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HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



BREVIORA %° *° 



MUSEUM OF COMPARATIVE ZOOLOGY 
Harvard University 



Numbers 296-325 
1968- 1969 



CAMBRIDGE, MASS. U.S.A. 

1969 



CONTENTS 

BREVIORA 

Museum of Comparative Zoology 

Numbers 296-325 

1968 

No. 296. The genus Dysderina (Araneae, Oonopidae) in Central 
America and the West Indies. By Arthur M. Checker- 
ing. 37 pp. October 11. 

No. 297. Population structure of the Asthenes flammulata super- 
species (Aves: Furnariidae). By Francois Vuilleumier. 
21 pp. October 1 1. 

No. 298. Monograph of the Cuban genus Viana (Mollusca: 
Archaeogastropoda: Helicinidae). By William J. 
Clench and Morris K. Jacobson. 25 pp. October 11. 

No. 299. What is Lumbricus eiseni Levinsen, 1884 (Lumbricidae, 
Oligochaeta)? By G. E. Gates. 9 pp. October 11. 

No. 300. Contributions to a revision of the Lumbricidae. III. 
Eisenia hortensis (Michaelsen) (1890). By G. E. 
Gates. 12 pp. October 1 1. 

No. 301. New echymid rodents from the Oligocene of Patagonia, 
and a synopsis of the family. By Bryan Patterson and 
Rosendo Pascual. 14 pp. October 11. 

No. 302. Geomyseria glabra, a new genus and species of scincid 
lizard from Bougainville, Solomon Islands, with com- 
ments on the relationships of some lygosomine genera. 
By Allen E. Greer and Fred Parker. 1 7 pp. October 1 1 . 

No. 303. Review of the genera of the tribe Loberini (Coleoptera: 
Languriidae). By T. Sen Gupta. 27 pp. December 31. 



No. 304. Redescriptions of Anachis avara (Say) and Anachis 
translirata (Ravenel) with notes on some related 
species (Prosobranchia, Columbellidae). By Amelie 
H. Scheltema. 19 pp. December 31. 

No. 305. Lytechinus williamsi, a new sea urchin from Panama. 
By Richard H. Chesher. 13 pp. December 31. 

No. 306. .Anew species of Eleutherodactylus (Amphibia, Salientia) 

from the (inavana region. Edo. Bolivar. Venezuela. 
B\ Juan A. Rivero. 11 pp. December 31. 

No. 307. A new species of Hyla (Amphibia, Salientia) from the 
Venezuelan Guayana. By Juan A. Rivero. 5 pp. 
December 3 1. 

No. 30S. The molluscan fauna of an unusual Bermudian pond: a 

natural experiment in form and composition. By 
Stephen Ja\ Gould. 13 pp. December 31. 

No. 309. Variability ol rodent incisor enamel as viewed in thin 
section, and the micro-structure o\ the enamel in fos- 
sil and recent rodent groups. By John H. Wahlert. 
1 S pp. December 31 . 

1 969 

No. 310. A shovel-tusked Gomphothere from the Miocene of 

Kenva. Bv Vincent J. Maglio. 10 pp. March 31. 

No. 311. The genera Apenesia and Dissomphalus in Argentina and 
Chile (Hymenoptera, Bethylidae). By Howard E. 

Evans. 23 pp. March 31. 

No. 312. Ecological observations on Anolis occultus Williams and 
Rivero (Sauria, iguanidae). By T. Preston Webster. 
5 Pp. March 31. 

No. 313. Louis Agassiz's numbers for Steamer Blake Stations of 
1X77-78, 1878 79. By Myvanwy M. Dick. 11 pp. 

March 31. 

No. 314. The cranial anatomy of the Permian amphibian Pantylus. 
By Alfred Sherwood Romer. 37 pp. March 31. 



No. 315. Stellicola dentifer n. sp. (Copepoda, Cyclopoida) as- 
sociated with a starfish in Jamaica. By Arthur G. 
Humes. 11 pp. March 31. 

No. 316. Cytotaxonomic studies on some unusual iguanid lizards 
assigned to the genera Chamaeleolis, Polychrus, Poly- 
chroides, and Phenacosaurus, with behavioral notes. 
By G. C. Gorman. R. B. Huey, and E. E. Williams. 
17 pp. April 30. 

No. 317. Relationships of two Cretaceous lizards (Sauria. Teiidae) . 
By Richard Estes. 8 pp. April 30. 

No. 318. Leucolepidopa sunda gen. nov., sp. nov. (Decapoda: 
Albuneidae), a new Indo-Pacific sand crab. By Ian 
E. Efford. 9 pp. April 30. 

No. 319. Competitive exclusion among anoles (Sauria: Iguanidae) 
on small islands in the West Indies. By A. Stanley 
Rand. 16 pp. April 30. 

No. 320. Taimanawa, a new genus of brissid echinoids from the 
Tertiary and Recent Indo- West-Pacific with a review 
of the related genera Brissopatagus and Gillechinus. 
By Robert A. Henderson and H. Barraclough Fell. 
29 pp. June 10. 

No. 321. Observations on the agonistic and breeding behavior of 
Leptodactylus pentadactylus and other amphibian spe- 
cies in Venezuela. By Juan A. Rivero and Andres 
Elroy Esteves. 14 pp. June 10. 

No. 322. The fossil record of amphiumid salamanders. By Richard 
Estes. 11 pp. June 10. 

No. 323. On the earthworms of Ascension and Juan Fernandez 
Islands. By G. E. Gates. 4 pp. June 10. 

No. 324. Polymorphism and evolution of the Hispaniolan snake 
genus Uromacer (Colubridae). By Henry S. Horn. 

23 pp. September 15. 

No. 325. The genus Phenacosaurus (Sauria, Iguanidae). By James 
D. Lazell, Jr. 24 pp. September 15. 



INDEX OF AUTHORS 

BREVIORA 

Museum of Comparative Zoology 

Numbers 296-325 

1968 1969 

No. 

Chesher. Richard H 305 

Chicki ring, Arthur M 296 

Clinch. William J 298 

Die k. Myvanwy 313 

Eei-ord. Ian I 318 

Esns. Richard 317, 322 

I steves, Andres Elroy 32 1 

1 \ ans. How ard E 311 

Fell, H. Barrac lough 320 

Gates, G. E 299, 300, 323 

Gorman, G. C 316 

Gould, Stephen Jay 308 

Greer. Allen E 302 

Henderson, Robert A 320 

Horn, Henry S 324 



No. 

Huey, R. B 316 

Humes, Arthur G 315 

Jacobson, Morris K 298 

Lazell, James D., Jr 325 

Maglio, Vincent J 310 

Parker, Fred 302 

Pascual, Rosendo 301 

Rand, A. Stanley 319 

Rivero, Juan A 306, 307, 321 

Romer, Alfred Sherwood 314 

scheltema, amelie h 304 

Sen Gupta, T 303 

VUILLEUMIER, FRANCOIS 297 

Wahlert, John H 309 

Webster, T. Preston 312 

Williams, E. E 316 



BREVIORA 

Museum of Comparative Zoology 

Cambridge, Mass. 11 October, 1968 Number 296 



THE GENUS DYSDERINA (ARANEAE, OONOPIDAE) IN 
CENTRAL AMERICA AND THE WEST INDIES 

Arthur M. Chickering 



Abstract. A total of twenty-four species of the genus Dysderina are 
recognized in this paper. Seventeen species are described as new. Nine 
of these are from Panama, two are from Costa Rica, two are from Trini- 
dad, W. I., one is from Jamaica, W. I., two are from St. Vincent, B. W. I., 
and the remaining one is from Dominica, B. W. I. Dysderina principalis 
Simon from St. Vincent, B. W. I. is not D. principalis (Keyserling) from 
Colombia and, therefore, is described as a new species, D. soltina sp. nov. 
Dysderina antillana Bryant, described from St. Croix, U. S. Virgin Islands 
in 1942 and reported from Hispaniola in 1948, has been shown to be 
Ischnothyreus peltifer (Simon) and is treated in another publication. 

The Oonopidae include a group of very small spiders usually oc- 
cupying concealed habitats such as leaf litter, debris, especially 
grass and weed debris, and other similar habitats. They are par- 
ticularly numerous in tropical and subtropical regions but are now 
known from many other parts of the world. The distinctive features 
of the family are treated in such publications as the following: 
Simon, 1892-1895; Petrunkevitch, 1939; Comstock, 1940; Kas- 
ton, 1948, and others, and will not be treated in this paper. Since 
early in my work of collecting and studying spiders in Panama, I 
have continued to be interested in these minute members of the 
order Araneae. For the past several years I have made a special 
effort to collect members of the family Oonopidae in Central 
America and the West Indies. As a result of this effort I have 
accumulated a rather large number of species of several genera 
belonging to this family, and the time has arrived for me to put 
the results of these years of study and collecting into a permanent 
record. It had been my intention to publish the results of these 
studies in a single monograph. Recently, however, the decision 
was made to publish a series of shorter papers each dealing with a 
single genus or a group of genera as conditions seem to warrant. 
The genus Dysderina is one of a group of genera to be treated early 
in the series. 



2 rreviora No. 296 

My appreciation and gratitude are again expressed for the con- 
tinued aid and encouragement in the pursuit of my studies ex- 
tended by the staff of the Museum of Comparative Zoology. My 
thanks are also extended to Dr. W. J. Gertsch, American Museum 
of Natural History, Dr. G. Owen Evans and Mr. D. J. Clark, De- 
partment of Zoology, British Museum (Natural History) for the 
loan of important species of the genus Dysderina. Grants GB-1801 
and GB-5013 from the National Science Foundation have made 
it possible for me to continue my work in the Museum of Com- 
parative Zoology and to spend a total of nearly twelve months 
during the last lour years collecting in Panama, Costa Rica, and 
the West Indies. 

Except as otherwise stated in later parts of this paper, all types 
together with my entire collection oi the genus Dysderina will be 
deposited in the Museum of Comparative Zoology. 

Genus Dysderina Simon, 1891 

The type species is Dysderina principalis (Kevserling) by mono- 
tvpv. The genus was established on the basis of a male from 
1 ilombia identified by Keyserling as Oonops principalis. Simon 
(1891 ) correctly recognized that this species could not be regarded 
as belonging to the genus Oonops and. therefore, placed it in a 
new genus Dysderina. Simon also described two additional species 
from St. Vincent Island. B. W. I. Dysderina plena O. P. -Cam- 
bridge was described from Mexico in 1894. Dysderina antillana 
Br\ant was described from St. Croix, U. S. Virgin Islands in 1942, 
and also reported from Hispaniola in 1948, but is now known to be 
Ischnothyreus peltifer (Simon) as will be shown in another paper. 
In 1951 I reported four species of Dysderina from Panama. Since 
that time I have collected these small spiders at every opportunity 
and as a result I now have a rather large collection consisting of 
numerous species from parts of Central America and the West 
Indies but none has appeared east of Dominica, W. I. The most 
important features of this genus observed during my study of the 
group may be stated as follows: Total length varies from about 
1.6 mm to 2.75 mm, with females usually somewhat larger than 
males. There is a remarkable similarity of general appearance 
among the species recognized in this paper. Males can be readily 
separated into species on the basis of the distinctive features of the 
palpal tarsi. Females, on the other hand, are very difficult to 
separate into species with any degree of certainty. In my treatment 
of this sex I have placed great emphasis on the epigynal areas 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 3 

which occasionally are quite distinctive but more frequently are at 
best only obscurely so. Nearly all outer parts are strongly chitin- 
ized. The carapace is moderately high; regularly arched from pos- 
terior eyes to beginning of posterior declivity; about three-fourths 
as wide as long nearly opposite second coxae; surface irregularly 
granulate; usually with no indication of a median groove or pit. 
Eyes: six in two rows; AME lacking; posterior row gently recurved 
as viewed from above; quite compactly arranged; little difference in 
size of eyes but shape often varies (long diameter always used for 
measurements); clypeus typically heavily bordered. Chelicerae 
moderately developed; usually vertical, parallel; with fang typically 
slender and evenly curved; fang groove with a single tooth, at least 
in certain species. Maxillae with a peculiar terminal notch seen 
only in males and thus far only in a few species (Fig. 2). Lip: 
strongly chitinized; considerably wider than long; transversely con- 
cave in middle. Sternum: strongly chitinized; usually deeply 
grooved with conspicuous lobules opposite the coxae; continued 
laterally between coxae and united with a sclerite surrounding the 
cephalothorax to make a strong enclosure; sternal suture strongly 
procurved; fourth coxae usually widely separated. Legs: usually 
4123 in order of length but with minor variations occasionally; 
first and second legs with conspicuous ventral spines which vary 
somewhat among different species; third and fourth legs without 
true spines. Male palp: all segments except tarsus simple and un- 
modified; apparently without true spines; tarsus somewhat inflated 
and with more or less distinctive terminal structures. Abdomen: 
ovoid; with pedicel strongly chitinized and deeply corrugated; dor- 
sal scutum varies somewhat in size but typically covers most of 
dorsum; epigastric scutum continued around pedicel and far dor- 
sally (Fig. 8); ventral scutum covers much of remainder of venter 
but varies in extent; a sclerite typically partially surrounds the 
spinnerets on ventral side; black bristles may mark the position of 
the reduced colulus; openings of book-lungs, tracheal spiracles, and 
genital organs more or less distinct. Genital area somewhat dis- 
tinctive but usually obscure. Where the species under consideration 
agrees fully with the stated features of the genus no mention will 
usually be made of the specific characteristics in the description of 
the species. 

As a result of my study of the genus Dysderina presented in this 
paper I am obliged to recognize a total of twenty-four species from 
the region under study (exclusive of D. principalis (Keyserling) 
from Colombia). The number is undoubtedly inflated because of 
the great difficulty in accurately matching up males and females. 



4 breviora No. 296 

Thirteen of these species are from Panama; two are from Costa 
Rica; one is from Mexico; two are from Trinidad. W. I.; one is 
from Jamaica. \V. I.; four are from St. Vincent, B. W. I.; and one 
is from Dominica. B. W. I. Fifteen of these species, ten of which 
arc regarded as new. are represented by males; nine are represented 
only b\ females of which seven are regarded as new; six of the dif- 
ferent kinds of males are accompanied by what are believed to be 
properly assigned females. It should be remembered, however, 
that matching females with the proper males is a very uncertain 
task. 

Key to the males of 1)">si>i kina from Central America 
and the n est Indies 

la. Species in which the palpal tarsus terminates in a pair of slender 
distal apophyses, one sickle shaped and one somewhat feather- 
shaped (Fig. 43) D. obtina, p. 2 1 

lb. Species without such palpal, tarsal, distal Structures as given above 
2 

2a. Species with B group of four or five distal, terminal apophyses on the 
palpal tarsus (Fig. ; " i D. globina, p. 14 

2 ). Species without such palpal, tarsal, distal structures as given above . .3 
Species with three definite, distal, palpal, tarsal apophyses (Figs. 41, 
77 ) ( meridina, spinigera ) 4 

3b. Species without such distal, palpal, tarsal structures as given above . .5 

4i Species with distal, palpal, tarsal apophyses as in Figure 41 

D. meridina, p. 20 

4b. Species with distal, palpal, tarsal apophyses as in Figure 77 

D. spinigera, p. 3 1 

Sa Species with a single, short, broad, distal, palpal, tarsal apophysis 

divided into an opaque half and a transparent half ( 1 ig. 4) 

D. abdita, p. 5 

5b. Species without such a single distal apophysis as given above 6 

Species with a long, slender, distal, palpal, tarsal spine and a relatively 
massive, somewhat twisted, closely associated apophysis (Figs. 20-22) 
D. craneae, p. 1 

6b. Species without such a pair of distal, palpal, tarsal structures as 
given above 7 

7a. Species in which the distal, palpal, tarsal apophysis turns at a right 
angle near its base and then divides into a pair of flattened structures 
( Figs. 37-38 ) D. intempina, p. 1 ( > 

7b. Species without such a distal, palpal, tarsal apophysis as given 
above 8 

8a. Species with a single, distal, palpal, tarsal extension divided into a pair 
of relatively short, incurved apophyses (Figs. 46-47) . . /). plena, p. 23 

8b. Species without such a pair of apophyses as given above 9 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 5 

9a. Species with two distal, palpal, tarsal structures, a slender curved spine 

and a relatively large, broad, compound apophysis (Fig. 63) 

D. seclusa, p. 27 

9b. Species without such a pair of distal, palpal, tarsal structures as given 
above 10 

10a. Species with palpal, tarsal, distal apophysis terminating in a broad, 
angular, somewhat concave structure (Fig. 16) . . . .£>. concinna, p. 9 

10b. Species without such a distal, palpal, tarsal structure as given above 
11 

11a. Species with palpal, tarsal, distal apophysis a single robust structure 
divided distally into a short, slender spine and a sharply pointed robust 
spine (Fig. 65) D. simla, p. 29 

lib. Species without such a palpal, tarsal, distal apophysis as given above 
12 

12a. Species with a main palpal, tarsal, distal apophysis sharply bent mid- 
way, enlarged and flattened distally and with a minute spine (Figs. 
58-59) D. recondita, p. 25 

12b. Species without such palpal, tarsal, distal structures as given above 13 

13a. Species with a robust palpal, tarsal, distal apophysis terminating in a 
series of short, pointed structures obscured by a cluster of hairs (Figs. 
69-70) D. soltina, p. 30 

13b. Species without such a palpal, tarsal, distal apophysis as given above 
14 

14a. Species with a pair of palpal, tarsal, distal apophyses curved toward 
one another; clypeus about three-fourths as high as diameter of ALE 
(Chickering, 1951, figs. 2, 3; and Fig. 24 this paper) . .D. dura, p. 12 

14b. Species with a long, curved, palpal, tarsal, distal apophysis together 
with a slender spine and a short, blunt process between the two; 
clypeus about as high as diameter of ALE (Fig. 81 ) . .D. watiua, p. 33 

DYSDERINA ABDITA Sp. IIOV. 

Figures 1-8 

Holotype. The male is from El Volcan, Republic of Panama, 
August, 1950. The name of the species is a Latin adjective re- 
ferring to its concealed habitat. 

Description. Total length 2.28 mm. Carapace 1.04 mm long; 
0.85 mm wide opposite second coxae where it is widest; well 
rounded along ventral border from opposite base of palp to pos- 
terior margin; 0.52 mm tall; quite evenly arched from PME to 
beginning of posterior declivity which begins opposite interval be- 
tween second and third coxae; with surface unevenly granulate 
with median arched area nearly devoid of granulations; without a 
median, longitudinal thoracic groove or pit; with sparsely situated 
short recurved hairs or fine bristles. Eyes: six in two rows as 
usual; posterior row occupies about five-sixths of width of carapace 



6 breviora No. 296 

at that level; viewed from above, posterior row gently recurved; 
viewed from in front, posterior row gently procurved (Fig. 1). 
Ratio of eyes ALE : PME : PLE = 10 : 9 : 9; boundaries of 
ALE somewhat irregular but nearly circular, others somewhat oval. 
ALE separated from one another by nearly their radius, from PLE 
by one-sixth of their diameter, from PME by a little less than one- 
third of their diameter. PME contiguous only for a short distance 
and separated from PLE by about one-ninth of their diameter. 
Height of clypeus (Fig. 1 ) slightly less than the diameter of ALE. 
Chelicerae: vertical; essentially parallel; basal segment 0.29 mm 
long; apparently without a basal boss; fang slender and evenly 
curved; fang groove apparently with a very small tooth, probably 
promarginal. Maxillae: moderately long; slender; distinctly con- 
vergent and almost meeting anterior to lip; palp inserted into basal 
third; with a well developed distal scopula; distal end divided 
(Fig. 2). Lip: strongly chitinized; extended posteriorly into sternal 
area and with an obscure groove separating the horizontal part 
from the more vertical portion; wider at base than long in ratio of 
about 9:7; anterior border with a row of stiff bristles. Sternum: 
scutiform in general; as wide as long; widest between second coxae; 
strongly chitinized and distinctly but less intricately grooved than 
in several other species (Fig. 3); posterior border recurved and not 
continued between fourth coxae which are separated by about 1.5 
times their width; coxae three and four short and stout; coxae one 
and two somewhat more elongated; sternal suture procurved. Legs: 
4213 in order of length; tibial index of first leg 10, of fourth leg 9; 
trichobothria present but exact number and placement undeter- 
mined. Tarsal claws apparently as recorded for D. seclusa 
(Chickcring. 1951 ). First femur with two pairs of ventral spines 
in distal half; first tibia with five pairs of ventral spines, first very 
long but diminishing distally so that last pair is hardly more than a 
pair of bristles; first metatarsus with three pairs of ventral spines, 
irregularly spaced. Second femur apparently with only one con- 
spicuous spine on prolateral surface at beginning of distal third; 
second tibia with four pairs of ventral spines; second metatarsus 
apparently with only two pairs of ventral spines, irregularly spaced. 
Third and fourth legs lacking true spines. Palp: only tarsus in- 
flated; important features, more or less distinctive, shown in Figures 
4-6. Abdomen: with a well developed and strongly corrugated 
pedicel; 1.24 mm long exclusive of pedicel; 0.9 mm wide near 
middle; spinnerets as usual in the genus; genital aperture as in 
Figure 7; dorsal scutum covers nearly the entire dorsum; epigastric 
scutum continued around anterior end of abdomen; boundaries be- 
tween epigastric and ventral scuta not clear; ventral scutum covers 



1968 



DYSDERINA IN C. AMERICA AND W. INDIES 




Figs. 1-8. Dysderina abdita sp. nov. Fig. 1. Eyes and clypeus; from in 
front. Fig. 2. Right maxilla; nearly lateral view (from dissected paratype). 
Fig. 3. Sternum; from below. Figs. 4-5. Left palpal tarsus; nearly ventral 
and retrolateral views, respectively. Fig. 6. Tip of left palpal tarsus; nearly 
dorsal view. Fig. 7. Genital aperture. Fig. 8. Abdomen; left side (from 
paratype.) 

nearly the entire venter except for a small area at posterior end 
and laterally where the two scuta nearly meet (Fig. 8). Color in 
alcohol: in general, a rich reddish brown, essentially as described 
for D. dura and others (Chickering, 1951 ). 

Records. Seven paratype males from El Volcan, Panama, Au- 
gust, 1950. It seems probable that one of the two kinds of females 
described from the same locality and collected during the same 
period should be matched with these males but this is not at present 
possible to do with any certainty. 



Dysderina Belinda sp. nov. 
Figures 9-12 

Holotype. The female is from Boquete, Panama, August, 1950. 
The name of the species is an arbitrary combination of letters. 



8 



BREVIORA 



No. 296 



Description. Total length 2.5 mm. Carapace: 1.08 mm long; 
0.9 mm wide opposite posterior border of second coxae where it is 
widest (Fig. 9); 0.49 mm tall opposite interval between second and 
third coxae where it is tallest; distinctly arched from PME to 
beginning of steep posterior declivity; surface distinctly granulate 
and with granulations often in rows; with a recurved groove just 




Figs. 9-12. Dysderina belinda sp. nov. Fig. 9. Carapace; dorsal view. 
Fig. 10. Posterior end of sternum; from below. Fig. 1 1. Venter, from below. 
Fig. 12. Epigynal area; from below. Figs. 13-19. Dysderina concinna sp. 
nov. Fig. 13. Carapace; left side. Fig. 14. Eyes and clypeus; from in front. 
Fig. 15. Sternum; from below. Figs. 16-18. Distal end of left palpal tarsus; 
nearly prolateral, nearly ventral and nearly dorsal, respectively. Fig. 19. 
Genital aperture. 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 9 

behind posterior eyes; thoracic part regularly rounded along ventral 
margin and sharply narrowed shortly behind eyes. Eyes: six as 
usual in a fairly compact group; posterior row occupies about two- 
thirds of width of carapace at that level; viewed from above, 
posterior row slightly recurved, measured by posterior borders. 
Ratio of eyes ALE : PME : PLE = 9.5 : 9 : 8.5. ALE separated 
from one another by about one-half their long axis; separated from 
PME by about one-fourth of their long axis and from PLE by a 
line at one point. PME contiguous for nearly one-fourth of their 
circumference; separated from PLE by a line at one point. Pos- 
terior row only slightly wider than anterior row. Height of clypeus 
equal to fully three-fourths of the long axis of ALE. Chelicerae, 
maxillae and lip typical of females of the genus as far as observed. 
Sternum: rather strongly convex; with the usual intricate grooves 
and lobes typical of the genus; posterior end very bluntly termi- 
nated and only extended between bases of fourth coxae which are 
separated by nearly 1.5 times their width (Fig. 10). Legs: tibial 
index of first and fourth legs 9; spines essentially as described for 
D. silvatica Chickering (1951) with no very important differences. 
Abdomen: essentially as described for D. silvatica Chickering 
(1951); scuta and sclerite partly surrounding spinnerets quite typi- 
cal of the genus (Fig. 11). Epigynal area: very simple as usual 
(Fig. 12). Color in alcohol: essentially as described for other 
species; with no important differences. 

This species appears to be closely related to D. silvatica Chicker- 
ing, 1951 (Fig. 6) but differs from that species definitely with 
respect to the features of the epigynal area. In the former species 
the epigynal area is a nearly semicircular area with a minute dot 
near the posterior border. In D. belinda sp. nov. the epigynal area 
(Fig. 12) appears as a nearly semicircular area with a complete, 
conspicuous border and a faint central, longitudinal stripe. 

Records. Three females taken in the same locality with the 
holotype and during the same short period of nine days in Boquete, 
Panama, August, 1950, appear to belong here as paratypes. 

Dysderina concinna sp. nov. 
Figures 13-19 

Holotype. The male holotype is from El Volcan, Panama, Au- 
gust, 1950. The name of the species is a Latin adjective referring 
to its neat, pleasing appearance. 

Description. Total length 2.27 mm. Carapace 1.01 mm long; 
0.86 mm wide opposite interval between first and second coxae 



10 breviora No. 296 

where it is widest; 0.44 mm tall, and therefore, about half as tall 
as wide (Fig. 13); otherwise essentially as described for D. abdita 
sp. nov. Eyes: posterior row occupies about seven-ninths of width 
of carapace at that level; gently recurved as seen from above. ALE 
separated from one another by about five-eighths of their diameter; 
separated from PLE by one-eighth of their diameter and from 
PME by about three-eighths of their diameter. PME contiguous 
for about one-fifth of their circumference; only slightly separated 
from PLE. Posterior row wider than anterior row in ratio of 9 : 7. 
Height of clypeus equal to about nine-eighths of the diameter of 
ALE. Viewed from in front, posterior row gently procurved (Fig. 
14). Chelicerae: basal segment 0.33 mm long; fang slender and 
evenly curved; otherwise essentially as recorded for D. abdita sp. 
nov. No teeth observed along fang groove. Maxillae and lip: ap- 
parently as described for D. abdita sp. nov. but no distal notch 
observed on maxillae (scarcity of paratypes prevents dissection for 
closer examination ). Sternum: scutiform in general; slightly longer 
than wide; intricately and obscurely grooved (Fig. 15); not con- 
tinued between fourth coxae which are separated by twice their 
greatest width; right first coxa abnormally small. Legs: tibial index 
of first leg 9, of fourth leg 10; several trichobothria observed but 
exact number and placement not determined; two tarsal claws. 
Spines: first leg with femoral ventral spines 0-lr-lr-2-2, with 
last two on retromargin reduced to little more than bristles; tibia 
with ventral spines 2-2-2-2-2-0; metatarsus with five ventral 
spines, three along promargin and two along retromargin, all ir- 
regularly placed. Second tibia with five ventral spines along pro- 
margin and four along retromargin; second metatarsus with two 
pairs of ventral spines; no true spines observed on legs three and 
four or on palp. Palp: essential features shown in Figures 16-18. 
Abdomen: 1.17 mm long; 1.04 mm wide near middle; pedicel, 
scuta, tracheal spiracles, openings to book lungs and spinnerets all 
essentially as described for D. abdita sp. nov.; genital aperture as 
indicated in Figure 19. Color in alcohol: essentially as described 
for other species in this paper with slight variations. 

Records. Two male paratypes were taken at El Volcan, Pan- 
ama, August, 1950. 

Dysderina craneae sp. nov. 
Figures 20-23 

Holotype. The male is from Simla, Arima Valley, Trinidad. 
W. I. in the close vicinity of the William Beebe Tropical Research 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 11 

Station, April 25, 1964. The species is named after Miss Jocelyn 
Crane, Director of the Station. 

Description. Total length 2.2 mm. Carapace 1.05 mm long; 
0.79 mm wide opposite second coxae where it is widest; 0.42 mm 
tall; surface conspicuously but very irregularly granulate. Eyes: 
eye group occupies fully four-fifths of width of carapace at level 
of PE; viewed from above, posterior row gently recurved; ratio of 
eyes ALE : PME : PLE =10:9:9; positions and relationships 
essentially as stated for D. globina sp. nov. Chelicerae, maxillae 
and lip: all essentially typical of the genus as observed in this 
study. Sternum: deeply grooved in general pattern usually found 
in this genus; anterior third with a conspicuous transverse groove 
but otherwise nearly smooth; posterior two-thirds intricately 
grooved with primary and secondary grooves; fourth coxae sepa- 
rated by about six-fifths of their width. Legs: tibial index of first 
and fourth legs 9; spines almost identical with those recorded for 
D. globina sp. nov. Palp: all segments typical of males of the 
genus except the enlarged tarsus with its embolus and associated 
structures (Figs. 20-22); the degree of twisting of these structures 
seems to vary somewhat among the observed male paratypes. Ab- 
domen: typical of males of the genus with respect to scuta, indis- 
tinctive genital region, spinnerets, book lungs, tracheal openings, 
etc.; spinnerets partly surrounded by a ventral semicircular 
sclerite. Color in alcohol: carapace brown with heavily granulate 
areas darker than middorsal and nongranulate areas; dorsal ab- 
dominal scutum lighter brown; other parts as usual with variations 
yellowish and light brown. 

Female paratype. Total length 2.45 mm. Carapace 1.12 mm 
long; 0.88 mm wide; 0.45 mm tall; granulate essentially as in 
male. Eyes: eye group occupies about three-fourths of width of 
carapace at level of PE; seen from above, posterior row gently 
recurved; ratio of eyes ALE : PME : PLE = 10.5 : 10 : 9; ALE 
separated from one another by slightly less than their radius; other 
relationships typical of females of the genus as seen in this study; 
clypeus heavily bordered; height equal to about seven-tenths of the 
diameter of ALE. Mouth parts essentially typical of females of the 
genus. Sternum: essentially as in male except that anterior region 
is more irregular because of secondary grooves. Legs: 41=23 
in order of length; tibial index of first and fourth legs 10; first leg 
with femoral ventral spines 0-0-lr-2-2-0, tibial ventral spines 2-2- 
2-2-2-0, metatarsal ventral spines 2-2-2-0; second leg essentially 
as in first; third and fourth legs devoid of true spines. Abdomen: 
dorsal and ventral scuta nearly cover this part of body; chitinous 



12 



BREVIORA 



No. 296 



scleritc completely surrounds spinnerets and anal tubercle but is 
very narrow dorsally. Epigynal area quite distinctive (Fig. 23). 

Records. Described female paratype taken with holotype from 
hay and weed debris by sifting. Simla, Arima Valley, Trinidad, 
W. L, April 25, 1964. Five male and seven female paratypes 
taken in vicinity of Simla, Arima Valley, or along roadside to 
Blanchesseusc, Trinidad, W. I., April, 1964. 




I igs. 20-23. Dysderina craneae sp. nov. Fig. 20. Left male palpal tarsus; 
nearly prolatcral view. Figs. 21-22. Tip of left male palpal tarsus; dorso- 
retrolateral view and nearly retrolateral view, respectively. Fig. 23. Epigynal 
area; from below. Figs. 24-25. Dysderina dura Chickering. Fig. 24. Tip of 
left palpal tarsus; Dearly prolatcral view. Fig. 25. Genital aperture of male. 
Figs. 26-27. Dysderina fitrtiva sp. nov. Fig. 26. Eyes from above. Fig. 27. 
Epigynal area; from below. 



Dysderina dura Chickering 
Figures 24-25 

Dysderina dura Chickering, 1951: 208. The male holotype is in the Museum 
of Comparative Zoology; collected on Barro Colorado Id., Panama 
Canal Zone, July, 1936. 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 13 

A detailed description of this species was published in 1951 to- 
gether with two figures illustrating certain features of the palp. 
Two more figures are added here to still further clarify the de- 
scription. About three dozen specimens have been added to the 
collection since the recognition of the holotype. All of these have 
come from the vicinity of the Panama Canal Zone. The species is 
by far the most numerous in the collection from Panama. The 
female is still not certainly known but is suspected of being D. sil- 
vatica Chickering. 

Dysderina furtiva sp. nov. 
Figures 26-27 

Holotype. The female holotype is from Jamaica, W. I., St. 
Catherine Parish, 3 mi. north of Spanishtown, Oct. 16, 1957. The 
name of the species is a Latin adjective referring to its concealed 
habitat. 

Description. Total length 1.98 mm. Carapace 0.84 mm long; 
0.69 mm wide opposite second coxae where it is widest; well 
rounded from just behind PE to posterior border which is only 
slightly notched; gently rising along median region from PME to 
beginning of steep posterior declivity; 0.25 mm tall; with no tho- 
racic groove; with a sparse covering of black hairs most conspicu- 
ous at beginning of posterior declivity. Eyes: posterior row 
occupies about eleven-fifteenths of width of carapace at that level; 
seen from above, posterior row very gently recurved measured by 
posterior borders but nearly straight (Fig. 26); ratio of eyes 
ALE : PME : PLE = 7.5 : 7 : 6.5; ALE separated from one 
another by a little less than one-third of their diameter, from PLE 
by about one-fifth of their diameter and from PME by about one- 
third of their diameter; PME contiguous for a short distance, 
barely separated from PLE; posterior row wider than anterior row 
in ratio of about 22 : 17; viewed from in front, posterior row defi- 
nitely procurved; height of clypeus equal to about four-fifths of the 
diameter of ALE. Chelicerae: basal segment 0.25 mm long; fang 
groove with a minute tooth on each margin near tip of the slender 
fang; otherwise essentially typical of females of the genus. Maxil- 
lae: somewhat more robust than described for D. silvatica Chicker- 
ing (1951) but essentially as in that species. Lip: wider at base 
than long in ratio of about 13 : 10; slightly concave along distal 
border; sternal suture gently procurved. Sternum: scutiform in 
general; as wide as long; strongly convex; only faintly lobulated 
opposite coxae; grooves and ridges lacking; this last feature very 



14 breviora No. 296 

unusual in the genus; posterior end not extended between fourth 
coxae which are separated by four-thirds of their width. Legs: 
4123 in order of length; tibial index of first leg 8, of fourth leg 7; 
first leg with only one spine on prolateral side about two-thirds from 
base of femur; first tibia with ventral spines 2-2-2-2-1 p; first meta- 
tarsus with ventral spines 2-2- lp; second leg with no femoral 
spines; second tibia with ventral spines 2-2-2-2-0; second metatar- 
sus with three spines along promargin and two along retromargin 
with irregularity in placement; legs three and four probably to be 
considered lacking spines but some of the numerous bristles might 
be considered weak spines especially on fourth tibia; palp without 
terminal claw but with main still bristles resembling weak spines 
Abdomen: pedicel shorter and less corrugated than in typical spe- 
cies; dorsal scutum covers about nine-tenths of dorsum; ventral 
scuta extend about seven-ninths of distance from pedicel to spin- 
nerets; without a chitinous band guarding the spinnerets as in typi- 
cal species; position of colulus barely indicated. Epigynal area: 
very simple as usual in the genus but its form (Fig. 27) seems to 
be more or less distinctive. Color in alcohol: essentially typical of 
the genus except that the highly chitinized cephalothorax and scuta 
are somewhat lighter in color. 

This species also appears to be closely related to D. silvatica 
Chickering. 1951, but the epigynal area is unlike that seen in any 
other species. In /). furtiva sp. nov. this region is a simple, broad, 
slit-like aperture with a narrow border surrounding a plain white 
area. 

Records. Paratype females were taken with the holotype and 
numerous specimens are in the collection from the following locali- 
ties: St. Andrew Parish. Stony Mill, May 26, 1956 (C. C. Hoff); 
Coopers Hill, Feb. 10. 1955 (P. F. Bellinger); Jack's Hill Road, 
Dec. 6, 1957; St. Ann Parish, vicinity of Moneague, Nov. 7, 1957; 
St. Catherine Parish, Evarton, Nov. 29, 1957. Numerous females 
from Trinidad, W. I., for some time regarded as representing a new 
species are now referred to this species. No males have yet been 
associated with these numerous females. 

Dysderina globina sp. nov. 
Figures 28-31 

Holotype. The male holotype is from Dominica, B. W. I.. 
Windward Islands, June 12, 1911. It will be deposited in the 
American Museum of Natural History, New York, N. Y. The 
name of the species is an arbitrary combination of letters. 



1968 



DYSDERINA IN C. AMERICA AND W. INDIES 



15 








I/// - 



34 



33 



Figs. 28-31. Dysderina globina sp. nov. Fig. 28. Eyes from above. Figs. 
29-30. Male palpal tarsus; nearly prolateral and retrolateral views, respec- 
tively. Fig. 31. Genital region from below. Figs. 32-34. Dysderina Humph- 
rey i sp. nov. Fig. 32. Eyes from above. Fig. 33. Sternum from below. Fig. 
34. Epigynal area from below. 



Description. Total length 1.83 mm. Carapace 0.91 mm long; 
0.73 mm wide opposite interval between first and second coxae 
where it is widest; 0.4 mm tall; other features essentially typical of 
males in the region under study. Eyes: eye group (Fig. 28) occu- 
pies about three-fourths of width of carapace at level of PE; seen 
from above, posterior row moderately recurved; ratio of eyes 
ALE : PME : PLE = 9 : 9 : 10 (some irregularities in outline 
noted); ALE separated from one another by three-tenths of their 
diameter and barely separated from PLE and PME; PME con- 
tiguous for more than one-fourth of their circumference and barely 
separated from PLE; height of clypeus equal to about three-fifths 
of the diameter of ALE. Chelicerae typical of males in the region 
under study; maxillae appear to be grooved and possibly divided 
distally. Lip wide at base and much narrowed distally. Sternum: 



16 breviora No. 296 

granulate; with a deep transverse groove shortly behind procurved 
sternal suture; with grooves, intricately developed in mam species, 
here very much reduced and simplified; fourth coxae separated by 
nine-sevenths of their width. Legs: tibial index of first and fourth 
legs 9; 4123 in order of length; first femur with ventral spines 
0-0-lr-lp-lp; first tibia with ventral spines 2-2-2-2-2 (last two little 
more than stiff bristles); first metatarsus with ventral spines 2-2- 
lp-O; second femur apparently only with ventral spines 0-0-1 p-0; 
second tibia and second metatarsus essentially as in first with re- 
spect to ventral spines; third and fourth legs lacking true spines. 
Palp: all segments as usual in the genus except tarsus with its 
complicated embolus and associated apophyses (Figs. 29-30). 
Abdomen: essentially as described for /). meridina sp. now; genital 
region obscure but apparently distinctive (Fig. 31 ). 

Records. One male paratype. somewhat damaged, accompanies 
the holotype. A female, recently moulted, is also with the holotype 
but is not in good condition for description. 

Dysderina humphreyi sp. nov. 
Figures 32-34 

Holotype, The female holotype is from Boquete. Panama, Au- 
gust, 1954. The species is named after Richard L. Humphrey, 
M.D., who, as an undergraduate student, was my assistant during 
a period of field work in Panama in the summer of 1954. 

Description. Total length 2.24 mm. Carapace 0.97 mm long; 
0.75 mm wide opposite second coxae where it is widest; 0.41 mm 
tall; somewhat less regularly arched along median dorsal region 
than usual in females of the genus; otherwise essentially typical of 
the genus. Eyes: posterior row occupies about three-fourths of 
width of carapace at that level; almost straight but very slightly re- 
curved as viewed from above (Fig. 32); ratio of eyes ALE : 
PME : PLE = 8.5 : 8.5 : 7.5 (boundaries of eyes, especially of 
ALE. are quite irregular); ALE separated from one another by 
about their radius; ALE barely separated from PLE and from PME 
by about one-fourth of their diameter. Posterior row of eyes very 
closely crowded together; PME contiguous for one-fourth of their 
circumference; barely separated from PLE. Posterior row of eyes 
wider than anterior row in ratio of about 6:5; viewed from in 
front, posterior row of eyes definitely procurved; height of clypeus 
equal to about five-eighths of the diameter of ALE. Chelicerae, 
maxillae, and lip essentially as described for D. silvatica Chickering 
(1951) as far as observed. Sternum: general features as usual in 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 17 

the genus; fourth coxae separated from one another by a little less 
than 1.5 times their width; pattern of grooves essentially as repre- 
sented in Figure 33. Legs: 4123 in order of length; tibial index of 
first leg 10, of fourth leg 9. Leg spines: first femur with five 
ventral spines, two on promargin and three on retromargin, not 
regularly paired; first tibia with five pairs of ventral spines; first 
metatarsus with five ventral spines irregularly placed; second femur 
with four ventral spines, one on promargin and three on retro- 
margin; second tibia with four pairs of ventral spines; second meta- 
tarsus with five ventral spines, three on promargin and two on 
retromargin; legs three and four apparently without true spines. 
Abdomen: scuta, pedicel, tracheal spiracles, etc., all essentially 
typical of females of the genus. Epigynal area: very simple as 
usual; essentially as represented in Figure 34. Color in alcohol: 
essentially typical of the genus with no unusual features. 

This species appears to be closely related to both D. plena 
O. P.-Cambridge and D. silvatica Chickering but the shape of the 
epigynal area is somewhat different than in the two previously 
known species and the dots along the posterior border are com- 
pletely lacking. 

Records. Eight paratype females collected August 4-11, 1954, 
in the close vicinity of Boquete, Panama. 

Dysderina improvisa sp. nov. 
Figures 35-36 

Holotype. The female holotype is from Barro Colorado Island, 
Panama Canal Zone; collected in a Berlese funnel by Dr. James 
Zetek, May-October, 1946. The name of the species is a Latin ad- 
jective meaning unexpected. 

Description. Total length 2.26 mm, exclusive of the slightly 
extended spinnerets. Carapace 0.95 mm long; 0.77 mm wide op- 
posite second coxae where it is widest; 0.4 mm tall; otherwise 
essentially as in D. silvatica Chickering (1951). Eyes: posterior 
row occupies about three-fourths of width of carapace at that level 
(Fig. 35); viewed from above, gently recurved. Ratio of eyes 
ALE : PME : PLE = 8.5 : 7.5 : 7. ALE separated from one 
another by a little more than their radius; contiguous to PLE; 
separated from PME by nearly one-fourth of their diameter; PME 
contiguous for about one-fourth of their circumference; barely sepa- 
rated from PLE; posterior row wider than anterior row in ratio of 
6:5; viewed from in front, posterior row definitely procurved. 
Height of clypeus equal to about three-fourths of the diameter of 



18 



HREVIORA 



No. 296 



ALE. Chelicerae, maxillae and lip essentially as described for D. 
silvatica C flickering ( 1951 ). Sternum: general features as usual in 
the genus; pattern of grooves rather complicated; fourth coxae 
separated from one another by about seven-fourths of their width. 
Legs: 4123 in order of length; tibial index of first leg 9, of fourth 
leg 10. Leg spines: first femur with irregularities of ventral spines 
on right and left; first tibia with five pairs of ventral spines; first 
metatarsus with three pairs of ventral spines; second femur with 
ventral spines lr-lr-2-lp; second tibia with ventral spines 2-2-2- 
2-1 p; second metatarsus with two pairs of ventral spines; third and 
fourth legs with few weak spines. Abdomen: essentially typical of 
females of the genus. Epigynal area: \er\ simple as usual; essen- 
tially as shown in Figure 36. Color in alcohol: also essentially 
typical of the genus with minor variations. 






37 



38 



40 



I igs. 35-36. Dysderina improvise sp. now Fig. 35. Eyes from above. Fig. 
36. Epigynal area from below. Figs. 37-39. Dysderina intempina sp. nov. 
Figs. 37-38. Distal end of left male palpal tarsus; prolateral ;ind retrolateral 
views, respectively. Fig. 39. Genital area from below. Figs. 40-41. Dysderina 
meridina sp. nov.; distal end of left male palpal tarsus; prolateral and retro- 
lateral views, respectively. 



D. improvisa sp. nov. is also regarded as closely related to D. 
silvatica Chickering but its epigynal area is distinctly oval in 
shape, as compared to the semicircular shape of this region in the 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 19 

previously known species. The pattern of grooves on the sternum 
appears to be somewhat distinctive also. 

Records. Seven female paratypes from Barro Colorado Id., 
Panama Canal Zone, were collected in a Berlese funnel by Dr. 
James Zetek, May-Oct., 1946. Six females were taken in the same 
locality as follows: June-Aug., 1936; April-May, 1942 (Zetek); 
June-Oct., 1946 (Zetek); Aug., 1954; January, 1958. One female 
was taken in the Canal Zone Forest Preserve, February, 1958. 

Dysderina intempina sp. nov. 
Figures 37-39 

Holotype. The male holotype is from Boquete, Republic of 
Panama, August, 1950. The name of the species is an arbitrary 
combination of letters. 

