xiii with a bifid ventral typhlosole 
and numerous low, rounded, irregular ridges that are gorged 
with blood, without discrete calciferous glands. Intestinal origin, 
apparently in xv. Caeca, intestinal typhlosoles and supra- 
intestinal glands, lacking. 

Dorsal blood vessel, single, complete, bifurcating under the 
brain, the branches uniting over subpharyngeal ganglion to be- 
come the ventral trunk. Extra-esophageal trunks, blood filled 
and recognizal)le alongside the ventral trunk in three segments, 
posterior portions empty and invisible. Supra-esophageal trunk, 
recognizable only in x-xiii. Subneural trunk, presumably lacking 
as no trace of one was found. Hearts, of x-xii are apparently 
lateral as no bifurcations to supra-esophageal trunk were seen, 
hearts of xiii latero-esophageal. 

Nephridia, in at least six longitudinal ranks posteriorly, those 
of the median rank on each side larger than the others (or median 
rank composite 1 ) . 

Male funnels, rather small and thick, without folds, in x-xi. 
Seminal vesicles, finely acinous, soft, in xii, none recognizable in 
coagulum of xi. Prostates, 3-4 mm. long, an ectal portion about 
one half mm. long and with a muscular sheen presumably being 
the duet. Penial and copulatory setae, not found. GM glands, 
apparently lacking. 

Spermathecae, fairly large, adherent to posterior faces of septa 
and to parietes. Diverticulum, nearly spheroidal, sessile on the 
duct just below the ampulla. Ovaries, fan-shaped, each with 
several (juite long egg strings. 

Reproduction. Spermatozoal iridescence is lacking in the 
opaque coagulum of spermathecal diverticula and probably also 
on the male funnels. Ovaries are fully mature but the clitellar 
epidermis is only slightly tumescent. Postsexual regression may 
have been under way and if so biparental reproduction is, per- 
haps, to be anticipated. 

Inyesta. Soil, brownish. 

Remarks. The type was received in May 1958 for identification. 
Better material, fully clitellate if possible, was requested from 
the collector. Directions for preservation were not followed and 
subsequent specimens from the type site (received in August, 
1958) were not identifiable as to genus or family. A search in 



1962 NEW SPECIES OF TRIGARTER 3 

the cave diirino' the dry soason of "Freedom Year" (1959) by 
the same person was fruitless. Collecting in the immediate vicin- 
ity of Tueva de Colon at the appropriate season was then recom- 
mended as the species is unlikely to be confined to caves. 
Unfortunately, receipt of further collections in the near future 
now seems to be as unlikely as during the past three years. 

Though the characterization above leaves much to be desired 
the species is distinguishable from its congeners by location of 
the spcMMuatbeeal pores. 

Key to species of Trigaster 

1. Gizzards, two 2 

Gizzards, three 3 

2. Rpeimathecal pores at (or slightly behind?) intersegmental fur- 

rows, spermathecae adiverticulate tolteea (1) 

Spermathecal pores equatorial, 

spermathecae diverticulate cavernicola 

3. Pigmentation, dense, red, resistant to alcohol and formalin. . rufa (2) 
Pigment, lacking or not as in rufa 4 

4. Spermathecae adiverticulate and with pores in AB, gizzards in 

vii-ix lankesteri (3) 

Spermathecae diverticulate and with pores at (unpaired?) or 

close to mV, gizzards in v-vii intermedia (4) 

(1) T. tolteea Bisen, 1900, Proc. California Acad. Sci., (3) 2, p. 203. 
Known only from the original description of a single specimen from Toluca, 
Mexico, at 8000 feet. Type (lost in San Francisco earthquake?), abnormal, 
preservation ' ' indifPerent. ' ' Testes are said to be present in x. 

(2) T. rufa nom. nov. for Trigaster sp. Gates, 1954, Bull. Mus. Comp. 
Zool., Harvard College, 111, p. 244. Known only from the very meager 
description of two macerated specimens from Luquillo Forest, at 1800 feet, 
Porto Rico. 

(3) T. lankesteri Benham, 1886, Quart. Jour. Micros. Sci., 27, p. 94. 
Known only from the original description of the anterior portion of a single 
specimen from St. Thomas, West Indies. Whereabouts of the type, if still 
extant, is unknown. Errors in text as well as figures and/or abnormality are 
suspected. Testes, male funnels and seminal vesicles were not found (lost 
from sections?) but the amputee was clitellate and the record of sperma- 
thecal sperm (the only one for the genus) enables an assumption that repro- 
duction is biparental. Michaelsen assigned Benham 's form the rank of 
typical subspecies, and named two additional subspecies. 

(4) T. lankesteri intermedia and mlwoodi Michaelsen, 1900, Das Tier- 
reich, 10, p. 235. Erected on three (or more?) specimens from St. Thomas, 
one juvenile, one clitellate but stretched and presumably macerated, and 
one that may have been aclitellate and not quite so much stretched. Types, 
if extant, presumably in the Hamburg Museum but information as to their 
fate and condition as vet unavailable. Such information about the two taxa 



/^'t 



4 BREVIORA No. 178 

as was vouchsafed by Michaelsen is in the definitions of his Tierreich mono- 
graph. Differences between intermedia and calwoodi now seem unimportant 
but specific distinction from lankesteri does seem i)ossible if Benluun 's 
characterizations of gizzards, spermathecae, clitelhim, etc., are correct. 



DTRCIT8ST0N 

In Tngaster a simple digestive system without calciferous and 
supra -intestinal glands, intestinal typhlosoles and caeca but with 
two or three esophageal gizzards is associated with a nieroic 
excretory system possibly much like that of other octoehaetid 
genera such as Eutifphoeus and Eudichogasto-. Four prostates 
and four spermatliecae, supposedly ancestral in the classical 
phylogenies, have l)een retained throughout but unassociated 
with penial and eopulatory setae. Although testes were said to 
be present in x of one species (cf. notes above), absence of all 
seminal vesicles except those of xii suggests that each species is 
functionally metandric.- 

None of the trigasters seems to be common and all may be 
restricted to more or less remote habitats difficult of access from 
routes usually travelled by man. None of the species is known 
to have been transported and all are believed to be endemic. 

The generic range is from Mexico through Cuba^ and Porto 
Kico to the American Virgins. The new Cuban species appears to 
to be more closely related to tlu> Mexican one. The Porto Rican 
species seems to be closer to those of St. Thomas. 

Earthworms (of course after exclusion of the transported 
forms), because of their v^-ontinement to, breeding in, and slow 
migration through the soil, long have been believed to be second 
in paleogeographical importance to no other group of animals. 
Trigaster, perhaps more than any otiier North American genus, 
now seems likely to provide especially interesting data if the 
native faunas of the Mexico-Virgin axis can be studied before 
they are too profoundly affected I)y human interference with 
their habitats. 



2 Male funnels of x, in other octoehaetid genera, frequently are retained after 
the testes of that segment ceased to mature sperm. 

3 Little is known of the oligochaete faunas of the West Indies, including His- 
paniola. Indeed, nothing- at all is known of the oligochaetes of Santo Domingo- 



I 



Date Due 



Mff^*M99V 



Harvard MCZ Libra 






3 2044 066 302 720 



nocc MOT /"•if^/^i II A ■*•!