Description. Total length 2.15 mm. Carapace 0.99 mm long; 
0.79 mm wide opposite second coxae where it is widest; 0.53 mm 
tall; otherwise essentially typical of males of the genus. Eyes: pos- 
terior row occupies about five-sevenths of width of carapace at that 
level; slightly recurved as seen from above. Ratio of eyes ALE : 
PME : PLE = 8.5 : 8.5 : 9. ALE separated from one another by 
their radius; almost in contact with PLE; separated from PME by 
their radius. PME contiguous only for a short distance; only sepa- 
rated from PLE by a narrow line. Posterior row of eyes wider than 
anterior row in ratio of about 5 : 4. Height of clypeus equal to 
slightly less than the diameter of ALE. Viewed from in front, 
posterior row of eyes moderately procurved. Chelicerae: basal seg- 
ment 0.31 mm long; essentially typical of males of the genus; no 
dissection conducted because of scarcity of paratypes. Maxillae and 
lip: apparently typical of males of the genus. Sternum: grooves 
quite clearly delineated; very similar to those in D. obtina sp. nov.; 
fourth coxae separated by slightly more than their width. Legs: 
4123 in order of length; tibial index of first leg 11, of fourth leg 9; 
trichobothria observed but exact number and placement not deter- 
mined. Leg spines: first femur with ventral spines 0-lr-lr-2-lp-0; 
first tibia with five pairs of long slender spines; first metatarsus 
with three pairs of ventral spines; second femur apparently with one 
weak ventral and two weak prolateral spines; second tibia with 
four pairs of ventral spines; second metatarsus probably with two 
pairs of ventral spines; these spines tend to be offset prolaterally or 
retrolaterally. Palp: apparently typical of males of the genus ex- 
cept for the tarsus whose distinctive features are shown in Figures 
37-38. Abdomen: pedicel, scuta, etc., apparently typical of males 



20 breviora No. 296 

of the genus; genital region as shown in Figure 39. Color in alco- 
hoi: essentially typical of the genus with minor variations. 

Records. One paratype male was taken during the same period 
as the holotype and in the same locality. Females taken at the 
same time and in the same locality may also belong here but there 
can be no certainty at this time. 

Dysderina meridina sp. nov. 
Figures 40-4 1 

Holotype. The male holotype is from San Jose, Costa Rica (En- 
rique Schmidt). No date oi collection is given but. presumably, 
the specimen was taken fairly recently. The name of the species is 
an arbitrary combination of letters. The holotype will be deposited 
in the American Museum of Natural History, New York, N. Y. 
The abdomen of the holotype is detached from the cephalothorax 
but otherwise is in a oood state of preservation. 

Description. Total length 2 mm. Carapace 0.94 mm long; 0.75 
mm wide opposite second COXae where it is widest; about 0.4 mm 
tall; rises gradualk from PMF to opposite posterior border of sec- 
ond coxae and then descends steeply to posterior border where 
there is a narrow upturned shelf; surface finely granulate; median 
thoracic groove lacking. Eyes: si\ as usual; seen from above, 
posterior row gentl) recurved. Ratio of eyes ALE : PME : PLE =z 
8.5 : 8 : 7.5. ALE separated from one another by slightly more 
than their radius; barely separated from I'll ■; separated from 
PME bv a little more than one-half their radius. PME contiguous 
for about one-fourth of their circumference: separated from PLE 
by a line. Height of clypeus equal to slightly less than the diameter 
of ALE; clypeus only moderately lobed in the middle of the ventral 
border. C'helieerae. maxillae and lip: all apparently typical of 
males of the genus from the regions under study. Sternum: also 
essentially typical of the genus in this general region; sternal 
grooves less intricate than in several other species; sternal suture 
procurved; fourth coxae separated by about four-thirds of their 
width. Legs: 42=13 in order of length; tibial index of first leg 
11, of fourth leg 10; as far as observed, claws and trichobothria 
essentially as described for D. seclusa Chiekering (1951). Leg 
spines also essentially as described for D. seclusa. Palp: all seg- 
ments except tarsus simple and w ithout special modifications; tarsus 
with distinctive embolus and associated structures (Figs. 40-41). 
Abdomen: in general typical of the genus; dorsal and ventral scuta 
cover nearly the entire surface; spinnerets retracted and nearly in- 
visible; genital aperture typical of the genus. Color in alcohol: 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 21 

carapace, sternum and abdominal scuta all a rich reddish brown; 
body of palpal tarsus light yellowish. 

Records. One poorly preserved male paratype accompanies the 
holotype. No date of collection is given. The female is unknown. 

Dysderina obtina sp. nov. 
Figures 42-45 

Holotype. The male holotype is from El Volcan, Panama, Au- 
gust, 1950. The name of the species is an arbitrary combination of 
letters. 

Description. Total length 2.54 mm, including extended spin- 
nerets. Carapace 1.1 mm long; 0.88 mm wide opposite second 
coxae where it is widest; 0.52 mm tall; otherwise essentially typical 
of males of the genus. Eyes: posterior row occupies about two- 
thirds of width of carapace at that level; only slightly recurved as 
seen from above. Ratio of eyes ALE : PME : PLE = 9 : 8.5 : 8. 
ALE separated from one another by five-ninths of their greatest 
diameter; separated from PLE by about two-ninths of their di- 
ameter and from PME by one-third of their diameter. PME 
contiguous for about one-sixth of their circumference; separated 
from PLE by about one-eighth of the diameter of the latter. Pos- 
terior row of eyes wider than anterior row in ratio of about 5 : 4. 
Height of clypeus equal to about eleven-ninths of the diameter of 
ALE. Chelicerae, maxillae and lip essentially as described for 
D. abdita sp. nov. Sternum: only slightly longer than wide between 
second coxae; grooved in a distinct manner closely similar to the 
pattern shown by D. abdita sp. nov. and D. recondita Chickering 
(Fig. 42). Fourth coxae separated by nearly five-thirds of their 
width. Legs: 4123 in order of length; tibial index of first leg 10, 
of fourth leg 9; trichobothria observed on first femora and tibiae 
but number and placement not accurately determined. Leg spines: 
first femur with ventral spines 0-0-2-2-0 unevenly placed; first tibia 
with five pairs of ventral spines and a pair of distal bristles; first 
metatarsus with three pairs of ventral spines; second femur with two 
small ventral spines on promargin; second tibia with four pairs of 
ventral spines; second metatarsus as in first. Third and fourth 
legs devoid of true spines. Palp: general features as usual in males 
of the genus; specific tarsal features shown in Figures 43-44. Ab- 
domen 1.32 mm long exclusive of extended spinnerets; 0.92 mm 
wide slightly behind the middle; pedicel, scutta, etc., essentially as 
described for D. abdita sp. nov.; genital aperture as shown in 
Figure 45. Color in alcohol: also essentially as described for D. 
abdita sp. nov. with minor variations. 



->-> 



HREVIORA 



No. 296 



Records. \o paratypes have appeared in niv collection and the 
female is unknown. For some time the holotype was regarded as a 
deviate of />. recondita Chickering but careful attention to the male 
paip has convinced me that it must be considered to represent a 
new species as presented here. 




47 



. 42-45. Dysderina obtina sp. nov. Fig. 42. Sternum from below. 
I igs. 43-44. Distal end of left male palpal tarsus; nearly prolateral and 
retrolateral views, respectively. Fig. 45. Genital aperture from below. Figs. 
4h-49. Dysderina plena O. P.-Cambridge. Figs. 46-47. Left male palpal 
tarsus; nearly prolateral and retrolateral views, respectively. Fig. 48. Distal 
end of male palpal tarsus; nearly ventral view. Fig. 49. Epigynal area from 
below. Figs. 50-52. Dysderina potena sp. nov. Fig. 50. Eyes from above. 
Fig. 51. Ventral scutum from below. Fig. 52. Epigynal area from below. 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 23 

Dysderina plena O. P.-Cambridge 
Figures 46-49 

Dysderina plena O. P.-Cambridge, 1894: 143; 1896: 192, pi. 17, fig. 7; 
pi. 24, fig. 4. Male and female syntypes from Teapa, Mexico, are in 
the British Museum (Natural History). F. P.-Cambridge, 1899: 44; 
Banks, 1909: 196; Petrunkevitch, 1911: 125; Roewer, 1933: 185; 1942: 
283; Chickering, 1951: 211; Bonnet, 1956: 1638. 

As pointed out in 1951, the specimens reported from Panama by 
Banks (1929) and referred to this species really belong to other 
genera and species. The specimens reported by Williams (1941) 
and identified by Dr. W. J. Gertsch as Dysderina plena O. P.- 
Cambridge appear to be Dysderina silvatica Chickering (1951 ). 

The female from Teapa, Mexico, on loan from the British Mu- 
seum (Natural History), has an epigynal area (Fig. 49) very 
similar to that of D. silvatica Chickering. The intricate grooves on 
the sternum are somewhat different from those on D. silvatica 
Chickering but the two seem to be closely related. 

Male. From the British Museum (Natural History). Total 
length about 2.1 mm. Carapace 0.98 mm long. 0.78 mm wide 
opposite second coxae where it is widest; about 0.52 mm tall oppo- 
site interval between second and third coxae where it is tallest; 
descent nearly precipitous from highest point to posterior border; 
surface finely granulate; without a median thoracic groove or pit. 
Eyes: posterior row occupies a little more than three-fourths of 
width of cephalic region at this level; gently recurved as seen from 
above. Ratio of eyes ALE : PME : PLE = 10 : 9.5 : 10. All eyes 
are somewhat irregular in outline; PME and PLE are more oval 
than ALE. ALE separated from one another by nearly one-third 
of their diameter. PME contiguous for about one-fourth of their 
circumference; PLE nearly contiguous to PME posteriorly but 
separated anteriorly; PLE and ALE nearly contiguous; posterior 
row longer than anterior row in ratio of about 5:4; viewed from in 
front, posterior row procurved. Height of clypeus a little more than 
two-thirds of the diameter of ALE; border of clypeus strongly 
chitinized. Sternum finely granulate and intricately grooved about 
as usual in the genus. Genital aperture a narrow slit surrounded by 
a broad margin. Important features of the palp shown in Figures 
46-48. Other features essentially typical of males of the genus. 



24 breviora No. 296 

Dysderina potena sp. nov. 
Figures 50-52 

Holotype. The female holotype is from El Volcan, Panama. 
August. 1950. The name of the species is an arbitrary combination 
of letters. 

Description. Total length, including spinnerets. 2.8 mm. Cara- 
pace 1 .08 mm long; 0.97 mm wide opposite second coxae where it 
is widest: 0.4S mm tall; gradually arched from PLE to posterior 
border; finely but irregularly granulate; without any visible thoracic 
groove or pit. Eyes: seen from above, posterior row straight if 
measured b\ posterior borders; posterior row occupies about two- 
thirds of width of carapace at that level; seen from in front, pos- 
terior row gentl) procurved. Ratio oi eyes ALE : PME : PLE = 
9.5 : 9 : 8. ALE separated from one another by a little more than 
their radius; barely separated from PLE (Fig. 50). Height of cly- 
pens equal to a little more than the diameter of ALE. Chelieerae. 
maxillae and lip; essential!) typical of females of the genus as ob- 
served in this studs. Sternum only slightly longer than wide; with 
the usual intricate grooves; quite convex; fourth coxae separated 
b\ about 1.5 times their width, legs: 4123 in order of length; 
tibial index of first and fourth legs 9; spines on legs essentially as 
described for />. silvatica Chickering (1951). Abdomen: essen- 
tialK typical oi females of the genus with minor variations; ventral 
scutum as shown in Figure 5 I . Epigynal area: appears to have cer- 
tain obscure but distinctive features (Fig. 52). Color in alcohol: 
carapace irregularly darkened along posterior declivity and lateral 
sides; generally a rich reddish brown. 

This species appears to be closely related to /). plena O. P. 
Cambridge but differs from that species most conspicuously with 
respect to its epigynal area which is in the form of a modified oval 
opening ( Fig. 52 | with a distinct anterior border and a series of 
faintly indicated granules along the posterior border. The height 
of the clypeus is somewhat greater than usual in the genus. 

Ren >nls. Two paratype females taken August, 1950, in vicinity 
of El Volcan, Panama. One female from Boqucte. Panama, Au- 
gust, 1954 is somewhat doubtfully assigned to this species. 

Dysderina princeps Simon 

Figures 53-54 

Dysderina princeps Simon, 1891, 557. The female holotype from St. Vin- 
cent. B. W. I., is in the British Museum (Natural History). Simon, 
1893: 290, 304; Petrunkevitch, 1911: 125; Roewer, 1942: 283; Bonnet, 
1956: 1638. 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 25 

So far as I have been able to determine, the species has never 
been correctly reported since its first recognition. In 1893, how- 
ever, Simon published his figure 260 which is a drawing of a male 
palp and it is labelled D. princeps. The figure closely resembles the 
original drawing (fig. 1 ) accompanying the brief description of 
D. principalis Simon. 

The following facts are derived from the study of a female from 
St. Vincent, B. W. I. on loan from the British Museum (Natural 
History) : Total length 2.66 mm (the author of the species gave the 
length as 4 mm). Carapace 1.17 mm long; 0.97 mm wide oppo- 
site second coxae where it is widest; otherwise essentially typical 
of females of the genus as seen during this study. Eyes in general 
as usual in the genus (Fig. 53) ; ratio of eyes ALE : PME : PLE = 
9 : 8.5 : 9.5. Clypeus somewhat higher than that in D. spinigera 
Simon but otherwise closely similar. Sternum intricately but not 
deeply grooved. Legs 1 and 2 with the usual long ventral spines 
essentially typical of the genus. Abdomen with the typical scuta 
and related parts. Epigynal area more distinctive than usual (Fig. 
54). 

Dysderina principalis (Keyserling) 
Figures 55-56 

Oonops principalis Keyserling, 1882: 296, pi. 11, fig. 16. The male holotype 
from Colombia is in the British Museum (Natural History). 

Dysderina principalis,Simon, 1891, 557, pi. 42, fig. 1. Simon, 1893: 304; 
Petrunkevitch, 1911: 126; 1928: 87; Roewer, 1942: 283; Bonnet, 1956: 
1638. (Not D. principalis Simon from St. Vincent, B. W. I.) 

Simon was correct in transferring Oonops principalis Keyserling 
to his new genus Dysderina but he was in error when he identified a 
species of this new genus from St. Vincent, B. W. I., as being D. 
principalis described from Colombia. Apparently Keyserling had 
only the male upon which to base his species. This specimen has 
been on loan from the British Museum (Natural History). It is 
dismembered, but the right palp is in good condition and estab- 
lishes Simon's error very clearly. In order to aid in clarifying this 
confusion I am offering two figures of the palpal tarsus (Figs. 
55-56). Keyserling's small figure 16 closely resembles those I am 
providing. 

Dysderina recondita Chickering 
Figures 57-59 

Dysderina recondita Chickering, 1951: 217, fig. 4. The male holotype and 
two paratypes from Boquete, Panama, July, 1939, are in the Museum 
of Comparative Zoology. 



26 



BREVIORA 



No. 296 



During my brief visit to Boquete, Panama, in August, 1954, I 
collected eleven additional males. The female is unknown but it 
seems probable that it is among the different females now recog- 
nized from the mountainous regions of Panama. Figures 58-59 are 
added to the original drawing of the male palp; Figure 57 shows 
the essential features of the sternal grooves. 




v 53-54. Dysderina princeps Simon. Fig. 53. Eyes from in front. 
I ig, 54. Epigynal area from below. Figs. 55-56. Dysderina principalis 
< Keyserling). Two views of right m;ile palpal tarsus. Figs. 57-59. Dysderina 
recondita Chickering. Fig. 57. Sternum from hclow. Figs. 58-59. Two views 
of distal end of left male palpal tarsus. 



Dysderina rigida sp. nov. 
Figures 60-61 

Holotype. The female holotype is from El Volcan, Panama, Au- 
gust, 1950. The name of the species is a Latin adjective referring 
to its strongly chitinized scuta, pedicel and other outer parts. 

Description. Total length 2.49 mm. Carapace 1.05 mm long; 
0.88 mm wide opposite second coxae where it is widest; 0.44 mm 
tall; other features essentially as described for D. silvatica Chicker- 
ing (1951). Eyes: posterior row gently recurved, seen from 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 27 

above; occupies about five-sevenths of width of carapace at that 
level. Ratio of eyes ALE : PME : PLE = 9:9:8. ALE separated 
from one another by slightly more than their radius; only slightly 
separated from PLE and from PME by about one-fourth of their 
diameter. PME contiguous along about one-fourth of their circum- 
ference and longer than wide in ratio of about 9 : 7.5; separated 
from PLE by one-sixth of their long diameter. Posterior row wider 
than anterior row in ratio of about 13 : 11. Height of clypeus 
slightly greater than the diameter of ALE. Chelicerae: vertical, 
essentially parallel; outer margin with a shallow groove at end of 
upper third and a slight swelling near middle; a single tooth along 
margin of fang groove appears to be definitely promarginal (ob- 
served on paratype); otherwise essentially as reported for D. sil- 
vatica (1951). Maxillae, lip and sternum also essentially as re- 
ported for D. silvatica (1951) except for the system of grooves on 
sternum (Fig. 60). Legs: 4123 in order of length; tibial index of 
first and fourth legs 9; trichobothria observed on palpal tibia and 
tarsus but number and placement not accurately determined. Leg 
spines: first femur with five ventral spines not evenly paired; first 
tibia with five pairs of ventral spines; first metatarsus with three 
pairs of ventral spines. Second leg with ventral spines nearly as on 
first leg; with only minor differences. Abdomen: spinnerets with a 
lightly chitinized ventral semicircular sclerite partially surrounding 
these organs; other features essentially typical of females of the 
genus. Epigynal area: simple as usual but apparently with obscure 
distinctive features (Fig. 61). Color in alcohol: essentially typical 
of the genus with minor variations. 

This species is also regarded as being rather closely related to 
D. plena O. P.-Cambridge. It differs from that species most con- 
spicuously with respect to its epigynal area which is in the shape 
of a modified oval region with a narrow, transverse area along its 
posterior border (Fig. 61). The pattern of grooves on its sternum 
also appears to be somewhat distinctive (Fig. 60). 

Records. Several female paratypes taken with the holotype, El 
Volcan, Panama, August, 1950. 

Dysderina seclusa Chickering 
Figures 62-64 

Dysderina seclusa Chickering, 1951: 213, fig. 5. The male holotype and 
five male paratypes from Barro Colorado Island, Panama Canal Zone, 
are in the Museum of Comparative Zoology. All of these were taken 
in a Berlese funnel by Dr. Zetek, July, 1943 -March, 1944, and June- 
October, 1946. 



28 



HREVIORA 



No. 296 



Since the original description was written, I have added four 
males to the collection as follows: June-October, 1946 (Zetek); 
June-August, 1949 (Zetek); August, 1954. I have been unable to 
match these males with females in any satisfactory manner. With 
the hope of further clarifying the status of the species three addi- 
tional figures are offered. 

\^ U, 1 fT\ W- 




1^-Ceo 





63 



67 



•■ 



.->— ,- 




\ 



62 



x ^MW,/// 



Figs. 60-61. Dysderina rigida sp. nov. Fig. 60. Sternum from below. Fig. 
61. Epigynal area from belou. Figs. 62-64. Dysderina seclusa (bickering. 
Fig. 62. Sternum from below. I i;^. 63. Left male palpal tarsus: ventral view. 
Fig. 64. Distal end of left male palpal tarsus, nearly dorsal view. Figs. 
65-68. Dysderina simla sp. nov. Fig. 65. Left male palpal tarsus; retrolatcral 
view. F - i lt . 66. Distal end of left male palpal tarsus; nearly ventral view. Fig. 
67. Genital area of male holotype. Fig. 68. Epigynal area of female paratype. 



Dysderina silvatica Chickering 

Dysderina silvatica Chickering, 1951: 217, fig. 6. The female holotype and 
numerous paratypes from several localities in the Panama Canal Zone 
and El Valle, Panama, are in the Museum of Comparative Zoology. 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 29 

Since my report in 1951 I have added numerous females of this 
species to the collection from the following localities: Barro Colo- 
rado Island, Canal Zone Forest Preserve and Summit Gardens all in 
Panama Canal Zone. Although the evidence is not conclusive, I 
think it probable that these females belong with D. dura Chickering. 

Dysderina simla sp. nov. 
Figures 65-68 

Holotype. The male holotype is from Simla, Arima Valley, 
Trinidad, W. I., April 5, 1964. The name of the species is a noun 
used in apposition after the type locality. 

Description. Total length 1.69 mm. Carapace 0.88 mm long; 
0.75 mm wide opposite second coxae where it is widest; 0.47 mm 
(all; median thoracic groove lacking; smooth along broad, median 
area from PLE to beginning of posterior declivity; irregularly 
granulate along lateral sides and much of posterior declivity; fits 
closely to anterior end of abdomen which is unusual in this genus. 
Eyes: six as usual in a compact group which occupies nearly 
three-fourths of width of carapace at level of PE; viewed from 
above, posterior row slightly recurved. Ratio of eyes ALE : PME : 
PLE = 8:9:9. ALE separated from one another by slightly 
more than their radius; other relationships of eyes typical of the 
genus in the region under study. Height of clypeus equal to about 
three-fourths of diameter of ALE. Chelicerae, maxillae and lip all 
appear to be typical of the genus. Sternum: granulate; intricately 
but not deeply grooved and lobed; the usual anterior, transverse 
groove is complete, conspicuous and only slightly behind procurved 
sternal suture; only slightly longer than wide just behind second 
coxae; fourth coxae separated by nearly twice their width. Legs: 
4123 in order of length; tibial index of first leg 10, of fourth leg 11. 
Spines on legs about as usual in males of the genus. Palp: all seg- 
ments simple and without special modifications except the tarsus 
which has distinctive features (Figs. 65-66). Abdomen: typical 
of the genus in general; with dorsal and ventral scuta less strongly 
chitinized than usual (all paratypes appear to be the same in this 
respect); genital region quite clear and rather distinctive (Fig. 
67). Color in alcohol: carapace essentially as described for D. 
craneae sp. nov.; legs and mouth parts a variable yellowish; 
sternum a deeper yellowish; abdomen with dorsal and ventral scuta 
a light yellowish in contrast to the usual much darker coloration; 
epigastric scutum somewhat darker; the free areas not covered by 
scuta are whitish. 



30 breviora No. 296 

Female paratype. Total length 1.98 mm. Carapace 0.9 mm 
long; 0.84 mm wide opposite interval between second and third 
coxae where it is widest; 0.36 mm tall; surface granulate essen- 
tially as in male. Cephalothorax closely contiguous to abdomen as 
in male. Eyes essentially as in male. Chclicerae, maxillae and lip: 
all essentially typical of females of the genus. Sternum: essentially 
as in male but with granulation more extensive; fourth coxae sepa- 
rated b\ neark twice their width. Legs: 41 23 in order of length; 
tibial index of first and fourth legs 1 1; spines essentially typical of 
the genus. Abdomen: general appearance as usual in the genus; 
dorsal scutum somewhat less extensive than usual; epigastric 
scutum about as usual: ventral scutum much smaller than usual 
and reaches slightly less than half way from genital groove to spin- 
nerets; a narrow, ventral, semicircular sclcrite partly surrounds the 
spinnerets. Epigynal area: quite distinctive but difficult to repre- 
sent accurately in a drawing (Fig. 68). Color in alcohol: all parts 
except the abdomen as in male; because of the smaller scuta much 
more o\~ the abdomen appears nearly white than in the male; 
posterior to the ventral scutum are several vague darker spots which 
in some paratvpes are quite distinct brownish, irregular spots. 

Records. The described female paratype was taken with the 
holotype. Three male and seven female paratvpes are in the col- 
lection from Simla. Arima Valley, and along the side of the road 
from Simla to Blanchesscuse. Trinidad, W. I., March 31 -April 25, 
1964. 

Dysderina soltina sp. nov. 

Figures 69-75 

Dysderina principalis, -Simon, 1891: 557, pi. 42, fig. 1. Male and female 
syni\pes from St. Vincent B. W. I. are in the British Museum (Natural 
History I. (Not D. principalis (Keyserling) from Colombia.) 

In his study of the genus Dysderina from St. Vincent, B. W. I., 
Simon selected what he considered to be a male and a female of 
D. principalis (Keyserling), gave a brief description and furnished 
a figure of the male palp. As a result of a study of these specimens 
together with an example of Keyserling's species from Colombia 
on loan from the British Museum (Natural History), I am obliged 
to disagree completely on the identification of these specimens. 
They seem to me to represent a new species which is described 
below. 

Holotype. The male holotype is from St. Vincent, B. W. L, and 
will be returned to the British Museum (Natural History). The 
name of the species is an arbitrary combination of letters. 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 31 

Description. Total length 1.91 mm (recorded as 3.5 mm in 
Simon's description). Carapace 0.92 mm long; 0.73 mm wide 
opposite second coxae; 0.35 mm tall; only gradually narrowed be- 
hind eyes; surface irregularly granulate as usual; otherwise typical 
of the genus. Eyes: six as usual; seen from above, posterior row 
gently recurved; ratio of eyes ALE : PME : PLE = 8 : 7 : 7 (out- 
lines poorly defined). ALE separated from one another by their 
radius. PME contiguous for about one-fourth of their circum- 
ference; barely separated from PLE. Height of clypeus equal to 
about five-eighths of the diameter of ALE. Chelicerae, maxillae 
and lip typical of the genus as far as observed. Sternum: typical of 
the genus in general; major and minor grooves and depressions 
moderately complex; extended as usual between fourth coxae and 
articulated with pedicel; fourth coxae separated by five-thirds of 
their width. Legs: 4123 in order of length; tibial index of first leg 
10, of fourth leg 9; ventral spines on first leg as shown in Figures 
71-73; fewer ventral spines on second leg. Palp: all segments ex- 
cept tarsus simple and unmodified as usual; tarsus considerably 
inflated and with distinctive terminal structures difficult to observe 
clearly (Figs. 69-70). Abdomen: all parts in general typical of 
males of the genus; scuta cover nearly the entire surface. Color in 
alcohol: also typical of the genus with few minor variations. 

Female paratype. This is apparently the specimen which Simon 
regarded as the female of D. principalis (Keyserling). For the 
present I am regarding it as the female of the new species D. sol- 
tina. Total length 1.98 mm (Simon gave length as 4 mm). Cara- 
pace 0.9 mm long; 0.73 mm wide; 0.4 mm tall; general features 
typical of females of the genus. Leg spines: the first femur has five 
ventral spines, two on promargin and three on retromargin (Fig. 
74); other segments with spines much as in male. Abdomen with 
scuta and other parts essentially typical of the genus. Epigynal 
area: obscurely distinctive as usual (Fig. 75). 

Records. As far as I have been able to determine the two 
specimens described here are the only members of the species 
known at present. 

Dysderina spinigera Simon 
Figures 76-79 

Dysderina spinigera Simon, 1891, pi. 42, figs. 2-3. Male and female syn- 
types from St. Vincent, B. W. I., are in the British Museum (Natural 
History). Simon, 1893: 304; Petrunkevitch, 1911: 126; Roewer, 1942: 
283; Bonnet, 1956: 1639. 



32 



BREVIORA 



No. 296 




I igs. 69-75. Dysderina soltina sp. nov. Fig. 69. Left male palpal tarsus; 
nearly dorsal view. Fig. 70. Distal end of left male palpal tarsus; retrolatcral 
view (more enlarged). Figs. 71-73. Ventral spines on first left femur, tibia 
and metatarsus, respectively. Hg. 74. Ventral spines on first left femur of 
female paratype. Hg. 75. Epigynal area of female paratype from below. 
Figs. 76-79. Dysderina spinigera Simon. Fig. 76. Eyes from in front. Fig. 
77. Left male palp; retrolateral view. Fig. 78. Left male palpal tarsus; dorsal 
view. Fig. 79. Epigynal area from below. 



Simon reported both sexes of this species from St. Vincent and 
Venezuela where he regarded it as widespread. I have not seen the 
specimens from Venezuela but. on the basis of my study of this 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 33 

genus, I consider it unlikely that they are the same as these from 
St. Vincent. I was disappointed in my failure to collect any speci- 
mens of the genus Dysderina during my brief visit to this island in 
October, 1966. 

The following facts relating to this species have been recorded 
as a result of my study of a male and two females on loan from 
the British Museum (Natural History). 

Male. Total length 1.61 mm (Simon gave length of male as 4 
mm). Carapace 0.86 mm long; 0.66 mm wide opposite second 
coxae where it is widest; otherwise essentially typical of the genus 
as observed in this study. Eyes: essentially as shown in Figure 76; 
ratio of eyes ALE : PME : PLE = 10 : 9 : 9 (outlines irregular); 
ALE closer together than usual in the genus; clypeus with strongly 
chitinized ventral border and conspicuously lobed ventrally in 
front (Fig. 76). Sternum moderately grooved in fairly typical 
manner. Legs essentially typical in respect to spination and rela- 
tive lengths; trichobothria observed but exact distribution unde- 
termined. Palp: quite distinctive; appearance very different from 
Simon's figure 3, 1891; essentials shown in Figures 77-78. Abdo- 
men: with typical dorsal and ventral scuta; genital aperture a 
simple, minute slit. 

Female. Total length, exclusive of projecting spinnerets and 
slightly exposed chelicerae 2.09 mm; including posterior spin- 
nerets and slightly exposed chelicerae, length is 2.2 mm (Simon 
gave length as 4.5 mm). General form and external features essen- 
tially like those of male. Abdomen: with dorsal and ventral scuta 
somewhat less extensive than in male; epigynal area very distinc- 
tive; unlike that seen in other species (Fig. 79). 

Dysderina watina sp. nov. 
Figures 80-84 

Holotype. The male holotype is from Turrialba, Costa Rica, 
July 25-Aug. 15, 1965. The name of the species is an arbitrary 
combination of letters. 

Description. Total length 2.24 mm. Carapace 1.03 mm long; 
0.86 mm wide opposite second coxae where it is widest; 0.45 mm 
tall opposite second coxae; moderately grooved immediately be- 
hind PE; regularly arched from groove to beginning of moderately 
steep posterior declivity; surface irregularly granulate; otherwise 
essentially typical of the genus. Eyes: six in two rows as usual; 
posterior row gently recurved, seen from above; posterior row 
occupies about seven-ninths of width of carapace at that level. 



34 breviora No. 296 

Ratio of eyes ALE : PME : PLE =10:8:9. ALE separated 
from one another by about their radius, from PLE by nearly one- 
fifth of their diameter and from PME by about three-tenths of their 
diameter. PME barely separated from one another and from PLE 
by about three-sixteenths of their diameter. Height of clypeus 
nearly equal to long diameter of ALE. Chelicerae. maxillae and 
lip typical of the genus as far as observed. Sternum: clearly but 
less intricately grooved than in several other species; posterior 
coxae separated from one another bv about 1.5 times their width; 
otherwise essentially typical of the genus. Legs: 4123 in order of 
length; tibial index of first leg 8, of fourth leg 7; spines con- 
spicuous! v long and slender but otherwise essentially tvpical of 
males of the genus. Palp: all segments except tarsus simple and 
unmodified; tarsus with distinctive distal features as shown in Fie- 
ures 80-81. Abdomen: essentially typical of males of the genus. 
Color in alcohol: lighter than usual in the genus; carapace, pedicel 
and sternum a medium yellowish brown; abdomen and legs yellow- 
ish with variations. 

Female paratype. Total length 2.75 mm exclusive of slightly 
projecting spinnerets and chelicerae; including these parts total 
length is 2.9 mm. Carapace 1.1 mm long; 0.92 mm wide opposite 
interval between second and third coxae; about 0.44 mm tall; 
rather deeply grooved immediately behind PE and then gently 
arched to posterior declivity; otherwise typical o\' the genus. Eves: 
PME contiguous for neark one-fourth of their circumference; 
ratio of eyes ALE : PME : PLE = 10 : 10 : 9.5; other features 
essentially as in male. Height of clypeus immediately below ALE 
equal to slightly less than the diameter of these eyes; clypeus con- 
spicuously lobed in middle between bases of chelicerae. Cheli- 
cerae, maxillae and lip essentially typical of females of the genus. 
Sternum: with the usual pattern of major grooves and other incon- 
spicuous irregularities; fourth coxae separated by five-thirds of 
their width. Legs: 4123 in order of length; tibial index of first leg 
9, of fourth leg 10; spines of first and second legs conspicuously 
long but essentially typical of females of the genus with minor 
variations. Abdomen: scuta, pedicel, spiracles, etc. nearly typical 
of females of the genus (Figs. 82-83); a very clear narrow sclerite 
partially surrounds spinnerets on ventral side; what appears to be 
a chitinous dot marks the position of the reduced colulus. Epigynal 
area: obscurely distinctive (Fig. 84). Color in alcohol: much more 
colorful than in male; carapace, sternum and abdominal scuta all 
a rich reddish brown with lateral sides of carapace irregularly dark- 
ened by granulations; areas of abdomen not covered by scuta clear 
white; legs brownish yellow with variations. 



1968 



DYSDERINA IN C. AMERICA AND W. INDIES 



35 



Records. The described female paratype together with three 
other females were all taken from weed and hay debris in the same 
general locality from which the male holotype was taken, Turrialba, 
Costa Rica, July 25-August 15, 1965. It seems reasonable to as- 
sume that the male holotype and females belong together but there 
can be no absolute certainty. 




86 



80 



Figs. 80-84. Dysderina watina sp. nov. Figs. 80-81. Distal end of left male 
palpal tarsus; prolateral and retrolateral views, respectively. Figs. 82-83. 
Dorsal and ventral views of abdomen of female paratype. Fig. 84. Epigynal 
area of female paratype, from below. Figs. 85-86. Dysderina zinona sp. nov. 
Fig. 85. ALE and clypeus from in front. Fig. 86. Epigynal area from below 



Dysderina zinona sp. nov. 
Figures 85-86 

Holotype. The female holotype is from St. Vincent, B. W. I., 
Mt. Soufriere, June 1, 1965 (R. T. Bell). The name of the species 
is an arbitrary combination of letters. The specimen serving here 
as the holotype of a new species is somewhat damaged but its most 
important features are still in good condition. 



36 breviora No. 296 

Description. Total length about 2.28 mm. Carapace 1.08 mm 
long; 0.86 mm wide opposite second coxae where it is widest; 
0.54 mm tall; irregularly granulate as usual; otherwise essentially 
typical of females of the genus. Eyes: in general, as usual in the 
genus; viewed from above, posterior row gently recurved and occu- 
pies about seven-tenths of width of carapace at that level. Ratio of 
eyes ALE : PME : PLE = 9 : 8.5 : 8. ALE separated from one 
another by nearly their radius; from PLE by nearly one-half their 
radius and by slightly more than this from PME, which are con- 
tiguous for about one-fourth of their circumference and barely 
separated from PLE. Clypeus conspicuously lobed in front (Fig. 
85); height immediately below ALE equal to nearly two-thirds of 
the diameter of these eves. Chelicerae. maxillae and lip typical of 
females of the genus as far as observed. Sternum: fairly typical 
also; major grooves and secondary depressions make a rather intri- 
cate pattern; fourth coxae separated by a little more than their 
width. Legs: 4123 in order oi length; tibial index of first leg 
10. o\' fourth leg l ); first leg with five ventral spines on femur as in 
Figure 74; first tibia with five ventral spines on promargin and 
tour on retromargin; first metatarsus with three \entral spines on 
promargin and two on retromargin. Second leg with somewhat 
fewer ventral spines than occur on first leg. Third and fourth legs 
without true spines. Abdomen: considerably damaged but ap- 
parently quite typical oi females of the genus. Epigynal area: un- 
damaged; rather distinctive (Fig. 86). Color in alcohol: carapace a 
rich reddish brown; sternum somewhat lighter; legs yellowish 
brown with variations; abdominal scuta probably much like cara- 
pace but somewhat lighter like the sternum. 

D. zinona sp. nov. is regarded as closely related to D. princeps 
Simon. The epigynal area (Fig. 86) is quite different from that 
region in D. princeps Simon (Fig. 54) and much simpler. The 
clypeus is also conspicuously lobed in front (Fig. 85). 

Records. No paratypes have appeared in the collection. 

BIBLIOGRAPHY 

Banks, Nathan 

1909. Arachnida from Costa Rica. Proc. Acad. Natur. Sci. Phila- 
delphia, April, 1909: 194-234, 2 pis. 
1929. Spiders from Panama. Bull. Mus. Comp. Zool.. 69: 53-96, 4 
pis. 
Bonnet, Pierre 

1956. Bibliographia Araneorum. Toulouse. Vol. 2 (2). 



1968 DYSDERINA IN C. AMERICA AND W. INDIES 37 

Bryant, Elizabeth 

1942. Notes on the spiders of the Virgin Islands. Bull. Mus. Comp. 

Zool., 89: 317-363, 3 pis. 
1948. The spiders of Hispaniola. Bull. Mus. Comp. Zool., 100: 331- 

447, 12 pis. 

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

1889- Arachnida-Araneida. Vols. I-II. In: Biologia Centrali-Ameri- 

1905. cana. Dulau & Co., London. 
Chickering, Arthur M. 

1951. The Oonopidae of Panama. Bull. Mus. Comp. Zool., 106: 
207-245,31 figs. 

COMSTOCK, J. H. 

1940. The Spider Book. Revised and edited by W. J. Gertsch. Double- 
day, Doran & Company, Inc., New York. 

Gertsch, W. J. 

1941. Report on some arachnids from Barro Colorado Island, Canal 
Zone. Amer. Mus. Novit., No. 1146: 1-14, 4 pis. 

Kaston, B. J. 

1948. Spiders of Connecticut. State of Connecticut. State Geological 
and Natural History Survey, Bull. 70: 1-874. 
Keyserling, Graf Eugen von 

1881. Neue Spinnen aus Amerika. III. Verhandl. Zool. Bot. Ges. 
Wien, 31,269-314, pi. 11. 

Petrunkevitch, Alexander 

1911. A synonymic index-catalogue of spiders of North, Central, 

South America, etc. Bull. Amer. Mus. Natur. Hist., 29: 1-809. 
1928. Systema Araneorum. Trans. Connecticut Acad. Arts and Sci., 

29: 1-270. 
1939. Catalogue of American Spiders. Pt. 1. Trans. Connecticut 

Acad. Arts and Sci., 33: 135-338. 

ROEWER, C. FR. 

1933. Araneen aus mexikanischen Ameisenwohnpflanzen. Zool. Anzei- 
ger, 102: 183-188. 

1942. Katalog der Araneae. 1: 1-1040. Bremen. 

Simon, E. 

1891. On the spiders of the island of St. Vincent. Pt. 1. Proc. Zool. 

Soc. of London, Nov. 17, 1891: 549-575. 
1892- Histoire naturelle des Araignees. Deuxieme Edition. 2 Vols. 
1903. Librarie Encyclopedique de Roret, Paris. 

Williams, Eliot C, Jr. 

1941. An ecological study of the floor fauna of the Panama rain 
forest. Bull. Chicago Acad. Sci., 6: 63-124. 

(Received 27 September 1967.) 



BREVIORA 



Museunri of Comtipsiratlve Zoology 

Cambridge, Mass. 11 October, 1968 Number 297 

POPULATION STRUCTURE OF THE 

ASTHENES FLAMMULATA SUPERSPECIES 

(AVES: FURNARNDAE) 

Francois Vuilleumier 1 



INTRODUCTION 

While conducting a survey of speciation phenomena in twenty- 
five genera of Andean birds, I found that the existing classifications 
of these birds were often insufficient for my purposes. I had to un- 
dertake systematic studies in each genus before I could proceed to 
an analysis of species formation. In view of the groundwork already 
laid by Hellmayr, Chapman, Zimmer, Peters, and others, it may 
seem surprising that a student of speciation should find it necessary 
to do so much systematic work. However, these earlier systematists 
were primarily concerned with the purely formal aspects of assign- 
ing samples to species and subspecies, rather than interested in the 
process of speciation; their works contain little information on 
population structure (in the sense of Mayr, 1959:294) or species 
relationships. My own taxonomic work on Andean birds was de- 
signed to fill this gap. 

This is the first in a proposed series of papers on these Andean 
birds. My purpose has been to gain an understanding of the process 
of speciation in each genus sampled through study of population 
structure and interspecies relationships, and then to synthesize this 
information, arriving at a general picture of speciation phenomena 
in Andean birds. In the present paper, an effort has been made to 
uncover the evolutionary relationships of the component members 
of the Asthenes flammulata superspecies by means of an analysis of 
the distribution and geographical variation of these spinetails. 

This study was undertaken at the Museum of Comparatize Zool- 
ogy of Harvard University. My investigations were based largely on 



1 Research Fellow, Department of Ornithology, American Museum Of 
Natural History, New York, New York 10024. 



2 BREVIORA No. 297 

the collections of this institution, but I have also examined speci- 
mens from other museums. When referring to individual specimens, 
the following abbreviations are used: MCZ = Museum of Com- 
parative Zoology; ANSP = Academy of Natural Sciences of Phila- 
delphia; CM — Carnegie Museum. Pittsburgh; FMNH = Field 
Museum of Natural History, Chicago; MVZ = Museum of 
Vertebrate Zoology, Berkeley; MP = Museum National d'Histoire 
Naturelle, Paris. 

ACKNOWLEDGMENTS 

I am most indebted to Ernst Mayr for his guidance, advice, and 
stimulation throughout my studies of speciation of Andean birds, 
which were becun at his suggestion. 

I wish to extend mv sincere thanks to the following persons, who 
have helped me during mv examination of specimens, who have 
arranged loans for me. or who have provided me with valuable in- 
formation: Raymond A. Paynter, Jr. (Museum of Comparative 
Zoology), James Bond (Academy of Natural Sciences). Dean Ama- 
dou and Charles O'Brien (American Museum of Natural History), 
Kenneth C. Parkcs (Carnegie Museum), Emmet R. Blake (Field 
Museum of Natural Histor\ ), Ned K. Johnson (Museum of Ver- 
tebrate Zoology), I. C. G. Galbraith (British Museum [Natural 
History]). Jean Dorst and Jean Prevost (Museum National dllis- 
toire Naturelle ). 

Financial assistance to delra\ the costs of field work was pro- 
vided through the Chapman Fund of the American Museum of 
Natural History, National Science Foundation funds from grants 
G-19729 and GB-3167 to Harvard University's Committee on 
Evolutionary Biology, and the Bourse Federale de Voyages de la 
Societe Helvetique des Sciences Naturelles in Switzerland. A visit 
to the Academy of Natural Sciences in Philadelphia was financed 
by the University of Massachusetts — Boston. I am very grateful 
to these agencies for their generous support of my work. 

Permission to reproduce maps published by the Department of 
Geography of the University of Chicago (Goode Base Maps Series. 
Copyright by the University of Chicago) was obtained through the 
courtesy of Mrs. Carole J. Major. 

THE ASTHENES FLAMMULATA SPECIES GROUP 

The genus Asihenes Reichenbach, 1853, consists of about 18 
species of small spinetails (Furnariidae) living in western and 



1968 ASTHENES FLAMMULATA SUPERSPECIES 3 

southern South America. At least four species groups can be rec- 
ognized in Asthenes. Since this arrangement of the genus will be 
published elsewhere, it is sufficient to mention here that what I call 
the Asthenes flammulata species group comprises four nominal 
species: Asthenes flammulata (Jardine), A. virgata (Sclater), A. 
maculicauda (Berlepsch), and^4. urubambensis (Chapman). 

These four forms were considered full species by Hellmayr 
(1925), Peters (1951 ), and de Schauensee (1966). They all have 
a diagnostic pattern of streaking — a narrow stripe along the 
feather shaft contrasting with the remainder of the feather. 

Within the A . flammulata species group, A . urubambensis can be 
diagnosed by the combination of a lack of streaking on the dorsum 
and a lack of rufescent color on the forehead and crown. Further- 
more, A. urubambensis is sympatric with two of the three other 
nominal species of the flammulata species group: with A. flammu- 
lata in Peru (see Zimmer, 1930:345) and with A. maculicauda in 
Bolivia (see Bond and de Schauensee, 1942:334). There is there- 
fore no doubt that A. urubambensis is a good biological species. 

THE ASTHENES FLAMMULATA COMPLEX 

The other three nominal species of the A. flammulata species 
group can be described collectively as the A. flammulata complex. 
They are morphologically rather similar to one another, and are 
largely allopatric, although there are two apparent exceptions: (1) 
in southern Peru ("contact" between virgata and maculicauda) and 
(2) in central Peru (alleged sympatry between flammulata and 
virgata ) . 

Sympatry 

Asthenes virgata was described by Sclater, in 1874, on the basis 
of one specimen collected by Jelski in "Junin," in the Andes of 
central Peru. In 1930, M. A. Carriker, Jr., secured five specimens 
of A. virgata in the Rock Forest area west of Lake Junin (Bond, 
1945:33; 1956:245). Although Jelski did not give precise locality 
data, it is possible that "Junin" refers to the village of that name 
situated southeast of the lake at the edge of the flat depression form- 
ing the present-day Lake Junin and adjoining marshes (see Fig. 1 ). 
Junin and Rock Forest are about 50 km apart. 

These are the only published localities where A. virgata has been 
taken, at least to my knowledge. However, C. Koford collected a 
single female Asthenes in the Department of Puno, southern Peru, 
in 1951 (MVZ 124302). This bird is morphologically much closer 



HRI \ lORA 



No. 297 




^ localities cited 
in the text 



" Junin uncertain 
type loc. of vi rga t a 



.' A R G K N T I N A 



# 



Fig. 1. Collecting localities of the Asthenes flammulata complex 



1968 ASTHENES FLAMMULATA SUPERSPECIES 5 

to the geographically distant A. virgata from central Peru than to 
its nearest neighbor, A. maculicauda. As far as I know, this speci- 
men has not been previously reported in the literature, although it 
was known to Zimmer, who identified it as A. virgata (pencilled 
comment on the label) . 

"Contact" between A. virgata and A. maculicauda appears pos- 
sible when the collecting localities of Koford's specimen of virgata 
and of the northernmost record of maculicauda are examined. Ko- 
ford's bird was collected eight miles SSW of Limbani at 15,250 
feet (label data). The northernmost specimen of maculicauda ap- 
pears to be a bird collected by Carriker at Huancarani, 13,000 feet 
(ANSP 103442). According to Bond (1945:19), Huancarani is 
"about ten miles above Limbani." Limbani is a village on the left 
(west) bank of the valley, and is accessible by only one road or 
trail from the southwest. Thus it is likely that "above Limbani" and 
"SSW Limbani" are very close to each other (Fig. 1). However, 
even if these two localities are not separated by a significant linear 
distance, they are separated by an altitudinal gap of 2250 feet 
(about 700 m). There is thus no direct evidence for sympatry be- 
tween A . virgata and A . maculicauda, but in view of their morpho- 
logical differences, their sympatry in southern Peru would not be 
surprising. 

Sympatry between A . virgata and A . flammulata in central Peru 
was made apparent by the mention of four specimens of A. flam- 
mulata taczanowskii collected at Rock Forest by Dorst (1956: 
268). But as I shall show below, Dorst's specimens are actually 
referable to virgata; sympatry is therefore non-existent, since only 
one taxon occurs at Rock Forest. 

Geographical Variation 

Altogether about 140 specimens have been examined from lo- 
calities covering the entire range of the three nominal species of the 
A. flammulata complex (Fig. 1). Specimens were grouped into 
"populations," which are mapped on Figure 2. The grouping was 
based on several considerations, primarily (a) whether or not cer- 
tain localities appeared to be isolated from others, thus representing 
potential interruptions in gene flow, and (b) whether or not the 
birds from several more or less adjacent localities were morphologi- 
cally uniform. Thus specimens from the widely-spaced collecting 
locatities of the Eastern Andes of northern and central Peru were 
considered as one "population" (H) because of their considerable 



BRLVIORA 



No. 297 




BOLIVIA 






r 



': 



5C*LC 



ARGENTINA I 

l 



II Eat tern Andes 

I ( : j j .1 m a r c a 

J (Jum v ilc j. I.ibcrtad 

K Rock Foren 

I. ssw Limban i, Puno 

\l I In. i in j r .1 n i. I'ti n 

\ La Pa / ( !o< habamba 



O Aconquija 

— 



&INUMOA4. ftoJCCTON 





Fig. 2. The populations of the Asthenea flammulata complex. 



1968 ASTHENES FLAMMULATA SUPERSPECIES 7 

morphological uniformity; on the other hand, those from the West- 
ern Andes of northern Peru were placed in two "populations" (I 
and J) because of geographical variation in some characters. 

Despite this grouping into populations, the samples available 
were small. Under these circumstances, statistical treatment of the 
mensural characters was not attempted. 

TABLE 1 

Geographical variation of bill length in the 
Asthenes flammulata complex 

Populations Range of mean bill length (in mm) 

Males Females 

Colombia, Ecuador 
northern Peru (A-L) 17.0-17.9 16.0-19.0 

Southern Peru, Bolivia, 
northern Argentina 

(M-O) 15.0-16.5 15.3-15.5 



TABLE 2 

Geographical variation of tarsus length in the 
Asthenes flammulata complex 

Population Tarsus length of males (in mm) Tarsus length of females (in mm) 





Range 


(Mean) 


N 


Range 


(Mean) 


N 


A 


24.0-27.0 


(25.3) 


3 


23.0-24.0 


(23.7) 


3 


B 
C 
D 


25.0-27.0 


(25.9) 


11 


23.5-26.0 


(24.7) 


6 


24.0-27.0 


(25.8) 


12 


25.0-25.5 


(25.1) 


4 


E 


25.0-26.0 


(25.6) 


9 


23.0-26.0 


(24.8) 


14 


F 


25.0-25.5 


(25.2) 


3 




24.0 


1 


G 


24.5-26.0 


(25.3) 


2 


24.0-25.0 


(24.5) 


2 


H 


23.0-24.0 


(23.9) 


7 


23.0-24.0 


(23.8) 


4 


I 


23.0-24.0 


(23.0) 


2 




25.0 


1 


J 


24.0-25.0 


(24.3) 


3 








K 


23.0-27.0 


(24.6) 


7 


23.0-24.0 


(23.5) 


2 


L 

M 




23.0 


1 




26.0 


1 


N 


22.0-24.0 


(22.8) 


4 


22.0-22.0 


(22.0) 


2 





22.0-23.5 


(22.8) 


3 


20.0-21.5 


(20.8) 


2 



8 breviora No. 297 

Mensural characters. Geographical variation in bill length, tar- 
sus length, and wing length, respectively, is summarized in Tables 1 
and 2 and in Figure 3. Populations M, N, and O (from southern 
Peru to northern Argentina) are distinguishable from their neigh- 
boring populations by a gap (step) in either absolute size or in 
mean length of all three characters. By contrast, steps between any 
two of the populations from A through L (Colombia to southern 
Peru) are much less significant (see especially Fig. 3). 

Population Males (N) Females (N) 

« so u n is mm mm » '0 mm 

•— t — i — i — i I I I ) — | I | I | | l i I | l i— t — l — l I l — l t I I I I I I I 

\ —I- (!) + (31 

B 1 — do) —I (i) 

C + (0 

1) 1 (!l) 1 (I) 

I 1 (•) —I (II) 

I + (?) 

(, -4- (?) -f (I) 

II 1 (I) —I («) 

I -I- i? i + (') 

J -HO) + 0) 

k 1 (1) + (J) 

I. + (0 

\l + (I) 

N (- («) -HO) 

() —l hi -I- in 

Fig. 3. Geographical variation of wing length in the Asthenes flammulata 
complex. 

From the variation in mensural characters, one might conclude 
that the A. flammulata complex consists of two units, each an in- 
terbreeding species. The first would include populations A through 
L (nominal species A. flammulata and A. virgata), and the second 
populations M through O (nominal species A. maculicauda) . 

Color characters. Geographical variation in the color of the 
gular patch, the color of the forehead, the amount of streaking on 
the abdomen and crissum, the color of the rump and upper tail 
coverts, and the markings of the median pair of rectrices is sum- 
marized in Table 3. Variation in wing pattern is illustrated in 
Figure 4. Detailed notes on variation in these characters are in- 
cluded in Appendix I; only the major trends will be discussed here. 



1968 



ASTHENES FLAMMULATA SUPERSPECIES 



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10 breviora No. 297 

Characters such as the color of the gular patch, the color of the 
forehead, and the intensity of streaking on the underparts vary 
rather gradually. There is a trend, from north to south, toward a 
decrease both in saturation of pigmentation and in the amount of 
streaking. Exceptions to this trend are provided by populations K 
and L (gular patch color) and populations M, N, and O (forehead 
color). 

The other characters seem to vary more discontinuously. These 
patterns of discontinuous variation permit one to distinguish mor- 
phologically several populations or groups of populations within 
the A. flammulata complex. The best-marked group includes popu- 
lations from southern Peru to northern Argentina (M, N, O). 
These birds arc differentiated from their neighboring populations by 
gular patch color, forehead color, and streaking on the rump. The 
steps observed ill the variation of these color characters correspond 
to those noted in the mensural characters. 

Other groups of populations can also be separated morphologi- 
cally, but on the basis of fewer characters. Thus the birds of 
populations A and B (Colombia) can be distinguished by the color 
of the gular patch and the color of the forehead. Similarly, birds 
from populations K and L (Rock Forest and SSW Limbani) are 
separable from others by the color of the gular patch and the gen- 
eral absence of streaks on the rump and upper tail coverts (if these 
streaks are present, they are obsolescent). 

However, it must be pointed out that such a neat separation of 
populations cannot be done using some other characters. For ex- 
ample, populations K and L are not distinguishable from popula- 
tions M, N, and O on the basis of color or streaking of the 
underparts. Also, populations K and L are indistinguishable from 
populations H through J in forehead color. 

When variation in color characters (as in mensural characters) 
is considered as a whole, the A. flammulata complex appears to 
comprise two units. Geographical variation within populations A 
through L, and within M through O, could be interpreted as that 
normally to be found in an interbeeding species. However, unlike 
what is found in mensural characters, the gap in color characters 
separating populations A-L and iM-O does not seem to be as well 
marked as the gap separating some populations within A-L. 

Population Structure 

The dual type of geographical variation (gradual and discon- 
tinuous) exhibited by the A. flammulata complex is often found in 



1968 ASTHENES FLAMMULATA SUPERSPECIES 11 

a single polytypic species with a partially continuous and partially 
discontinuous range. 

Although the number of specimens examined is not large enough 
to reach definite conclusions, it seems probable that the gradual size 
trends (especially in tarsus and wing length) detected are positively 
correlated with altitude. Thus the largest birds seem to live at 
slightly higher elevations (in north-central Ecuador and central to 
southern Peru) than the smallest birds (in southern Ecuador and 
northern Peru. Altitudinal variation in Andean birds has been 
demonstrated by Traylor (1950) and myself (unpublished). How- 
ever, the correlation of increase in size with increase in altitude is 
not the only possible one. In Peruvian birds, for example, the 
north-south size increase along the Western Andes could be due to 
latitude or aridity (Hamilton, 1961), rather than to altitude. 

The decrease in the intensity of pigmentation from north to south 
cited earlier is also detectable in several species belonging to other 
Andean genera (unpublished). Such variation may be correlated 
positively with the decrease in rainfall along the Andes from north 
to south (see, e.g., Troll, 1959); thus it corresponds to Gloger's 
rule. 

The apparent trends of gradual geographical variation are clear- 
est in populations A through L, which have the earmarks of repre- 
senting a single species, as already noted. It is, therefore, of interest 
that populations M-O, which are separated from populations A-L 
by a marked morphological gap in some characters, have a very 
different type of population structure. Although populations M, 
N, and O are apparently disjunct (see Fig. 2), the birds I have 
examined are all remarkably uniform and display only slight geo- 
graphical variation — in wing length, for example (see Fig. 3). 
This lack of significant geographical variation can be explained 
either by free gene flow, or by recency of occupation of the vast 
geographical range, or possibly by both. Whatever the reason, the 
differences in population structure would tend to confirm the pre- 
vious impression of specific distinctness between populations A-L 
(flammulata-virgata) and M-O (maculicauda) . Since, as noted 
previously, there is no apparent barrier between the two groups of 
populations in southern Peru, I would conclude that their differen- 
tiation does not reflect the result of evolution in situ (unless, of 
course, a barrier once present has subsequently been removed). 
This question cannot yet be answered, but the problem will be 
taken up in a later paper in this series. 

To study the possible correlation between discontinuous variation 
and geographical barriers in the A. flammulata complex, we must 
turn to populations A-L, ranging from Colombia to southern Peru. 



12 hreviora No. 297 

Discontinuous Variation and Geographical Barriers 

The presence of a geographical barrier between avian popula- 
tions does not ipso facto mean that gene flow between the isolates 
is interrupted. Similarly, the fact that geographical isolates are not 
differentiated morphologically, or are so only slightly, does not 
necessarily mean that gene flow takes place between them. Conse- 
quently. Statements made in the following paragraphs about dif- 
ferentiation and gene Bow across barriers must be taken, not as 
expositions of known facts, but as cautious expressions of proba- 
bilities. 

The correlations between morphological differentiation and geo- 
graphical barriers in populations A-L of the A. (lammulata complex 
will be mentioned in geographic sequence from north to south. 

(1) In Colombia, the birds from the Eastern Andes (multo- 
striata) are separated from those oi the Central Andes (quindiana) 
by the Magdalena Valley (Fig. 5). Morphological differences be- 
tween these two isolated populations (A and B on Fig. 2) include 
wing length, the width of the streaks on the underparts, the pres- 
ence or absence oi a slight bulTv wash on the undertail coverts, the 
extent o\' the gular patch, and the intensity of the coloration of the 
forehead. The two populations are certainly sufficiently differen- 
tiated to be recognized taxonomically. although they are more simi- 
lar to each other than they are (either as a group or separately) to 
populations farther south in Colombia and Ecuador (C-G). 

i 2 i Between the Eastern or the Central Andes of Colombia and 
the Andes of Nariho in southern Colombia is an apparent distribu- 
tion gap (Fig. 5 ). The only specimen from Nariho (C) I have seen 
(Mayasquer, ANSP 149893) appears somewhat intermediate in 
both postocular stripe and cheek color between Colombian birds 
(A-B) and Ecuadorian specimens (D-G). However, since these 
differences are slight, it is not possible to assess the real extent of 
intermediacy. In other characters, furthermore, the Narino speci- 
men falls fully within the range of variation of Ecuadorian speci- 
mens. The vast geographical gap between Colombian populations 
A-B and southern Colombian-Ecuadorian populations C-G is there- 
fore correlated with a relatively well-marked morphological gap. 
Taxonomic recognition of populations C-G (subspecies flammu- 
lata) is thus warranted. 

(3) The Interandean Depression of Ecuador (Fig. 5) is a low 
and relatively dry region separating the Eastern from the Western 
Andes. Only a slight difference in wing length and tarsus length 



1968 ASTHENES FLAMMULATA SUPERSPECIES 13 

between birds of the Eastern (D) and Western (E) Andes was de- 
tected, suggesting only a slight restriction in gene flow; taxonomic 
subdivision within populations C-G (fiammulata) would be un- 
justified. 

(4) There is a gap between the northern Peruvian populations 
of the Eastern (H: taczanowskii) and Western (I: pallida) Andes 
and those of southern Ecuador (G). For population H, the gap 
corresponds to the eastward bend of the Maranon River; for popu- 
lation I it corresponds to the generally low altitude and small area 
of suitable habitat of the Andes of northwestern Peru (Northern 
Peruvian Low: Fig. 5). The morphological differences between 
Ecuadorian and Peruvian birds are not large. They include the in- 
tensity of streaking on the underparts and the extent of rufous on 
tail feathers. The lack of major differences suggests either some 
gene flow over the gap, or that the separation of Ecuadorian from 
Peruvian birds is of recent origin. 

(5) The Upper Maranon Basin (Fig. 5) separates the northern 
Peruvian populations of the Eastern Andes (H: taczanowskii) 
from those of the Western Andes (I and J: pallida). Only slight 
morphological differences can be noted between them, however: 
these include characters such as tarsus length, wing length, and 
intensity of streaking of the rump and upper tail coverts. I could 
find no differences in the other characters examined. In view of 
this minor morphological differentiation, subspecific separation of 
the two populations may not be warranted; pallida may be consid- 
ered a synonym of taczanowskii. As Bond (1945:32) has re- 
marked, pallida "is not a very well-marked race." 

(6) The specimens collected in the Rock Forest area of central 
Peru (K: virgata) are distinguishable morphologically from the 
birds living in both the Eastern (H) and Western (I, J) Andes. 
The differences are in wing length, color of gular patch, color of 
central pair of rectrices, amount of streaking on rump and upper 
tail coverts, wing pattern, and color of the lower abdomen and 
crissum. These differences are better marked between Rock Forest 
specimens (K) and birds from the Eastern Andes (H), than be- 
tween Rock Forest birds and specimens from the Western Andes 
(I, J) . An approach to Rock Forest birds is clearly seen in Western 
Andean birds (especially population J) in size (Fig. 3) and in wing 
pattern (compare nos. 5 and 6 on Fig. 4). 

Geographical isolation between Rock Forest birds {virgata) and 
Eastern Andean birds {taczanowskii) is apparently provided by the 
Lake Junin Basin (Fig. 5). But since the first specimen of virgata 
may have come from "Junin" in the center of the basin, a more 



14 



BREVIORA 



No. 297 





® 





© 






Fig. 4. Geographical variation of wing pattern in the Asthenes flammu- 
lata complex. The feather illustrated is the sixth outer primary. 

Key: 1 = Volcan Pichincha, Ecuador; 2 = Huanuco, northern Peru; 3 = 
Maraynioc, central Peru; 4 and 5 = Quirivilca, Libertad, northern Peru; 
6 = Rock Forest, central Peru; 7 — SSW Limbani, southern Peru; 8 and 9 = 
Incachaca, Cochabamba. Bolivia 



1968 ASTHENES FLAMMULATA SUPERSPECIES 15 

likely barrier may be the high Andes farther east. In any event, 
although the barrier is not clear-cut at present, morphological dif- 
ferentiation is fairly sharp. 

Isolation between Rock Forest birds (virgata) and populations 
found farther north along the Western Andes (J: "pallida") is not 
clear. The closest specimens come from Quirivilca, Libertad, about 
375 km from Rock Forest. No specimens are known to me from 
the intervening area, although apparently suitable habitats occur 
there. A possible barrier might be the Santa River Canyon, which 
makes a cut across the Andes between the Cordillera Blanca and 
the Andes of Libertad. 

(7) The vast Andean area between central and southern Peru 
(about 800 km) seems to represent a distribution gap in the range 
of the A. flammulata complex. However, suitable habitat occurs 
there, and further collecting might reveal the presence of the com- 
plex there (? on Fig. 5). The specimen from SSW of Limbani is 
practically identical with Rock Forest specimens {virgata), except 
in the pattern of the central pair of rectrices. 

Taxonomic Conclusions 

The classification of Hellmayr (1925), Peters (1951), and de 
Schauensee (1966) is as follows. Colombian, Ecuadorian, and 
northern Peruvian populations (A-J) are treated as one species, 
Asthenes flammulata. The two disjunct populations from central 
and southern Peru (K and L) would constitute a second species, 
A . virgata. Finally, the populations from southern Peru to northern 
Argentina (M, N, O) make up a third species, A. maculicauda. A 
slight modification of this classification was made by Bond and de 
Schauensee (1942:334), who considered maculicauda a subspecies 
of flammulata. (Later, however, Bond, 1945:32, cited maculi- 
cauda as a full species without any comment.) This classification 
finds justification in the fact that there is a morphological and 
partial geographical gap between flammulata and virgata (see (6) 
above), and because there is a morphological gap between flammu- 
lata-virgata and maculicauda. 

I believe, however, that the evidence presented in this paper 
allows one to construct a classification of the A. flammulata com- 
plex which is at the same time a simplification and an improvement 
over the older scheme. 

The Asthenes flammulata complex constitutes a superspecies 
(sensu Mayr, 1963:501) having two component species. The first 



16 



BREVIORA 



No. 297 





Northern Peruvian Low 

c 

Upper Mar a Eon Valley 

ARI \ 

WHERE \ Lake Jum'n Has 

INTER- V?, 

GR tDATION ? £ 

BETW I I N 

FLAMMULA1 \ 

WD VIRGA1 \ 

IS POSSIBLE 



PROBABLE CONTACT 
BET w I I N VIRGATA 



: ///. AM) MAC I 1. 1 CALL) A 



A t I j ni in ii I i i i 



A m j i u I u j u d .i 





Fig. 5. Taxonomic and evolutionary interpretation of distribution and 
geographic variation in the Asthenes flammulata superspecies. 



1968 ASTHENES FLAMMULATA SUPERSPECIES 17 

species is composed of several isolated and semi-isolated popula- 
tions ranging from Colombia to southern Peru (Asthenes flammu- 
lata). Fully isolated populations of A. flammulata which have 
reached a moderate to marked degree of differentiation (incipient 
species) are the Colombian multostriata and quindiana. These two 
isolates are perhaps best considered as a subspecies group with two 
subspecies. The specimens from Rock Forest and SSW Limbani 
fall within the range of variation of a single taxon (virgata). 1 Some 
gene exchange between Rock Forest birds and birds farther north 
in the Western Peruvian Andes is suggested by the variation of 
characters such as wing length (Fig. 3). I consequently recom- 
mend that virgata be treated as a subspecies of A . flammulata. 

The second component species of the A. flammulata super- 
species, Asthenes maculicauda, is composed of three apparent iso- 
lates (M, N, O), but it does not exhibit much geographical varia- 
tion. 

The taxa of the A. flammulata superspecies and their relation- 
ships can be listed as follows: 

Asthenes flammulata superspecies (two component species which 
may have achieved reproductive isolation) 

( 1 ) Asthenes flammulata ( Jardine, 1 850) 

(a) Colombian subspecies group (one or two incipient 
species): A. flammulata multostriata (Sclater, 
1858), A. flammulata quindiana (Chapman, 
1915) 

(b) Ecuadorian-Peruvian subspecies group (two or 
three incipient species): A. flammulata flammu- 
lata (Jardine, 1850); ,4. flammulata taczanowskii 
(Berlepsch and Stolzmann, 1894; includes A. f. 
pallida Carriker, 1933); A. flammulata virgata 
(Sclater, 1874) 

(2) Asthenes maculicauda (Berlepsch, 1901) 
Shows insignificant geographical variation 

SUMMARY 

The Andean spinetails of the Asthenes flammulata complex in- 
clude three nominal species, flammulata, virgata, and maculicauda, 
which, together with a fourth, A. urubambensis, constitute the A. 
flammulata species group. 



1 The specimens reported by Dorst (1956) as A. flammulata taczanowskii 
from Rock Forest are therefore referable to virgata. 



18 brlviora No. 297 

Specific status of A. urubambensis is shown by sympatry with 
two nominal species of the A. fiammulata complex and bv morpho- 
logical distinctness. 

The evolutionary relationships of the three nominal species of the 
A. fiammulata complex were analyzed by a study of geographical 
\ariation and its correlation with geographical barriers. Geographi- 
cal variation is largely discontinuous in populations ranging from 
Colombia to southern Peru (nominal species fiammulata and vir- 
gata). By contrast, populations from southern Peru to northern 
Argentina show great morphological uniformity (nominal species 
maculicauda ) . 

It is concluded that the Asthenes fiammulata complex is best con- 
sidered as a superspecies with two component species. The first 
species, A. fiammulata, contains several isolated groups of popula- 
tions (incipient species) showing various degrees of morphological 
differentiation across barriers. A. virgata should be included as a 
subspecies of A. fiammulata. The second species, A. maculicauda, 
may come in contact with A. fiammulata virgata in southern Peru. 
Gene exchange between the two species i^ not suggested, but there- 
are only two specimens mi which to base a decision. 

REFERENCES CITED 

Bond, J. 

1945. Notes on Peruvian Fumariidae. Proc. Acad. Nat. Sci. Philadel- 
phia, 97: 17-39. 

1956. Additional notes on Peruvian birds II. Proc. Acad. Nat. Sci. 
Philadelphia. 108: 227-247. 
Bond, J., and r. m. df. Schalensee 

1942. The birds of Bolivia. Part I Proc. Acad. Nat Sci. Philadelphia. 
94: 307-391. 
Carriker, M. A., Jr. 

1933. Descriptions of new birds from Peru with notes on other little 
known species. Proc. Acad. Nat. Sci. Philadelphia, 83: 1-38. 

DORST, J. 

1956. Etude d*une collection d'oiseaux rapportee du Perou central. 
Bull. Mus. Paris, 2e Ser., 28: 265-272. 
Hamilton, t. h. 

1961. The adapti\c significance of intraspecific trends of variation in 
wing length and body size among bird species. Evolution, 15: 
180-195. 
Hellmayr, C. E. 

1925. Catalogue of birds of the Americas. Part IV. Furnariidae- 
Dendrocolaptidae. Field Mus. Nat. Hist. Zool. Ser., 13: 1-390. 



1968 ASTHENES FLAMMULATA SUPERSPECIES 19 

MAYR, E. 

1959. Trends in avian systematics. Ibis, 101: 293-302. 
1963. Animal Species and Evolution. Harvard Univ. Press, Cam- 
bridge. 
Peters, J. L. 

1951. Check-list of Birds of the World. Vol. VII. Mus. Comp. Zool., 
Cambridge. 

SCHAUENSEE, R. M. DE 

1966. The Species of Birds of South America and their Distribution. 
Livingston Publ. Co., Narberth, Pennsylvania. 

SCLATER, P. L. 

1874. Descriptions of three new species of the genus Synallaxis. Proc. 
Zool. Soc. London, 1874: 445-447. 
Traylor, M. A. 

1950. Altitudinal variation in Bolivian birds. Condor, 52: 123-126. 
Troll, C. 

1959. Die tropischen Gebirge. Bonner Geogr. Abhandl., Heft 25: 
1-93. 

ZlMMER, J. T. 

1930. Birds of the Marshall Field Peruvian expedition, 1922-1923. 
Field Mus. Nat. Hist., Zool. Ser., 17: 233-480. 

(Received 23 November 1967.) 



APPENDIX I: NOTES ON GEOGRAPHICAL VARIATION IN NON- 
MENSURAL CHARACTERS 

Gular patch. The gular patch varies in color intensity from whitish 
and yellowish to rich ochraceous, and in extent from a small, roughly tri- 
angular patch more or less restricted to the chin, to a wider patch com- 
prising also the throat and upper chest. 

Colombian specimens (populations A and B) have a richly ochraceous 
gular patch. Central Andean birds (B) have a rather broad patch, while 
in Eastern Andean birds (A) it is more restricted. 

Ecuadorian specimens and the Nariho specimen from Mayasquer (popu- 
lations C-G) have relatively narrow and more or less triangular gular 
patches, varying from whitish-yellow to pale ochraceous. 

Peruvian specimens (populations H-J) have a broad, ill-defined gular 
patch merging into the upper chest. The patch varies in color from dirty 
white to yellowish-cream. Usually the chin area is somewhat paler than the 
throat. Some specimens of Ecuadorian birds (e.g. MCZ 199104, female, 
and MCZ 199099, male) have a more ill-defined gular patch than do most 
other Ecuadorian specimens, thus showing an approach to Peruvian 
specimens. 

Peruvian specimens from Rock Forest and SSW Limbani (K and L) 
have a relatively small, well-defined gular patch, usually of a rich ochraceous 
color. 



20 BREVIORA No. 297 

Specimens from southern Peru to northern Argentina (M, N, O) lack a 
buffy-ochraceous gular patch: the gular area is of about the same color as 
the chest. 

Forehead color. In Colombian specimens from both the Eastern and 
Central Andes i A and B) the forehead is an almost solid rufous, especially 
in males. The forehead of most Ecuadorian specimens (C-G) is streaked 
with rufescent, although in some birds (e.g. MCZ 138553) it is a solid 
rufous. In Peruvian birds from populations H, 1. and J the forehead streaks 
are less rufescent. more yellowish, but here again at least one specimen 
i I MNH 65849, male) has a rufous forehead, where the streaks almost join 
to form a solid color. In Peruvian specimens from Rock Forest and SSW 
LimbanJ (K-L) the streaks are relative!) yellowish, not verv rufescent. 
Finally, specimens from southern Peru, Bolivia, and northern Argentina 
(M, N. O) have rufescent foreheads, distinctly more pigmented than those 
of Peruvian birds in general, and of Rock Forest-SSW I imbani birds in 
particular. 

Coloration of under parts. Colombian specimens are the most beavil) 
streaked; Bolivian and Argentine specimens are almost unstreaked. The 
birds from Rock forest. SSW 1 imbani. Huanearani. Bolivia, and northern 
Argentina (k-Ot are the least streaked ol all populations within the Asthenes 
ftammulata complex I be additional fact that in the birds from these same 
populations the lower abdomen and crissum are buffy with obsolescent 
streaks, rather than whitish with rather conspicuous streaks, permits one to 
distinguish them immediately from all other populations (A-J). There is. 

in particular, a rather well marked difference in streaking intensity between 
the crissum of Peruvian birds from the I astern (H) and Western Andes 
i I. J t. and that of birds from Rock Forest iki. 

Specimens fiom southern Peru to northern Argentina (M, N, O) have, 
alone among the populations of the A. tlammuhitu complex, a series of 
ill-marked streaks on the chest converging toward the center to form a 
ring. 

Color of rump and upper rail coverts. I he birds from Rock Forest and 
ssW Limbani (K and 1.) have slight or obsolescent streaking on the rump 
.mA upper tail coverts. (This was already noticed by Bond [1956: 245] for 
Rock Forest specimens.) By contrast, specimens from all other populations 
have generally much heavier streaking on the rump and upper tail coverts. 
These streaks are relatively well marked in Peruvian populations H, [, and J, 
as well as in Peruvian, Bolivian, and Argentine populations M, N, and O. 
Rock Forest and SSW Limbani ( K.-E ) specimens are thus distinguishable 
as a group from neighboring populations. 

Wing pattern. The primaries of birds in all populations are more or 
less rufous with a dark brown to blackish tip. I have studied interpopulation 
variation in pattern by comparing the patterns of the sixth outer primary. 
which was chosen arbitrarily. 

Except for a few Colombian specimens, which have only a relatively 
narrow rufous edge to both vanes of the sixth primary, in most other speci- 
mens (including those from Colombia) the tip of the feather is more or 



1968 ASTHENES FLAMMULATA SUPERSPECIES 21 

less wholly dark brown, while its base is rufous (see Fig. 4, Nos. 1, 2, 3, 4, 
and 9). 

The most significant population variation is in the presence or absence 
of a thin dark brown stripe along the outer vane of the primary (com- 
pare Nos. 4 and 5 of Fig. 4, for example). This pattern is seen in some 
specimens from Quirivilca, Libertad (population J), every specimen from 
Rock Forest (K), the single specimen from SSW Limbani (L), and most 
specimens from southern Peru to northern Argentina (M, N, O). The 
presence of the two major patterns in the Libertad and Cochabamba popu- 
lations is illustrated in Fig. 4 (Nos. 4 and 5, and Nos. 8 and 9, respectively). 

Marking of rectrices. Analysis of variation in this character is difficult 
because a rather large number of specimens have one or several rectrices 
in molt or missing and because many specimens have extremely worn 
rectrices. Nevertheless, the detectable variation appears as follows. 

There is a tendency for Peruvian specimens (including the specimen from 
SSW Limbani, but excluding Rock Forest specimens) to have more rufous 
on outer tail feathers than do Ecuadorian or Colombian specimens. 

Rock Forest specimens lack entirely or almost entirely the narrow 
rufescent or pale brownish outer edge to the outer vane of the central pair 
of rectrices, which is present in specimens from all other Peruvian localities 
including SSW Limbani. 

In birds from southern Peru to northern Argentina (M, N, O), the 
central pair of rectrices is marked with irregular dark brown designs on a 
rufous or rufescent background. No other specimen from anywhere in the 
range of the Asthenes flammulata complex has similar markings on the 
median rectrices, with one exception: in the central pair of rectrices of the 
specimen from SSW Limbani, the outer part of the outer vane is rufescent, 
with a thin and irregular dark brown line running alongside the rufescent 
color. This pattern, although reminiscent of that found in birds from 
populations M, N, and O, is not really similar to it. 

In rectrices other than the central pair, a pattern reminiscent of the one 
found in specimens from southern Peru to northern Argentina (M, N, O) is 
noticeable in specimens from Rock Forest (ANSP 92428, ANSP 92427, MP 
CG1956 No. 942, MP CG1956 No. 940), as well as in other populations. 
But the number of specimens in which this pattern occurs tends to become 
smaller north of central Peru. 



BREVIORA 



Museum of Conn para tive Zoology 

Cambridge, Mass. 11 October, 1968 Number 298 

MONOGRAPH OF THE CUBAN GENUS VIANA 
(MOLLUSCA: ARCHAEOGASTROPODA: HELICINIDAE) 

William J. Clench and Morris K. Jacobson 

Abstract. The genus Viana in the family Helicinidae is limited in its 
distribution to the Sierra de los Organos in Pinar del Rio, the westernmost 
province of Cuba. All members are found only on limestone substrate. 
The genus consists of one species, regina Morelet, with two subspecies, 
laevigata Pfeiffer and subunguiculata Poey, that form many races which 
are not clearly discrete morphologically and which show no coherent 
geographical distribution. 



INTRODUCTION 

This study of the genus Viana is based upon specimens housed 
in the Museum of Comparative Zoology which include the very 
large collections made by Carlos de la Torre, Pedro Bermudez and 
Julio Armenteros. Much additional material was contributed by 
Thomas Barbour, William J. Clench, Paul Bartsch, Carlos G. 
Aguayo, Miguel L. Jaume, Morris K. Jacobson, and numerous 
other Cuban and American collectors. 

The strikingly handsome shells of the helicinid genus Viana are 
confined to the Sierra de los Organos, Pinar del Rio Province, in 
the western end of Cuba. Their range is limited in the west by the 
sierras and mogotes 1 about Mendoza, and in the east by the Sierra 
Rangel, about 75 miles away. The species of Viana are purely 
calciphilous, living exclusively on the limestone mogotes and 
sierras which emerge from the surrounding red, iron-bearing soil. 
In such locations during the rainy season they can be collected as 
readily as littorinids on rocky sea coasts. Unlike the related genus 
Emoda, they are found in no other part of the island. 

Within their range they inhabit practically every outcrop of 
limestone of any size which enjoys sufficient moisture and shade. 



1 In western Cuba a mogote is any isolated limestone block separated 
from the main sierra. 



HRI-VIORA 



NO. 298 



Thus they are common on high sierras like those of Guane, Paso 
Real, Vinales, San Carlos, San Andres, Guajaibon, and Rangel, 

and on the small mogotes in the \alle\ oi \ ihales. the Laguna de la 
Piedra, and around Luis La/o. Isabel Maria, and between the 
Estrechura of San Carlos and Guane ( Teneria. Punta de la Sierra. 
1 .1 Muralla, and Los Portales). 




Map I Showing the greater part of the Province of Pinar del Rio, Cuba, 
outlining the areas containing ihe sierras and mogotes which are occupied 
In the three subspecies of Viana regina Morclet. 

A. liana retina .uilutnguiculata (Poev). B, V. r. laevigata (Pfeiffer). 
C. V. r. regina (Morelet). 



(hies and towns 


Sierras and Mogotes 


1. Guanaja) 


X. 


Sierra Rangel 


2. Artemisa 


9. 


Pan de c ruajaibon 


3. ( onsolacion del Norte 


10. 


Sierra San Andres 


4. \ ihales 


1 1. 


Sic i ra \ ihales 


5. Consolacion del Sur 


12. 


Mogotes at "Km. 14" 


ft. Pinar del Rio 


13. 


Ski ra del Quemado 


7. Guane 


14. 


Sierra Paso Real 



The genus Viana appears in a series of differentiated forms. 
These vary in sculpture from smooth and shining to dull and spi- 
rally grooved; in color from vcllow to reddish and purple, orna- 
mented at times with red. yellow, or diaphanous bands, or with 






CUBAN GENUS VIANA 



whitish mottling; and in size from the large forms of Sierra Rangel, 
(29.5 mm in width) to the small forms (18.2 mm in width) which 
inhabit small mogotes such as those at Kilometer 14, Vinales. In 
most cases, specimens of Viana collected on small mogotes are 
dwarf forms about one-half the size of typical specimens. It is 
apparent that the size of individual specimens is a function of the 
area of the habitat and may very well be of selective value for 
survival, based upon the supply of food. In addition to other 
variations, the margin of the peristome may be simple and unre- 
flected as in V. regina regina and V. regina laevigata, or more or 
less strongly reflected as in V. regina subunguiculata. 

From the scant field notes we assembled we find that vianas are 
largely satisfied with the presence of limestone, shade, and mois- 
ture. We have never collected a live specimen other than on lime- 
stone rock. Nothing has been published about their food, but they 
probably eat the lichens found on the rocks on which they live. 

They have an enemy in the larval form of the large Cuban fire- 
fly, Alecton sp.; pictures have been taken of such a larva devouring 
a Viana through the aperture (PI. 1). Probably the firefly larva 




PLATE 1 

Viana regina (Morelet) 

Viana regina Morelet being attacked by a lampyrid beetle larva (Alecton 
sp.). Photo by F. de Zayas (about 6 X). 



4 BREVIORA No. 2 C >K 

attacks its pre) when the snail is relaxed. The predator may wait 
for the Viana to relax the operculum and then attack, much like 
the marine predators of the genus Melongena, among others. 

Baker described the radula of Viana ( l l )22) and its anatoim 
(1926). We could find nothing in print regarding its natural 
history. The following notes are translated from a personal com- 
munication from Oscar Alcalde l.edon who. until he left the 
countr) in 1963 was in charge of the Division oi Malacology in 
the r.cwk formed Acadcnna de Ciencias de la Rcpublica de Cuba. 
Alcalde tells us that he unfortunately left his notes behind and 
writes from recollection only. 

I I he eggs] are subglobular in shape, the size of a capital letter 
( )' m normal typewriter print, or perhaps a hit larger. I found them 
under a slab of roek ami the) were covered with the reddish soil on 
which the rock rested Vfter being washed, the) proved to be wax 
colored [whitish]. I did not know at tit si thai the) were Viana eggs, 
but later upon breaking them. I found the ncpionic whorls of a snail 
which under microscopic examination proved to be iden'ical to the 
nepionic whorls of Viana. The operculum was ver) small and thin."' 

According to Thiele ( 1929:81 ). all helicinids lack a male copu- 
lator) organ, and the manner of transfer of sperm from male to 
female is unknovt n. 

Several of the forms of Viana mentioned above were given 
fames b\ the late Dr. Carlos de hi I orre. but almost all remain in 
manuscript. Under these manuscript names the) have been widck 
distributed b\ Cuban collectors, and one of the purposes of the 
present Stud) was to examine the validity i^i these forms. After 
carefull) localizing a large number of Viana specimens on an en- 
larged map of the Sierra de los O.ganos region, we find that we 
are amply justified in rejecting these names in i<>to. 

The confusion in this genus is very much like that of Polymita 
and Liguus. That is. unless the localities of the various forms are 
carefull) plotted, the worker would find it easy to attach names to 
the many color forms and forms with only minor morphological 
differences. A review of the literature covering Polymita and 
/ iguiu has demonstrated amply the confusion which results from 
such a proliferation of names. We choose to follow a more con- 
servative course and to refer to striking varieties, which blend into 
one another, bv vernacular names such as marbled, liratc. smooth, 
banded, roseate, yellow, dwarf, shining, dull. etc.. instead of using 
Latin infraspecific taxa which might be misunderstood as tax- 
onomic designations. 






CUBAN GENUS VIANA 5 

In doing so we follow the procedure generally adopted for such 
a variable species as the intertidal marine mollusk from the north- 
east coast of North America, Thais lapillus (Linnaeus). The 
analogy between the two groups mentioned is quite close, since in 
both cases we are dealing with prosobranch mollusks living in more 
or less isolated colonies on rocks, and hence displaying a rich 
variety of sculpture, size, and color. The common practice today 
is to designate all the lapillus varieties under the simple name 
Thais lapillus. We propose to do the same for Viana regina 
(Morelet). The distribution of the various populations as revealed 
on our maps nevertheless seems to justify the recognition of sub- 
unguiculata Poey and laevigata Pfeiffer as valid subspecies. 

V. subunguiculata extends southwestward from a line running 
between Sierra del Quemado Sierra Vihales (with a very strong 
salient reaching into the Sierra Celadas branch of the Sierra Vinales 
complex) and Sierra Cabezas. Southwestward from this line, sub- 
unguiculata is found reaching to the extreme western limit of the 
distribution of the genus Viana at Sierra de Guane and Sierra 
Paso Real near Guane. This range includes the sierras around 
Luis Lazo (San Carlos, Los Acostas), at Isabel Maria and 
Cabezas, and at the isolated mogotes between Luis Lazo and 
Guane. 

The subspecies laevigata is centered about the complex of 
mountain ranges which go under the name of Sierra de Vihales, 1 
with disconnected outrunners at the Cayos de San Felipe, La- 
gunillas, Matahambre, Pan de Azucar, Cabezas, Isabel Maria, and 
some colonies on the Sierra del Quemado. The line separating laevi- 
gata from regina, sensu stricto, appears to run south between the 
Banos de San Vicente on the west and the Sierra San Andres 
(with La Jagua) to the east. East of this line, and reaching to the 
eastern limit of the range of the genus at Rangel, the nominate 
subspecies regina regina can be found. The mogotes at Kilometer 
1 4 and El Guama also are inhabited by regina regina, as are many 
mogotes in the Vinales valley (Dos Hermanos, Mogote La Vega, 
Hoyo de los Santos, Mogote de los Muertos), whereas others in 
the same area (Mogote Pequeno, Mogote Alto, Hato Morales, El 
Cuajani) have populations of laevigata. 

We realize, of course, that certain populations are sufficiently 
distinct to enable the student to recognize the locality from which 



1 Sierra del Infierno, Sierra Penitencia, Sierra Pan de Azucar, Sierra 
Martillo, and others. 



6 breviora No. 298 

they come. Nevertheless, there arc many specimens for which this 
is not possible, nor did we find it easy to establish fixed dividing 
lines between the various characteristics, with the exceptions noted 
below. Throughout the range of the genus there is much dupli- 
cation o^ the characteristics upon which de la Torre based his 
manuscript names. 

This separation of the genus into three subspecies and numer- 
ous, if not clearly defined, races presupposes an early continuous 
limestone area which, bv subsequent lifting and corrosion, was 
broken up into a series of discontinuous ranges (sierras) and iso- 
lated blocks (mogotes). There is a small amount of territory at 
the southwestern end of Sierra de Vihalcs and at Sierra del Oue- 
mado where laevigata and mbunguiculata overlap with very little 
fusion of characters. However, mure collecting is needed in these 
two areas. I his is also true of the small territorial overlap in 
Vihalcs at El Ouecjue between laevigata and regina. Mechanical 
dispersal may be responsible for this overlap. Gene How. how- 
ever, between members of the same subspecies must be vary rare, 
particularly between isolated populations, such as those oi Tan de 
Guajaibon and Sierra Rangel. Hurricanes could be the means of 
such mechanical dispersal, carrying specimens or their eggs. 

Writing about a large mogote near Vihalcs. called previously T.I 
I umbadero but now El Quequc. Henderson ( 1916:234) states: 

"The rock is discolored and considerabl) altered bj metamorphic 
process, it is |um possible thai the limestone of this southern range 
of the Organos Mountains is ;i shade older than that of the northern 
system (the ( ostanera), hut the folding upward of the two was cer- 
tainl) contemporaneous. All the mogotes of the Viriales valley, in- 
eluding the large Tumbadcro. the Dos Hermanos. and the many 
smaller ones scattered about the valle\ floor are clearly but fragments 
of the main sierra now wholly detached and isolated by erosion 
[corrosion]. The ammonites, which Dr. Torre has been collecting for 
several /ears, come principally from the rocks of this southern range 
and indicate probably Jurassic origin of the limestone. 1 As hereto- 
fore noted, the mogotes about Kilometer 14' do not belong to the 
Vinales range, but together with the Cerro de Cabras hills farther 
west, represent either remnants of a third line of Jurassic sierra that 
once followed a course parallel with the existing ranges, or possiblv 
a later Tertiary deposit upon the flanks of the older mountains." 



'According to We\l (1966: 42-47), the Vinales limestone (Kalk) is of 
Upper Jurassic age and has come by three overthrusts from the north. 



CUBAN GENUS VIANA 7 

Some of the localities cited here must be taken provisionally. 
There are no reliable detailed maps of the Sierra de los Organos 
region. Even the map composed by the Cuban army engineers in 
1933-1941 is too general, not always accurate, and in several in- 
stances illegible. Coupled with this difficulty is the haphazard way 
in which the Cuban peasant assigns names to the various features 
of his local geography. Many mogotes, cerros, vegas, hoyos, and 
arroyos have the same name, but on the other hand, many of 
these features have several. Frequently the name of a mogote is 
that of the farmer near or on whose land it stands; hence, when 
this land changes hands, these names frequently change. There is 
no certainty that such names were ever officially recognized or 
published, or even that they appear on local land records. A final 
difficulty lies in the matter of spelling. The unique Cuban pro- 
nunciation of Castillian is reflected at times in the orthography. 
Thus, ayua, the local name of a tree {F agar a sp.), which is also 
associated with the name of a hill and cove in the Sierra San 
Andres, appears as ayuda, allua and alluda on field labels. In 
one instance a mogote called muela de la vieja (Old Woman's 
Tooth) also appears on a collecting label as colmillo de la vieja 
(Old Woman's Fang). 

The most useful local maps where those published by de la Torre 
and Bartsch (1938) and field maps prepared by Oscar Alcalde 
Ledon and Julio Armenteros. The gazetteer on Cuba published 
by the United States Board on Geographic Names (1957) was 
also very useful. In some cases, however, we have had to be 
satisfied with reporting localities as they appear on the collection 
labels. We do so in the hope that local inquiry at a later date will 
enable the collector to find these localities. 

ACKNOWLEDGMENTS 

We are deeply indebted to Ruth D. Turner for the dissection of 
the soft anatomy and of the radula, and to Kenneth J. Boss and 
Ruth D. Turner for having read the manuscript and having offered 
many valuable suggestions. 

This work was supported largely by National Science Foun- 
dation grant GB-1004. 

SYSTEMATIC DESCRIPTION 

Genus Viana H. and A. Adams 

Viana H. and A. Adams, March 1854, The Genera of Recent Mollusca, 
2: 305 [as subgenus of Trochatella Swainson] (type-species, Helicina 
regina Morelet 1849, subsequent designation, Pilsbry and Brown 1910, 



8 BREVIORA No. 2 l )S 

Proc. Acad. Nat. Sci. Philadelphia, f>2 525); non Walker 1869 [Lepi- 
doptera] 

Hd/uini Gray, November 1856, Ann. Mag. Nat. Hist., (2) IK: 414 (type- 
species, Trochatella regina (Pfeiffer) [sic, Morelet], original designa- 
tion ). 

Rhynchocheila Shuttleworth IS77. Notitiae Malacologicae, 2: 15 (type- 
species, Helicina regina Morelet 1849, monotypic). 

Fitzia Guppj 1895, Proc. Victoria Ihm rrinidad, I. pt. 2: 74 (type-species, 
Helicina regina Morelel 1849, monotypic). 

Description. Shell large, reaching 29 mm in diameter, imper- 
forate, trochiform, generally solid, taint 1 \ to rather strongl) sculp- 
tured, and lacking periostracum in adult specimens. Slid Is uni- 
eolored. or ornamented with bands of varying width, or with a 
marbled or mottled design; a thin reddish subsutural line is present 
in some specimens. I ip simple and occasional!) sinuous or Baring, 
male shell with a notch mid\va\ between the outer lip and its 
insertion in the parietal area. ProtOCOnch \Vl whorls, smooth or 
irregularl) pitted, sharpl) raised above the succeeding whorls as in 
Eutrochatella, generall) lighter in color than the rest of the shell 
except the base 

Operculum calcareous, auriculate, white externall) with a thin, 
light brown, internal corneous layer and darker at the periphery. 
I he outer margin is widel) rounded, the columellar margin con- 
cave above, generall) straighl and descending almost vertical!) 
below. It is rounded at the base, generall) on a single plane, but 
sharpl) raised basall) and less so along the- columellar margin. 
I here is a long, rounded, low. more or less vertical protuberance 
along the inner margin, thickest centrally. Outer margin very 
slightK thickened; a thin, opaque, transverse area reaches internally 
across the mid-line. Nucleus lateral and central near the columellar 
margin, with concentric lines of growth which follow the outline o\ 
the shell. 

Remarks, The notch on the peristome of the male, which we 
hereafter call the peristome notch, was one cause o\' the repeated 
renaming which this genus underwent. Morelet ( 1849: 19) writes 
"Peristome supero. . . profunde emarginato." Gray ( 1856: 414) 
likewise thought the notch was characteristic of the entire group 
and created his genus Hapata. H. and A. Adams (1854: 305) 
also thought this to be true of their genus Viana. Fischer 
(1885: 796), citing Pfeiffer, says "les coquillcs d'une mcnic 
especc ont leur labre tantot echancre. tantot a peine sinueux et 
cette difference serait en rapport avec la sexe." Arango (1878: 
41 ) took the notched shells to be females. Wagner ( 1908: 129) 
correct!) stated that it was the male which possessed the notch, but 
then changed his opinion (1910: 1X5). He was corrected by 



CUBAN GENUS VIANA 9 

Baker (1926: 52), and the dissections performed for this study 
by Turner l support Baker's contention that the notched shells are 
male. Athough Thiele (1929: 83), correctly describes the situa- 
tion, Wenz (1938: 438) and Keen (1960: 288) state that the 
entire genus is characterized by the notch. Wagner (1910: 184) 
noted other features of sexual dimorphism in the shell which our 
examinations did not seem to justify. 

Baker (1922: 62) considered Ampullina De Blainville a syn- 
onym of Eutrochatella sensu stricto, a Jamaican group. However, 
the only helicinids of Jamaica that compare in size with De Blain- 
ville's figure of the type, A. striata (1827: pi. 35, fig. 4a-c), are 
Eutrochatella tankervillei (Gray) and Alcadia major (Gray). But 
neither the description (1824: 441) nor the rather poor figure 
indicate either of these. There is no mention of the strong sculp- 
ture of tankervillei, and the aperture of Alcadia major is quite 
different. 

Ampullina striata De Blainville (1824: 567) (locality un- 
known) has been considered to be in the genus Viana, but a re-ex- 
amination of the figure published by De Blainville in his Manuel de 
Malacologie (1827, pi. 35, fig. 4a-c) indicates that it is not a 
member of this genus. The operculum figured (4b-c) is quite dif- 
ferent from that of Viana. De Blainville shows a strong ridge on the 
inner face from the center margin at the parietal area and extend- 
ing horizontally to the opposite margin. This morphological struc- 
ture does not appear in any of several hundred opercula examined 
in the three subspecies of Viana. The whereabouts of the figured 
specimen (type) is unknown to us. 

The range of Viana extends in a broad arc from the Sierra 
Rangel near San Cristobal, west and south to the immediate vicin- 
ity of Guane, all within the Province of Pinar del Rio, Cuba. 

Viana regina regina (Morelet) 
Plate 2, figures 1 -4 

Helicina virginea d'Orbigny 1842 [in] Sagra, Hist. Tile de Cuba, Mol- 
lusques, 1: 241 (Interieur de l'lle de Cuba); non Lea 1834. 

Helicina regina Morelet 1849, Testacea Novissima Insulae Cubanae et 
Americae Centralis, Paris, 1: 19 (Ad Montes Guajaibon, Insulae Cuba). 
[Paratypes, MCZ 146706, ex Putzey's collection.] 

Helicina maculosa 'Newcomb' Jay 1850, A Catalogue of the Shells Con- 
tained in the Collection of John C Jay, 4th ed., New York, p. 261 
[nomen nudum]. 



1 Dr. Ruth D. Turner is in the process of preparing a detailed anatomical 
investigation of Viana. which will appear later in this series of reports. 



10 BREVIORA No. 298 

Helicina multistriata 'Velasquez' J;i\ 1850 ibid., p. 2M [nomen nudum]. 
Eutrochatella {Hapatd) regina Morelet. Wagner l ( 'i!S, [in] Conchylien- 

( abinet, (2) I. Sect. is. pt. 2: 130, pi. 25, figs. 12. 13, IS. 19. 
Viana regina (Morelet). Baker l l >22. Proc. ^cad Nat. Sci. Philadelphia, 

74: 63, pi. 7, figs. 34-36. 
liana regina marmorata (lone MS) Webb I'MS. Foreign I and and Fresh 

Water shelN. St. Petersburg, Florida, p. 143. pi. 65, fig. 6. 

Description. Shell moderate!) large, turbinate, sculptured with 

spiral cords oi varying degrees of strength, from deeply lirate to 

cjuite smooth, crossed by weak, oblique, irregular growth lines. 
Color basicalK yellow, whitish, or light reddish purple, at times 
with faintly outlined spiral bands. Shell with some degree o'i mot- 
tling, occasionalK covering the entire surface or else being confined 
to the spue. When such a shell is held to the light, the whitish or 
yellowish markings are seen to be more opaque than the base color 
o\' the shell, and the) apparent!) constitute an added layer o\' shell 
matter on the surface. I hese markings, however, are SO closel) 
bound to the rest o\ the shell that they cannot be removed even by 
the most diligent scraping. Whorls 6V£, moderately inflated, regu- 
larl\ widening; bod\ whorl rounded. Aperture widely lunate, white 
near the edge, yellowish or light reddish purple internally. Outer 
lip entire, not reflected and not flaring, except vcr\ weakly near the 
insertion with the columella, and deflected downward at the base ol 
the aperture. Peristome notch rather deep. Y-shaped, narrowly 
rounded at the base. Parietal lip smooth, without folds, gcntl\ 
convex above, sharply and almost verticall) descending below. 
barel) rounded at the juncture with the basal part of the lip. Colu- 
mella without a fold but with a low. oblique, rounded and slightly 
raised margin at the umbilical area, appearing as a slight tubercle 
on the outer basal edge. Shell imperforate, but with a shallow 
excavation just above the base o\ the columella. Sutures moder- 
ately impressed, faintly and irregularis scalloped by the edges of 
the growth lines. Operculum as in the generic description. 



Pan de (iuajaibon. Syntypc. 
Sierra Rangel. 

Pico Grande. Sierra San Andres. 
Sierra de Guacama\a. San Andres. 
Kilometer 14, Vinales. 



Height 


Width 




mm 


mm 




20.5 


23.0 


9 


24.5 


29.5 


6 


21.5 


23.5 


i 


23.0 


25.0 


9 


15.5 


IS. 2 


6 


15.2 


17.5 


S 


16.5 


18.5 


9 






CUBAN GENUS VIANA 1 1 

Remarks. There are many intergrading forms throughout the 
range of this subspecies. Some populations are composed largely of 
a single variety, while others are a mixture of several forms. At 
Guajaibon, the type locality, there exists a pale yellow form, 
occasionally ornamented with a more or less distinct reddish sub- 
sutural line. The shell here is moderately lirate, dull, and rather 
coarse to the touch. The characteristic marbling is confined to the 
upper whorls. At San Andres a rather spectacular form is found. 
The early whorls are pale yellow but this soon gives way to a light 
reddish purple or roseate ground color, variously ornamented with 
irregular white or pale yellow marbling. This marbling is occasion- 
ally arranged in broad spiral bands distinguished by the differing 
nature of the spots. The ground color, as well as the marbling de- 
signs, becomes obsolete near the aperture and at the base, both of 
which are white. The lirations are very faint, not perceptible to the 
touch and barely visible under a magnifying glass, but the shell, 
though smooth, does not possess the brilliant sheen which occurs 
in some of the populations of the subspecies laevigata. In some col- 
onies in the San Andres region (e.g., El Aserradero near Caigua- 
nabo and Puerto Escondido), yellow shells occur together with pur- 
plish ones, as well as some that have varying mixtures of purple and 
yellow. The marbling in both color forms covers practically the 
entire shell. The bands sometimes consist of a solid color and hence 
resemble the prominent spiral bands of the subspecies laevigata. 
Occasional populations, as at La Sierra, have both the yellow and 
purple colors, which, however, are very pale and appear faded, and 
the marbling is widely scattered over the surface. At some isolated 
mogotes (Bella Maria and Talavera near La Palma, Hoyo de los 
Mogotes near Vinales, La Catalina near San Diego de los Banos, 
Mogote del Rojero between Cayo San Felipe and Isabel Maria, and 
some of the mogotes about Kilometer 14) a dwarf form occurs, 
generally in some shade of yellow. 

These remarks are not to be construed as having at all ex- 
hausted the range of variations of regina. Variations of all sorts 
occur, in sculpture, color, and size, sometimes in localities quite 
close to one another (as near Isabel Maria), sometimes even on a 
single mogote (as El Queque). It is not the purpose of this study 
to give a full account of all the forms in which regina or either of 
the other two subspecies occurs, or to give a detailed description 
of each population. Such an undertaking would lead us far be- 
yond our original purpose. 

Specimens examined. CUBA: Pinar del Rio. Vinales: Mogote 
W of Kilometer 14; Mogote E of Kilometer 14; Abra de la Colmena; 



12 MRI-MORA No. 298 

\rrovo Melindre; Encinar Alto; La Laguna, Potrero Constancia; 

Mogote de Jose Maria Garcia; Hoyo de la Sierra; Hato Morales; 
Hoyo de los Mogotes; \\o\o do los Santos; Mogote la Canona; 
Constancia; Mogote Capon; Mogote Dos Hermanns. Vinalet (/'<//- 
marito)'. Ensenada de Antonio Miranda; Vega de Lorenzo Mar- 
line/; Hoyo de Maja; Vegas de Pahnarito; Ensenada Martin 
Miranda; II Grillo; Vega de Lorenzo. Vinales (El Queque [Tum- 
badero]): Ensenada del Valle; lower part of Id Queque; Ensenada 
del Rio; Hoyo de los Muertos; I nsenada de la Bandera. ,S<//; Andres 
I nsenada de la Avua; Pasada de la Auia; Colmillo (or Muela) de 
la Vieja; Mogote 1 a Paloma. N'W ot La Paloma; entrada a la A\ua; 
Sitio de la Sierra; Mogote de la Tumha. Puerto I seondido; En- 
senada del Corojal, Puerto I SCOndido; Mogote del Puerto I seon- 
dido; l a Pastora, Puerto I seondido; Ensenada del Cafetal; Mogote 
Smi.Mi. Mogote del Indio; Pico Grande; Pico Chico; I OS Caracoles; 
Mono del Intierno; Mogote Puis Diaz, Falda de Pico Chico; Mo- 
m>tc las Cuevitas; Hoyo Paige de San Antonio. San Andres (Can- 
ulac). Los Hoyos; Mogote del Abra de Canalete; Ensenada de 
Borges; I nsenada de los Colorados; Mogote de Abraham; Ense- 
nada de Canalete. San .-Indies (Caiguanabo) : El Aserradero; 
Mogote Large Mogote Pucrtccitas; Mogote Andres Diaz; Mogote 
de Caiguanabo; Abra de Caiguanabo. s</// Andres (Consolacidn 
del Norte or La Palnta): Sierra de Guacamaya; Rio de la Puerta, 
Guacamaya; I as Vulicas; Faralldn de las Avispas; Mogote de la 

Palma; Mogote Bella Maria. S ( //; Audit's (Galaldn)'. Mogote Col- 

mena de Piedra; Cueva del Chino; Pinalito; Pas Calabazas; Mogote 
de Galaldn; Mogote Fuerte, S\v of Sierra de Galaldn; Mogote 
Campamento; Mogote Delicias. San Diego de los Bancs. Cueva del 
Indio; Pa Cumbre; Mogote de los Portalcs de San Diego; Mogote 
Colorado; Los CayitOS, Catalina; Mogote de los Indies; Bcrmejales, 
Sierra de la dura; 1 a Catalina. N of San Diego; Pagunita. Sierra 
la (iuira; Hoyo de la Jutia. I km I: of San Diego; Mogote el Bosque; 
(amino de Galaldn, Sierra de la Cumbre; Soroa. I'an de (iuajai- 
hon: San Jose de Sagua. Pan de duajaibon and Sierra Chica. S of 
Pan de Guajaibon. Rangel: Rancho Mundito; El Retire; El lace; 
La Plata; about 2 km S el Rancho Mundito. San Cristobal; Poma 
Sabicu; El Aspire; Guanibadro; about 3 km N of Santa Cruz de los 
Pinos. 

VlANA REGINA LAEVIGATA ( PfeifTer) 

Plate 3. figures 1-4 

/ ochatella regina Morelet. \ar. PfeifTer 1X64. Malakozoologische Blatter, 

II: l()f) (CayOS de San Felipe [Charles Wright); Pan de Aziicai 
[Rafael trango] [both Vinales. Pinar del Rio|. Cuba); PfeifTer 1865, 



Height 


Width 




mm 


mm 




21.5 


25.5 


5 


22.0 


23.0 


$ 


21.0 


24.0 


$ 


21.0 


23.0 


9 



CUBAN GENUS VIANA 13 

Novitates Conchologicae, 2: 253, pi. 44, figs. 4, 5. [Holotype prob- 
ably destroyed.] We here limit the type locality to the Cayos de San 
Felipe close to the town of Vinales. 

Trochatella regina laevigata Pfeiffer 1865, Monographia Pneumonopomo- 
rum Viventium, 3: 211 (Cuba). 

Eutrochatella regina lyrata Torre. Hand 1927, Nautilus, 40: 89 [nomen 
nudum]. 

Description. Shell similar to regina regina, but lacking mottling 
or marbling. It differs from regina subunguiculata in having the lip 
of both male and female specimens simple and not flaring. 



Sitio del Infierno, Vinales. 
La Chorrera, San Vicente, Vinales. 
Cayos de San Felipe, Vinales. 
Sierra Celadas, Vinales. 

Remarks. As is the case with the other subspecies, laevigata is 
variable in size, sculpture, and color pattern. This subspecies seems 
to have more populations in which pronounced, and occasionally 
quite spectacular, banding occurs, as for example the populations in 
several of the sierras of the Sierra del Infierno complex, at the 
Sitio del Infierno and at Cayos de San Felipe. The last two locali- 
ties have some of the most beautiful shells in the genus. They are 
gleaming roseate, purplish, or bright lemon-yellow and generally 
have three rather broad alternate whitish bands. Many of the 
lemon-colored shells have a thin, dark-reddish subsutural line. As 
is the case with the other two subspecies, some populations consist 
of shells differing in color, ornamentation, and sculpture, whereas 
others are more or less uniform. 

At Sierra del Quemado and Sierra Celadas there are populations 
of subunguiculata as well as laevigata. At El Queque, besides 
laevigata, populations of regina also occur. See remarks under 
subunguiculata. 

Specimens examined. CUBA: Pinar del Rio. Vinales: Sitio 
del Infierno; Mogote del Cejanal, El Abra, Sierra de Vinales; La 
Penitencia; Pan de Azucar; Sierra de Pan de Azucar; Sierra de 
Vinales; Cayos de San Felipe; Sierra Derrumbada; El Cuajani; Hoyo 
del Majagual; Sierra Serrucha; Hoyo de la Jutia; Mogote de los 
Muertos; Mogote de la Vega; Mogote Pequefio; Potrero de Manuel 
Sanchez (near Capon). Vinales (San Vicente): Las Cuevitas, Finca 
Ancon; Liana Manacas; Mogote La Canona (between Laguna de la 
Piedra and Constancia); La Guasasa; Ensenada de los Banos; San 
Vicente de los Banos; Puerta del Ancon; Costanera de San Vicente 



14 BREVIORA No. 298 

i or Ancon ) ; Valle de Delicias; Finca Ancon; La Chorrera; Cejanal, 
El Abra; Punta de la Costanera; Ensenada de los Banos; Mogote 
Pequeno, Costanera de San Vicente (or Anc6n); Cueva de Jose" 
Miuucl (ionic/; \lo\o de Magdalena, Costanera de San Vicente; 
Motiotc del Cao, Lacuna de la Piedra; Moeote de Justo; Cueva del 

DC 

Rio San Vicente; Sierra Gorda, Ancon; Sierra del Aneon. Isabel 
Maria: Monotc Isabel Maria; boundar\ between Isabel Maria and 
( abe/as. 

\ I \N \ Kl ois \ si in NGI ICULATA | Poe\ ) 
Plate 4. figures 1-4 

Helicina subunguiculata Poe) is s ". Memori s sobre la Historia Natural «.lc 
la (sla de ( uba, Habana, 2: 14 (Sierra de Guane). [Lectotype, here 
selected, M< Z 73672.] 

Description. Shell similar to the typical form but differing in the 

nature of the lip. which in the female is expanded and slightly re- 
flected, in the male is extended Into a claw-like structure which is 

sometimes expanded, especiall) at the base where it joins the COlll- 

mellar margin. 



Siena de ( mane. S\nt\pc. 

Punta de Sierra de ( mane. 

Siei r.i San Carlos. I.uis 1 .a/o. 
Sierra Celadas. \ males. 

Remarks. In si/e. the shells var) from the medium-sized ones 

found at the type localit) and near Mendoza to the rather large 
ones from Luis I. a/o and especiall) on the Sierra Celadas of the 
Siena Yirialcs complex. I he shells at the type locality are white 
or pale \cllow in color, and sculptured with spiral lirations of mod- 
erate strength so that the) arc moderately rough to the touch. \i 
La Muralla on the road to Luis I. a/o the shells are larger, quite 
smooth, although with \cr\ taint. wideb spaced luations. pale 
lemon-yellow, and frequent!) ornamented with a red subsutural 
band. These shells also display some degree of whitish banding 
and mottling on the spire as in some populations o! regina, but 
their large si/e. and especially the Baring lip in the female, and the 
strong and at times flaring ungulation in the male, reveal their true 
subspecific standing. At Luis Lazo the shells arc similar, the lira- 
's somewhat stronger, and many purplish or roseate specimens 



Height 


Width 




mm 


mm 




24.0 


24.5 




22.0 


24.0 




23.5 


26.5 


9 


25.0 


28.5 





CUBAN GENUS VIANA 15 

appear. At Gramales and especially on the Sierra Celadas a popu- 
lation of large, heavy, smooth, generally pale yellow shells occurs, 
with a strongly flaring lip and white bands that vary from moder- 
ately broad to quite narrow. Occasionally a purplish specimen 
occurs. This population seems to share the Sierra Celadas with a 
typical broadly banded, purplish colony of laevigata. In all likeli- 
hood the two subspecies, as at Sierra del Quemado, live in isolated 
areas in the sierra. It must be borne in mind that the sierras of the 
Sierra Vifiales complex do not necessarily constitute a continuity 
of ecological conditions. Isolation of populations sometimes occurs 
as the result, among other things, of gaping chasms or hoyos that 
present effective barriers even to calciphiles like Viana. A popula- 
tion of stenomorphs, light purplish in color, of this subspecies lives 
on a mogote at the border between Isabel Maria and Luis Lazo. 

Specimens examined. CUBA: Pinar del Rio. Guam: Sierra 
Paso Real; Sierra de Guane; Mogote 1 km N of Mogote Punta de la 
Sierra; Punta de la Sierra; Mogote near Punta de la Sierra. Luis 
Lazo (Sumidero): Entrada a Sumidero; Ensenada de los Barrios; 
La Picapica; Ojo de la Palma, Finca "La Giiira/' Luis Lazo: La 
Estrechura; Valle de San Carlos; Sabanas Lianas; Sierra Los Acos- 
tas; Valle de Luis Lazo (Valle de San Carlos [El Potrerito]); La 
Claraboya; La Giiira; Mogote Central; El Junco. Isabel Maria 
(Guamd): El Guama; Hoyo El Guama; Mina El Guama. Isabel 
Maria (Cabezas): Ensenada Chica; Valle Cabezas; Mogote del 
Cementerio; Ensenada los Burros. 

REFERENCES CITED 

Adams, H., and A. Adams 

1854. The Genera of Recent Mollusca. London, 2: 300-309; 3: pi. 87. 

Arango, Rafael 

1878-1880. Contribution a la Fauna Malacologica Cubana. Havana. 
280 pp. (-)- 35, index and errata). 

Baker, H. Burrington 

1922. Notes on the radula of Helicinidae. Proc. Acad. Nat. Sci. 

Philadelphia, 74: 29-67, pis. 3-7. 
1926. Anatomical notes on American Helicinidae. Idem, 78: 35-56, 

pis. 4-8. 

Blainville, H. M. Ducrotay de 

1824. Dictionaire des Sciences Naturelles. Strassbourg and Paris, 
32: 567. 

1825. Manuel de Malacologie, Strassbourg, 8 + 632 pp.; 2 folded 
tableaux, 87 pis. (1827). 



16 BREVIORA No. 298 

BOWDICH, T. E. 

1822. Moments of ( onchologv. Paris. 31 pp.. pi. 9, fig. 2. 

CLfcNCH, W. J.. AND M. K. JACOBSON 

1966. On Ampullina Guppy. Nautilus. 80: 71. 

I is* ut-R. P. 

I S 8 5 . Manuel de Conchyliologie, Paris. Helicinidae, pp. 794-798, 
figs. 554-556. 

Gray. J. E. 

1856. Notice of a new subgenus oi Helicinidae. Ann. Mag. Nat. 
Hist., (2) 18: 414. 

Hi ND1 ksiin. Inns H 

1916. The ( raise of the rbmas Barrera. Ci P. Putnam's Sons. 
I ondon and New York, 320 pp . 42 pis., i map. 

KlIS, \1\ R\ 

i960. Helicinidae. [in] I leatise on Invertebrate Paleontology, Moi- 
lusca. I: 285-288, Ggs. 186-187. 

M< 'Kill I . AKI 111 H 

I S49. restacea Novissima Insulae ( ubanae el Americae Centralis, 
Pans. 1 : 1-31. 

Pi i ii i i r. I in is 

1864. Zur Molluskenfauna von Cuba. Malakozoologische Blatter, 
l l : (02-109. 

1X65. Monographia Pncumonopomorum Viventium, Suppl. 2 ( assel. 
240 pp 

I III! 1 I . Joll \NM s 

1929. Handbuch der Weichtierkunde. Jena. I: 80-91, figs. 57-67. 

1 orri . Carlos di la, and Paui Bartsch 

1938. rhe ( uban operculate land shells of the subfamily C hondro- 
pominae. Proc. U. S. Nat. Mus., 85: 193-403, pis. 7-39, figs. 
71-101. 

Unii i ii si \ 1 1 s Board on Geographic Names 

1957. Gazetteer No. 30, Cuba. Washington, DC. 

Wagnlr, Am on 

1907-1908. [in] Martini and Chemnitz, Systematisches ( onchylien- 

Cabinet ( 2 ) . 1 : Sect. 1 8, pt. 2. pp. 46- 1 38. pis. 7-25. 

1910. Uber die Formenunterschiede der Gehause bei mannlichen 
und weiblichen Indi.iduen der Heliciniden. Abhandl. Senckenb. 
Naturforsch. Ges.. Frankfurt-am-Main. 32: I81-186.pl. 16. 






CUBAN GENUS VIANA 



17 



Wenz, W 

1938. Gastropoda, [in] Handbuch der Palaozoologie, Berlin, 6(1) 
435-448, figs. 1071-1118. 

Weyl, Richard 

1966. Geologie der Antillen. Berlin. 410 pp.. 24 pis., 130 text-figs. 

(Received 17 November 1967.) 




5KM 



O 

5 

N 




Map 2. Sierra Paso Real to Sierra San Carlos. 1 

1. Guane (town); 2, Sierra Paso Real; 3, Sierra de Guane; 
4, Los Porta!es; 5, La Muralla; 6, Teneria. 
A. Sierra San Carlos. 



' Maps 2 to 5 were redrawn from those given by Torre and Bartsch 
(1938) and the military (topographic) maps of Cuba (1933-1934). 



18 



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20 HRKVIORA No. 29S 



Map 4. Siena del Quemado to Sierra Guacamava. 

H, Sierra del Quemado. I. Siena Celadas; 11, Santo Tom&s 
(town). J. Sierra Pan de Azucar. K, Siena Martillo. L, Siena 
del Infierno; 12, Cayos de San Felipe'; 13. El Guama; 14, Mogotes 
de Km 14. M. Sierra Penitencia. N, Sierra El Abra. O, Sierra 
Vinales; 24. Mogote Palmarito; 25, Hoyo Magdalena. P, El 
Queque; 17. Mogote Vigil; IS. Mogote Rojas; 19, Abascal; 20. Dos 
Hermanos; 21. Vinales (town); 22, Hoyo de los Santos; 23. 
Ensenada Miranda; 26, Puertecites; 27, Mogote Capon. Q, 
1 a Chorrera; 28, Hoyo de Jaruca; 29. Laguna de la Piedra; 
30, MogOte Jutia; 31, Fogon de los Negros; 32, West ridge of 
the Siena San Vicente; 33. East ridge of the Siena San 
Vicente; 34. Ratios de S.m Vicente. R. Sierra Cialeras; 15. Pan 
de Azucar. S. Sierra Ancon; 61, San Cavetano (town). T, 
Sierra San Andres; 35, Mogote Mina; 36, Rinconada; 37, La 
Jagua: 38, AsientO de la lamia; 39, Mogote de la Jagua; 40. 
Hoyo Corto de San Antonio; 41, Canalete (town); 42, San 
\ndres ( town ) . 43, Mogote funic; 44. Mogote de Canalete; 
45. MogOtes de Canalete; 46, Ensenada de la Ayiia; 47, Pasada de 
las Escaleras; 48, Puerto de San Andres; 50. Pico Grande; 51, 
c olmillo de la Vieja; 53, Sitio de la Sierra; 54, Mogote Bella 
Maria. U. Sierra Gaucamava; 55, Mogote Grande; 56, Caiguan- 
abo; 57, Pinalito; 58, Mogote I argo; 59, Mogote Angelena; 60. 
Mogote Mamey. 

I ocalities aumbered 49 and 52 represent place names that could not be 
located on an\ maps available to us. 



1 Not to be confused with the small islands "Cayos de San Felipe" off 
the northwest coast of the Isla de Pinos. 



CUBAN GENUS VIANA 



21 




~>-> 



HRF.VIORA 



No. 298 




N 

i 



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M 



• »4 



Map S Sierra Guacamaya to Sierra Rangel. 

U. Sierra Guacamaya. V, Sierra Cialalon; <> I . Caracoles; r>2. 
Mogote Colmena de Piedra; 63, I os Portales. W, Sierra La 
Giiira; '4. San Diego de los Banos (town); 65, Los Cayitos de 
Catalina. 66. Mogote Bosque; 67. I a < timbre. X, Pan de 
Guajaibon. \ . Sierra RangeL 



CUBAN GENUS VIANA 



23 




PLATE 2 
Viana regina regina (Morelet) 

Fig. 1. Pan de Guajaibon, Consolacion del Norte, Pinar del Rio (para- 
type, MCZ 146706 9 ). 

Figs. 2, 3. Pico Grande, Sierra de San Andres, Consolacion del Norte 
Pinar del Rio (MCZ 256282(4 ). 

Fig. 4. La Catalina, San Diego de los Banos, Consolacion del Sur, Pinar 
del Rio (MCZ 126684 9 ) (all 2 X). 



24 



HRHVIORA 



No. 298 




PI ME 3 
I mini regina laevigata (Pfciffer) 

I k,s i. : ( ,i\ds de San Felipe, Vinales, Pinar (.Id Km (Topo types, 
\K / 126716 ). 
Fic ; sun., Celadas, Vinales, Pinar del Rio I \l( / 256173 |. 
lh, 4. Pan de Azucar, Vinales, Pinar del Rio (MCZ 126703 I (all 

: \). 



CUBAN GENUS VIANA 



25 




PLATE 4 
Viuna regina subunguiculata (Poey) 

Fig. 1. Siena Celadas, Santo Tomas, Vinales, Pinar del Rio (MCZ 

256342 & ). 

Fig. 2. Sierra Celadas, Santo Tomas. Vinales. Pinar del Rio (MCZ 
256342 6 ), to show the outer peristome notch in the male. 

Fig. 3. Valle San Carlos, Luis Lazo, San Juan y Martinez, Pinar del 
Rio (MCZ 256335^ ). 

Fig. 4. Sierra de Guane, Guane, Pinar del Rio (Lectotype MCZ 
73672 6 ) (all 2 X). 



BREVIORA 



Museum of Coeip&irative Zoology 

Ca mbridge, Mass. 11 October, 1968 Number 299 

WHAT IS LUMBRICUS EISENI LEVINSEN, 1884 
(LUMBRICIDAE, OLIGOCHAETA)? 1 

G. E. Gates 2 

Abstract. Levinsen's species, eiseni, after being in the synonymy of an 
unrelated form and in three lumbricid genera, is excluded from Bimastos, as 
defined by characters less liable to rapid evolutionary modification than 
those of the genitalia on which the classical system of the Oligochaeta 
(Michaelsen, 1900 — Stephenson, 1930) was mainly based. The species, now 
more adequately characterized, cannot go into any other lumbricid genus 
because of current ignorance of somatic anatomy in their type species. 
Relationships suggested by available anatomical data are with Allolobophora 
chlorotica and, less closely, with Dendrobaena rubida and Lumbricus spp. 

As the references below show, no agreement has been reached 
as to the status of Levinsen's species and as to its generic affilia- 
tions. The taxon, like other more or less widely spread lumbricid 
anthropochores as well as most endemics of the family, never was 
adequately characterized. Although much less material could be 
procured than was desired, study of new material has enabled 
the following contribution toward a solution of the problem. 

"Lumbricus eiseni" Levinsen, 1884 

Lumbricus eiseni Levinsen, 1884, Vidensk. Meddel. Naturhist. Forhandl. 
Copenhagen, (4) 5:311, 241. (Type locality, Deer Garden, Copenhagen. 
Types, 5, in the Copenhagen Museum.) 

Allolobophora (Dendrobaena) mew'.-Friend, 1892, Jour. Linnean Soc. Lon- 
don, (Zool.), 24:302. 

Helodrilus (Bimastus) mem.-Michaelsen, 1900, Das Tierreich, 10:503. 

Bimastus eiseni, Friend, 1911, Zoologist, (4) 15:145. 

Eisenia parva f. typica (part) Pop, 1950, An. Acad. Romane, Sect. Sti. Geol. 
Geogr. Biol., (A) 1 (9):89. 

Eisenia eiseni, -Graff, 1953, Regenwiirmer Deutschlands, p. 27. 



1 From research financed by the National Science Foundation. 

2 Zoology Department, University of Maine, Orono, Maine, U.S.A. 



2 breviora No. 299 

SPECIMENS EXAMINED 

Channel Islands. Herm. under rock above tide mark, June 24, 1925. 1-0-0. 

G. E. Pickford. 
South Africa. Cape Province. Cape Peninsula. Kirstenbosch, rotting stump, 

October 29, 1925, 5-0-4. Table Mountain, Window Gorge, rotting log at 

400 feet, July 30, 1926, 0-0-9. Brockenhurst. Queen's Bower, April 2, 

1925.0-0-1. G. E. Pickford. 

Interceptions by U.S. Bureau of Plant Quarantine 

i ire. From soil with twenty-nine narcissus bulbs and two bundles of lily 
plants, in baggage on plane, arriving at Boston, June 22, 1964, 0-0- 1. 

Scotland. From heather plants, in plane, arriving at New York, October 26, 
1949.0-0-1. 

Germany. From cabbage, in ships stores on Norwegian M S I ancing, ar- 
riving at Savannah, Georgia. October 2 l >. 1964, 0-0 1. 

Portugal. From soil with five azaleas, Bve ferns, two orchids, in baggage 
on plane, arming at Boston. M,i\ 21, 1965, 1-0-0. 

Azores. From soil with one begonia, in baggage on plane, arriving at Bos- 
ton, June 18, 1965,0-0-1. 

From roots of plains o\ ( 'Una sp.. in baggage on plane, arriving at Bos- 
ton. June 18. 1965. 2-0-0 

DESCRIPTION 

External characteristics. Segments. 79-1 12 (cf. Table 1 ). Color, 
brown, slate to almost black but never red, restricted to dorsum 
except in ii-vi or vii, most obvious anterior to the clitcllum, usually 
lacking or very sparse in buccal and anal segments. Body sub- 
circular in cross section posteriorly. Prostomium, tanylobous (all). 

Setae, all present in ii. the d setae at or below mL anteriorly but 
posteriorly above that level. BC usually < AA throughout post- 
clitellar region where DD < AC. 

Nephropores, obvious. On one South African worm they are 
above B on the left sides of x-xiii, xv, xvi, xix-xx, xxvi, on the right 
sides of ix-xi. xv, xxii-xxvi, xxix. Other pores of that worm, 
in the region of iv-xxix. are well above D. Pores of xv-xvi, except 
on one other worm (left pore of xv not found), were lateral to B. 
Pores may be at the same level on one side for three, four, or five 
consecutive segments, but on the opposite side asymmetry is oc- 
casional. Pores of xii have been seen (rarely) just below C. The 
arrangement throughout most of the axis can be characterized as: 
irregularly alternating, usually between two levels, one slightly 
lateral to B and another about midway between D and wD.with 
occasional intrasegmental asymmetry. 



1968 LUMBRICUS EISENI LEVINSEN 3 

First dorsal pore, at 5/6 (21 specimens), occasionally smaller 
than the one at 6/7. 

Female pores, slightly lateral to B, equatorial in xiv. Male pores, 
minute, at or median to mBC, each laterally in a small cleft at 
eq/xv. Male tumescences, confined to xv and to median half of BC. 

Clitellum, red (formalin preservation), saddle-shaped, reaching 
ventrally to or nearly to A, or mostly annular, in xxiv-xxxii but 
restricted to DD in xxiv and xxxii (13), xxiv-xxxiii/n (1), xxv- 
xxxi (2, possibly not fully developed). Even at apparent maximal 
development the clitellum is thinner in AA than laterally and inter- 
segmental furrows still are recognizable ventrally from mBC. 

Genital tumescences, often conspicuous, without distinct boun- 
daries, transversely elliptical, including a,b, in xvi (27), xxiv (2), 
xxiv-xxv (l),xxv (26). 

Internal anatomy. Septa, none thickly muscular, 10/11 bulged 
anteriorly and 11/12 posteriorly by the calciferous gland, the 
coelomic cavity of xii, xiii and xiv quite small. Pigment, lacking 
in peritoneum, present anteriorly in the longitudinal muscle band 
at mD but disappearing behind the clitellum, associated with or in 
circular muscle layer, apparently brown, yellow brown, or dark 
brownish red. Pharyngeal glands to 8/9. 

Brain, circumpharyngeal connectives and subpharyngeal gan- 
glion all left in iv (19), sometimes well posteriorly, by a transverse 
section exactly along 3/4. 

Ventral bundles of the longitudinal musculature, according to 
Prof. Harman (in litt.), can be characterized as fasciculate, the 
pinnate arrangement so characteristic of some species of Lumbricus 
being lacking. The cells form compact "Kastchen" ventrally and a 
compact band dorsally. Ventrally the bundles resemble those of 
A. chlorotica. 

Calciferous sacs, in x, elongated horizontally rather than ver- 
tically, anteriorly or anterolaterally directed, reaching to or nearly to 
9/10, apparently opening posteriorly into gut lumen at level of 
insertion of 10/11. Calciferous lamellae, extending nearly to 
anterior end of sac but in x small and of squarish shape. Gut 
much widened, rather bead-shaped and with vertically slit-like 
lumen in xi, narrower (half as thick as in xi) and of nearly uni- 
form calibre through xii-xiv. A deep internal constriction (or 
vertical groove) on one or both sides at or slightly behind insertion 
of 11/12. Calciferous lamellae, large and rectangular in xi, but 
behind insertion of 11/12 quite small, not reaching 12/13 (if 
present posteriorly too small to be recognized in dissections). The 
inner lining of the gut was rotted away in slightly macerated speci- 
mens so as to reveal lamellae with diagrammatic clarity. Intestinal 



4 breviora No. 299 

origin, in x\ (20). Gizzard, in xvii (20). the layei with brilliant 
muscular sheen in horizontal sections narrowing behind insertion 
of 17/18 and not reaching 17/18. A postgizzard valve, it" present, 
is relaxed in each worm but the gut is somewhat narrowed at 
18/19 or in xix. Tvphlosole. beginning gradually in region of xx- 
\\i\. at first a low horizontal band, then becoming vertically and 
rather thickly lamelliform. A slight longitudinal groove may be 
recognizable on the ventral face at mV. or, if not. a slight longi- 
tudinal flange ma) be visible on each side. (Condition does not 
allow a more precise characterization.) The tvphlosole ends, 
abruptly in unamputated specimens, in xc-xevii (Table 1 ), leaving 
8-15 segments (proctodeal?) atyphlosolate. 

Dorsal blood vessel, continued onto pharyngeal bulb. Sub- 
neural, adherent to nerve cord, l:\tra-oesophageal trunks, median 
to hearts, posteriori) in xii passing up to dorsal trunk (21). 
Hearts, present in viii-xi (22). Quite slender vessels at sites (in 
other species) of hearts o\ vii were traceable to ventral trunk only 
in three worms in each of which onlv the dorsalmost quarter is 
heart-like. 

Nephridia, vesiculate. Bladders, each in shape oi a long U- 
shaped loop, from A well into DD, both limbs of the loop of about 
the same length, posterior limb passing into parietes near H. The 
one other major loon is longer than the bladder loop. 

Holandric. Testes, flattened horizontal discs sometimes rather 
fan-shaped or saccular and pyriform. Male funnels, crenellate. 
large. Male deferent ducts, rather slender, almost straight, slightly 
sinuous, with several, very short, u-shaped loops, once a single long 
hairpin-loop, or coiled into a ball, loops or coils up to a millimeter 
awa) from funnel septum. Ducts of a side, uniting posteriorly in 
xii. continued to cq w across top of atrial gland (latter developed 
after penetration of duct into parietes'.') or disappearing from 
sight at edge oi gland. Seminal vesicles, rather small to small but 
not vestigial, vertical or dorsal, soft, without obvious lobing, in 
xi-xii (15). Anterior vesicles, flattened in the space between 
septum 10 11 and the anterior face of the calciferous gland. 

Ovaries, often large and then reaching up to level of dorsal face 
of gut. filled except in proximal quarter or so with large ova, each 
with a single egg string containing one to three ova. Ovisacs, with 
one to five ova which were easily released by teasing, especially 
those in a distal portion. Oviducts, rather thick passing into 
parietes lateral to D. 

Atrial glands (14 clitellate worms), confined to median portion 
of BC or reaching nearly to C, flat or of a rather low dome-shape, 
usually with a cleft at eq/xv and another at insertion of 15/16. 



1968 LUMBRICUS EISENI LEVINSEN 5 

reaching to or well toward 16/17. TP glands, lacking (all). GS 
follicles, large, conspicuously protuberant into coelomic cavities, 
genital setae elongated and otherwise modified. Follicles of ventral 
couple of xv not as much enlarged as the others. Supraparietal 
glands around GS follicles, lacking (all). Follicles of all ventral 
couples, from the clitellum anteriorly, larger than those of the 
lateral setae. 

Variant specimen. Clitellum, light pink rather than red, reach- 
ing little below C, but of near maximal thickness in region of mD 
except at intersegmental furrows which are still obvious even there. 
Male funnels, smaller than usual but crenellate, with very slight 
iridescence. Male deferent ducts pass into parietes at eq/xv. 
Atrial glands lacking (no rudiments recognizable in longitudinal 
musculature). GS follicles smaller than usual. Distal two-thirds 
of ovaries with apparently mature ova. No ova in ovisacs. 

The clitellum probably had not attained full development. 
Absence of atrial glands does not, however, appear to be attrib- 
utable to immaturity. Hence the worm may be of a strain (possibly 
with obligatory parthenogenesis and markedly reduced spermato- 
genesis) in which the atrial glands already have been lost and in 
which genital setae as well as tumescences are disappearing. Elim- 
ination of genital tumescences and genital setae would result in a 
state similar to that of Bimastos species. 

Reproduction. Spermatophores were present externally in 
clitellar region of two worms but no iridescence was recognized in 
the opaque matter within the transparent cases. Spermatozoal 
iridescence was lacking on funnels of one worm but on those of 
other clitellate worms does not seem to be as brilliant as in species 
with obligatory cross-breeding, and was not recognized in male 
deferent ducts. Sparse maturation of sperm is associated, in some 
lumbricids, with parthenogenesis. Pyriform testes are unusually 
soft, much of the interior filled with a watery fluid in which there 
are many morulae but only a very few sperm. Uniparental repro- 
duction has been demonstrated for this species by Evans and Guild 
(1948). 

Bimastos eiseni is diploid, 2n = 32, according to Muldal 
(1952:61). Parthenogenesis is not, however, contraindicated by 
the diploidy, that method of uniparental reproduction being obliga- 
tory (idem:66) in diploid individuals of Octolasion tyrtaeum 
(Orley, 1881). 

Remarks. Soil particles of any sort were completely absent from 
guts of each of these worms, the lumen being filled with brown 
organic matter. 



6 BREVIORA No. 299 

Distribution. Ireland, Scotland, England, Channel Islands. 
Sweden. Denmark. Germany. Poland. Belgium, Czechoslovakia, 
I ranee, Switzerland, Austria. Italy (including Sicily). Jugoslavia. 
Albania. Bulgaria. Portugal. 

Azores, Madeira. Canary Islands, South Africa, St. Helena. 
India (Western Himalayas only), New Zealand (including Stewart 
Island ). 

The extra-European records show that the species has been fre- 
quently transported by man with man) successful colonizations. 
Absence from the Americas where so main other lumbricid anthro- 
pochorcs are well established is puzzling. In Europe, presence 
above the southern limit of Ouaternar\ glaciation may also be a 
result of unintentional human carriage. 1 he original home of the 
species is unknown. It must have been somewhere in Eurasia but 
how much oi the distribution in southern Europe is fortuitous, 
remains to be learned. 

I ( »cies has been called cosmopolitan < Omodco, 1950:5) 
and Euro-American (Omodeo, 1961, table). The first adjective 
hardl) seems applicable to a species absent from all Americas. 
most o\' Africa and all oi Asia excepl for a couple oi localities in 
the western Himalayas. The second characterization is in- 
applicable. 

Systematic relationships. The clitellum is shorter posteriorly 
than in English definitions (Cernosvitov and Evans, L947; (icrard, 
1964). Paucity oi data about individual variation in eiseni at 
present ob\ iates further discussion of the differences. 

The species docs not belong in Bimastos as that genus now 
((iates. in press) is defined. The calciferous sacs of segment x 
suggest relationships with / umbricus, Dendrobaena rubida and 
related species (but certainly not 1). octaedra and related species), 
as well as Allolobophora chlorotica. Further evidence is provided 
by the similarity, in each taxon, of nephridial vesicles and ncphro- 
pore locations. Main characters provided by those and other 
somatic systems now seem resistant to rapid evolutionary modi- 
fication and so are oi greater systematic importance than the more 
easily changed genitalia on which classical specialists so much 
relied. 

The purple red pigment by which Dendrobaena, Eisenia and 
Lumbricui were defined (Pop, 1941 ) is lacking in chlorotica and 
eiseni. The longitudinal musculature of Lumbricui spp. and of D. 
rubida (also closely related species?) is pinnate in cross-section 
rather than fasciculate as in eiseni and chlorotica. Atrial glands 
(ignored by classical authorities) are indeed present in D. rubida 



1968 LUMBRICUS EISENI LEVINSEN 7 

and related species just as in eiseni and chlorotica but are lacking 
in all species of Lumbricus studied by the author. Those differ- 
ences seem to suggest that chlorotica and eiseni may be more 
closely related to each other than either is to any of the other 
above-mentioned taxa. 

Then, should eiseni be transferred to Allolobophora of which 
chlorotica recently was made the type species? If the two species 
do belong in the same genus, that is where eiseni seemingly should 
go. A. chlorotica, however, differs from every one of the common 
allolobophoras by its calciferous sacs, by the presence of well 
developed atrial glands (lacking in every one of the common 
allolobophoras), as well as by other characters. Information now 
available indicates that: 1) Enterion chloroticum Savigny, 1826, 
should not have been designated the type of Allolobophora. 2) All 
of the common allolobophoras, if that type designation in some 
way cannot be nullified, must go into another and possibly new 
genus. 

The common species of the classical Allolobophora are now 
widely distributed throughout much of the world. They success- 
fully colonized many areas to which they were introduced and 
even now are spreading, and being spread both accidentally and 
deliberately, through considerable areas of Australia and New 
Zealand. Those same species also have been widely used in experi- 
ments. A. chlorotica and B. eiseni, on the contrary, were much 
less often transported, have not colonized so extensively, are much 
less common, and only rarely have been studied experimentally. 

Attention was directed on two previous occasions (Gates, 
1958a, b) to the fact that eiseni does not belong in the American 
genus Bimastos, primarily in the hope that some interest in the 
problem would thereby be aroused on the part of those who have 
access to more plentiful material. Because of our ignorance of so 
much essential data for a large majority of lumbricid species, no 
nomenclatural changes are made herein. 

The single North American anthropochore, Bimastos parvus 
(Eisen, 1874), is not of the same genus as eiseni and just as ob- 
viously does not belong in Eisenia. What Eisenia parva of sundry 
authors was, probably can only be determined from an examination 
of the specimens. 

Parasites. Nematodes from coelomic cavities of the Scotch 
worm, according to Dr. S. Prudhoe of the British Museum, prob- 
ably are Dicelis filaria Dujardin 1845. Nematodes from the ventral 
blood vessel in x-xiv of a Channel Islands juvenile were identified 
by Dr. Maybelle Chitwood as larvae of Spiroptera turdi Molin by 
Cori, 1898. 



8 breviora No. 299 

REFERENCES CITED 

Cernosvitox . I ... and A. C. Evans 

1947. Lumbricidae. Linnean Societj of l ondon Synopses of British 
Fauna, No. 6: 1-36. 

Evans, A. C. and W. J. di ild 

l l >48. Some notes on reproduction in British earthworms. Ann. Mag. 
Nat. Hist . (1MI 1:654-659. 

Gatks. G. E. 

1958a. On endemicity of earthworms in the British Isles \nn. Mag 
Nat. Hist . ( 13) 1:33-44. 

1 958b. Contribution to a revision of the earthworm family Lum- 
bricidae. II Indian species. Breviora, Mus. Comp. Zool., No. 
<>l:l-16. 

In press. On tWO American genera of the earthworm family I um- 
bricidae. Jour. Nat. Hist 

(,l RAKD, B. M 

1^64 Lumbricidae. linnean Societj of London, Synopses oi British 
i auna, 2nd ed . No. 6:1-58. 

Ml I DAI . S 

1952 I he chromosomes of the earthworms. I. I he evolution of poly- 
ploidy. Heredity, 6:55-76. 

( )M()DI (). P. 

1950. Oligocheti. \nn. Inst Mus. Zool. Univ. Napoli, 2(10): 1-12. 

1961. l e peuplemenl des grandes iles de la Mediterranee par lesoligo- 

chetes terricoles, Colloques Internatl. Centre Natl. Rech. Sci., 

No. 94:127-133. 

Pop. \. 

1941. /ur phylogenie und Systematik der Lumbriciden. Zool. Jahrb. 
Sysl Okol.Geogr. Tiere, 74:487-522. 

(Received 16 January 1968.) 



968 



LUMBRICUS EISENI LEVINSEN 



TABLE 1 

Typhlosole termination and segment number 
in "Lumbricus eiseni" 







Atyphlo- 


Number 




Serial 


Typhlosole 


solate 


of 




Number 


ends in 


segments 


Segments 


Remarks 


1 


70 


9 


79 


Posterior amputee* 


2 


74 


7 


81 


Posterior amputee* 


3 


82-85 


3 


88 


Posterior amputee* 


4 


80-86 




86 


Posterior amputee* 


5 


88 


11 


99 


Channel Islands 


6 


90 


9 


99 




7 


90 


13 


103 




8 


91 


13 


104 




9 


92 


9 


101 




10 


92 


12 


104 


Posterior amputee* 


11 


92 


15 


107 




12 


93 


12 


105 




13 


95 


8 


103 


Posterior amputee* 


14 


96 


13 


109 




15 


97 


15 


112 





Specimens from South Africa unless otherwise indicated. 

* After loss of the posterior end, the presently terminal metamere had been 
reorganized and now looks much like a normal anal segment. Proof of its 
original, preterminal position is provided by vestiges of former apertures 
of setal follicles. The latter were lysed after dehiscence of their setae. 



BREVIORA 

Museumi of Comparative Zoology 

Cambridge, Mass. 11 October, 1968 Number 300 

CONTRIBUTIONS TO A REVISION OF THE LUMBRICIDAE. 
III. EISENIA HORTENSIS (MICHAELSEN) (1890). 1 

G. E. Gates 2 

Abstract. The classical system of the Oligochaeta (Michaelsen, 1900- 
Stephenson, 1930) and proposed neoclassical revisions, did not permit defini- 
tive conclusions as to disposition of a taxon variously known as var. or 
/. hortensis which has been in each of five lumbricid genera. Characters 
less liable to rapid evolutionary modification than those of the genitalia, on 
which previous systematics primarily had been based, allow inclusion of the 
taxon, as a distinct species, in a genus recently redefined in terms of formerly 
neglected somatic anatomy. Parthenogenesis may be facultative in some 
lines. Evolutionary developments now under way in hortensis parallel some 
of the degradations in genital anatomy associated, in D. rubida, with parthe- 
nogenesis and male sterility. Characteristics of an undegraded amphimictic 
morph are predicted. Abnormalities are characterized. To the distribution, 
as previously known, are added Ireland and Greece. 

Disagreements as to systematic status and generic affiliations of 
various species (one example provided by the synonymy below) 
indicate a need for revision of lumbricid classification. Even widely 
spread anthropochores, that usually, though mistakenly, are be- 
lieved to be adequately characterized systematically, need a search- 
ing reinvestigation. 

A single Irish specimen that arrived on St. Patrick's day provided 
a clue to a possible solution of some of the problems hitherto asso- 
ciated with "dendroidea." 

LUMBRICIDAE 

Eisenia Malm, 1877 
Eisenia hortensis (Michaelsen, 1890) 

Allolobophora subrubicunda f. hortensis Michaelsen, 1890, lahrb. Ham- 
burgischen Wiss. Anst., 7: 15. (Type locality, Hamburg. Types, orig- 
inally in the Hamburg Museum.) 



1 From research financed by the National Science Foundation. 
- Zoology Department, University of Maine, Orono, Maine. 



2 BREVIORA No. 300 

Allolohophora (Notogama) veneta f. hortensis Michaclsen. 1900, Abhandl. 

Naturwiss. Ver. Hamburg, 16: 12. 
Helodriliu (Eisenia) venetus \ar. hortensis.-Michaelstn, 1901. Ann. Mus. 

Zool. Acad. Sci. St. Petersbourg, 15: 37. 
Allolohophora {Eisenia) veneta dendroidea Friend. 1909, Gardener's Chron- 
icle. 46: 243 (Type locality, St. James Garden. Malvern, Worcestershire, 

England. T\ pes. in the British Museum.) 
Allolohophora ilu.se/iiii) veneta var. hortensis Michaelscn, 1922. Cap. Zool. 

1 (3): 13. 
Eisenia veneta var. dendroidea Friend. I l >23, British Earthworms, London, 

p. 30. 
Eisenia veneta van dendroidea 4- /■.". v. var. hibernica f. dendroidea Cer- 

aosvitov, 1M42. Proc. Zool. Soc. London. Ill: 240. 241, 274. 
/ isenia veneta \ar. hibernica f. dendroidea 4- E. V. var. hortensis Cernosvitov 

and Evans, 1947, Linnean Soc London, Synopses of the British Fauna. 

No. 6: 25. 
Dendrobaena hortensis (part '.'» 4- D. h. dendroidea Omodeo. 1955, Ann. 

1st. Zool. Univ. N.ipoli. 7: 6. 8. (Excluding />. hibernica.) 
I tenia veneta ^ar. hibernica f. dendroidea dates. 1958, Ann. Mag. Nat. 

Hist., (13) 1: 34. (Name Of the ta\on should have been enclosed by 

quotation marks' ) 
Dendrobaena veneta var. hibernica f. dendroidea 4- D. v. var. hortensis 

Gerald. 1V64. linnean Soc. 1 ondon. Synopses of the British fauna. 

No. 6 (2d ed.): 19 
Bimastos veneta Causey. 1953. Proc. Arkansas Acad. Sci. 6: 47. 

SPECIMENS EXAMINED 

Oregon, Yamhill County. McMinnville, kitchen drain, August 1 1. 
1944, 0-o_f3. Backyard, March 12. 1945, 0-1-1. Backyard, 
brought out bv wash water. October 31, 1945. 0-0-1. Chicken 
yard, under cow manure, November 14. 1945, 0-1-0, Novem- 
ber 23. 1945. 0-4-10. Under wet paper carton on back porch, 
January 10. 1946, several specimens. Backyard, December 
1,1947.0-1-7. D. McKey-Fender. 

California. San Francisco County. Arboretum of Golden Gate 
Park, March 28, 1951, 0-0-2. H. B. Leech per D. McKey- 
Fender. 

Ohio. Trumbull County. Masury, greenhouse, 0-0-2. W. R. Mur- 
chie. 

Virginia. Giles County. Goldbond, sawdust, August 21, 1955, a 
number of specimens. W. A. Harman. 

Italy. Naples, November 23, 1925. 0-0-2. Posillipo, October 7, 
1925, 0-0-7. G. E. Hutchinson per G. E. Pickford. 

India. Darjiling, Northpoint, at ca. 6250 feet, June 1945 0-1-38. 



1968 EISENIA HORTENSIS MICHAELSEN 3 

Interceptions by U.S. Bureau of Plant Quarantine. 

Ireland. From 25 pounds of wood duff with seeds of Crataegus 
oxycanthus in mail arriving at Hoboken, November 1, 1963, 
(?)-l-0. 

Greece. From soil with 25 unidentified herbs in baggage on plane 
arriving at Boston, August 28, 1962, 6-0-0. 

DESCRIPTION 

External characteristics. Size, 26 by IV2 mm (Ireland), 26-32 
by 2-2^ mm (Maine, New York), to 22 by V-A mm (Greece), to 
37 by 2+ mm (Ohio), 42-54 by 2*/2-3 mm (Naples), width 
measured in the clitellar region not always at maximal tumescence, 
thickness of adults near hind end W2 mm. The body is dorso- 
ventrally compressed behind the clitellum and almost oblong in 
transverse section, ventral and lateral sides flat, the dorsum slightly 
convex. The b and d setae are at the four corners of the body. 
Segments, 42, 50, 55, 56, 60 (2), 64 (2), 69 (2), 70 (2), 74, 76, 
77, 81, 82, 83, 84 (2), 85, 86 (2), 88, 90 (3), 92 (3), 94 (3), 
95 (7), 96 (4), 97 (5), 98 (4), 99 (2), 100 (3), 101 (2), 102 
(3), 103 (4), 104 (2), 105, 108, 115 (2), 124, 126, 130. The 
mean number of segments for 69 of the specimens is 90.7536, with 
a standard deviation of 17.4266 and a standard deviation of the 
mean of 2.0979. A large majority of unamputated worms have 
segments in a range of 90-105 which may be more useful systemati- 
cally than the computerized figures. Worms with 1 15-130 segments 
were from Maine greenhouses in which they might almost be said 
to have been force-fed. Color, unrecognizable after alcohol preser- 
vation, otherwise light to bright red, reddish, restricted to dorsum 
and lacking below C except in front of xv and near the hind end 
where the ventrum may also be colored. In some Greek worms, 
color is in discontinuous unpatterned areas. Prostomium, epilobous, 
tongue open (all). 

Setae, present from ii where none usually are lacking though 
sometimes hard to recognize, not closely paired, width of AB and 
CD somewhat variable but CD slightly < or > AB < BC < or > 
A A < DD which is of course < Y2C, the c setae often seeming to 
be about at mL. In some worms, AB is ca. = V2AA and slightly 
< BC. Nephropores, inconspicuous, in xv-xvi at or just above B 
( 39 specimens ) , one pore of xv at D ( 1 ) , one pore of xvi at D ( 1 ) , 
both pores of xiv at D (4), at B (10), one of xiv at B the other at 
D (22), usually unrecognizable in iii-vi or vii. Nephropores, at B on 
left sides of 17th, 19th, 21st, 24th, 26th, 29th, 30th, 33d, 34th, 
and on the right sides of the 17th, 23d, 25th, 27th-29th, 31st-33d 



4 breviora No. 300 

segments (No. 1), on left sides of 14th-20th, right sides of 15th- 
17th. 19th. 20th segments (No. 2). On the leftside of No. 3 the 
following locations were noted, at B, 35th-40th, 52d. 57th. 60th. 
61st, 64th, 65th, at /) in the 34th. 42d-51st. 53d-56th, 58th-59th, 
62d-63d. Pores on the opposite sides of those same segments more 
often than not were in the alternate location. Both pores of vi 
were in CD, one Italian worm on which the right pores of 7th, 
8th. and 1 Ith segments were near B, of 12th at D, but those of 
1 3th- 1 7th in BC. Nephropores. just above B, on left sides of 7th. 
8th. 11th, 14th-16th. lSth-22nd. 24th-32d. on the right side of 
8th, 9th, 12th-15th. 17th-34th (Naples, No. 4). on the left of 
7th- 10th. 12th. 13th. 15th-22d. 24th. 27th, right sides of 7th-8th, 
10th-l 2th. 15th-19th. 22d. 24th. 26th (Naples. No. 5). At pres- 
ent, about the only seemingly feasible characterization is: pores 
usually alternate irregularly and with asymmetry between two ma- 
jor locations along most of the anteroposterior axis. First dorsal 
pore, at 5 6 (22 specimens), at 7 8(2). 

Spermathecal pores, minute, superficial, at 9/10-10/1 1, close to 
mD in an area where coloi often is faint or quite unrecognizable. 
I emale pores, minute, superficial, equatorial in xiv and just lateral 
to B. Male pores, minute, not superficial, within a cleft laterally 
and seemingly about at ////>'( Male tumescences, almost confined 
to median half of BC but obliterating 14 15-15/16 and reaching 
well into w and xvi. 

Clitcllum, saddle-shaped, at maximal tumescence reaching to or 
nearly to B, dorsal pores occluded, intersegmental furrows oblit- 
erated, setae unrecognizable, xxvii-xxxii (15 specimens), xxvii- 
xx.xiii (14). eq xxvii-xxxiii, eq (2). eq 'xxvii-xxxiii (2), xxviii- 
x.xxii (29), xxviii-xxxiii (3), eq xx\ iii-xx.xiv/cq (9). Tubercula 
pubertatis, usually longitudinal bands of greyish transluccnce just 
lateral to B, with straight median margins but with lateral margins 
slightly convex, in xxx-xxxi but occasionally extending slightly into 
\\\ii. rarely reaching eq/xxxii. A deep groove occasionally de- 
marcates a tubcrculum laterally. Even when a red coloration 
(post-preservation artifact) of the clitellum is best developed, none 
is recognizable in the tubercula. 

Genital tumescences, often rather indistinct and especially so in 
relaxed material, include setal couples as follows: a, b of xi ( 1 
specimen), xii (4), xvi (2), xxii (1 ), xxiii (7), xxiv (7), xxv (1 ). 
xxvi (1), xxix (26), xxx (35), xxxi (35), xxxii (9), c,d of x 
( 1 ), xi (22). Condition obviated recording locations on a number 
of worms but dissections indicate that ventral setae of xxx and xxxi 
always were associated with tumescences. Follicles opening 



1968 EISENIA HORTENSIS MICHAELSEN 5 

through tumescences are thickened and have slightly enlarged 
apertures. The b,c and d follicles project conspicuously into the 
coelomic cavities but a follicles obviously are shorter and protrude 
only slightly above the parietes. 

Internal anatomy. Septa, 13/14-14/15 often thickest and quite 
muscular, 10/11 near the gut separated into two lamellae the an- 
terior of which passes straight to the gut, the other inserted on it 
more posteriorly so as to leave a space filled by a watery fluid with- 
out corpuscles or other solid matter. This curious situation pre- 
sumably arises as a result of an ontogenetically late elongation of 
a small section of the esophagus. The anterior lamella seems to 
be at the front end of the calciferous gland. Setal gaps in longi- 
tudinal musculature, obviously eight. Special longitudinal muscle 
band at mD, present from 5/6. Pigment, red, in circular muscle 
layer, lacking immediately under intersegmental furrows. Pigment 
also is visible on or in the coelomic face of the parietes, dorsally 
throughout but also ventrally in front of xv, in flecks, or spots or 
rarely even in larger patches. Location of the brain is variable, a 
section exactly along 3/4 sometimes left the brain anteriorly in iii, 
sometimes anteriorly in iv. Pharyngeal glands, through iv-v to 5/6. 

Esophagus of x has the shape of a truncate cone, narrower an- 
teriorly. The wall is dark (gorged with blood?) and un thickened 
(no sacs or lamellae). The calciferous gland, insofar as could be 
learned from dissection, is between 10/11 and 12/13, sometimes 
not reaching the latter, usually not constricted at 11/12 but of the 
same thickness through both segments. Rarely the portion in xi is 
moniliform. Intestinal origin, in xv (62). Gizzard, superficially 
appearing to extend through xvii-xix. Actually, the muscular layer 
becomes much thinner just behind apparent level of insertion on 
gut of 17/18. Gut often valvular in region of insertion of 19/20. 
Typhlosole, beginning rather abruptly in region of xxi-xxv, thickly 
lamelliform, height variable, when gut is empty sometimes reaching 
or nearly touching intestinal floor, ending abruptly, as shown in 
Table 1. 

The typhlosole does not end abruptly in specimens 1-5, 
8-9, and 33 (cf. Table 1) but becomes much lower or even 
vestigial in the indicated regions, i.e., in lxi-lxv of No. 9. Posterior 
amputation had been involved in each of those cases. The typhlo- 
sole, in a very large majority of the unamputated individuals, ended 
in a region comprising lxxiv-xcv. The typhlosole of the above- 
mentioned specimens, on the contrary, ends in a region comprising 
xxxix-xlv, leaving 1-10 intestinal segments atyphiosolate. Very 



6 BREVIORA No. 300 

probably, the typhlosole had completely disappeared in some seg- 
ments of each oi those worms. In unamputated worms with abrupt 
typhlosolar termination. 10-15 segments were atyphlosolate, 10 
(3), 11 (5), 12 (7). 13 (13), 14(6), 15 (2). [f, as some have 

thought, the atyphlosolate portion of the gut is of proctodeal origin, 
then that part of the gut extends through more segments than the 
embr\ ologistS indicated. 

Extra-esophageal trunk, passing up to dorsal \essel in \ii. but 
often unrecognizable (because empty) behind 10 II. In \ii (one 
specimen) the trunk bifurcated, the larger branch joining the dorsal 
\esse!. the other branch passing \entrall\ and shortly disappearing. 
\ branch from the trunk OCCasionall) passes up to the dorsal trunk 
in \. When that connective is distended with blood ami the pos- 
terior portion o( the trunk is empt\ the cxtra-csophagcal could be 
thought to end in x. I he ventral trunk is complete, bifurcating 
over the subpharyngeal ganglion. The dorsal trunk is also com- 
plete, having been traced forward to the brain when blood is 
present. I he subneural trunk probabl) always is complete but 
USUall) is not traceable throughout because of lack o\ blood. 
Hearts, present onl\ in \u-\i (62 specimens). 

Ncphridia. present from in . presence or absence in ii not de- 
mined), vesiculate. Bladders, sausage-shaped (ellipsoidal), 
transversel) placed, reaching to or nearb to ( or even beyond ( . 
joined lateralis by the tubular portion of the ncphridium. passing 
into p.: it or near B. 

restes, fan-shaped. Male funnels, plicate, size normal with ref- 
erence to that oi the body. Sperm ducts, without epididymis. 
Seminal vesicles, in ix. xi and \ii. Atrial glands, large, dcepb cleft 
at ei| x\. extending through xi\ as well as x\i. 

TP and atrial glands, acinous, supraparietal. CiS glands, acinous, 
each lobe narrow ing to a slender thread-like duct that passes 
straight down in the parietes. associated only with the </./' follicles 
oi xii ( once ). the h follicles of xxx and xxi but usually lacking even 
in those two segments. 

Spermathecae. not sessile, ducts slender, coclomic portions 
shorter than, to longer than, the ampullae. Spermathecae. in ix, x 
( 5d specimens) but the following divergences were recorded; all in 
\ ( twice ). on^: oi the anterior pair in x (once), one of the posterior 
pair in xi ( twice ), in x-xi (once). 

Ovaries, in xiii, each with one egg string which may contain 1-6 
ova. Oviducts, passing into parietes lateral to B. Ovisacs, small, 
lobed. always present. 



1968 EISENIA HORTENSIS MICHAELSEN 7 

The peritoneum frequently is blistered away from the muscula- 
ture. Intervening spaces are occupied by a flocculent white ma- 
terial. The blistering seems not to be so rigidly localized as 
anteriorly in D. octaedra. Areas recorded (perhaps not the only 
ones) from median margin of TP glands to just median to A in 
xxxi-xxxii or xxxiii, the surface more or less deeply cross-hatched, 
in poor preservation not easily distinguishable from the TP glands. 
Blistering, continuous through AA from one side to the other 
(once, Italy), between A and B of each side through xxvi-xxxii 
(Naples), in DD through a region containing the spermathecal 
pores. 

Reproduction. Iridescence on male funnels and in spermathecae, 
of some worms, was so brilliant as to suggest profuse maturation as 
well as massive copulatory transfer. Spermatophores, some of 
which contained sperm, also were noted, in /45/xxviii. As a result, 
until nearly at the end of the present study, laboratory records 
contained the following comment, "Reproduction presumably is 
biparental." Although copulation usually had taken place in worms 
of appropriate age and stage, there does now seem to be some 
reason for suspecting facultative parthenogenesis. 

Number of choromosomes in hortensis is 36 (2n) according to 
Muldal (1952:59) as well as Omodeo (1952:188). The latter 
says spermatogenesis is normal. Neither author mentioned any evi- 
dence indicative of parthenogenesis. Nor was polyploidy detected. 

Breeding, in favorable conditions, may be year round. 

Biology. Feeding appears to be selective, as guts contained only 
humus. Shaking ingesta in water produced no turbidity. 

Male tumescences and tubercula pubertatis are obvious before 
other epidermal modifications are recognizable in the clitellar re- 
gion. 

Many of the worms were posterior amputees. Tail regenerates 
of the usual epimorphic sort never were seen. Regeneration caud- 
ally appears to be restricted (at least usually) to reorganizing 
posterior segments so that external evidence of the amputation no 
longer is recognizable. In such cases, if lysis has not yet proceeded 
too far, the typhlosole sometimes can provide proof of the ampu- 
tation. 

Brown bodies, in No. 36, were present in segments xc-xciii, one 
to three on each side of each segment. Coelomic cavities of xc- 
xcviii in No. 39 were also filled with similar structures, but in this 
worm there was only one brown body on each side in a segment, 
though the coelomic cavities were thereby almost completely filled. 



8 BREVIORA No. 300 

Similar brown matter was not found in coelomic cavities an- 
teriorly. The bodies may then have been formed in situ. 

Distribution. E. hortensis had not been recorded previously 
from Ireland and Greece. 

The single record for Ohio is of a specimen found in a toilet 
bowl of a citv buildine. 

Commenting on the Nearctic distribution of E. hortensis, Mrs. 
D. McKey-Fender wrote {in lift.) "the paucity of North American 
records of this species may be due to the filthy places it inhabits. 
It takes some fortitude to collect it. The Oregon specimens are 
from friable black soil saturated with septic tank effluvium. They 
seem to be between the wettest zone, where E. foetida is even 
more numerous, and the wet. relatively uncontaminatcd soil where 
asea, A. chlorotica and the caliginosa complex predominate." 

Abnormality. Mctameric abnormality, just behind clitellum 
(once), further posteriorly (two specimens); other instances seen 
during counting of segments not recorded. 

No. 1. Right ovary, rudimentary (clitellate adult). 

No. 2. Lett anterior spermatheea. bifid entallv. One ampulla is 
in i\ ami the other is in \ ( Naples). 

No. 3. Left female pore, anterior to eq/xiv but not quite half 
wa\ toward 13 14. Extra pair of male clefts, asso- 
ciated with small male tumescences, in xvi. No pores 
were found in the posterior clefts and no sperm ducts 
were found in their vicinity. Atrial glands of x\i. large, 
each with a deep equatorial cleft 

No. 4. (Ireland). Tubereula pubertatis. markedly protuberant, 
subcircular, two pairs, those of a side completely sepa- 
rated from each other by an uninterrupted intersegmen- 
tal furrow, 30/31. Testes, one pair, in vertical testis 
sacs that reach up to level of dorsal face of gut in xi. 
Seminal vesicles, one pair, medium-sized, not filling 
coelomic cavity of xii. Spcrmathecae. lacking. Irides- 
cence on male funnels, none. The worm obviously is 
male sterile. 

Abnormalities in metamerism are common and their frequency 
indicates the case with which development of normal segmentation 
can be influenced, in a single small region, in a much larger axial 
portion, in several more or less widely separated areas, or, occa- 
sionally, even throughout nearly all of the axis. Divergences from 
normal of specimens 1-3 appear to have resulted from a rarer sort 
of interference with development. So rare, in fact, that exact repeti- 
tion is unknown. 



1968 EISENIA HORTENSIS MICHAELSEN 9 

The male sterility of No. 4, being known at the moment only 
from a single specimen of hortensis, can also be of fortuitous origin. 
However, the aberrations of that worm exactly parallel conditions 
characterizing some male sterile morphs of various other megadrile 
species. In such taxa, where reproduction necessarily is partheno- 
genetic, the associated anatomy is exactly repeated again and again, 
always without spermatogenesis or receipt from another individual 
of copulatory sperm. The male sterility is inherited. 

Knowledge of the evolutionary changes associated with male 
sterility and parthenogenesis in other megadriles enables charac- 
terization not only of a probably amphimictic norm for E. hortensis 
but also of current evolutionary trends therein. 

Polymorphism. The amphimictic norm now postulated for E. 
hortensis is as follows. Male tumescences, large, obliterating 
14/15-15/16 and extending through xiv-xvi. Male cleft deep but 
confined to parietes. Seminal vesicles, three pairs, in ix, xi, xii, all 
functional. Atrial, TP and GS glands, large, supraparietal, acinous. 
The atrial glands reach 13/14 as well as 16/17 and are deeply cleft 
at eq/xv 

The more common evolutionary trends now recognizable in 
sperm-maturing and sperm-exchanging individuals are as follows. 
Reduction in size of male tumescences, confinement to xv with 
furrows 14/15-15/16 no longer obliterated though one or the 
other or both may be slightly displaced, restriction to slight swell- 
ings of immediate margins of male pore clefts. Finally, there will 
be no tumescence at all and clefts will have disappeared leaving 
the male pores superficial as in certain species of Lumbricus. 
Correlated with that trend is decrease in size of the associated 
atrial glands, limitation to xv, representation only by a few fila- 
ments slightly protuberant into the coelom from gaps between 
bands of longitudinal musculature. Slight protuberances of body 
wall, without interruptions of the muscular layer, over sites of 
male pores, may indicate presence of last rudiments of the atrial 
glands. Similar trends involve the TP and GS glands, one or the 
other or both of which have disappeared in certain lines. 

Seminal vesicles of ix and xi, but never of xii, seem to be dis- 
appearing, more especially in ix. There the vesicles no longer be- 
come as large as in xi, and may retain a juvenile texture through 
maturity or may even be unrecognizable on one side or the other. 

Inclusion of testes in paired, vertical testis sacs is known today, 
in the Lumbricidae, only in association with male sterility. The 
metandric reduction (deletion of tests and gonoducts of x), on 
the contrary, has been acquired in various families by amphimictic 



10 BREVIORA NO. 300 

taxa as well as by male sterile morphs. Associated therewith is loss 
of all vesicles in ix and xi. Many male sterile morphs. in various 
families, are atheeal but spermatheeae have been lost by individuals 
that still produce mature sperm. 

Evolutionary trends in hortensis may parallel, in part, those of 
D. rnhida. That species has male fertile morphs (usually called 
subspecies, variety or f. subrubicundd) as well as male sterile 
morphs with obligatory parthenogenesis. Degradation of genital 
structure may have progressed further in hortensis than in ruhida. 
F.ven more drastically modified morphs than now known are antici- 
pated for hortensis, especially if its parthenogenesis was acquired 
earlier. 

Disappearance of atrial and TP glands results in a state that 
parallels conditions characterizing the genus Lumbricus. 

Remarks. Relaxed material furnished by Mrs. McKe\ -Pender 
provided beautiful demonstrations of nephropore locations and of 
uncollapsed Dephridial bladders. Such information often is unpro- 
curable from contracted specimens. Nevertheless, on her worms, 
genital tumescences were unrecognizable and clitellar boundaries 
probabl) could have been determined accurately, if at all, only 
from microtome sections. 

Generic relationships. The following characters were found to 
be invariant in all samples examined. Calciferous gland, without 
sacs, opening directly into esophageal lumen behind insertion of 
10,1 1 through a circumferential circle of small pores. Intestinal 
origin, in x\. Gizzard, mostly in \\ii. Hearts, m vii-xi. Pxcrctory 
organs, holoic. Nephridial bladders, sausage-shaped (ellipsoidal), 
transversely placed. Nephropores, inconspicuous, alternating ir- 
regularly and with asymmetry between a level just above H and 
another above /). Setae, paired. Longitudinal musculature, pin- 
nate (as seen in transverse section). Pigment, red. Ovaries, in 
xiii. each terminating distallv in a single egg string. Ovisacs, 
present in xiv. Some of these characters are now expected to be 
definitive at the family level. Others require inclusion of Michael- 
sen's hortensis in the genus Eisenia (of. Gates, in press). 

REFERENCES CITED 

Gates, G. E. 

(in press) On two American genera of the earthworm f;miilv I umbricidae. 
Jour. Nat. Hist. 



1968 EISENIA HORTENSIS MICHAELSEN 11 

MULDAL, S. 

1952. The chromosomes of the earthworms. I. The evolution of 
polyploidy. Heredity, 6: 55-76. 
Omodeo, Pietro 

1952. Cariologia die Lumbricidae. Caryologia 4: 173-275. 

(Received 16 January 1968.) 

TABLE 1 
Typhlosole termination and segment number in Eisenia hortensis 



1. 36-39 3 42 

2. 46-47 3 50 

3. 47-48 7 55 

4. 52-54 10 64 

5. 57-59 1 60 

6. 58 6 64 

7. 59 10 69 

8. 61-62 8 70 

9. 61-65 9 74 

10. 62 7 69 

11. 64 13 77 Greece, juvenile 

12. 65 5 60 Posterior amputee 

13. 68 8 76 Greece, juvenile 

14. 71 10 81 Posterior amputee? Greece, juvenile 

15. 72 11 83 

16. 74 8 82 

17. 76 9 85 Posterior amputee 

18. 77 11 86 

19. 78 6 84 

20. 78 10 88 

21-22. 78 12 90 Ohio (1) 

23. 80 14 94 

24. 80 15 95 

25. 81 13 94 Greece, juvenile 

26. 82 14 96 

27. 83 12 95 

28. 83 13 96 

29. 83 14 97 



12 






BREVIORA 


No. 300 


30-31. 


84 


11 


95 


San Francisco ( 1 ) 




32. 


84 


12 


96 


Ohio 




33. 


80-84 


8 


92 






34-35. 


85 


13 


98 






36. 


86 


8 


94 






37-38. 


86 


12 


98 


Ohio ( 1 ) 




39.40 


86 


13 


99 


Maine, Italy 




41. 


87 


12 


99 






42. 


88 


13 


101 


Great Britain 




43. 


88 


14 


102 






44. 


89 


11 


100 






45. 


89 


13 


102 






46-47. 


89 


14 


103 






48. 


89 


15 


104 






49-50. 


90 


13 


103 


Ohio ( 1 ) 




51. 


91 


13 


104 






52. 


92 


10 


102 


Posterior amputee? 




53. 


92 


13 


[05 






54. 


95 


13 


108 


Naples 





BREVIORA 



Museum of Comparative Zoology 

CAMBRroGE, Mass. 11 October, 1968 Number 301 

NEW ECHIMYID RODENTS FROM THE OLIGOCENE OF 
PATAGONIA, AND A SYNOPSIS OF THE FAMILY 

Bryan Patterson and Rosendo Pascual 1 

Abstract. The living and extinct echimyid rodents are briefly reviewed, 
and placed in five subfamilies: Echimyinae Murray, Dactylomyinae Tate, 
Heteropsomyinae Anthony, Myocastorinae Ameghino, and Adelphomyinae 
nov. Prospaniomys and Spaniomys are placed in the Myocastorinae. Two 
new genera and species, Xylechimys obliquus, Deseadan, and Paradelphomys 
fissus, Colhuehuapian, are described; both are referred to the Adelpho- 
myinae. 

A SYNOPSIS OF THE ECHIMYIDAE 

Echimyids today form the largest of the numerous families com- 
prising the suborder Caviomorpha. Fifteen living genera are cur- 
rently recognized, all but one wholly tropical or subtropical in 
distribution. Extinct genera, including those described below, 
number only eighteen and of these four are Antillean. We are 
hence far from a really adequate understanding of the phylogeny 
of the group. The living forms have usually been divided into two 
subfamilies, Dactylomyinae and Echimyinae (e.g. Ellerman, 1940; 2 



1 Museo de la Plata, La Plata, Argentina. 

2 Ellerman, while recognizing these subfamilies, also included in his Echi- 
myidae, with the rank of subfamily, the Octodontidae (with Ctenomys), 
Abrocomidae, Capromyidae, Plagiodontia, Myocastor and even the African 
Thryonomyidae and Petromuridae. As will appear below, we agree as to 
recognition and inclusion of the Myocastorinae. Further work may require 
that the brachyodont to hypsodont Echimyidae and the hypselodont Capro- 
myidae be merged; in that case the subfamilies of each would become 
tribes. The octodontids surely had a common origin with the echimyids in 
pre-Deseadan time, but the two groups diverged thereafter and merit fa- 
milial rank. Abrocomids and ctenomyids are derivatives of the Octodon- 
tidae, the Ctenomys group being only a subfamily of it. The African 
families have nothing to do with this assemblage, being descended from the 
Fayum Phiomyidae (Lavocat, 1963; Wood, in press). 



2 BRKVIORA NO. 301 

Cabrera, 1960), with the extinct forms placed in the latter (e.g. 
Simpson, 1945 ). There can be little doubt that Dactylomys and its 
relatives constitute a single suprageneric group, but the same can 
hardly be claimed for the Echimyinae, particularly il the extinct 
forms are taken into account. Wood and Patterson ( 1959:301-2), 
on the basis of work in manuscript by Patterson and Kraglicvich. 
divided the familv into five groups: the Proechimys-Eumysops, the 
Dactylomys, the Echimys, the Spaniomys and the Adelphomys. 
1 he Echimys and Dactylomys groups are so far unrepresented, or 
at least unrecognized, in the fossil record. The largest group and 
the one with the most continuous record is the ProechUnys- 
Eumysops, which can be traced back to the C'olhuehuapian Oli- 
gocene Protacaremys; it includes almost half o\ the known genera. 
I he Echimys group, the Echimyinae proper, comprises only three 
genera. Hllerman (1940:106-108) pointed out the distinctions 
between these three and the rest o^ his Echimyinae but did not 
separate them formally. The extinct West Indian Heteropsomys, 
Homopsomys, Brotomys and Boromys appear to be closer to the 
Proechimys-Eumysops group than to am other division oi the 
famil) (Miller. 1930; Williams and Koopman, 1951), and Krag- 
lievich (1957, 1965) has extended Anthony's subfamily Heterop- 
sonninac 1 (1917) to include the whole assemblage. As regards 
two other extinct Antillean genera, fsolobodon and Aphaetreus, 
that are often placed in the Echimyidae, we are inclined for the 
present to follow Miller and Kellogg ( 1955:644-5) and associate 
them with Plagiodontia as a sublamil;. oi the C'apromyidae. 

Wood and Patterson placed Spaniomys and Prosponiomys in a 
separate group on the basis oi cheek tooth structure, but did not 
comment as to possible relationships to later forms. We now oiler 
the suggestion that Myocastor may be a Burvivor of this group. 
This rather specialized genus does not fit comfortably into the 
Capromvidae — otherwise a purely Antillean group so far as 
known" — with which it is frequently associated. It also stands 
somewhat apart from the later Echimyidae in such characters as 
the very long paroccipital process, the prominent lateral process of 
the supraoccipital. the greatl) reduced coronoid process, the pres- 
ence of four fully developed lophids in all lower molars and five 

1 Proposed by Anthonv on the supposition that he was defining a sub- 
family of the Dasyproctidae. This is the oldest available name, antedating 
Fumvsopidue Rusconi 1935 (Eum\sopinac Kraglievicb 1945). 

1 The Venezuelan record of "Procapromys" was surely based on erroneous 
locality data. 



1968 OLIGOCENE ECHIMYID RODENTS 3 

lophids and a small anterior fossettid in dm 4 . A tendency toward 
reduction and loss of the metalophid in the molars is characteristic 
of the family. Dactylomyines, echimyines and most later members 
of the Adelphomys group (see below) lack it, while heterop- 
somyines may retain a vestige of it, or, alternatively, acquire a 
neolophid, on anterior lower molars, in the form of a small crest, 
connecting anterolophid and hypolophid, that isolates a small 
anterointernal fossettid. The members of the Spaniomys group do 
not display this tendency to the same degree. In the Colhue- 
huapian Prospaniomys the metalophid is present in Mi. 2 and in- 
terrupted or absent in M :i ; in the Santacruzian Spaniomys it is 
present in M t . 2 and absent in M 3 . The cheek teeth of Spaniomys 
are quite variable, however, and two specimens in the Ameghino 
Collection display a tendency toward reacquisition of a fourth 
lophid in M :! . In one of these there is a pronounced spur project- 
ing posteriorly from near the center of the hypolophid, and in the 
other (the type of Graphimys provectus Ameghino, a synonym of 
Spaniomys modestus Ameghino) this spur is larger, extending to 
the posterointernal corner of the tooth. The end result of this, if 
continued, would have been an M 3 with four lophids, the third of 
which, while resembling the third (the metalophid) of M^o, would 
have been a neomorph. 

Unworn and, especially, little worn upper and lower molars of 
Myocastor (there is much change in crown proportions and some 
in structure as wear proceeds) resemble the corresponding teeth of 
Prospaniomys and Spaniomys more closely than they do those of 
other echimyids. The fourth crest of M 3 could have been inherited 
either from ancestors like Prospaniomys, in which the crest was 
still present in certain individuals, even if somewhat reduced, or 
from ancestors resembling Spaniomys, in which the tendency to- 
ward a neomorph crest had become fixed. Dm 4 of Myocastor is 
fully molariform as is that of Prospaniomys; Spaniomys differs in 
having only three lophs. Dm 4 of Myocastor is a complex tooth 
with five lophids, the anterior two united lingually to isolate the 
small anterior fossettid. Spaniomys has four lophids, the anterior 
pair uniting lingually with wear to isolate one or two shallow fos- 
settids; in Prospaniomys the two anterior lophids are united at or 
shortly following eruption. These crests correspond topograph- 
ically to the anterolophid and metalophid of the molars. The small 
extra lophid of Myocastor (the second in the series) is an addition 
to the crown. Acquisition of this crest by the ancestors of the 
living form seems likely. In one species of the late Pliocene 
Isomyopotamus the rather large anterior fossettid is partially 



BRIVIORA 



No. 301 



divided bv a spur directed internally from the united anterolophid 
and metalophid (Reig, 1950:1 13, fig. 2c); this suggests a method 
of origin of the extra lophid. Myoca.stor is specialized as regards 
the progressive enlargement of the check teeth from dm j back- 
ward and the progressive obliquity of the lophids in the lower 
series. In the late Pliocene Isomyopotamus and Paramyocastor 
(Ameghino, 1904:252) these tendencies are less marked. 

An imperfect skull of Spaniomys is known (Scott. 1905:410- 
411). This displays certain resemblances to Wyocastor, The ros- 
trum is relatively long and its ventral border somewhat elevated 
above the bony palate, the tip of the upper incisor is approximately 
at the level of the cheek teeth, the paroccipital processes are longer 
and the lateral processes are freely projecting (fide Scott'). The 
coronoid process of the mandible is not reduced, but sueh reduc- 
tion may have been a late event in the group. Ameghino describes 
the coronoid o\ Paramyocastor as being "mucho mas larga que en 




Fig. 1. Spaniomys modestm Ameghino. x 4. 
15577. Dorsal view of incomplete left mandible. 



Princeton University No. 



'The specimen (American Museum No. 9529) does not now show these 
last two features clearly; possibly it has suffered some damage since 1905. 



1968 OLIGOCENE ECHIMYID RODENTS 5 

Myocastor." Myocastor has a very large, wide masseteric shelf 
that begins beneath M] and runs back to the angle. Spaniomys 
possesses a smaller but nevertheless rather comparable structure so 
far as preserved (Fig. 1 ) . Both forms have a muscle scar running 
from the coronoid process to the front of the shelf; and in both the 
grinding surface of the upper cheek teeth faces somewhat out- 
wardly, that of the lowers inwardly. 

These resemblances in skull and dentition are collectively rather 
impressive. We believe a relationship to be possible and accord- 
ingly for the present include the Spaniomys group in the Myo- 
castorinae. The test of this hypothesis will come with the dis- 
covery of late Miocene and earlier Pliocene forms ancestral to the 
later Pliocene myocastorines Isomyopotamus and Paramyocastor. 

The Adelphomys group remains for consideration. It was the 
earliest of the several subdivisions to appear in the record and is 
the only one that, so far as can be judged on present evidence, has 
no surviving representatives. Deseadomys (Wood and Patterson, 
1959:303-312) and Xylechimys gen. nov. of the early Oligocene 
are higher crowned than the late Oligocene Protacaremys and 
Prospaniomys, and are more advanced as regards obliquity of lophs 
and lophids and (in Deseadomys) reduction or loss of the meta- 
lophid in the lower molars. Deseadomys and Xylechimys differ 
from all later echimyids in the replacement of dm^ by P*. As 
pointed out by Wood and Patterson (1959:301), retention of the 
milk molars in the family presumably came about between Desea- 
dan and Colhuehuapian time. There appears to have been an early 
dichotomy into the Adelphomys group, on one hand, and the an- 
cestors of the rest of the family, on the other (Wood and Patterson, 
1959, fig. 3), and it follows from this that retention of dm| has 
occurred more than once in echimyid history. 1 The group is 
distinct from the time of its earliest appearance, survived into the 
early Pliocene, and includes a third of the extinct genera. It is 
worthy of subfamilial rank and may be defined as follows: 

ADELPHOMYINAE subfam. nov. Cheek teeth sub-mesodont 
to mesodont, no unilateral hypsodonty; lophs and lophids tending 
to become markedly oblique, sometimes lamellar; upper molars 
with metaloph and posteroloph uniting early to convert metaflexus 

1 The selective advantage of retention of dm* , especially in groups that 
retained brachyodont teeth throughout their subsequent history, is not too 
clear. A possible explanation might be that this afforded a simpler, more 
rapid means of attaining crown complexity than progressive molarization 
of P^, dm* being of course more complex than the replacing tooth. 



6 BREVIORA NO. 301 

to metafossette. other flexi persistent; lower molars with marked 
tendenc) toward reduction and loss of metalophid 

Distribution: Earl) Oligocene to early Pliocene, South America. 

Genera: Adelphomys, Stichomys, Santacruzean; Paradelphomys 
gen. qov., Colhuehuapian; Xylechimys gen. nov., Deseadomys, 
Deseadan. There is. in addition, an undescribed genus in the 
Chasicoan I Pascual, MS). 

1 he distinctive characters oi the cheek teeth oi the other sub- 
families ma\ be summarized, with a listing o( the included genera: 

/ himyinae Murray ISM->.' Mesodont, relatively long; lophs 
and lophids rather stout, tending to form transverse lamellae; flexi 
and flexids (or transverse valleys) fairly persistent. Echimys, 
Diplomys, Central and South America, hothrix, South America. 
Recent. 

Dactylomyinae late 1 *-> 3 5 . Brachyodont, wide, large relative to 

e oi skull: lophs and lophids rather thin, sharp and with ten- 
dencies toward acquisition k)\ lamellae; flexi and flexids persistent. 
Dactylomys, Thrinacodus, Kannabateomys, South America. Recent. 

Heteropsomyinae Anthom 1917. Brachyodont to hypsodont, 
with some unilateral hypsodonty; upper cheek teeth tending to 
become subcircular to quadrate in outline; flexi and flexids not 
\er\ peis, stent, converting to Eossettes and fossettids after moderate 
wear; small antcroinicrnal fossettid may be present in lower 
molars. Proechimys, Central and South America. Recent. Puerto 
Rico, uncertain age; Hoplomys, Central and South America, Re- 
cent; Euryzygomatomys, Clyomys, Mesomys, Lonchothrix, South 
America. Recent; Carterodon, South America. Pleistocene to Re- 
cent; Cercomys, South America. Pliocene to Recent; Heterop- 
somys, Homopsomys, Puerto Rico, latest Pleistocene or early Re- 
cent; Brotomys, Hispaniola, latest Pleistocene or early Recent; 
Boromys, Cuba, iatest Pleistocene or early Recent; Eumysops, 
South America, late Pliocene to Pleistocene; Protacaremys, South 
America, late Oligocene. Protadelphomys, South America, late 
Oiigocene. is tentatively placed here, and there is an undescribed 
genus in the Santa Cruz Miocene of Patagonia. 

Myocastorinae Ameghino 1904.- Brachyodont to hypsodont; 
labial flexi and lingual flexids of later forms converting to fossettes 
and fossettids after considerable wear; later forms showing an in- 
crease in size and in obliquity of lophs and lophids from front to 

'As Echinnina ( = Echymyna Gray 1825, Echsmidae Bonaparte 1X45). 
-'Proposed as a family ( = Myopotamini Tullhcrg 1900. Myiopotamyina 
Bonaparte 1850). 



1968 OLIGOCENE ECHIMYID RODENTS 7 

back; drn 4 with a small anterior fossettid, with five lophids in later 
forms; M 3 usually with four lophids, metalophid either retained or 
a third lophid reacquired. Myocastor, South America, Pliocene?, 
Pleistocene to Recent (and currently doing only too well in the 
feral state in Europe and North America); Paramyocastor, Iso- 
myopotamus, South America, late Pliocene; Spaniomys, South 
America, early Miocene; Prospaniomys, South America, late Oli- 
gocene. Tramyocastor does not appear to be separable from 
Myocastor. 

The new genera to be described, one of them Deseadan, the other 
Colhuehuapian, are both adelphomyines and appear to be more 
closely related to each other than to any of the previously known 
forms. 

DESCRIPTION OF NEW FORMS 
Xylechimys 1 gen. nov. 

Type species: X. obliquus 1 sp. nov. 

Distribution: Deseadan, Patagonia. 

Diagnosis: Reduced metalophid present on all molars, joined to 
metaconid to isolate antero-fossettid but continuing on into meso- 
flexid to define small anteroflexid, metalophid smallest in M 3 , sub- 
equal in Mi_ 2 ; hypolophid and posterolophid very oblique, espe- 
cially on M^; posterolophid terminating lingually near median 
line of tooth row; principal flexids deep, all of nearly equal depth 
on M 2 _3, metaflexid shallower on Mi; metaconid sharply angulate 
posteriorly. 

Xylechimys obliquus sp. nov. 

Type: Museo de La Plata No. 59. II. 26. 81, portion of right 
ramus with Mi_ 3 , base of I and alveoli of P 4 . 

Hypodigm: Type only. 

Horizon and locality: Deseadan Oligocene; Laguna de los 
Machos, Campo Belisario Escobar, Departamento Paso de los 
Indios, Chubut, Argentina. 2 

Diagnosis: As for the genus. 



1 £v\ot, wood, plus Echimys. Named for Professor Albert E. Wood in rec- 
ognition of his contributions to our knowledge of the order. The specific 
name refers to the obliquity of the posterior lophids. 

2 This is a new locality that has yielded a small faunule. Found with 
Xylechimys were specimens of Asteromys, Archaeohyrax, Plagiarthrus, Pro- 
pachyrucos, Prohegetotherium, Trachytheriinae indet., which are sufficient 
to reveal the age. 



8 



BREVIORA 



No. 301 



Description: The anterior cheek tooth is unfortunately repre- 
sented only by its broken alveoli. X-rays reveal no trace of a tooth 
below these alveoli. It is virtually certain that the missing tooth 
was P 4 , as in Deseadomys. 

The molars are mesodont and exhibit no tendency toward uni- 
lateral hypsodonty. M L . is the largest, and M : , the smallest of the 
scries, but the size differences are not great. All are broader rela- 
tive to length than in Deseadomys and are wider anteriorly than 
posteriorly, most aotabl) as regards M :t . The anterior portions of 
the molars are short anteroposterior^. 

The anterolophids are Dearly transverse. A reduced metalophid 
is present on all teeth, that of M :: being the shortest, those of Mi. 2 
sub-equal. Near the middle of its course each metalophid is united 
to the metaconid to isolate a small anterofossettid, and then con- 
tinues linguallv beyond this junction for a short distance. A short 





B 



I ig. 2. Xylechimys obliquus gen. et sp. nov. x 10. Type. Museo de La 
Plata No. 59. II. 26. 81. A, RM, ,, crown view; B, RM, . lingual view. 



1968 OLIGOCENE ECHIMYID RODENTS 9 

and shallow anteroflexid is thus present in addition to the antero- 
fossettid, a marked contrast to Deseadomys arambourgi. The 
hypolophid is decidedly oblique, being almost in a direct line with 
the ectolophid. On Mi. 2 , but not on M 3 , this crest is slightly 
swollen lingually, the swelling marking the position of the ento- 
conid. The posterolophid is essentially parallel to the hypolophid 
and terminates posteriorly a short distance lingual to the mid-line 
of each tooth. The metaconid is sharply angulate posteriorly. 
Hypoflexid and mesoflexid are of equal depth, extending nearly to 
the base of the enamel. The metaflexid progressively deepens 
posteriorly through the series until it is as deep as the hypoflexid 
and mesoflexid on M 3 . The walls of these three valleys are steep. 
In Mo. 3 , especially in the latter, these flexids deepen toward the 
mid-line. The anterofossettid is shallower than the principal flexids, 
and the anteroflexid is little more than a conspicuous groove. The 
mesoflexid and anteroflexid together constitute the largest valley in 
Mi.o. On M 3 the connection between the posterolophid and the 
ectolophid had not yet been fully established by wear, which sug- 
gests that in the unworn state a gap existed between these lophids. 
On M 3 a slight notch in the anterior wall of the anterofossettid had 
not yet been obliterated by wear. 

The base of the incisor is poorly preserved. All that can be said 
concerning it is that the enamel extends about half way around the 
lateral face. 

The mandibular ramus, so far as preserved, is robust and deep, 
about as in Deseadomys. As in that form, the ascending ramus 
arises near the front of M 3 . The masseteric crest begins, as a 
small shelf, at a point beneath the anterior portion of M x . 

Paradelphomys 1 gen. nov. 

Type species: Paradelphomys fissus 1 sp. nov. 

Distribution: Colhuehuapian, Oligocene, Patagonia. 

Diagnosis: Lophids very oblique, posterolophid forming an iso- 
lated lamella; metalophid vestigial on dm 4 , absent on molars; 
anterofossettid of dm t elongate. 

Paradelphomys fissus sp. nov. 

Type: Museo municipal de Mar del Plata No. 125, fragment of 
right mandible with dm 4 — M x . 



1 7rapa, beside, plus Adelphomys. The specific name alludes to the isola- 
tion of the posterolophid. 



10 



BREVIORA 



No. 301 



Hypodigm: Type only. 

Horizon and locality: Colhue-Huapi Formation; barranca south 
of Lake Colhue-Huapi, Chubut. Argentina. 

Diagnosis: As for the genus. 

Description: Dm, is nearly oval in outline and longer than M,. 
The anterolophid is gently convex anteriorly and very obliquely 
directed posterolinguallv. Near its extremity it is connected to the 
hypolophid by a short bridge, a vestige o<i the metalophid, which 
isolates an elongate, sinuous anterofossettid. The mesoflexid. which 
is in consequence short, extends down nearly to the base of the 
crown. Protoconid. ectolophid and hypolophid are in a nearly 
straight line. The posterolophid is an isolated, somewhat slipper- 
shaped lamella that preserves no vestige of a former connection to 





B 

Fig. 3. Paradelphomys fissus gen. et sp. nov. x 10. Type. Museo mu- 
nicipal de Mar del Plata No. 125. A, Rdm, — M,, crown view; B, Rdm, — 
Mi, lingual view. 



1968 OLIGOCENE ECHIMYID RODENTS 11 

the hypolophid-ectolophid. The combined hypoflexid-metaflexid 
is a narrow, deep, parallel-sided cleft extending diagonally across 
the tooth. Not as deep as the mesoflexid, it is somewhat deeper on 
the lingual than on the labial side; at an extreme stage of wear the 
bases of the protoconid and hypoconid would be joined. 

Mx is square in outline anteriorly and tapers posterolingually at 
the crown surface, although not at the base. The anterolophid is 
nearly transverse anteriorly and curves posteriorly in the region of 
the metaconid. The mesoflexid is a diagonal slit that extends al- 
most to the anterior extremity of the tooth and reaches downward 
nearly to the base of the enamel. The hypolophid-ectolophid is 
slightly more sinuous than in dm 4 . The posterolophid is also an 
isolated lamella, but, unlike that of dm 4 , it is decidedly convex 
anteriorly, the convexity marking the site of an earlier junction 
with the hypolophid. Posteriorly, its wall slopes outward and 
downward in a postero-labial direction so that its area would in- 
crease considerably with wear. The hypoflexid-metaflexid is deeper 
at the center than at the extremities; at an extreme stage of wear 
an elongate, narrow fossettid would be isolated. 

Enough of the ramus is preserved to reveal that the masseteric 
crest began beneath the anterior end of Mi. 

TABLE 1 

Tooth measurements (in mm) of type specimens of 
Xylechimys obliquus and Paradelphomys fissus 







X. obliquus 


P. fissus 


drru. 


antero-posterior 


— 


4.0 




width, anterior 


— 


2.1 




width, posterior 


— 


2.7 


M,. 


anteiio-posterior 


3.1 


3.7 




width, anterior 


2.5 


2.9 




width, posterior 


2.5 


2.7 


M,. 


antero-posterior 


3.2 


— 




width, anterior 


2.8 


— 




width, posterior 


2.8 


— 


M 3 , 


antero-posterior 


2.9 


— 




width, anterior 


2.8 


— 




width, posterior 


2.3 


— 



12 BREVIORA No. 301 

DISCUSSION 

Xylechimys invites comparison with its contemporary, Desea- 
domys. The former is more primitive as regards retention of the 
metalophid and anterofossettid on all molars and in the lingual 
extension o( this lophid beyond the connection with the metaconid. 
It is more advanced in the greater obliquity of the hvpolophid and 
posterolophid and in the nearly median position of the lingual end 
of the posterolophid. Wood and Patterson ( 1959:306, 309, 312) 
called attention to various characters of Deseadomys arambourgi 
and of D. loomisi that suggested close relationship. If not actual 
ancestry, to the Santa-Cruzean Stichomys and Adelphomys, re- 
spectively. This view still seems reasonable. Xylechimys does not 
resemble these later forms to the same extent. 

Paradelphomys is decidedly specialized in the loss oi the meta- 
lophid in M,. the very marked obliquity o\ the lophids and the 
complete separation of the posterolophid. It is. in fact, in these 
features the most specialized member of the Adelphomyinae so far 
known. There are. ho\\e\er. resemblances between it and Xyle- 
chimys, such as a nearly comparable degree of obliquit) o\ the 
hvpolophid and posterolophid. the lingual end of the posterolophid 
not extending much be\ond the mid-line of the tooth until after 
heaw wear and the tendene\ toward separation of the postero- 
lophid present in Xylechimys. Elimination o\ the metalophid. 
straightening o\ the protoconid-cctolophid-hvpolophid crest, and 
further separation oi the posterolophid would convert molars of 
Xylechimys structure into those oi Paradelphomys. It is, of course, 
quite uncertain if the two stood in ancestor-descendant relationship 

— the more exact resemblance between the species of Deseadomys 
and Stichomys and Adelphomys would argue against this possibility 

— but on present evidence it seems likely that they were more 
closely related to each other than to any of the genera previously 
known. 

Its name notwithstanding. Protadclphomys Ameghino of the 
Colhuehuapian is neither ancestral to Adelphomys nor a member 
of the Adelphomyinae. Ameghino ( 1902: 112) based his concept 
of relationship on the lower incisors, which are relatively large, 
flat anteriorly, and have the transverse (mesio-distal) and antero- 
posterior diameters nearly equal in both. This is the only close 
similarity between the two forms and it is more than offset by the 
structure of the molars, which in Protadelphomys are short and 
quadrate rather than elongate, retain reduced metalophids on M,. : . 
have the hypolophid directed lingually and the posterolophid 
deeply separated from the hypolophid. The last character is a 



1968 OLIGOCENE ECHIMYID RODENTS 13 

point of resemblance to Paradelphomys but it is the only one and, 
in view of the tendency toward lamellar structure that crops up in 
several of the echimyid subfamilies, we do not consider it to be in- 
dicative of any close affinity. Protadelphomys is known only from 
one incomplete ramus with broken or deeply worn teeth. What can 
be seen of the structure suggests that the genus represents an early 
and specialized offshoot of the Heteropsomyinae, and more than 
this cannot at present be said. 

ACKNOWLEDGMENTS 

Each of us is indebted to The John Simon Guggenheim Memo- 
rial Foundation for the opportunity of working in the country of 
the other. For the privilege of studying and describing specimens 
in their care we are indebted to the authorities of the Museo 
Argentino de Ciencias Naturales; Sr. Galileo J. Scaglia, Director 
of the Museo Municipal de Ciencias Naturales y Tradicionales de 
Mar del Plata; Professor Glenn L. Jepsen, Princeton University; 
and Dr. Richard H. Tedford, The American Museum of Natural 
History. The drawings are the work of Mr. Arnold D. Clapman. 
The X-rays were taken by Mr. Joseph J. Murphy. 

REFERENCES 

Ameghino, F. 

1902. Premiere contribution a la connaisance de la faune mamma- 
logique des couches a Colpodon. Bol. Acad. Nac. Cien. 
Cordoba, 17:71-138. 
1904. Nuevas especies de mamiferos cretaceos y terciarios de la 
Republica argentina. An. Soc. Cien. Arg., 58:225-290. 
Anthony, H. E. 

1917. New fossil rodents from Puerto Rico, with additional notes of 
Elasmodontomys obliquus Anthony and Heteropsomys insulans 
Anthony. Bull. Amer. Mus. Nat. Hist., 37:183-189. 
Cabrera, A. 

1960. Catalago de los mamiferos de America del Sur. Rev. Mus. Arg. 
Cienc. Nat., Cienc. Zool., 4:309-732. 
Ellerman, J. R. 

1940. The families and genera of living rodents. Volume I. Rodents 
other than Muridae. London: British Museum (Natural His- 
tory) : i-xxvi, 1-689. 
Kraglievich, J. L. 

1957. Revision de los roedores extinguidos del genero "Eumysops" 
Ameghino 1888. (Nota preliminar.) Ameghiniana, Rev. Asoc. 
Paleont. Arg., 1:37-39. 



14 BREVIORA NO. 301 

1965. Speciation phyletique dans les rongeurs fossiles du genre Eumy- 
sops Amegh. (Echimyidae, Heteropsomvinae). Mammalia, 29: 
258-267. 
Lavocat, R 

1963. Reflexions sur I'origine el la Structure du groupe des rongeurs. 
Problemes actuels de la Palcontologie, Colloques Internat.. 
Centre Nat. Recherches Sci., No. 104:287-299. 
Miller, G. S. 

1^30. Three small collections of mammals from Hispaniola. Smith- 
son. Miscell. Coll.. 82(15): 1-10. 

and J. W. Gidley. 

1918. Synopsis of the supergenerie groups of rodents. Jour. Washing- 
ton Acad. Sci., 8:431-448. 
and R. Kellogg. 



1955. List of North American Recent mammals Bull. U.S. Nat. Mus., 
205:i-\ii, 1-454. 
RJBIG.0 V 

[950. Sohre nuevos restos de roedores del genero Isomyopotamm 
Rov. An. SOC. Cien. Arg.. 119:108-118. 

Scott, w. b. 

1905. Paleontology. Part 111. dines Rcpts Princeton Univ. Expeds. 
Patagonia, 5:384-487. 
Simpson, G. G. 

1945. The principles ol classification and a classification of mammals. 
Bull. Amu Mus. Nat Hist, 85:1-350 
Williams, E. E. and K. F. Koopman 

1951. A ncu t\)ssil rodent from Puerto Rico. Amer. Mus. Novit. No. 

1515:1-9. 
Wood. A. E. 

In press. Early Cenozoic mammalian faunas, Fayum Province, 

I jypt United Arab Republic. Part II. The African Oligocene, 

Rodentia. Bull. Yale Peabody Mus 

and B. Patterson 

1959. The rodents of the Deseadan Oligocene of Patagonia and the 

beginnings of South American rodent evolution. Bull. Mus. 

Comp. Zool., 120:281-42X 

( Received 26 February 1968.) 



BREVIORA 



Museum of Comparative Zoology 



Cambridge, Mass. 11 October, 1968 Number 302 

GEOMYERSIA GLABRA, A NEW GENUS AND SPECIES OF SCINCID 
LIZARD FROM BOUGAINVILLE, SOLOMON ISLANDS, WITH COM- 
MENTS ON THE RELATIONSHIPS OF SOME LYGOSOMINE GENERA 

Allen E. Greer and Fred Parker 1 

Abstract. A new genus and species of skink, Geomyersia glabra, is de- 
scribed from Bougainville, Solomon Islands. Notes on the reproduction and 
ecology of the species are presented along with a generic diagnosis and de- 
scription. 

The large group of skinks centering on Leiolopisma (as used in its broad- 
est context ) is divided into two major subgroups on the basis of the relation- 
ships of the bones in the secondary palate; the alphas are wide ranging 
throughout the Old and New World whereas the betas, which appear to be 
derived from the alphas, are confined to subsaharan Africa and the Aus- 
tralian Region. Geomyersia glabra is a beta skink. The relationships be- 
tween the African and Australian Region betas are obscure as yet, but the 
relationships of the taxa within these two groups are somewhat clearer. 
G. glabra appears to be a part of the radiation of the betas in the Australian 
Region. 

INTRODUCTION 

Recently one of us (Parker) obtained in the highlands of Bou- 
gainville a new species of cryptic skink that possesses the follow- 
ing combination of characters unique in lygosomine skinks: trans- 
lucent spectacle in the movable lower eyelid; prefrontals absent; 
single frontoparietal; a single median scale situated between the 
single pair of nuchal scales; minute external ear opening; and 
pentadactyl limbs. 

As the proper generic allocation of the new species was not 
clear from these external characters, a skull was prepared for 
comparison with the skulls of more than 350 species of skinks. 
The skull morphology immediately allowed us to recognize the 
nearest relatives of the new skink and determine that skull char- 
acters in combination with the external characters warrant separate 
generic status for the species. 

1 Kundiawa, Chimbu District, Territory of New Guinea. 



2 breviora No. 302 

SYSTEMATIC DESCRIPTION 

The new genus and species may be known as: 

(ii omyi rsia glabra 1 new genus and species 

Holotype: Museum o\ Comparative Xoologv l >3714. collected by 
natives lor Fred Parker at Mutahi. Bougainville ( elevation 2200- 
32(H) feet) on IS Ma\ 1966. 

Paratypes (7 specimens): NIC'/. 8761 I. 93710, 93712, Natur- 
historiska Riksmuseet (Stockholm) 67-0001, same data as holo- 
type. but collected in the period 10-19 Ma\ 1966. MCZ 93713, 
Melilup, Bougainville (elev. 3000 - 500 feet) on 14 Ma\ 1966. 
MCZ93711,Ramazon River, Bougainville (elev. L 600-2400 feet) 
on 19 May 1966. MCZ 93708, Turiboiru, Bougainville (elev. 
500 feet) on 21 March 1966. 

Generic diagnosis: The genus is similar to an assemblage of 
lygosomine skinks inhabiting the Australian Region and subsaharan 
Africa. This assemblage mav be characterized by the following 
suite ol skull characters: palatal rami of pterygoids somewhat ex- 
panded and deeply emarginated posteriori) and separated mediall) 
bv medioposterior processes from the palatines, which project into 
the interpterygoid vacuity (beta palatal pattern, see p. 10 and 
Figure 4); 11-15 teeth on the premaxillae; no postorbital bone; 
SUpratemporal fenestra absent or minute; and Meckel's groove 
obliterated bv the overlapping and fusion of the dentary. 

I he genus Geomyersia mav be distinguished from all other ta\a 
in this assemblage (see p. II) bv its depressed body form, the 
complete absence o\' prefrontal scales, and the presence of a single 
median scale between the single pair of nuchal scales. All other 
taxa in the assemblage possess well-defined prefrontals and lack an 
"internuchal" scale. No other skink, in fact, is known to have such 
an internuchal scale as a normal feature (Figure 1 ). 

Species diagnosis: The species diagnosis is the same as that for 
the genus. 

Description (Figure 1): In general appearance Geomyersia 
glabra is a small (32-36 mm in snout-vent length), very dark, 
brownish black skink with a markedlv depressed body form. The 



1 The genus is named for Professor George Sprague Myers of the Division 

of Systematic Bioloi:\. Stanford University, who stimulated the early interest 
of the senior author in systematic and evolutionary hiology. The specific 
name (glabra) refers to the extremely smooth appearance of the species. 



1968 NEW SKINK FROM BOUGAINVILLE 3 

limbs are pentadactyl and fail to meet when adpressed to the 
body. 

Head noticeably depressed; rostral about twice as wide as deep, 
forming an almost straight suture with the very large frontonasal; 
external naris centered in a rectangular nasal; a single loreal be- 
tween the nasal and two superposed preoculars; prefrontals entirely 
lacking; frontal short, as long as the medial longitudinal length of 
the fused interparietals, and forming a shallow, but broad, con- 
cave suture with the frontonasal; frontal in contact with the two 
anteriormost of the four supraoculars; lower eyelid movable with 
a large translucent spectacle; small scales of eyelid separated from 
supralabial scales by complete row of suboculars; 6-6 supra- 
labials, 4th supralabial below center of eye; frontoparietals fused 
into a single scale; interparietal distinct, shorter than medial longi- 
tudinal length of the single frontoparietal; parietals large, forming 
a suture approximately two-thirds the length of interparietal; each 
parietal bordered posteriorly by a single large nuchal and temporal; 
a small, median scale separates the pair of nuchals. 




Figure 1. Dorsal view of the head of the holotype of Geomyersia glabra 
(MCZ 93714). Notice the absence of prefrontal scales and the presence of 
the single frontoparietal and internuchal. 



Tympanum sunk below level of epidermis; external ear opening 
present but minute and without auricular lobes; body scales smooth, 
slightly iridescent, and disposed in 22-24 longitudinal rows around 
midbody; a median pair of enlarged preanal scales bordered laterally 



4 BREVIORA No. 302 

by two subcqual scales; median row of subcaudal scales transversely 
enlarged. 

Dorsal surface of digits covered by a single row of scales (Group 
1 of Brongersma, 1942); 11-15 subdigital lamellae beneath 4th 
(longest) toe; lamellae folded into a median and two lateral ridges. 

(Dior: The dorsum and sides are dark grayish brown to brownish 
black; a median series of anteriorly-directed dark brownish black 
chevrons beszin on the base of the tail and extend forward to the 
nape; three or four horizontal streaks composed oi long dark 
dashes extend along the centers of the lateral scale rows between 
the fore and hind legs. Color does not appear to alter with preser- 
vation. 

The head is as dark as the dorsum, except for light cream stripes 
along the sutures between the supralabials. The venter is light 
with thickly clustered dark spots which become more confluent 
and dense posteriorly. The underside of the tail is so heavily 
spotted as to be almost uniformly dark. 

Distribution (Figure 2): The species is known only from middle 
elevations (500-3500 feet above sea level) in the highlands of 
northern and southern Bougainville. 

Reproduction: Geomyersia glabra is oviparous and apparently 
lays only a single egg at a time. Three o\ the seven individuals 
known were gra\id. MCZ 87611 and MCZ 93712 (snout-vent 
lengths 35.5 and 34 mm respectively) each contained 1 oblong. 
leathery shelled egg. MCZ 93714, with a snout-vent length of 33 
mm. contained a single, large volkv egg in the oviduct. 

Ecological observations: (icomxersiu glabra has been collected 
between 500-3500 feet. Only one specimen (MCZ 93708) has 
any precise ecological information associated with it. This indi- 
vidual was found in decayed vegetable matter in secondary forest. 
All the specimens of G. glabra were brought in by native collectors 
along with large numbers of Sphenomorphus (especially solomonis 
and tanncri). Apparently the natives do not distinguish Geomyersia 
glabra from the small svmpatric Sphenomorphus (see Morphologi- 
eal Comparisons with Other Small Solomon Islands Skinks). 

The few (six) specimens of Geomyersia glabra obtained in the 
Mutahi-Melilup area, in proportion to the thousands of individuals 
of other species collected in the same area, indicate that the species 
is either extremely rare or is more or less restricted to a peculiar 
habitat that was not widely sampled during the overall collecting. 



1968 



NEW SKINK FROM BOUGAINVILLE 



•6°S 



7°S 



156° E 



30 miles 



o 
155 E 



SHORTLAND 



MONO I (^J> 



6°S 




156° E 



Figure 2. Bougainville and neighboring islands showing the known locali- 
ties from which Geomyersia glabra has been collected. 



The apparent rarity is not solely the result of size, as many juveniles 
of the small skinks are collected and the natives were offered in- 
centives to collect the species. (Interestingly enough the largest 
skink in the Mutahi-Melilup area, Sphenomorphus taylori, is as 
infrequently collected as is Geomyersia glabra.) 

Morphological Comparisons with Other Small Solomon Islands 
Skinks: No close relatives of the monotypic Geomyersia are known 



6 breviora No. 302 

to occur naturally in the Solomon Islands, 1 although there are three 
small to medium sized skinks with a more or less uniform dark 
dorsal coloration in the Solomons with which the new species might 
be confused. Two of these three, Sphenomorphus solomonis and 
S. tanneri, occur together with G. glabra on Bougainville, while the 
third. S. bignelli, is known at present only from the southern Solo- 
mon Islands. The distinguishing features of the four species are 
outlined in Table 1. 

Discussion: Among lygosomine skinks there is a large group cen- 
tering on the genus Leiolopisma, as used in its broadest sense, that 
may be distinguished on the basis of ( I ) a single pair of nuchal 
scales behind the parietals. each o\' which is in contact with a single 
large temporal scale laterally, (2) the upper sides of the digits 
covered by a single row of scales (Group I of Brongersma, 1942), 
( 3 ) the lower eyelid usually, but not invariably, with a clear spec- 
tacle, and (4) preanal scales often subequal or at least not greatly 
enlarged. 

Correlated with these external characters are the following skull 
characters: ( 1 ) palatal rami of pterygoids separated on the midline 
— often by medioposterior projections from the palatines into the 
interpterygoid vacuity. (2) II or more premaxillarv teeth. (3) a 
small or minute supratemporal fenestra. (4) no postorbital bone. 
and (5 ) Meckel's groove obliterated by the overlapping and fusion 
of the dentary. 

Within this large group, two subgroups may be recognized on 
the basis of the presence or absence of a medial, posteriorly pro- 
jecting process from the palatal ramus of the pterygoid and a 
concomitant emargination in the palatal ramus (Figures 3 and 4). 

One group, the alpha group for the purposes of this discus- 
sion, has the palatal rami of the pterygoids gradually and smoothly 
diverging postcriorlv from the midline (Figure 3). In the other 



( arlia fusca, a close relati\c of Geomyersia glabra (see below), is known 
to occur in the Solomon Islands only on tiny Sohano Island in the strait be- 
tween Buka and Bougainville, on I-aisi Island and at Buin, Bougainville. The 
species is as yet unknown from any other localities in the Solomons, and it 
has almost certainly been introduced in the known localities. C. fusca is very 
common around the town of Rabaul, New Britain, the port from which 
Bougainville receives most of its supplies. The ecology of the species in 
Rabaul. the Solomons, and New Guinea is similar. It is a grass-dweller, pre- 
ferring the thick tussock grass of open areas. 



1968 



NEW SKINK FROM BOUGAINVILLE 



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HRl.VIORA 



No. 302 







Figure 3. The alpha palatal pattern (ventral view): (A) Mabiiya poly- 
tropi.s (MCZ B103), Kribi. Cameroon; (B) Leiolopisma me tallica (MCZ 
67129), Mt. Toolbewong, Healesville, Victoria; (C) Emoia samoense (MCZ 
16931 ), Fiji Islands; (D) Eumecia anchictac (MCZ 41562), Kaimosi, Kenya. 
Not drawn to scale. Abbreviations: E, ectopterygoid; P, palatine; PT, 
pterygoid. 



968 



NEW SKINK FROM BOUGAINVILLE 







Figure 4. The beta palatal pattern (ventral view): (A) Carlia bicari- 
nata (MCZ 64315), Port Moresby, New Guinea; (B) Leptosiaphos bloch- 
manni (MCZ untagged). Upper Mulinga, Idjwi Island, Democratic Republic 
of the Congo; (C) "Ablepharus" smithi (MCZ 42880, paratype), Nyonga, 
Democratic Republic of the Congo; (D) Geomyersia glabra (MCZ 87611, 
paratype), Mutahi, Bougainville. Not drawn to scale. Abbreviations as in 
Figure 3. 



10 BREVIORA No. 302 

group, here called the beta group, 1 the palatal rami of the pterygoids 
have medial, posteriorly projecting processes set off bv emargina- 
tions lateral to them. In addition, whereas medial, posteriorly pro- 
jecting processes from the palatines are present only in some alphas, 
they are a constant feature of the beta palatal pattern (Figure 4). 

The functional significance of the deeply emarginated palatal 
ramus oi the pterygoid in the beta group (Figure 4) is not pcr- 
fectl) clear. The emarginatioa may serve as a "catch" for the 
basipterygoid process as the quadratal ramus o\' the pterygoid 
slides backward along the outer edge of the process. Such a 
movement on the part oi the quadratal ramus is evident in Creshl) 
prepared skulls when the forward part of the skull is depressed 
(hinging at the transverse frontal-parietal suture), and the whole 
palatal complex is displaced backward. In freshly prepared skulls, 
the basipterygoid process can catch in the emargination, stopping 
the posterior displacement o\' the palatal complex, thus limiting the 
amount o{ strain put on the skull. 1 he skull is probably capable o\' 
such movement during active feeding, but the differences in feeding 
behavior or habits that would make this "catch" mechanism an 
advantage to the skinks o{ the beta group are unknown. 

The taxonom) of the alpha and beta groups is extremely com- 
plex and will be dealt with in future papers. It is of interest for the 
present discussion, however, to list, using current generic concepts, 
those skinks. with their distributions, which belong to the alpha and 
beta groups. I he number of species in each genus in each area is 
given in parenthesis. 



I he beta skinks ma) be of mono- or diphvletic origin. Present evidence. 
largelj geographic, suggests to us th;it the betas of subsaharan Africa and of 

the Australian Region base probably independent!) originated from an alpha 
ancestry, i.e.. they are diphyletic. if so, use of the word "assemblage," which 
carries no connotations of close relationship for zoologists, would perhaps 
be more appropriate than the word "group," which does connote close rela- 
tionships, when discussing the beta skinks as a unit. In the diagnosis of 
Geomyersia the word "assemblage" was used for the beta skinks. hut in the 
following discussion, largeh as a matter of stylistic balance with the term 
"alpha gioup" (on present evidence the alphas are monoph\letic in origin), 
the term "beta group" will be used. It should be borne in mind, however, 
that the mono- or diphvletic origin of the beta skinks is still a problem 
needing further research. 



1968 



NEW SKINK FROM BOUGAINVILLE 



11 



alpha 

Ablepharus Australia (part, 11); 
Asia (8) 

Anotis New Caledonia (3) 

Didosaurus mauritianus, subfossil 
on Mauritius 

Emoia Indo-Australian archipel- 
ago, N. Australia, and Pacific 
islands (40) 

Eugongylus New Guinea, N. Aus- 
tralia, and Solomon Islands (5) 

Eumecia subsaharan Africa 
(1 or2) 

Leiolopisma Asia (17); North 
America (6); Indo-Australian 
archipelago (20); Australia (part, 
8) ; Lord Howe Is. ( 1 ) ; New Cale- 
donia (6); New Zealand (16); 
Mauritius (1) 

Mabuya Central and South America; 
to Indo-Australian archipelago 
(66) 

Sphenomorphns New Guinea (mi- 
nutus); Solomon Islands (bignelli); 
New Zealand (pseudornatus) 



beta 

Ablepharus Africa (6); Australia 
(greyi and burnetii) 



Carlia Australian Region (21 ) 



Geomyersia Solomon Islands ( 1 ) 



Leiolopisma Africa (7); Australia 
{weeksae, challengeri, mustelina, 
guichenoti, delicata) 



Leptosiaphos Africa (7 



Panaspis Africa (7) 



Within the beta group, the group to which Geomyersia belongs, 
relationships are complex and still incompletely worked out. For 
the moment, therefore, our purpose will only be to discuss some of 
the clearest groupings within the beta group and with this informa- 
tion assess the relationships of Geomyersia. 

(1) It is implicit in the list of alpha and beta skinks given 
above that the genus Ablepharus, characterized by "no movable 
eyelids, [and] a transparent disk covering the eye" (Boulenger, 
1887), is not monophyletic; indeed, the genus is an assemblage of 
species, derived from diverse stocks, with a clear spectacle in the 
movable lower eyelid (Smith, 1935 and 1937; Greer, 1967a; Fuhn, 
in an unpublished discussion of the polyphyletic origin of Able- 
pharus). 



12 BREVIORA No. 302 

The Australian A. greyi and A. burnetii (beta group) arc the 
only "Ablepharus" with a reduced digital formula (4 fingers and 
5 tecs), and significantly, they occur in the same zoogeographic 
region as the only other group of beta skinks with a similarly re- 
duced digital formula, i.e.. Carlia. The fused frontoparietal and 
spiny auricular lobules of A. burnetti are shared with Carlia and 
suggest that A. burnetii is simplv a Carlia in which the movable 
lower eyelid with its clear spectacle has fused almost completely 1 
to the upper edge o( the orbit. 

The 4-5 digital formula and fused frontoparietals of I. greyi 
also argue for the close relationship o\ this species with Carlia. but 
the unique supraocular arrangement found in A. greyi would sug- 
gest separate generic rank. The generic name Menetia Gray, 1845, 
originall) proposed for this species, is available. 

(2) I he other beta skinks in the Australian region, in addition 
to Carlia and its two ablepharine derivatives discussed above, are 
Geomyersia glabra and five species referred to Leiolopisma: week- 
sue, challenger i, mustelina, guichenoti, and delicata. In possessing 
5 lingers these five species of Leiolopisma, as well as Geomyersia, 
displa) the primitive morphological condition from which the 
4-lingered condition oi Carlia and its two ablepharine relatives must 
have been derived. 

Three of the 5 beta Australian Leiolopisma displa) yet another 
primitive character from which a more advanced character state 
displayed by all other beta Australian skinks must have evolved: 
Leiolopisma weeksae, chaUengeri and mustelina have paired fronto- 
parietals while all other Australian betas have fused frontoparie- 
tals. 3 



1 he onl\ other skinks with B reduced number of fingers ami iocs pre 
viously considered to be in the genus Ablepharus are six Australian species: 
elegans, distinguenda, orient alis, muelleri, timida, lineata. These species are 

now considered to be congeneric (genua Lerista) with the Australian skinks 
formerly referred to Rhodona (Greer. 1967a). and are only distantly related 
to any of the alpha or beta skinks discussed in this paper. 

In the single specimen of Ablepharus burnetii available to us (MCZ 
6486 ) there is a small palpebral slit remaining along the dorsal edge of the 
lower eyelid. The e\e is effective!) "ablepharine," however, as this free dor- 
s.il edge is surely too short to allow much movement of the eyelid. 

'Geomyersia grabra may be distinguished from the 2 beta Australian 
Leiolopisma with fused frontoparietals and 5 fingers {guichtnoti and delicata) 
by the complete absence of prefrontal scales and the depressed body form. 



1968 NEW SKINK FROM BOUGAINVILLE 13 

These 3 primitive beta Australian Leiolopisma cannot, at this 
stage of the study, be distinguished morphologically as a group 
from the African beta Leiolopisma, all of which have 5 fingers and 
separated frontoparietals. This similarity could be the result of 
either convergence or close relationship. For the present the 
widely disjunct distribution of the two groups implies to us that 
the groups have arisen independently in each area, but, admittedly, 
this is an unsolved problem and will be dealt with elsewhere. 

(3) Taken as a group the African betas are themselves taxo- 
nomically difficult at an infragroup level. Several characters might 
be used to distinguish taxa within the African betas: the presence 
of supranasals, presumably a primitive condition, serves to diagnose 
Panaspis; a scaly lower eyelid, minute ear opening and a group 
tendency toward an elongate body and reduced number of digits 
distinguishes Leptosiaphos; the ablepharine eye and reduced num- 
ber of supraocular scales (3 instead of 4) defines the African 
"Ablepharus," while an African group of Leiolopisma could be 
"diagnosed" negatively on the basis of showing none of these char- 
acters. Certain species, however, bridge these apparent taxonomic 
gaps. For example, Panaspis cabindae may have 3 instead of 4 
supraoculars (Parker, 1936), and in P. breviceps the supranasal 
may be fused to the nasal (Smith, 1937) ; in addition, Leptosiaphos 
kilimense and L. rhodurus are atypical of other Leptosiaphos in 
displaying a transparent window in the movable lower eyelid. Such 
difficulties suggest that on the basis of these characters, it might be 
better to resort to, at most, a subgeneric scheme of ranking taxa. 

(4) On the basis of geography, Geomyersia glabra would seem 
more closely related to Australian betas than to African betas. In 
addition the fused frontoparietals of G. glabra may be a morpho- 
logical indication of relationship with the Australian betas which 
possess this advanced character in 26 of the 29 species. The Afri- 
can betas have divided frontoparietals, presumably the more primi- 
tive condition, in all but one of the 27 species. Only in the African 
beta Ablepharus wahlbergi are the frontoparietals normally fused, 1 
a condition probably derived independently from that of the Aus- 
tralian betas with fused frontoparietals. 



1 The fusion of the frontoparietals and their fusion with the interparietal 
is a variable condition in A. wahlbergi (personal observation and de Witte, 
1953). 



14 hreviora No. 302 

Geomyersia glabra docs, however, have 5 fingers — a character 

which is predominant in the African betas and which might, there- 
tore, be thought to indicate greater likelihood of relationship with 
African rather than Australian beta skinks. However. 5 fingers is 
the primitive condition for both the African and Australian betas, 
and although in the Australian betas the 4 fingered species are a 
conspicuous element in that group's radiation, the 5 lingered G. 
glabra can be readily interpreted as a relic of an early stage in 
the evolutionary history of Australian betas. It would presumably 
represent a stage later than that of Leiolopisma weeksae, /.. chal- 
lengeri and /.. mustelina and approximating that of I., guichenoti 
and /.. delicata, but with specializations of its own. 

Certainly the island of Bougainville on the periphery of the 
Australian region would be a logical refuge in which to find a relic 
of an early stage of a radiation taking place in this region. 

A dendrogram of the Australian beta skinks. assuming on the 
basis of present evidence — mainl) geographic — that they are 
monophyletic, is presented in figure 5. 

I he radiations of the Australian and African beta skinks will be 
discussed further in forthcoming papers. 

( mparisons with Morphologically Similar Species: It is desir- 
able to mention brief!) those species in the Australian Region that 
are confusing!) similar to Geomyersia glabra on the basis of exter- 
nal morphology. Only those skinks displaying the external charac- 
ters of the alpha and beta skinks are discussed. 

Leiolopisma mccooyi (Lucas and frost. 1894) from Victoria 
lacks prefrontals and has a minute external ear opening as does 
G. glabra, but the frontoparietals of mccooyi are divided, the body 
is not depressed, and the palate is of definitely alpha type. 

/ ygosoma graciloides Lonnberg and Anderson, 1913 (=L\go- 
soma scharffi Boulenger, 1915) has been collected twice and is 
known from 4 specimens. Although described twice under the 
genus / ■///</. the species has the clear spectacle and other ex- 

ternal characteristics of the alpha and beta skinks outlined above. 
Like Geomyt rsia glabra the prefrontals are absent and the ear 
opening is minute, but L. graciloides differs from Geomxcrsia 
glabra in having paired frontoparietals and only 4 fingers. More 
important, however, are the palatal dilferences. for L. graciloides 
is an alpha skink. 

The three endemic Anotis (or Siaphos) on New Caledonia — 
mariae, gracile, and slevini — are undoubtedly closely related to 



1968 



NEW SKINK FROM BOUGAINVILLE 



15 



Ablepharus " bu rnetii 



Menetia greyi 



Prefrontals present 
Fused frontoparietals 
Four fingers 
Ablepharine eye * 



Prefrontals present 
Fused frontoparietals 
Four fingers 
Ablepharine eye * 
Unique supraocular scale 
arrangement * 
/ 



Geomyersia glabra 



/ 



Carlia 



Prefrontals present 
Fused frontoparietals 
Four fingers * 
Clear spectacle in movable 
lower eyelid 



Prefrontals lacking • 
Fused frontoparietals ' 
Five fingers 
Spectacle in movable 
lower eyelid 



Leiolopisma delicata 
L. guichenoti 



Prefrontals present 
Fused frontoparietals * 
Five fingers 

Clear spectacle in movable 
lower eyelid 



Leiolopisma weeksae 
L. challenged 
L. mustelina 



Prefrontals present 
Paired frontoparietals 
Five fingers 

Clear spectacle in movable 
lower eyelid 



Alpha ancestry 

Figure 5. A dendrogram of the Australian Region beta skinks. Changes 
in character states of a taxon over its immediate hypothetical ancestral con- 
dition are marked with an asterisk (*). 



16 BREVIORA No. 302 

each other and rather similar to Geomyersia glabra in gross appear- 
ance. These three species differ from G. glabra, however, in pos- 
sessing minute prefrontals and. judging from a single skull of mariae 
(MCZ 92393). the group has the alpha palatal pattern. 

Although the literature to date on these three species states or 
implies that there is a clear spectacle in the lower eyelid, the two 

ccimens oi mariae (MCZ 1960S, 4ol77) and the one specimen 
o\ rtevini (MCZ ( )2^5. holotype) in the Museum of Comparative 
(logy clear!) show sutures across the eyelid. In the single speci- 
men (.)! gracile (MCZ 46172) available, the situation in regard to 
the eyelid is equivocal. In G. glabra, on the other hand, there is ;i 
distinct single translucent window in the lower eyelid. 

Acknowledgements: Dr. Carl Edelstamm oi the Naturhistoriska 
Riksmuseet, Stockholm, verj kindly cooperated in an exchange ol 
specimens that permitted us to examine 1 of the 3 syntypes of 
ma graciloiaU s. 

Mrs. Patricia II. Kerfoot prepared the line drawing oi the holo- 
type o\ Geomyersia glabra for Figure 1. 

Dr. Ernest E. Williams oi the Museum oi Comparative Zoology 

has read the manuscript in several editions and has, as usual, offered 

main helpful eomm nts. We have also benefited greatly from main 

s with Dr. Williams concerning the reptile faunas of 

islands. 

Dr. Richard Fstcs has also provided valuable criticism oi the 
manuscript 

I' rt oi the research by Greer for this paper was supported by 
the Evolutionary Biology Fund which is administered for the Na- 
tional Science I oundation by the Evolutionary Biolog) Commit- 
»f the Biological laboratories. Harvard University. 

A grant from the Science and Industry Endowment Fund of the 
Commonwealth Scientific and Industrial Research Organization of 
Australia covered part o\ Parker's collecting expenses during his 
1966 field work in the Solomons. 

Publication costs were defrayed by funds from the National 
Science Foundation grant GB 6944 to Dr. Ernest E. Williams. 

REFERENCES CITED 

BOULF.NGER. G. A. 

1887. Catalogue of the lizards in the British Museum. Vol. III. Tay- 
lor and Francis. London, xii + 575 pp. 

1915. Descriptions of two new lizards from Australia. Ann. Mag. Nat. 
Hist., (8) 16:64-66. 



1968 NEW SKINK FROM BOUGAINVILLE 17 

Brongersma, L. D. 

1942. On the arrangement of the scales on the dorsal surface of the 
digits in Lygosoma and allied genera. Zoologische Medeelingen 
(Leiden), 24 (1-2) : 153-158. 

Gray, J. E. 

1845. Catalogue of the specimens of lizards in the collection of the 
British Museum. Edward Newman, London, xxviii -f 289 pp. 

Greer, A. E. 

1967a. A new generic arrangement for some Australian scincid lizards. 

Breviora, Mus. Comp. Zool., 267: 1-19. 
1967b. The generic relationships of the African scincid genus Eumecia. 

Breviora, Mus. Comp. Zool., 276:1-9. 

Greer, A. E. and F. Parker 

1967. A new scincid lizard from the northern Solomon Islands. 
Breviora, Mus. Comp. Zool., 275: 1-21. 

Lonnberg, E. and L. G. Anderson 

1913. Results of Dr. E. Mjoberg's Swedish scientific expeditions to 
Australia 1910-13. III. Reptiles. Kungl. Svenska Vetenskapsaka- 
demiens Handiingar, 52 (3) : 1-17. 

Lucas, A. H. S. and C. Frost 

1894. The lizards indigenous to Victoria. Proc. Roy. Soc. Victoria, 
(N. S.) 6 (2):24-92. 

Parker, H. W. 

1936. Dr. Karl Jordan's expedition to South-West Africa and Angola: 
herpetological collections. Novitates Zoologicae, 40:115-146. 

Smith, M. A. 

1935. The fauna of British India, including Ceylon and Burma. Rep- 
tilia and Amphibia. Vol. II — Sauria. Taylor and Francis, Lon- 
don, xiii-f 440 pp. 

1937. A review of the genus Lygosoma (Scincidae: Reptilia) and its 
allies. Rec. Indian Mus., 39 (3):213-234. 

Witte, G. F. de 

1953. Exploration du Pare National de 1'Upemba. Reptiles. Institut 
des Pares Nationaux du Congo Beige, Bruxelles, 322 pp. 

(Received 5 March 1968.) 



BREVIORA 

Museeiii of Comparative Zoology 



Cambridge, Mass. 31 December, 19 68 Number 303 

REVIEW OF THE GENERA OF THE TRIBE LOBERINI 
(Coleoptera: Languriidae) 

T. Sen Gupta 1 

Abstract. The subfamily Loberinae (Bruce 1951) and the tribe Lob- 
erini are fully characterized for the first time; the genera of Loberini are 
redefined; three new subgenera of Hapalips are introduced, and a key to the 
adults of the subgenera of Hapalips is given; a new species Hapalips acaciae 
and the larva of Hapalips prolixus are described; a key to the genera of the 
adults of the tribe Loberini is given. 

INTRODUCTION 

Bruce (1951) was the first author to point out that species of 
Telmatophilus Heer have male genitalia like those of Cryptophagus 
Herbst and unlike those of Hapalips Reitter, Loberus LeConte, and 
Toramus Grouvelle. He also noted that in the confusedly punc- 
tured elytra, Telmatophilus agrees with Cryptophagus and differs 
from the last three genera mentioned above, in which the punctures 
form regular rows. He proposed the name Loberinae for the old 
Telmatophilinae, excluding Telmatophilus but including Leuco- 
himatium Rosenhauer, which, despite the simple tarsi, have male 
genitalia and elytral punctation similar to those in Loberus. Sen 
Gupta and Crowson (1967) considered Pharaxonotha Reitter and 
related genera, including Leucohimatium and Xenoscelis Wol- 
laston, as comprising the tribe Pharaxonothini of the Loberinae. 
Recently, Martinez and Berrera (1966) established a new genus 
Loberopsyllus, in the family Cryptophagidae and related to Lo- 
berus, which is here considered to be in the Pharaxonothini be- 
cause of the simple tarsi, rather short trochanters, absence of 
femoral lines on the first ventrite, and narrowly open front coxal 



1 Department of Entomology, University of Alberta, Edmonton, Alberta, 
Canada. 



2 BREVIORA No. 303 

cavities. The genus Toramus and its allies are considered by me 
( 1967) as a subfamily Toraminae of the Languriidae. The charac- 
ters of the larvae of Hapalips, Pharaxonotha, and Bolerus Grou- 
velle, given by Rymer Roberts (1939) and Rymer Roberts and 
Van Emden (1958). and of the larva of Eicolyctus Sahlberg, de- 
scribed by Sen Gupta and Crowson ( 1967), support the classifica- 
tion here proposed. 

The chief characters in which the Loberinae differ from the 
Cryptophagidae are as follows: 

Elytral epipleura well defined up to the apex, and elytra] punc- 
tation usually in regular rows. Wing always with subcubital fleck 
and radial cell, never with five anal veins. Tarsal formula never 
5-5-4 in male. First ventrite not markedly longer than second. 
Aedeagus turned on one side in resting condition, with long, 
threadlike, double median struts. Sternal fitting between the meso- 
coxae witli a single knob (found only among the Atomariinae in 
the Cryptophagidae). Larvae never with annular spiracles, single 
jointed labial palpi, bifid apex, or serrated caudal margin of pros- 
theca. Larvae with five to six ocelli on each side of head, and 
granulated upper body sin face. 

The subfamily Loberinae may he defined by the following 
characters: 

With general characters oH Clavicornia, Languriidae (according 
to Crowson. 1955). 

1 ) Head often with stridulatorv files: Ironto-clypeal suture ab- 
sent; transverse line on occipital region sometimes present; anterior 
part of gular region with ( Fig. 20) or without a transverse groove, 
sometimes with a large transverse cavil' ( Fig. 23). 

2) Antennal insertions hidden bv the sides of frons; antenna] 
club three-jointed, sometimes ver) weakly developed. Mandible 
with well-developed mola (Fig. 13): maxillary lacinia with three 
apical spines, galea narrow and elongated (Fig. 8); labium with 
apical segment of palpi somewhat transverse, mentum sometimes 
with single or paired cavities on ventral side; ligula usually poorly 
developed. 

3) Front coxal cavities internally open behind (Figs. 2, 6, 24). 
externally moderately wide open (Fig. 6) to nearly (Fig. 24) or 
completely closed behind (Xenoscelis) . 

4) Elytra glabrous or pubescent, usually regularly punctured, 
scutellary stride often present. Wings (Figs. 18, 19) always with 
four anal veins, radial cell, r-m cross vein; anal cell sometimes 
present. 



1968 GENERA OF LOBERINI 3 

5) Mesocoxae usually closely situated, and sternal fitting be- 
tween them with a single knoblike projection (Fig. 9). Meso- 
sternal pockets (Fig. 9) rarely present, mesepisternal pockets usu- 
ally well developed (Fig. 9). 

6) Metendosternite with anterior tendons separated by slightly 
less than width of basal stalk, and lateral plates narrow (Figs. 
21,25). 

7) Tarsi lobed or simple; trochanters broadly elongate (Fig. 
26) or broad and short. 

8) Ventrite 1 with (Fig. 1) or without femoral lines. Ovipositor 
as in Figure 12, with styli attached at the apex of coxites, except 
in Leucohimatium, where the styli are attached slightly above the 
apex of coxites. Aedeagus of Erotylidae-Languriidae type (Figs. 
10,11). 

9) Larvae as far as known with mandibular prostheca large, 
translucent, and triangular; ocelli usually five to six on each side 
of the head, sometimes absent; dorsal surface granulated; two tar- 
sungular setae; larvae not endophytic as in Languriinae. 

TRIBE LOBERINI 

(Loberinae Bruce, 1951: 4, partim.) 

This group includes those members of the Loberinae that have 
lobed tarsi, front coxal cavities that are usually clearly open be- 
hind, and cryptophagid-like facies, but that have regular rows of 
punctures on the elytra. Previously these genera have been in- 
cluded in the family Cryptophagidae. According to present knowl- 
edge, the tribe includes seven genera: Loberus LeConte, Telma- 
toscius Sharp, Hapalips Reitter. Truquiella Champion, Pseudo- 
henoticus Sharp, Pseudhapalips Champion, and Bolerus Grouvelle. 
Of these, only the two largest, Loberus and Hapalips, occur in 
both the New and Old Worlds. The genera Telmatoscius, Tru- 
quiella, Pseudohenoticus, and Pseudhapalips are restricted to the 
New World, and the genus Bolerus to the Old World. The species 
occur mainly in tropical and subtropical climatic zones, a few 
species of Loberus and Hapalips extending into warm temperate 
climates. None have been recorded from Europe. The only pre- 
viously known larvae are those of Bolerus (=Platycladoxena) 
angulosus Arrow and a Mexican Hapalips, both described by Ry- 
mer Roberts (1939). The larva of Hapalips prolixus (Sharp) 
from New Zealand is described below. 

The tribe Loberini may be defined by the following characters: 
With general characters of Languriidae, Loberinae. 



4 bkhviora No. 303 

1 ) Head never with a transverse line on vertex. 

2) Transverse groove on anterior part of gular region (Fig. 
20) almost always present. 

3) A pair of stridulatory files on vertex often present (Fig. 22). 

4) Antennal club well developed and three-jointed. 

5) Front coxal cavities usually moderately wide open, some- 
times very narrowly so (Hapalips, Fig. 24). but never completely 
closed behind. 

6) Elytra rarely glabrous, usually with punctures in regular 
rows, and sometimes with scutellary stride. 

7) Wing with (Fig. 18) or without (Fig. 16) anal cell. 

8) Mesocoxae closely situated except in Pseudhenoticus; mes- 
episternal pockets well developed (Fig. 9). 

9) Tarsi pseudotetramerous and trochanters broadly elongated 
(Fig. 26). 

10) Femoral lines on first ventrite always present, and oviposi- 
tor of Loberinae-type (Fig. 12); stvli attached at the apex of 

COXltCs. 

1 1 ) Larvae as far as known with granules of tergites arranged in 
regular transverse and longitudinal rows (Fig. 34), frontale with- 
out endocarina (Fig. 27), and mandible without hairy appendages 
at the base of mola ( Fig. 29 ) . 

Genus Loberus LeConte 

Loherus LeConte. 1861: 98. Type species, by subsequent monotypy. 

Loberus impressiu LeConte, 1863: 70. 
Glisonotha Motschulsky, 1863: 430. 

LeConte (1861) established this genus in the family Crypto- 
phagidae but did not name a species of it until 1863. This is the 
largest genus of the tribe; Schenkling (1923) included 59 species, 
and Bruce has subsequently added another nine from Africa, Java, 
and Mexico. Grouvelle (1919) gave a key to the Old World spe- 
cies known at the time, and another for American species that he 
had seen. 

Description. With general characters of Loberinae, Loberini. 
Facies somewhat cryptophagid-like. Head usually with a pair of 
stridulatory files; eyes large and moderately coarsely faceted; cly- 
peus with straight apical margin. Antenna with scape longer than 
pedicel, which is shorter than segment 3; segments 4-8 equal in 
length, segments 9 and 10 semicircular and transverse, segment 9 
slightly narrower than 10, terminal segment slightly longer than 
broad, its apical margin rather asymmetrically rounded. Prothorax 
(Fig. 4) transverse, side margins weakly curved, more or less 



1968 GENERA OF LOBERINI 5 

smooth, front and hind margins almost equal in breadth, front 
angles obtuse, hind angles more acute; prebasal impressions on 
pronotum present; prosternal process narrow and its apex almost 
straight. Elytra usually pubescent, rarely almost glabrous, pubes- 
cence often double with recumbent setae and sparsely distributed 
long erect ones as in Toramus; punctation on elytra usually in more 
or less regular rows, rarely quite irregular; scutellary striole ab- 
sent; epipleura narrow, rather indistinct toward the apex as in 
Toramus. Wing without anal cell (cf. Fig. 15). Mesocoxae closely 
situated, mesepisternal pockets well developed; metasternum wider 
than its length, median impressed line extending half of its length. 
Tarsi with segment 1 very slightly longer than segment 2, segment 
3 lobed below, segment 5 almost equal to first three segments to- 
gether, excluding the lobes; tibiae narrow, not widened at apex. 
Ventrite 1 with intercoxal process broad at base, narrowed to a 
pointed apex. Aedeagus as figured (Fig. 11); ovipositor of Loberi- 
nae-type (cf. Fig. 12). 

Species examined. The species L. impressus LeConte and L. 
humeralis Reitter have been studied in detail for the characteriza- 
tion of the genus. 

Habitat. Nothing recorded, and larva undescribed. 

Geographical distribution. Very extensive, occurring in both the 
New and Old Worlds, species recorded from North and South 
America, Ethiopian and Madagascar regions, Indo-Malayan re- 
gion, and Australia, but absent from Palaearctic region except for 
the Far East, and not known from New Zealand. It is unfortunate 
that no authors have made critical comparisons between the New 
and Old World species of Loberus. 

Genus Telmatoscius Sharp 

Telmatoscius Sharp, 1900: 581. Type species, by monotypy, Telmatoscius 
claviger Sharp. 

Sharp (1900) established this genus for a single described spe- 
cies, placing it in the Cryptophagidae, close to Loberus. Grouvelle 
(Grouvelle and Raffray, 1912) added another species, but none has 
been described since. 

Description. With general characters of Loberinae, Loberini. 
General appearance as in Loberus and not very easily distinguish- 
able from it. Differs from Loberus in having comparatively long, 
dense recumbent setae on elytra; prothorax more narrowed in 
front, and segment 9 of antenna narrower in comparison to seg- 
ment 10 than in Loberus. Head as figured (Fig. 22), with a pair 



6 BR] mora No. 303 

of well-separated stridulatory files; eyes large, fairly coarsely 
faceted; clypeus with straight apical margin. Antennae with scape 
slightly larger than pedicel, which is as long as segment 3. segment 
4 slightly shorter than 3 and 5. segments 6-8 equal in length, seg- 
ment 10 distinctly larger than segment 9. terminal segment wider 
than long, its apex slightly depressed. Prothorax (Fig. 7) weakly 
narrowed in front, side margins curved and smooth, front angles 
more or less rounded, hind angles acute; prebasal impressions on 
pronotum present; prosternal process as in Loberus. Elytra with- 
out scutellary striole. Meso- and metasterna as figured (F ; ig. 9), 
metasternum longer than in Loberus, median impressed line ex- 
tending half o( its length. Metendosternite as figured (cf. Fig. 25). 
Legs and abdomen as in Loberus. Aedeagus and ovipositor as 
figured ( cf. Figs, l 1 . 12). 

Species examined. A male and female of T. claviger have been 
studied in detail lor generic characters. 

Habitat. Nothing recorded, and larva undeseribed. 

Geographical distribution. Central America. 

Genus Hapalips Reittcr 

Hapalips Reitter, 1877: 122. I \ pc species, In presenl designation, //<//></- 
lips mexicanus Rdtter. 

Reitter ( 1877) established this genus in the family Rhi/ophagi- 
dae. Gorham ( 1898) transferred it to Languriidae. howler ( 1908) 
retained it in Languriinac. Champion ( 1 1 > 1 3 ) and Grouvelle 
( 1919) placed it in C"r\ptophagidac. and Arrow | 1 l >2 c >a ) proposed 
that Hapalips, together with all Cryptophagidae having lobed tarsi. 
be plaeed in Languriidae. Rymer Roberts | 1939) placed Hapalips 
with Boleriu (=Platycladoxena) angulosus in Languriidae. Cla- 
doxeninae, alter studying larvae oi the two genera. Bruce 
(1951) noted that the aedeagus of Hapalips is similar to those 
oi Leucohimatium, Loberus, and Toramus. Crowson (1955) re- 
tained Hapalips in the Languriidae and considered it as a tran- 
sitional form between that family and Erotylidae. This is the 
second largest genus of the tribe Loberini. Reitter (1877) de- 
scribed eight species. Schenkling ( 1923) listed 47, to which Arrow 
(1927) added another one, and more recently, Bruce (1952, 
1963) added five more species and excluded one of his older 
species. //. spegazzini, because of its different type of aedeagus. 
Thus the genus at present consists of 54 valid species. There ap- 
pears to have been no previous designation of a type for this genus, 
so I here designate as type H. mexicanus, one of the original species 
described by Reitter ( 1877) and quite common in Mexico. 



1968 GENERA OF LOBERINI 7 

Description. With general characters of Loberinae, Loberini. 
General appearance more or less as in Xenoscelis Wollaston; form 
narrow, elongate, rather flattened, and more or less parallel sided. 
Head (Fig. 23) with broad clypeus, its front margin evenly 
rounded; antennal insertions completely hidden by the sides of 
frons. Stridulatory files rarely present and, if so, paired and well 
separated; sometimes files represented by a pair of longitudinal 
ridges without any striations (e.g. H. juscus and H. acaciae). 
Transverse groove on anterior part of gular region sometimes re- 
placed by transverse cavity (Fig. 23). Antenna of moderate 
length, scape small but longer than pedicel, which is often shorter 
than segment 3, segments 4-8 usually equal in length, segments 9 
and 10 transverse and equal in size, terminal segment equal in 
breadth to segment 10 and more or less rounded at apex. Pro- 
thorax (Fig. 24) usually elongate with side margins smooth, more 
or less parallel sided, front and hind margins equal in width, all 
the angles rather obtuse; prebasal impressions on pronotum often 
indistinct. Front coxal cavities usually with narrow, sometimes 
slitlike (Fig. 24) opening behind; prosternal process with its apical 
margin straight. Sometimes shape of pronotum differs between 
male and female. Elytra with regular rows of strial punctures, 
usually with a scutellary striole, and pubescence absent or fine and 
recumbent. Wing with (Fig. 18) or without (Fig. 16) anal cell, 
radial cell rarely without spur of radial sector (Fig. 17). Meso- 
coxae closely situated; mesepisternal pockets often obscured; meta- 
sternum elongate or as broad as long; median impressed line of 
metasternum usually extending % of its length. Tarsi short and 
compact, first three segments rather broad and equal in length, 
segment 3 lobed below, segment 4 minute, received in the lobe of 
segment 3, segment 5 about equal in length to first three together; 
tibiae rather short, broad at apex, which is obliquely truncate with 
two normal spurs. Ventrite 1 with intercoxal process pointed at 
apex. Ovipositor and aedeagus as figured (Figs. 12, 10). 

Species examined. The species H. mexicanus Reitter, H. eichel- 
baumi Grouvelle, H. cribricollis Gorham, H. grouvellei Gorham, 
H. nigriceps Reitter, H. nitidulus Champion, H. filum Reitter, H. 
scotti Grouvelle, H. prolixus Sharp, H. juscus Reitter, H. acaciae 
sp. n., and also H. taprobanae Grouvelle (external characters 
only), have been studied in detail for generic characters. 

Habitat. Little is known about the habitat of members of this 
genus. H. filum has been recorded from Cuba in corn (maize) 
stalks, and H. annulosus Grouvelle from Guadeloupe in flowers of 
the cactus Cereus triangularis. H. prolixus has been recorded from 



8 breviora No. 303 

New Zealand in tree ferns, and more recently R. A. Crowson col- 
lected both larvae and adults of this species under the sheathing 
bases of dead palm leaves. //. championi Grouvelle and //. scotti 
Grouvelle are recorded from the Seychelles, both from the bases 
of palm leaves. 

Geographical distribution. This genus is widely distributed in 
both the New and Old Worlds, mainly in tropical and subtropical 
regions; a few species extend into warm temperate areas. No spe- 
cies are yet recorded from Australia, and onlv one species. H. 
proli.xus, is known from New Zealand. 

Subgenus Hapalips \cnsit str'n to 

Description. With the general characters of Hapalips. Form 
more convex, less parallel-sided; elytra with scutellarv striole; wing 
(Fig. 18) with anal cell and radial cell with spur of Rs. Head 
sometimes with a pair o( stridulatoiy tiles; a transverse groove pres- 
ent on gular region | Fig. 20); eyes large, facets moderately coarse. 
Pronotum more coin ex. length and breadth more or less equal. 
Mctasternum not elongate, more or less equal in length and 
breadth. 

This subgenus includes the majority of the species. 

( \\ OPHORUS subgenus novum 
Type species. Hapalips fuscus Reitter 

Description. With the general characters oi Hapalips. General 
appearance as in Hapalips sensu stricto. Head with vestiges of 
Stridulatory tiles, represented by two longitudinal ridges without 
striatum (Fig. 23). On the gular region, in place of the usual 
transverse line, there is a large trilobed cavity (Fig. 23). opening 
ventrally and protected by hairs. Eyes large, as in Hapalips sensu 
stricto. Elytra, wing venation, shape of pronotum and mctasternum 
as in Hapalips sensu strict". 

According to present knowledge, this subgenus includes only 
two Central American species, H. fuscus Reitter and //. acaciae 
sp. n. 

Xenosceloides subgenus novum 
Type species. Hapalips proli.xus Sharp 

Description. With the general characters of Hapalips. Form 
more flattened and parallel-sided. Wing without anal ceil (1 
16), radial cell sometimes without spur of Rs (Fig. 17). Elytra 



1968 GENERA OF LOBERINI 9 

often without stridulatory files; gular region with a transverse 
groove as in Hapalips sensu stricto. 

This subgenus includes Hapalips prolixus Sharp, H. fdum Reit- 
ter, H. scotti Grouvelle, and possibly H. taprobanae Grouvelle. I 
have not been able to check in detail the characters of H. taproba- 
nae, which is less flattened than the other species; if it truly 
belongs in this subgenus, then Grouvelle's name Loberina will 
have priority over Xenosceloides. 

Key to the Subgenera of Hapalips 

1. Anterior part of gular region with a large cavity (Fig. 23) . .Cavophorus 
Anterior part of gular region without cavity but with a transverse 

groove (Fig. 20) 2 

2. Wing with anal cell (Fig. 18); body shape more convex and less parallel 

sided; elytra with scutellary striole. Prothorax and metasternum 
more or less equal in length and breadth. Stridulatory files on head 

sometimes present; eyes large with moderately coarse facets 

Hapalips 

Wing without anal cell (Fig. 16); body shape more flattened and parallel 
sided; elytra often without scutellary striole. Prosternum (Fig. 24) and 
metasternum longer than broad. Stridulatory files on head absent; 
eyes smaller with finer facets Xenosceloides 

Hapalips acaciae species novum 

Holotype, 9 , and para type $ , Cotaxtla Exp. Sta., Veracruz, 
MEXICO, VIII- 15-1 962, D. H. Janzen coll., seed pods of Acacia 
cornigera, deposited in the Museum of Comparative Zoology, 
Cambridge, Mass. (holotype MCZ No. 31604). Paratype (on 
slide), same data, deposited in the Department of Zoology, The 
University, Glasgow, Scotland. 

Measurements of holotype. Total length: 5.58 mm; width of 
head across eyes: 1.18 mm; length of antenna: 1.26 mm; length 
of prothorax along midline: 1.23 mm, width across middle: 1.40 
mm; length of elytra: 3.60 mm, width across middle: 1.45 mm. 

Description. With the general characters of Loberini, Hapalips 
(Cavophorus). Larger and more elongate than H. fuscus Reitter; 
upper surface uniformly reddish brown. Head (Fig. 23) trans- 
verse; eyes large and rather coarsely faceted. Vestiges of stridula- 
tory files represented by a pair of widely separated longitudinal 
ridges without transverse striations. Occipital region glabrous, an- 
terior half of dorsal side of head sparsely clothed with recumbent 
setae. A large transverse trilobed cavity (Fig. 23) present on 
anterior part of gular region, opening ventrally and protected by 



10 breviora No. 303 

fine hairs. Antenna with scape about double the length of pedicel, 
which is shorter than segment 3. segments 4-8 equal in length. 
segments 9 and 10 roughly semicircular, segment 9 slightly less 
wide than segment 10. terminal segment asymmetrically rounded in 
apical part and longer than preceding two segments. Prothorax 
slightly wider than its length, side margins more or less parallel 
sided, bordered by fine hairs, posterior angles pointed and an- 
terior angles obtuse. Punctation on pronotum coarser and closer 
than on head, setae minute and directed toward center. Elytra 
uniformly reddish brown, stria! punctures in rows, interstices with 
sparse fine punctures, scutellarv striole rather irregular and indis- 
tinct, pubescence short and recumbent. Scutellum minute, nar- 
rowed in front and weakly angulate on posterior margin, pubescent. 
Wing with anal cell. Mesocoxal lines present but very short; 
mesepisternal pockets obscured. Tarsi densely hairy on underside, 
first three tarsal segments more or less equal in length, segment 5 
about as long as first three segments together. Femoral lines on 
ventrite 1 very short; intercoxal process narrow and pointed. 

Key to the Species of Cavophorus 

l. Species larger and more elongate, ahum 5.80 nun in length; prothorax 

more or less equal in length and hreadth. Femoral lines on ventrite 1 

very short //.(<-.) acaciae sp. n. 

Species smaller and less elongate, aboul 3.80 mm in length; prothorax 

distinct]) transverse. Femoral lines on ventrite l fairly long 

//. (<) fuscus Reittei 

Description of a Larva op Hapalips prolixus Sharp 

One larva recorded from New Zealand under the base of dead 
palm leaves {Rhopalostylis sapidd) along with the adults, by R. A. 

Crowson, deposited in the Glasgow University Zoolog\ Depart- 
ment. 

Total length 3.48 mm; length of head including labrum 0.64 
mm. width across the middle 0.56 mm; width of prothorax across 
the middle 0.64 mm; width of 9th abdominal segment across the 
front margin 0.48 mm. 

General appearance narrow, elongated, somewhat flattened. 
tapered in front and behind. Dorsal surfaces bearing many gran- 
ules symmetrically arranged in a definite pattern of lines, except 
on pronotum and head, wartlike setiferous tubercles on cither side 
of each tergite. setae near middle line minute and blunt, becoming 



1968 GENERA OF LOBERINI 11 

longer towards sides. Setae on ventral surface rather short, fine 
and pointed. 

Head rather elongate, shape of head and arrangement of setae 
on dorsal surface as figured (Fig. 27), dorsal surface granulated. 
Frontal suture rather indistinct, as figured (Fig. 27); endocarina 
absent. At the base on either side of occipital foramen there are 
three microscopic peglike setae. Ocelli not distinct. Antennae 
short, narrow, length of the segments 1:2:2, sensory appendage 
lying ventrally, % of the length of segment 3. Mandible (Fig. 
29) with two equal teeth, mola well developed bearing transverse 
ridges and asperites. Between the apical teeth and mola the pros- 
theca translucent and pointed at apex. Ventral crushing tubercle 
well developed; single sensory pit present in the middle of dorsal 
side of mandible, another one on ventral side anterior to dorsal 
one. Maxillary mala (Fig. 31) acute at apex, with three apical 
spines as in Hapalips sp. larva described by Rymer Roberts 
(1939), a row of seven setae present on dorsal side of inner mar- 
gin of mala; at the base of these setae on dorsal side a group of 
small denticles, similar denticles present at the base of palpiger 
(Fig. 31). Cardo rather long, narrow, at right angle to stipes; 
maxillary articulating area well developed and oval. Labium (Fig. 
32) with two jointed palpi, palpiger not distinguishable; ligula 
rather indistinct; hypopharynx with well developed hypopharyngeal 
bracon. 

Pronotum slightly wider than head; granulation on dorsal sur- 
face irregular. Meso- and metathorax slightly shorter and progres- 
sively wider than prothorax; granulation on dorsal side in a regular 
symmetrical pattern as in abdominal tergites (Fig. 33). Abdomi- 
nal segments 1-6 equal in length and breadth and equal to meta- 
sternum, segments 7-9 progressively narrower. Arrangement of 
setae and granulation are similar on meso- and metanotum and 
first 8 abdominal tergites, on tergite 9 granules and setae are ar- 
ranged as in Figure 33. Each segment with two transverse rows 
of four minute blunt setae; two pairs of tubercles on either side of 
each segment, anterior pair carrying two blunt and comparatively 
short setae, posterior pair with single long and pointed seta (Fig. 
33). Urogomphi well developed, as figured (Fig. 33), not hooked 
or upturned, projecting posteriorly. A pair of setiferous tubercles 
(pregomphal process) present anterior to urogomphi. Pygopod 
small, rounded, and not projecting. 

All spiracles are bicameral (Fig. 34), lying on body surface, 
lateral air tubes directed posterolaterally. Legs fairly long, coxae 
closely situated, claws simple with two tarsungular setae. 



12 breviora No. 303 

This larva may be distinguished from that of Hapalips sp. de- 
scribed by Rymer Roberts ( 1939) by the following key: 

Urogomphi long, not hooked (Fig. 33). Mandibular prostheca not 
narrow or hooked at apex (Fig. 29). Ocelli obscured. Tubercles 
on each segment with pointed and blunt setae (Fig. 33) .... 

H. prolixus Sharp 

Urogomphi short, hooked. Mandibular prostheca rather narrow and 
hooked at apex. Ocelli 5 posterior to antenna. Tubercles on each 

segment with single long pointed seta Hapalips sp. 

Discussion. Although all the species of Hapalips are restricted to 
the warmer climatic zones, the two larger subgenera are repre- 
sented in both the New and Old Worlds; even within these sub- 
genera there are no obvious general differences between the New 
and Old World forms. The subgenus Cavophorus, with only two 
known species, both restricted to the New World, appears to repre- 
sent a specialized development from Hapalips sensu stricto, at 
least in respect to the vestigial stridulatorv files and the big cavity 
in the anterior part o( the gular region. apparentl\ developed from 
the transverse groove which is present in other Hapalips in the 
same position. The subgenus Xenosceloides is another group 
which may represent a specialized off-shoot o[ Hapalips sensu 
stricto; the main differences between Xenosceloides and Hapalips 
sensu stricto parallel those between the genus Hemipeplus Berth 
and its relatives in the Mvclcridac ( Heteromera ). Typical Hemi- 
peplus spp. as far as known occur under leaf bases of palms, as do 
Xenosceloides adults. I hus the distinguishing features ol Xenosce- 
loides ma) be adaptive to this mode of life and may have originated 
independently in the Old and New Worlds, in which case Xenosce- 
loides would not be a natural subgenus. Further research will be 
needed to establish whether this is in fact the case. The only spe- 
cificallv identified larva of this genus is that of //. prolixus from 
New Zealand, which shows considerable differences, as well as 
similarities, when compared with the larva of Hapalips sp. de- 
scribed bv Rymer Roberts (1939) from Mexico; the differences 
are such that the two larvae would be expected to represent at 
least different subgenera. If the Mexican larva is really of a 
Hapalips sensu stricto, then larval characters support the separa- 
tion of Hapalips sensu stricto and Xenosceloides as subgenera; if, 
on the other hand, the Mexican larva proves to be of a Xenosce- 
loides, this will strongly support the theory that the New and Old 
World forms of the subgenus are not really related. 



1968 GENERA OF LOBERINI 13 

Genus Truquiella Champion 

Truquiella Champion, 1913: 87. Type species, by monotypy, Truquiella 
gibbifera Champion. 

This monotypic genus was established by Champion (1913) un- 
der Cryptophagidae; I have found no subsequent references to it. 
Champion described it as having tetramerous tarsi, but I found its 
tarsi to be pseudotetramerous, as in other Loberini. The genus 
seems to be very closely related to Hapalips. 

Description. With general characters of Loberinae, Loberini. 
General fades more or less as in Hapalips. Head with fairly large 
and coarsely faceted eyes. Stridulatory files apparently absent. A 
pair of protuberances present on anterodorsal side of eyes (Fig. 
3) in both sexes, more prominent in male; clypeus broad with 
rounded apical margin as in Hapalips; antennal insertions com- 
pletely hidden by frons. Antenna moderately long, with scape 
small but larger than pedicel, which is shorter than segment 3; 
segments 4-8 equal in length and shorter than segment 3; segments 
9 and 10 weakly transverse, terminal one elongate with rounded 
apex. Prothorax weakly transverse, side margins smooth, front 
and hind margins more or less equal in breadth, front angles 
slightly projecting forward and weakly acute, hind angles obtuse. 
Prebasal impressions on pronotum obscured; front coxal cavities 
rather narrowly open behind; prosternal process narrow and 
truncated at apex. Elytra with regular rows of punctures, scutellary 
striole present. Wing with closed anal cell, venation as in Hapa- 
lips eichelbaumi Grouvelle (Fig. 18). Mesocoxae closely situated; 
mesepisternal pockets weakly developed; metasternum weakly 
transverse; median impressed line extending % of its length. 
Tarsi with first three segments equal in length, segment 3 lobed 
below, segment 4 minute and segment 5 about as long as first four 
together; tibiae weakly broadened at apex with two normal spurs. 
Intercoxal process of ventrite 1 narrow and pointed at apex. 

Habitat. Unknown, larva undescribed. 

Geographical distribution. Mexico. 

Genus Pseudhapalips Champion 

Pseudhapalips Champion, 1913: 112. Type species, by monotypy, Pseud- 
hapalips lamellifer Champion. 

Champion (1913) established this genus under Cryptophagidae, 
and described it as closely related to Hapalips, although he noted 
several dissimilarities from Hapalips, e.g. the extraordinary form 



14 BREVIORA No. 303 

of head, very prominent eyes and thorax, as in Platoberus, etc. 
He described the front coxal cavities as closed behind, but careful 
study of a slide preparation reveals that these cavities are distinctly 
open behind ( 1 ig. 6). Arrow (1929a) described Pseudhapalips 
as having stridulatory files on the head, but neither I nor Mr. R. D. 
Pope of the British Museum could find stridulatory tiles on the 
head of P. lamellijer. It seems that the genus may be related to 
Hapalips, as suggested by Champion and Arrow, but not very 
closely. Since Champion's description only one species has been 
added to this genus, and that was by Grouvelle (1919) from 
French Guiana. 

Description. With general characters of Loberinae, Loberini. 
I lead transverse, in male with a strong ridge between the eves; in 
female this ridge is less distinct and Battened in front; ebpeus 
broad with rounded apical margin. Eyes large, markedly project- 
ing and moderatel) coarsely faceted. Antenna moderately long 
with scape slightly longer than pedicel, which is slightly shorter 
than segment 3. segments 3-8 equal in length, club loose, segment 
9 very slightly wider than segment 10. terminal segment rather 
transverse and rounded at ape\. 

Prothorax (Fig. 6) strongly transverse, slightly narrowed be- 
hind, side margins weakl) undulate or dentate, front angles slightly 
projecting and obtuse, hind angles acute. Prebasal impressions on 
pronotum strongl\ marked; prosternal process weakly broadened 
posteriori) with straight apical margin (Fig. 6). Elytra glabrous, 
strial punctures in regular rows and with a scutellary striole. Wing 
as figured (Fig. 15), without anal cell. Mesepistcrnal pockets 
weakly developed; metasternum elongate and median impressed 
line extending to % of its length. Tarsi are unlike Hapalips, first 
three segments lobed below, segment 5 equal to length of first two 
segments together; tibiae broadened at apex. \ entrite 1 with inter- 
coxal process narrow and pointed at apex 

Habitat. Unknown, larva undescribed. 

Geographical distribution. Amazon. Surinam, and French 
Guiana. 

Genus Pseudhenoticus Sharp 

Pseudhenoticus Slmrp, 1900: 596. Type species, by monotypy, Pseudheno- 
ticus parallelus Sharp. 

Sharp ( 1900) established this genus under Cryptophagidae and 
placed it just before Henoticus, Grouvelle (1919) added 10 spe- 
cies to it, and more recently. Bruce (1943) described another 



1968 GENERA OF LOBERINI 15 

species from Madagascar. From Bruce's figure and description it 
seems very doubtful whether this specimen is a true Pseudhenoticus 
or even a member of Loberini; if it is a Pseudhenoticus, then the 
genus will manifest a distribution pattern unusual in Clavicornia, 
though it is known in some other organisms. 

Description. With general characters of Loberiinae, Loberini. 
Head transverse, clypeus less broad than in Hapalips, broad at 
base, narrowed in front, its apical margin rounded. Unlike Hapa- 
lips, frons scarcely projecting over antennal insertions. Stridulatory 
files apparently absent; eyes of moderate size and very finely 
faceted. Antenna rather short and stout, scape, pedicel and seg- 
ment 3 more or less equal in length, segments 4-8 slightly shorter 
than segment 3 and equal in size, segments 9 and 10 somewhat 
semicircular, and terminal segment elongated, about double the 
length of segment 10. Prothorax (Fig. 5) transverse, weakly nar- 
rowed in front, side margins undulate or more or less dentate, pre- 
basal impressions on pronotum present. Front coxal cavities 
rather narrowly open behind; prosternal process broad at apex as in 
Bolerus (cf. Fig. 2). Elytra with regular rows of punctures and 
without scutellary striole, pubescent. Meso- and metacoxae mod- 
erately widely separated, sternal fitting between the mesocoxae in 
a straight line. Metasternum transverse, narrowed in front, median 
impressed line extending % of its length. First three tarsal seg- 
ments more or less equal in length, segment 2 weakly and segment 
3 strongly lobed below and narrow, segment 5 almost equal in 
length to first three segments together; tibiae slightly broadened 
at apex. Ventrite 1 with intercoxal process moderately broad and 
its apical margin rounded. 

Habitat. Unknown, larva undescribed. 

Geographical distribution. Panama, Bolivia,? Madagascar. 

Genus Bolerus Grouvelle 

Bolerus Grouvelle, 1919: 93. Type species, by present designation, Crotchia 

minuta Fleutiaux, 1887: 68. 
Thallisellodes Arrow, 1925: 257. Type species, by original designation, 

Thallis transversus Gorham, 1895: 325. 
Platycladoxena Kraatz (partim), — Arrow, 1929a: 316. 
Crotchia Fowler (partim), — Fleutiaux, 1887: 68. 

The nomenclature of this genus is very complicated. The genus 
Platycladoxena (type P. castanea Kraatz) was established by 
Kraatz (1899), in the Languriidae. Arrow (1925) synonymized 
it with Microlanguria Lewis and erected a new genus Thallisellodes 
with four species. He later (1929a) stated that P. castanea and 



16 BREVIORA No. 303 

P . javanica of Kraatz are congeneric with Thallisellodes and sank 
the latter name. In the same paper, he listed 13 valid species in 
this genus. Grouvelle (1919) described the genus Bolerus, which 
he considered to be related to Hapalips and Loberus, and trans- 
ferred Crotchia minuta Fleuiiaux to the genus Boletus. Arrow 
( 1929a) pointed out that the species of Bolerus are congeneric 
with Thallisellodes. \ illiers ( 1961 ). having seen the type material 
of Platycladoxena castanea Kraal/, stated that the two specimens 
represented two distinct species, both of the genus Microlanguria; 
the lectotype should be named Microlanguria castanea (Kraatz), 
while the other specimen was described as M. angulosa Villiers. 
The above facts indicate that the genus Bolerus Grouvelle ( 1919) 
must have priority over Thallisellodes Arrow (1925). Unfortu- 
nately. Grouvelle did not specify a type o\ his genus Bolerus; there- 
fore I here designate Crotchia minuta Fleutiaux as type species. 
I Ins is one of the species originally included b\ (irouvelle ( 1919). 

The species Bolerus minimis, which 1 have studied in detail, 
seems to be \er\ similar to Thallisellodes angulosus Arrow and T. 
transversus (Gorham). I have found its characters more similar 
to those of the Loberini than to those of Cladoxenini (e.g. front 
coxal cavities internally open behind and ovipositor like other 
Loberini with st\li attached at the apex of coxites). Rymer Rob- 
erts (1939) described the larva of Bolerus ( -.Platycladoxena) 
angulosus Arrow and stated that it is very similar to the larva of 
Hapalips. The present study supports the view of Grouvelle and 
Rymer Roberts and shows that the genus is actually related to 
Hapalips and probabl) Pseudhenoticus. 

Description. With the general characters of Loberinae, Lobe- 
rini. General appearance less linear than in Microlanguria and 
prothorax Strongl) transverse. Head transverse, with a pair of 
closely situated stndulatory files; eyes fairly large and moderately 
coarsely faceted. Clypeus broad at base and narrowed in front, 
with straight or weakly rounded apical margin; antennal insertions 
less hidden under Irons than in Hapalips. Transverse groove on 
anterior gular region weak but distinguishable. Antenna with 
scape and segment 3 longer than pedicel, club rather compact and 
segments 9. 10. and 1 1 transverse and equal in length, apical seg- 
ment rounded at apex and slightly less transverse than segment 10, 
sometimes club loose and apical segment elongate and pointed at 
apex. Prothorax (Fig. 2) parallel sided, side margins more or less 
smooth or finely dentate, front angles rather rounded, hind angles 
somewhat acute, prebasal impressions on pronotum strongly 



1968 GENERA OF LOBERINI 17 

marked. Front coxal cavities rather narrowly open behind, pro- 
sternal process as figured (Fig. 2), sometimes sinuate at apical 
margin. Elytra glabrous, strial punctures in regular rows and with 
scutellary stride. Wing as figured (Fig. 19), with an anal cell. 
Mesocoxae moderately widely separated; mesepisternal pockets 
well developed; metasternum weakly transverse; mesocoxal lines 
short; median impressed line extending about % of its length. 
Metendosternite broad and short, as figured (Fig. 21). Tarsi as 
in Hapalips; tibiae broad at apex and obliquely truncated. Ventrite 
1 with intercoxal process slightly broad, shape as figured (Fig. 1); 
sometimes a second pair of lines present on outer sides of the nor- 
mal femora] lines (Fig. 1 ). 

Habitat. Adults and larvae of B. angulosus have been found on 
a lichen-covered rock in a cave in Malaya. 

Geographical distribution. Indo-Malayan region. 

Key to the Genera of the Tribe Loberini 

1. Trochanters short and broad; tarsi simple, except in Xenoscelis, where 

segment 3 is very slightly lobed below. Femoral lines on first ventrite 
absent; anterior part of gular region without transverse groove. Front 
coxal cavities usually narrowly open, rarely completely closed behind 
{Xenoscelis) Tribe Pharaxonothini 

 — trochanters broadly elongate (Fig. 26); tarsi distinctly lobed below, with 
minute segment 4. Femoral lines on first ventrite present; anterior 
part of gular region with a transverse groove (Fig. 20) or a cavity 
(Fig. 23). Front coxal cavities usually fairly widely, more rarely 
narrowly open behind Tribe Loberini .... 2 

2. Mesocoxae more widely separated and sternal fitting between them in a 

straight line. Prothorax (Fig. 5) with anterior part of side margins 
undulated. Intercoxal process of first ventrite broad with more or less 
rounded apical margin Pseudhenoticus 

— Mesocoxae more closely situated and sternal fitting between them with 
single knob (Fig. 9). Prothorax not as above. Intercoxal process of 
first ventrite narrow and pointed at apex (except in Bolerus) 3 

3. Prothorax as figured (Fig. 2), with prosternal process broad at apex. 

Elytra glabrous; mesocoxal line on metasternum present. Intercoxal 
process of first ventrite broad, as figured (Fig. 1 ) Bolerus 

— • Prothorax with prosternal process not as above. Elytra usually hairy; 
mesocoxal lines on metasternum absent. Intercoxal process of first 
ventrite narrow and pointed 4 



18 breviora No. 303 

4. Species larger, narrow and elongated, less Cryptophagidae-like in form. 

Elytra with scutellary striole (except in Hapalips scotti Grouvelle); 
uing often with anal cell (Fig. 18). Tibiae broad and truncate at 
apex; tarsal lobes broad. Metastcrnum less transverse 5 

— Species smaller, more elliptical and Cryptophagidae-like in form. Elytra 

without scutellary striole; wing without anal cell. Tibiae slender, not 
broadened at apex; tarsal lobes narrow. Metasternum more trans- 
verse 7 

5. Prothorax as figured (Fig. 6). side margins dentate or undulate, front 

angles projecting forward. Tarsal segments 2 and 3 lobed below. 
Anterior part of dorsal side of the head with transverse ridge .... 
Pseudhapalips 

— Prothorax not as above, side margin not dentate or undulate, front 

angles not projecting forward. Only tarsal segment 3 lobed below. 
WuAd not as abo\e 6 

I- Head with a pair of humps on anlerodorsal sides of eves dig. 3). Pre- 
basal impressions on pronotum indistinct. I lytra pubescent. Truquiella 

— Head without humps .is above. Prebasal impressions on pronotum 

iisualh distinct Elytra glabrous or pubescent Hapalips 

1. Prothorax narrowed in front, shape as figured (Fig. 7). Antenna with 
segmenl 9 considerably smaller than segment 10, which is markedly 
broad Telmatoscius 

— Prothorax not narrowed in front, shape as figured (big. 7). Antennal 

segment l > \er\ little smaller than segment 10. which is less trans- 
verse Loberus 



ACKNOWLEDGMENTS 

This research was carried out in the Department of Zoology, 
University of Glasgow. I am grateful to Prof. D. R. Newth for 
laboratory facilities. I am deeply indebted to Dr. R. A. Crowson 
of Glasgow University for his valuable advice and also for the 
larva oi Hapalips prolixus, and to Dr. J. F. Lawrence of the Mu- 
seum of Comparative Zoolog) for reading the manuscript and for 
the adults of Hapalips acaciae. I would like to thank the stall oi 
the Coleoptera section of the British Museum (Natural History), 
particularly Mr. R. D. Pope, who have made available to me the 
facilities to study material in the Museum collection. 



1968 GENERA OF LOBERINI 19 

LITERATURE CITED 

Arrow, G. J. 

1925. Fauna of British India, including Ceylon and Burma. Coleop- 
tera: Clavicornia: Erotylidae, Languriidae and Endomychidae. 
London, Taylor and Francis, viii + 416 pp. 

1927. Clavicornia and Lamellicornia. In: Insects of Samoa. Part 4, 
Coleoptera. Brit. Mus. (Nat. Hist.), pp. 35-66. 

1929a. On the families of Coleoptera related to the Erotylidae, with 
descriptions of a new family, two new genera and a few new 
species. Ann. Mag. Nat. Hist., (10) 4: 305-322. 

1929b. A revision of the African Coleoptera belonging to the family 
Languriidae. Proc. Zool. Soc. London, 1929, Part 1: 1-15. 
Batra, L. R. 

1963. Ecology of ambrosia fungi and their dissemination by beetles. 
Trans. Kansas Acad. Sci., 66(2): 213-235. 
Bruce, N. 

1940. Coleoptera — Cryptophagidae von Juan Fernandez. In: C. 
Skotsberg, ed., The Natural History of Juan Fernandez and 
Easter Island, Vol. Ill, Zoology: 682-688. 

1943. Drei neue tropische Cryptophagiden nebst Bemerkungen zur 
Synonymie der Cryptophagiden Familie (Coleoptera: Crypto- 
phagidae). Arb. Morph. Taxon. Ent. Berlin-Dahlem, 10: 56- 
61. 

1951. Cryptophagidae (Coleoptera: Polyphaga). Explor. Pare Nat. 
Albert. Mission G. F. De Witte (1933-1935). Fasc. 75, 26 pp. 

1952. Revision der im Deutschen Entomologischen Institut befindli- 
chen Hapalips-Arten. Beitr. Z. Ent., 2: 461-473. 

1963. Coleoptera-Cryptophagidae in Musee Royal de l'Afrique Cen- 
tral (V). Rev. Zool. Bot. Afr., 47: 203-220. 
Champion, G. C. 

1913. Notes on various Central American Coleoptera with descriptions 
of new genera and species. Trans. Ent. Soc. London, 1913, 
Parti: 87-113. 
Crowson, R. A. 

1955. The Natural Classification of the Families of Coleoptera. Lon- 
don, Nathaniel Lloyd, 187 pp. 
Fleutiaux, E. 

1887. Descriptions des coleopteres nouveaux de l'Annam. . . . Ann. 
Soc. Ent. France, Ser. 6, 7: 59-68. 
Fowler, W. W. 

1908. Languriinae. In: P. Wytsman, Genera Insectorum, 78: 1-45. 
Gorham, H. S. 

1895. List of the Coleoptera in the collection of H. E. Andrews, Esq., 
from India and Burma, with descriptions of new species and 
notes. Families: Malacodermata — Erotylidae — Endomy- 
chidae. Ann. Soc. Ent. Belgique, 39: 293-330. 



20 hreviora No. 303 

1898. Coccinellidae (part) and supplement to Erotylidae (part). In: 
F. Godman and O. Salvin, eds., Biologia Centrali-Americana. 
Insects. Coleoptera. Vol. 7, pp. 241-256. 

Gro< Mill. V 

l l >14. The Percy Sladen Trust Expedition to the Indian Ocean. 
Coleoptera: Cucujidae, Cryptophagidae. Trans. I. inn. Soc. 
London: Zool., 17: 141-159.' 
1919. Descriptions de genres et d'especes nouvelles de Cryptophagidae. 
In: A. Grouvelle. Memoires Entomologiques. Etudes sur les 
Coleopteres. Fasc. 2. pp. 70-203. 
Groi Mill. A., AND A. RAFFRAY 

1912. Supplement a la liste des Coleopteres de la Guadeloupe. Ann. 
Soc. 1 m. France, 81: 289-312. 

HlNTON, H. E. 

1945. A Monograph of the Beetles Associated with Stored Products. 
I ondon, British Museum (Natural History), vii f 443 pp. 
Kraatz, ( i. 

1899, Ueber die Languriiden Arten von Kamerun nebst einigen ver- 
wandten l ormea Deutsch. I at Zeit, 1899: 307-313. 

1 ECONTE, J. 1 . 

1861. Classification of the Coleoptera of North America. Part I. 
i 1st Part > Smithsonian Misc. ( oil., ix\ + 208 pp. 

1863 New Bpecies of North American Coleoptera. Part I. Smith- 
sonian Misc. c oil.. No. 167. pp. 1-86. 

I i NDBI BO, S. 

1966. Eicolyctus brunneua GylL (< oleoptera), nagot om hi. a. hiolo- 
gin. Int. ridskr., 87: 47-49. 

\1 \RIIM .'. V. \M> A. H\RKI R\ 

1966. Hallazgo de ( ryptophagidae anoftalmos y aptcros. associados 

a mamiferos. ( iencia, 25(1): 11-16. 
Mora in i sky. V. 

IS' ai dun catalogue des insectes de l'lle Ccylan. Bull. Soc. 

Imp. Nat. Moscou, 36 (1): 421-532. 

Rl I 1 1 1 R. I 

IS77. Hapalips, neue Gattung der Rhizophagidae. Verhandl. Naturf. 
Ver. Brunn. IS: 122-128. 
R\ mi R Robi are, A. w. 

1939. On the taxonomy of Erotylidae (Coleoptera). with special rcf- 
ence to the morphological characters of larvae. Trans. Roy. 
Ent Soc. London, 88: 89-118. 
Ri mi R ROBI Ris, \. W.. \nd F. Van Emden 

1958. On the taxonomy of the Erotylidae (Coleoptera I. with special 
reference to the morphological characters of the larvae. Trans. 
Roy. Ent. Soc. London. 110: 245-285. 
SCHENtXINO, S. 

1923 < ryptophagidae. In: W. Junk and S. Schenkling, eds.. Coleop- 
terorum Catalogus. Vol. XV. Pars 76, 92 pp. 



1968 GENERA OF LOBERINI 21 

1928. Languriidae. In: W. Junk and S. Schenkling, eds., Coleop- 
terorum Catalogus, Vol. XV, Pars 10, 40 pp. 
Sen Gupta, T. 

1967. A new subfamily to Languriidae based on 4 genera, with a 
key to the species of Toramus. Proc. Roy. Ent. Soc. London, 
Ser. B, 36 (11-12): 167-176. 
Sen Gupta, T., and R. A. Crowson 

1967. The systematic position of Eicolyctus Sahlberg (Coleoptera: 
Languriidae). Proc. Roy. Ent. Soc. London, Ser. B, 36 (5-6): 
87-93. 
Sharp, D. 

1900. Fam. Cryptophagidae. In: F. D. Godman and O. Salvin, eds., 
Biologia Centrali-Americana, Insecta, Coleoptera, Vol. II, Part 
1: 579-626. 

VlLLIERS, A. 

1942. Notes sur quelques Cladoxenitae Indo-Malais (Coleoptera: 
Erotylidae). Arb. Morph. Taxon. Ent. Berlin-Dahlem, 9: 
89-92. 

1943. Etude morphologique et biologique des Languriidae (Col. 
Erotylidae). Publ. Mus. Nat. Hist. Natur., No. 6, 98 pp. 

1961. Revision des Coleopteres Languriides Africains. Ann. Mus. 
Roy. Afr. Cent., Ser. 8°, Sci. Zool., No. 98, 385 pp. 

(Received 19 November 1967.) 



EXPLANATION OF LETTERING ON FIGURES 

ac — anal cell mt — metasternal knob or 

as — apical spine projection 

at — anterior tendon my — pocket or mycangium ( ? ) 

cc — mesepisternal pocket p — paramere 

cp — corpotentorium pa — peglike setae 

ct — coxite Pp — paraproct 

es2 — mesepisternum pr — prostheca 

ga — galea s2 — mesosternum 

hr — hypopharyngeal bracon s3 — metasternum 

hu — hump si — styli 

la — lacinia si — stridulatory file 

lp — lateral plate su — supratentorium 

Is — longitudinal line on tc — trochanter 

metasternum tg — transverse groove on anterior 
It — laminatentorium part of gular region 

m — mola ur — urogomphi 

ml — median lobe vf — valvifer 

mp — mesosternal pocket vp — sensory appendage 

ms — median strut vt — ventral crushing tubercle 



22 



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Mm senium of Comparative Zoology 

Cambridge, Mass. 31 December, 1968 Number 304 

REDESCRIPTIONS OF ANACHIS AVARA (SAY) AND 

ANACHIS TRANSURATA (RAVENED WITH NOTES 

ON SOME RELATED SPECIES 

(PROSOBRANCHIA, COLUMBELLIDAE) 1 

Amelie H. Scheltema- 



ABSTRACT 

Both Anachis avara (Say) and A. translirata (Ravenel) are 
found from Massachusetts to southern Florida. The shell of avara 
varies geographically from short and broad with many fine ribs in 
the north to tall and slender with a few heavy ribs in the south. 
A. translirata does not vary in shell morphology over its range. In 
New England the two species are very commonly found together 
at mean low water and have usually been confused with each other 
because of a superficial similarity in ribbing on the shell. Egg 
capsules and veliger larvae of northern and southern populations 
of avara are morphologically similar. Egg capsules, larvae, and 
radulae of translirata are different from those of avara. 

In southwestern Florida, avara is replaced by A. semiplicata 
Stearns, which appears to be a species endemic to that area. The 
relationships of an Anachis species in the western Gulf of Mexico 
are not clear. Egg capsules of A . floriclana Rehder show it to be a 
species distinct from avara. A. similis (Ravenel) is considered to 
be a nomen dubium. 



1 Contribution No. 1888 from the Woods Hole Oceanographic Institution, 
Woods Hole, Massachusetts. 

- C/o Woods Hole Oceanographic Institution, Woods Hole. Massachu- 
setts 02543. 



BRIVIORA NO. 304 



INTRODUCTION 



[wo very common gastropod species living along the shores oi 
southern New England arc the columbellids Ainu his avara (Say, 
1S22) and A. translirata (Ravenel, ISM). The shells and living 
animals of both arc quite similar, and the two species have a long 
histor) of being confused with each other. South of New England 
misidentifications have been fewer tor two reasons: first, translirata 
is much less common near the tidchne from New Jerse) south ( see 
Table I ): second, avara is highly variable in shell form, and those 
from southern localities are less similar to translirata than those 
from New England. 

Gould ( 1841 ). in Ins earl) work on New England invertebrates, 
recognized onl) one species, avara. Verrill ( 1873) noted correctl) 
that another species oi Anachis occurs in New England, which he 

called A. similh (Ravenel. 1861). llowe\er. I regard similh a 
nomen dubium (sec p. 12) and translirata the correct designation. 
\ errill figured similh with a shell of avara reproduced from Gould 
I 1870). From VerrhTs work until the present time, "avara" has 
been the name more USUall) applied to both species; when trans- 
lirata has been recognized .is a distinct form, the names "similis" 
and "translirata" have been used indiscriminately, often as varieties 

or subspecies ol avara (e.g.. Dall, 1889; Johnson. 1934). Abbott 
(1954) gave correct names and descriptions of the two species. 
but unfortunate!) the plate figures arc reversed and the geographic 
ranges are not correct. 

Recently, Scheltema and Scheltema (1963) described the egg 

capsules and \ehger larvae o\' translirata as those of avara. be- 
cause the adults from which egg capsules were obtained were com- 
pared with misidentified museum specimens. The error became 
evident when new ke\s to invertebrates in the Woods Hole region 
were being compiled (Smith, ed., 1964). Subsequently, avara were 
collected alive in Beaufort, North Carolina, and held in aquaria; 
these snails deposited egg capsules that were distinctly different 
from those of translirata. This fortunate circumstance made pos- 
sible the identification o\ similar capsules often found attached to 
eel >:rass m the Woods Mole area, and consequently larvae have 
been reared from both Beaufort and Woods Hole populations of 
avara. I hese larvae were morphologically the same, and distinct!) 
different from those of translirata in their soft external anatomy 
(Scheltema. MS in preparation). I hereforc. on the basis of cap- 
sules and larvae, it has been \erilied that the two New Faigland 
forms oi Anachis are distinct species, and also that the southern 
and northern forms of avara constitute a single species. 



1968 REDESCRIPTION OF ANACHIS SPECIES 3 

The necessity for redescribing the two species and for determin- 
ing their geographic ranges became obvious if common errors in 
identification were to be corrected. 

ACKNOWLEDGMENTS 

Material has been made available to me through the generosity 
of many individuals and museums. I would like particularly to 
thank Emma B. Richardson of the Charleston Museum, Joseph 
Rosewater and George Radwin of the U.S. National Museum 
(USNM), R. Tucker Abbott and Robert Robertson of the Acad- 
emy of Natural Sciences of Philadelphia (ANSP), and William J. 
Clench of the Museum of Comparative Zoology (MCZ). Robert 
H. Parker, Jack B. Pearce, and Edmund H. Smith, formerly of the 
Systematics-Ecology Program, Marine Biological Laboratory, 
Woods Hole, kindly allowed me to make use of their Anachis col- 
lections. I am extremely grateful to Louise Ridge for her many 
efforts and hours of collecting to provide me with live specimens 
of Anachis floridana, which she mailed from Florida. John R. Hall 
and Robert Robertson made the photographs of the shells on 
Plate I, for which I am very appreciative. Thanks are due also to 
Ruth D. Turner and Kenneth J. Boss, who gave this paper a criti- 
cal reading and made several helpful comments. 

Finally, I take pleasure in acknowledging the financial support 
of The Radcliffe Institute, a grant from which has made this work 
possible. 

Anachis translirata (Ravenel) 
Plate 1, figs. 1, 2, 6; Text-figs, lb, 2b; Table 1 

Columbella translirata Ravenel 1861, Proc. Acad. Nat. Sci. Philadelphia 

1861: 42 (no fig.) ("off Charleston bar" [South Carolina]). [Type 

specimen probably destroyed.] 3 
Columbella avara Say. Gould 1841, Invertebrata of Massachusetts, pp. 

313-314 (in part), fig. 197; Tryon 1883, Manual of Conchology, 5: 

159 (in part). 



3 Much of Dr. Ravenel's collection, which lay in the line of General 
Sherman's march through South Carolina in 1865, was destroyed; no labeled 
specimens of either A. translirata or A. similis remain in his collection at 
the Charleston Museum. Ravenel's description of translirata, although not 
figured, is sufficiently precise to enable one to ascertain the species to which 
he was referring. 



UR1 VIORA 



No. 304 



Inachis avara (Say). Perkins 1869, Proc. Boston Soc. Nat. Hist. 13: 113 
(in part); Sumner. Osburn. and Cole 1913, Bull. U.S. Bur. Fish. 
31(2): 710-711. 712 (in part): M. Smith 1945, last (oast Marine 
Shells, p. [19, pi. 46. fig. 24: Abbott 1954, American Seashells, pi. 25, 
lii:. ee (fig. only); Scheltema and Scheltema l l >63. Hydrobiologia 
22(1-2): 85-91, 13 text-figs, (egg capsules and larvae). 

Anachis similis (Ravenel). Verrill 1873, Kept. IS. Comm. 1 ish. and Fisher- 
ies 1871-1872: 644-645 (description only, not fig. I<> 1 >>. 

Anachis avara translirata (Ravenel). Dall 1889, Bull. U.S. Nat. Mus. 
.'{7: 116. 

Anachis avara similis (Ravenel). Dall 1889, Bull. U.S. Nat. Mus. :*7: 116; 
M. Smith 1945, last Coasl Marine Shells, p. ||>). p|. 4(>. fig. I l > (juve- 
nile). 




h-J 





t=3 




rext-Fig. I. Radular teeth of (a) Anachis avara and (b) Amu his transli- 
rata. Scale line equals 0.2 mm. 



Description. Shell elevated, conical, whorls llattcned and regu- 
larl\ ribbed. Height to 17.5 mm. width to 6.5 mm. median range 
in height 12.5 to 14.0 mm. Average width to length ratio 0.40. 
Aperture a little less than one-half total height of shell. Larval 
whorls (protoconch) three, smooth, rounded, and translucent. 



1968 



REDESCRIPTION OF ANACHIS SPECIES 



Post-larval whorls seven or eight, nearly flat, with distinct, regu- 
larly spaced, straight axial ribs and less distinct spiral striae. 
Suture indented. Ribs usually parallel to shell axis, and most ele- 
vated at the suture. Those on the spire extend from suture to 
suture and are about the same width as the interspaces. Ribs on 
the body whorl average 15 (range 11 to 20), are not as closely 
spaced as those on upper whorls, and extend from the suture to 
just below the periphery, except that those near the aperture extend 
only to the periphery. Periphery somewhat angulated. Faint axial 
ridges sometimes apparent below periphery, extending between 
ends of ribs of body whorl. Outer lip often thickened so as to oblit- 
erate one or more ribs. Spiral striae usually distinct between ribs. 




Text-Fig. 2. 



Egg capsules of (a) Anachis avara, (b) Anachis translirata, 
and (c) Anachis floridana. Scale line equals 1.5 mm. 



6 breviora No. 304 

The first spiral stria following the suture is the widest, uually the 
most deeply impressed, and crosses the ribs, producing the subsut- 
ural nodules that are the distinctive shell character of the species. 
Other striae may or may not cross ribs, giving a more or less can- 
cellate appearance: the first two post-larval whorls in particular 
are often beaded. Spiral striae least impressed at periphery. Below 
periphery, spiral striae are deeply impressed and form the dominant 
shell sculpture; they produce weak beading as they cross the axial 
ribs and ridges. Aperture elliptical. Outer lip thin or thick, re- 
curved anteriorly to form a short siphonal canal. When the lip 
is thickened it is recurved posteriorly into an anal notch. Teeth 
may be present on outer lip when it is thickened, and range from 
three to nine (average 6) in number, often indistinct, the largest 
posterior. Parietal callus, when present, is narrow, sharp-edged, 
and bears a number of teeth, which are the ends of the spiral striae. 
Columella is smooth. Shell is dull, color varying from straw-yellow 
to chestnut-brown, often with spiral streaks o\' white, especially at 
subsutural knobs and angulation o\ periphery, emphasizing these 
characters. Juveniles are without thickened lip or callus, and with 
the periphery sharpl) angulated. 

Operculum elliptical, concentric, with eccentric nucleus. 

The radula is rachiglossan; the median tooth is a flat plate; the 
lateral tooth has three cusps, with the proximal one rounded (Text- 
fig, lb) 

" Pigmentation oi head and loot similar to that of A. avara ((/.v. ). 
except that the white posterodorsal tip o\' the foot is small and 
grades into the mottling of the rest oi the loot. 

Egg capsule volcano-shaped, ringed b\ sculpture of concentric 
ridizes, with one ridge more pronounced than the rest (Text- 
fig.^2b ) . 

Remarks. The shell of A. translirata is differentiated from that 
of A. avara by its usually more flattened whorls and greater height, 
and by the spiral subsutural stria that crosses the axial ribs and 
is the widest stria. (This stria ma\ not be evident on all whorls, 
especially on an eroded or fouled specimen.) In avara, the spiral 
striae are usually not as pronounced and do not cross the ribs. 
The ribs of the first two post-larval whorls of avara are therefore 
smooth, whereas those of translirata are beaded. Juveniles of the 
two species are easily distinguished by the difference in width- 
length ratio, which is less variable in juveniles and therefore more 
obvious, and by the shape of the body whorl, which is sharply 
angulated in translirata but rounded in avara (PL 1. figs. 5, 6). 
The posterodorsal tip of the foot and the egg case of each species 



1968 REDESCRIPTION OF ANACHIS SPECIES 7 

also serve to differentiate them. The lateral teeth of the radulae 
have one small consistent difference between the two species: in 
avara the proximal cusp bears a denticle; in translirata it is 
rounded (Text-figs, la, b). 

The Pliocene fossil figured as Anachis (Costoanachis) avara 
translirata (Ravenel) by Gardner (1948, pi. 30, figs. 36, 37) is 
probably not this species. 

Habitat and distribution. This species lives on shelly bottoms, 
rocks, pilings, or other firm substratum, from below mean low 
water to 48 fathoms, from Cape Cod Bay and Georges Bank south 
to Florida (see Table 1). It is very common subtidally in New 
England, where it can also be found in beds of eel grass with A. 
avara. 

Specimens examined. The entire collections of translirata of 
the MCZ and the USNM were examined. Shell measurements and 
descriptions are based on specimens from the following localities: 
Nobska Point, Woods Hole, Mass. 

(Scheltema collection) 19 (living) 

Quicks Hole, Mass. 

(Scheltema collection) 54 (29 living) 

Sullivans Island, S.C. (MCZ 260887) 34 

Hadley Harbor, Mass. (R. Parker collection) 30 
Wellfleet, Mass. (MCZ 19372) 32 

Beaufort, N.C. (MCZ 256848) 1 (living) 

Radulae were examined from four of the specimens taken from 
Quicks Hole. Descriptions of the pigmentation of the head-foot 
are based on the 1 9 specimens from Nobska Point. 

Anachis avara (Say) 

Plate 1, figs. 3-5, 7-10; Text-figs, la, 2a, 3 and 4; Table 1 

Colombella [sic] avara Say 1822, Jour. Acad. Nat. Sci. Philadelphia 2: 230 
(no fig.) ("coast of the southern states"). [Lectotype, here selected, 
Academy of Natural Sciences of Philadelphia No. 16887, ex Mrs. 
Say's collection, "Florida."] Non C. avara Duclos 1840, Histoire 
Naturelle Coquilles Univalves Marines, pi. 1, figs. 1, 2. 

Columbella avara Say. Gould 1841, Invertebrata of Massachusetts, pp. 
313-314 (in part, not fig. 197); DeKay 1843, Natural History of New 
York 5, Mollusca: 139-140, pi. 8, fig. 179. 

Amycla (Astyris) avara (Say). H. and A. Adams 1858, Genera of Recent 
Molluscs 1: 187. 

Anachis avara (Say). Perkins 1869, Proc. Boston Soc. Nat. Hist. 1869- 
1871, 13: 113 (in part). 



8 



uriviora 



No. 304 



Anachis avara "Perkins.' Verrill 1873. Rcpi. U.S. Comm. Fish and Fish- 

eries 1 8 7 1 - 1 S ~ 2 : 643-644 (no fig.). 
Anachis similis (Ravenel). Verrill 1873. Kept. U.S. Comm. Fish and 
Fisheries 1871-1872: pi. 21. fig. 109 (fig. only, not description pp. 

644-645). 



25 



I II II I ! II II [I II I 



N = 50 

WOODS HOLE, MASS- 

R.PARKER COLL. 




25 



-I — n — n — n — n — n — n 




N=62 

BEACH HAVEN, N.J. 

ANSP 71291 




N=27 

BEAUFORT, N.C. 
MCZ 81921 



ii ii — 1 1 1 1 i 



A/--/8 
* BEAUFORT, N.C. 
MCZ 256849 



i — n — n — n — n — n — n — i 



25 



.111.1 



A/-"/ 7 

CHARLSTON.S.C 
MCZ 256850 



i — n — ii ii — i i i i — n — r-n — i — n — n — i — i 



25 r 




N=103 

LAKE WORTH, FLA. 

ANSP 221 188 



h — n — r I 1 I — n — n — rn 

CO O CM « 

V <n <n m 



WIDTH/LENGTH x 100 



rext-Fig. 3. Frequency distribution of width to length ratios of Anachis 
i. vara from selected localities between Massachusetts and Honda (expressed 
in percentage). N is sample size. Larvae were reared from the Beaufort, 
North Carolina, population indicated by an asterisk. 



1968 REDESCRIPTION OF ANACHIS SPECIES 9 

Anachis avara translirata (Ravenel). M. Smith 1945, East Coast Marine 

Shells, p. 119, pi. 46. fig. 23. 
Anachis translirata (Ravenel). Abbott 1954, American Seashells, pi. 25, 

fig. ff (fig. only). 

Description. Shell variable, elevated, conical, whorls slightly 
ventncose and ribbed. Height to 14.5 mm, width to 6.3 mm, me- 
dian range in height 10.5 to 12.0 mm. Width-length ratio ranges 
from 0.37 to 0.56; the more slender shells (ratio less than 0.42) 
occur in populations south of Cape Hatteras (PI. 1, figs. 7, 8, 9). 
Aperture about one-half total height of shell, slightly less in slender 
shells, slightly more in broad shells. Larval whorls (protoconch) 
three, smooth, rounded, and translucent. Post-larval whorls six 
to eight, with variable sculpture. Suture indented. Axial ribs on 
first two or three post-larval whorls; very rarely absent. Succeed- 
ing whorls ribbed (PI. 1, fig. 3) or smooth (PI. 1, figs. 4, 7, 8, 9). 
Ribs present on body whorls straight or curved, highest at the 
periphery; however, those near the lip disappear above the periph- 
ery. Outer lip often thickened so as to obliterate one or more ribs. 
Ribs on body whorl range in number from 7 to 21; populations 
north of Cape Hatteras range from 10 to 21 (PI. 1, figs. 4, 10), 
those south of Cape Hatteras from 7 to 14 (PI. 1, figs. 7, 8, 9). 
Ribs heaviest on shells from southern populations. Spiral striae 
faint to strong, but not crossing ribs, and strongly impressed below 
periphery. Aperture elliptical, wider in northern populations than 
in southern. Outer lip thin or thick, recurved anteriorly to form a 
short, slightly recurved siphonal canal; when lip is thickened, it is 
recurved posteriorly into an anal notch. Teeth often present on 
outer lip if it is thickened, and range from 4 to 12 (average 8 or 9) 
in number. Largest tooth usually penultimate one before anal 
notch. Parietal callus, when present, is narrow and sharp-edged, 
and bears a number of teeth, which are the ends of spiral striae. 
The columella is smooth. Juveniles (PI. 1, fig. 5) without thick- 
ened lip or parietal callus, body whorl with or without ribs. Shell 
color varying from straw-yellow to chestnut-brown, and usually 
with conspicuous white round or elliptical mottlings, although 
these may be lacking (PI. 1, fig. 4) in some populations; ribs of 
body whorl often tipped with white at suture. Shell may be dull or 
lustrous. 

Operculum elliptical, concentric, with eccentric nucleus. 

The radula is rachiglossan; the median tooth is a flat plate; the 
lateral tooth has three cusps, the proximal one ending in a sharp 
denticle (Text-fig. la). 



10 breviora No. 304 

Head and foot are mottled black and white. Tentacle and siphon 
tips are white, and usually have a black stripe around them. The 
anterodorsal part of the foot has two large white spots; the postero- 
dorsal tip of the foot is white, and is sharply demarcated from the 
mottling o( the rest of the foot. 

Egg capsule volcano-shaped, sculptured by fine striae running 
from base to apex (Text-tig. 2a ). 

Remarks. (See also remarks under A. translirata.) Northern 
and southern populations of A, avara arc sufficientl) different (PI. 
I. figs. 4. 7) in shell character to warrant investigating the possi- 
bility of the existence of subspecies. A number of populations 
from throughout the range of avara (listed below under Specimens 
examined) were examined in detail to determine whether there is 
any basis for differentiating subspecies b\ shell characters alone, 
lengths and widths of about 440 shells were measured, and ribs 
on the body whorl and teeth on the outer lip were counted. Shell 
color, thickness, luster, and number of post-larval whorls were 
noted. Width to length ratios and rib numbers show an essentially 
clinal change north to south, from short, broad shells with many 
ribs t*.i tall, slender ones with few ribs ( Text-figs. 3, 4). The more 
slender shells may have an extra whorl. The numbers of teeth seem 
to bear no relationship to other shell characters; for example, the 
range is 4 to 12 in the population from Iladlev Harbor. Massachu- 
setts, and 7 to 11 in one from Beaufort. North Carolina. Shells 
with hitih luster are in collections from Delaware Bay, Chinco- 
league Bay, and Lake Worth. The thinnest, darkest colored shells 
occur from New Jersey northward. As in many shelled mollusks 
of the eastern United States, the largest shells are from Chinco- 
teague Bay. 

These data on shell characters do not provide sufficient evidence 
for differentiating subspecies. Also, the length of larval life (up to 
5 weeks in the laboratory) is long enough for the larvae to be dis- 
persed oxer considerable distances, making genetic interchange 
likely between local populations. That the slenderest, heaviest 
shells and fewest specimens are from South Carolina. Georgia, and 
Florida (see Table 1) may be a reflection of greater isolation of 
populations, for suitable habitats (see below) may be separated 
by vast stretches of sand banks and large backwater areas with 
black, sulfurous muds bordered by marsh grass. 

Two shells from southern localities (the paralectotype from 
"Florida" [PI. 1, fig. 3] and a single shell from Sullivan's Island. 
South Carolina) have the shape and rib count of northern shells. 
They may have been collected from the ocean side of the outer 



REDESCRIPTION OF ANACHIS SPECIES 



11 




Text-Fig. 4. Range in rib number on body whorl of Anachis avara from 
selected localities between Massachusetts and Florida. Means are indicated 
by horizontal lines; sample sizes by numbers under the vertical bars. Those 
individuals with body whorl incompletely ribbed are not included. Letters 
above bars refer to locality: A, Woods Hole, Mass. (MCZ 203663); B-F 
as in Text-Fig. 3: B, Beach Haven; C and D, Beaufort; E, Charleston; 
F, Lake Worth. 



sand banks, rather than from the embayments from which most of 
the avara collections south of Cape Hatteras have been made. 

Habitat and distribution. A. avara is found chiefly in eel grass; 
it is also common below mean low water on a variety of firm sub- 
strata, often with translirata. It is distributed along the entire 
coast from Massachusetts Bay and Nantucket Island to Lower 
Matecumbe Key, Florida (Table 1). It is chiefly subtidal, but 
has been taken from the following depths: Massachusetts Bay, 5 
fathoms; Chesapeake Bay, 25 fathoms; off Cape Charles, 10 fath- 
oms; off Beaufort, N.C., 9 fathoms; off Cape Kennedy (Canav- 
eral), 10 fathoms. 

Specimens examined. The entire collections of avara in the 
MCZ, USNM, and ANSP were examined. The descriptions are 
based on specimens from the following localities: 

Gunning Point, Woods Hole, Mass. 

(Scheltema collection) 12 (living) 



12 HRKVIORA No. 304 

Nobska Point, Woods Hole. Mass. 

(Scheltema collection) 2 (living) 

Beaufort, N.C. (MCZ 256849) 20 (living) 
Hadley Harbor. Woods Hole. Mass. 

( R. Parker collection ) 50 

Woods Hole, Mass. ( MCZ 203663 ) 24 

Beach Haven, N.J. I WSP 71291) 85 

Barnegat Bay, N.J. (ANSP 106708) 23 

Delaware Bay, Cape May. N.J. (ANSP I 82645 ) 8 

Delaware Bay, Little Creek. Del. (MCZ 198032) 13 

Chincoteague Bay, Va. (MCZ 197795) 6 

Beaufort. N.C. (MCZ SI 921) 27 

Beaufort, N.C. (ANSP 145680) 21 

Charleston. S.C. (MCZ 256850) 17 

St. Augustine, Fla. (USNM 416015) 17 

St. Augustine, Fla. I WSP 140800) 8 

Pake Worth. I la. (ANSP 221 188) 107 
Radulae of three specimens from Gunning Point, two from 
Beaufort, N.C, and one from Lake Worth. Fla. (ANSP 221 188), 

were examined. 

NOTES ON RELATED SPECIES 

I he following remarks on some other western .Atlantic species 
oi Anachis are offered as notes. Anatomical studies, such as 
those ol Marcus and Marcus (1962, I 964 ) on Brazilian Colum- 
bellidae, are greatly needed for species along the entire coast and 
Gulf of Mexico. 

Anachis simii is I Ravenel I 

Columbella similis Ravenel 1X61. Proc. Acad. Nat. Sci. Philadelphia 
1861: 41-42 (no liu. ) ("common on the coasts of North and South 
( arolina" ). 

No type material has been found (see footnote 3 under A. trans- 
lirata), and it is impossible from the description to know precisely 
to what species Ravenel was referring. The description fits juve- 
nile A. avara as well as a number of small species of Columbelli- 
dae. Probably the small, ribbed columbellid common in shallow 
water in western Florida should not be referred to A. .similis (e.g., 
Perry and Schwengel, 1955). Cardner (1948) has figured a fossil 
A. avara similis (Ravenel) that is too large to lit Ravenel's de- 
scription. 

Anachis similis (Ravenel) should be considered a nomen (la- 
bium. 



1968 REDESCRIPTION OF ANACHIS SPECIES 13 

Anachis semiplicata Stearns 
Plate 1 , figure 1 3 

Anachis semiplicata Stearns 1873, Proc. Acad. Nat. Sci. Philadelphia 1873; 
344-347, 4 text-figs, (west coast of Florida); Perry and Schwengel 
1955. Marine Shells of the Western Coast of Florida, p. 159, pi. 51, 
fig. 345 (egg capsules); iwn avara semiplicata 'Stearns' of various au- 
thors (western and northern Gulf of Mexico). 
This appears to be a valid species that is endemic to southwest- 
ern Florida. The shell is higher and narrower (average width to 
length ratio 0.36, 20 specimens from ANSP 221 189) than that of 
avara and either lacks spiral striae or has only very faint ones; 
the egg capsules, as described by Perry and Schwengel (1955), 
seem to be different from those of avara, although unfortunately 
the figure is not clear. In addition, the juvenile shells are more 
elongate than those of juvenile avara. The radula has not been 
studied. 

The northern and western Gulf of Mexico species of Anachis 
found on jetties, pilings, and oyster beds (PI. 1, fig. 12) does not 
appear to be A. semiplicata (e.g., PufTer and Emerson. 1953; 
Pulley, 1952). It most closely resembles Beaufort, N.C., popula- 
tions of avara, but is smaller (median range in length, 9.1-10.0 
mm, 29 specimens from USNM 606003). The lateral radular 
tooth (1 specimen examined) is the same as that of avara. The 
status of this species is not at all clear; it may possibly be a relic 
population (see Deevey, 1950) of avara now differentiated at 
either a specific or subspecific level, or it may have affinities with 
populations of Anachis sp. to the south (see Weisbord, 1962: 313- 
315). Abbott (1954) has included both this species and A. 
semiplicata in his range for avara. 

Anachis floridana Rehder 
Plate 1 , figure 1 1 ; Text-figure 2c 

Anachis floridana Rehder 1939, Nautilus 53 ( 1 ) : 20, pi. 6, fig. 6 (near Cape 
Canaveral, Brevard County, Florida, in 30 feet of water; USNM 
473202). 
This species differs from A. avara in having a smooth rather 
than an indented suture and flattened whorls lacking spiral striae. 
Egg capsules obtained from animals held in the laboratory most 
closely resemble those of A. brasiliana (Marcus and Marcus, 
1962). The lateral radular tooth (1 specimen examined) is simi- 
lar to that of A. translirata and lacks the pointed denticle on the 
proximal cusp of avara. 



14 brlviora No. 304 

REFERENCES CITED 

Abbott. R. T. 

1954. American Seashells. New York, xiv + 541 pp. 

Dall, W. H. 

1 SS9. A preliminary catalogue of the shell-bearing marine mollusks 
and brachiopods. Bull. U.S. Nat. Mus., 37: 1-221. 

Dlevey, E. S. 

1950. Hydroids from Louisiana and Texas, with remarks on the 
Pleistocene biogeography of the western Gulf of Mexico. 
Ecology, 31: 334-367. 

Gardni r. J. 

1948. Mollusca from the Miocene and Lower Pliocene of Virginia 
and North Carolina. Part 2. Scaphopoda and Gastropoda. 
U.S. Geol. Surv. Prof. Pap. 1 99-11 : 179-310. 

Gould, A. A. 

IS41 \ Report on the Inverlebrata of Massachusetts. . . . Cambridge. 

M.iss., 373 pp. 
1870. Report of the Invertebrata of Massachusetts. 2nd ed. (W. G. 
Binney, ed.). Boston, v + 524 pp. 

Johnson, C. W. 

1934. List of marine Mollusca of the Atlantic coast from Labrador 
to texas. Proc. Boston Soc. Nat. Sci. Hist., 40 ( 1 ) : 1-204. 

Maro s. i . and I Marcus 

1962. Studies on Columbellidae. Bol. Fac. Filos. Cien. Let., Univ. 
Sao Paulo, 261 (Zool. 24 | 335-384. 

1964. On the dove-shell Anachis pulchella (Blainv.). An. Acad. 
Bras.l. Cien., 36 (3): 359-366. 

Pi kki . L. M.. and G. S. Sc nv% i NG1 i 

1955. Marine Shells of the Western Coast of Florida. Ithaca, NY., 
318 pp. 

I'i i i l k. 1 1 ... and W. K. 1 mi RSON 

1953. The molluscan community of the oyster-reef biotope on the 
central Texas coast. Jour. Paleont.. 27 (4):537-544. 

Pi LLF.Y, T. E. 

1952. An illustrated check list of the marine mollusks of Texas. 
Texas Jour. Sci., 4: 167-199. 

SCHELTfcMA, R. S.. and A. H. SCHELTEMA 

1963. Pelagic larvae of New England intertidal gastropods II. 
Anachis avara. Hydrobiologia, 22 (1-2) : 85-91 . 

Smith, R. I. (ed.) 

1964. Keys to marine invertebrates of the Woods Hole region. 
Contrib. No. 11, Systematics-Ecology Prog., Mar. Biol. Lab., 
Woods Hole. 208 pp. 



1968 REDESCRIPTION OF ANACHIS SPECIES 15 

Verrill, A. E. 

1873. Report on the condition of the sea fisheries of the south coast 
of New England in 1871-1872. XVIII. Report upon the inver- 
tebrate animals of Vineyard Sound and the adjacent waters, 
with an account of the physical characters of the region. 
Rept. U.S. Comm. Fish and Fisheries, 1871-1872: 295-778. 

Weisbord, N. E. 

1962. Late Cenozoic gastropods from northern Venezuela. Bull. 
Amer. Paleont., 42 (193): 1-604. 



(Received 2 February 1968.) 



TABLE 1 



Distributions of Anachis avara (Say) and Anachis translirata 

(Ravenel) based upon collections of the U.S. National Museum 

(U), Museum of Comparative Zoology (M), Academy of Natural 

Sciences of Philadelphia (A), and the author's (S).* 

Anachis Anachis translirata 
Locality or Region Collection avara Inshore Offshore 

? Maine — Grand Manan 



45 fms 



Island 


U 


+ 


+ 


F Georges Bank ( 1 juvenile, 








dredged ) 


M 






assachusetts 
Boston 


A 


+ 




Cape Cod Bay — Duxbury 


M, U 


+ (.U) 


+ (M) 


off Wellfleet (dredged) 


M 


5 fms 


5 fms 


Cape Cod, all shores 








(numerous localities) 


U, M, S 


+ 


+ 


Nantucket (several 








localities) 


U, M 


+ 


+ 


Martha's Vineyard 








(several localities) 


U, M 


+ 


+ 


off Gay Head (dredged) 


U 




+ 


Vineyard Sound (dredged) 


U 


+ 


+ 


Buzzards Bay (dredged) 


U 


7 fms 


7 fms 


Shores of Buzzards Bay 








(numerous localities) 


U, M. S 


+ 


+ 


Elizabeth Is. (dredged) 


S 


+ 


+ 



* Specimens from inshore that were obviously dead when collected are not 
included. A few "dead" or "drilled" specimens from offshore, so noted, are 
listed. Anachis avara has been collected only from inshore localities. 



16 


BREVIORA 


No. 304 






Anachis 


Anachis t runs li rata 


Locality or Region Collection 


avara 


Inshore Offshore 


Rhode Island 








Sakonnett Pi. 


\1 


+ 


+ 


Narragansett Baj (dredged, 








several localities) 


U 




8-19 fms 


Bristol 


M 


+ 


+ 


Westerly 


U, M 


+ 


+ 


Connecticut and New York 








Stonington 


U 




+ 


off Fisher's Is. (dredged) 


M 




+ 


long Island — Montauk Pt. 


U 


+ 


+ 


Green port 


U 


+ 


+ 


Peconic Bay ( ? dredged I 


u 




+ 


Long Island Sound 








( ? dredged ) 


u 




+ 


New Haven 


u 


+ 


+ 


New Jerse\ and Delaware 








Atlantic ( ii\ 


u 


+ 




Spraj Peach 


\l 


+ 




Barnegal Ba) 


u. \ 


+ 




Beach Haven 


\ 


+ 




( ape \la\ 


u 




+ 


off (ape \la\ ( dredged ) 


u 




+ 


Delaware Baj ( ape Ma) 


A 


+ 




1 ittle (reek 


\l 


+ 




Mai j land and Virginia 








( bincoteague 


U, M. \ 


+ 




Smith Is ( Va.) 


U 


+ 




Isaacs 


u 


+ 




off Cape Charles 


u 


10 fms 




off Cape Henry 


u 


4': fms 




Chesapeake Bay (dredged. 








several localities) 


u 


4-25 fms 


1 2 fms 


35 mi. east of Wallops Is. 








(shell drilled) 


M 




18 fms 



North Carolina 

48 mi. East of Currituck Sd 

(•dead" shell) M 

23 mi. east of Currituck Sd. 

Cdead" shell) M 

27 mi. east of Pamlico Sd. M 

off Cape Hatteras U 

Cape Hatteras Pt. U 



+ 



18 fms 

18 fms 

24 fms 
13, 16. 48 fms 



968 REDESCRIPTION OF ANACHIS SPECIES 17 







Anachis 


Anachis 


translirata 


Locality or Region 


Collection 


a vara 


Inshore 


Offshore 


Cape Lookout 


U 


+ 






Shackleford Is., Sound shore 


: M 


+ 






Beaufort region (several 










localities) 


M, U, A. S 


+ 


+ 




off New River 


U 


+ 


+ 




South Carolina 










Pawleys Is. 


M 


+ 






Winyah Bay (mouth) 


U 


+ 






Sullivans Is. 


M 


+ 


+ 




Charleston 


M 


+ 


+ 




Cooper R. 


M 


+ 






Ashley R. (dredged) 


M 


+ 






Beaufort 


U 


+ 






off Beaufort 


U 


6-9 fms 






Georgia 










Tybee 


u 


+ 






St. Simons 


u 


+ 






Florida 










St. Augustine 


A, U 


+ 






off Cape Canaveral 










(Kennedy) 


U 


10 fms 






Lake Worth 


A 


+ 






off Lake Worth (drilled 










shells) 


M 






83-92 fms 


Waveland, Dade Co. 


U 


+ 


+ 




Lower Matecumbe Key 


U 


+ 






15-35 mi. off Ft. Walton 


M 






13-19 fms 



18 breviora No. 304 



Plate i 

Figs. 1 and ~. Anachis translirata (Ravencl), Sullivans Island. South 
Carolina. MCZ 260887. 

Fig. 3: Anachis avara (Say), paralcctotype, "Florida." ANSP 306058. 
Note the similarity to the northern form shown in Fig. 4. 

Fig. 4: A. avara, WeUfleet, Massachusetts. MCZ 19372. 

Fig. 5: A. avara, juvenile, Woods Hole, Massachusetts. 

Fig. 6: A. translirata. juvenile. Woods Mole. Massachusetts 

Fig. 1: A. avara. Fake Worth. Florida. ANSP 221 188. 

FlOS. 8 and 9: AlUtt his avara ( Sa\ ). leetotype, "Florida." ANSP 16887. 

Fig. in I. avara, ( hincoteague, Virginia. MCZ 197795. 

Fig. 1 1 : Anachis floridana Rehder, Matanzas Inlet, Florida. 

Fig. 12: Anachis sp.. Aransas Bay. Texas. USNM 606003. 

Fig. 13: Anachis temiplicata Stearns, Venice Bay, Florida. ANSP 221189. 



1968 



REDESCRIPTION OF ANACHIS SPECIES 



19 




BREVIORA 

Mm sen Jim of Comparative Zoology 

Cambridge, Mass. 31 December. 1968 Number 305 



LYTECHINUS WILUAMSI, 
A NEW SEA URCHIN FROM PANAMA 

Richard H. Chesher 1 

ABSTRACT 

A fourth species of Lytechinus from the Atlantic, L. williamsi, is 
described as a Panama endemic. It differs from the other shallow- 
water species, L. variegatus, in having fewer than 15 wedges per 
primary spine and fewer than 14 interambulacral plates per series 
in adults. Variation in several taxonomic features of L. variegatus 
is examined. 

INTRODUCTION 

During an underwater survey of the Atlantic coral reefs of 
Panama in September of 1967, a new species of sea urchin was 
discovered living in beds of leaf coral (Agaricia agaricites (Lin- 
naeus) ). The specimens, which were collected at Buena Ventura in 
depths of 5 to 10 meters, belong to Lytechinus, primarily an 
American genus, which now has four Atlantic and four Pacific 
species (see key, below). Two of the Pacific species are sympa- 
tic: L. pictus (Newport Bay, California, to Gulf of California) 
and L. anamensus (St. Barbara to Cedros Island, California). 
Mayr (1954) omitted L. pictus (Verrill 1867) in his zoogeo- 
graphic discussion of the genus Lytechinus as a possible synonym 
of L. anamensus Clark 1912. Examination of specimens in the 
Museum of Comparative Zoology substantiates Mortensen's 
(1943) view that the two forms represent valid species. In the 
Atlantic, L. callipeplus (Carribean and Gulf of Guinea) is sym- 
patic with L. euerces (Caribbean) in depths of 100 to 500 meters. 
L. williamsi (Panama) inhabits a very small portion of the geo- 
graphic range of the shallow-water species L. variegatus (Brazil to 

1 University of Guam, Agana. Guam. 



2 BREVIORA No. 305 

Bermuda in the Western Atlantic and Cape Verde Islands in the 
I astern Atlantic). Mortcnscn ( 1943) separted L. variegatus into 
several allopatric subspecies (see ke\. below), which were cited 
by Mayr I 1954) as evidence for geographic speciation. The dis- 
tribution of species indicates that tropical America has been the 
major center of speciation for this genus and that recent migrations, 
possibl) through the Equatorial Undercurrent, are responsible for 
the presence of /.. callipeplus and /.. variegatus in the Eastern 
Atlantic (C neshcr 1966). 

ro provide statistical data for comparison of the new species 
with L. variegatus, specimens of the latter were measured from 
localities ranging from Brazil to Bermuda (Table 3). The sub- 
specific taxa were ignored, and it was found that the characters 
that were measured showed \cr\ little variation over the entire 
geographic range ( [able 2. Fig. 5). Swan (l l ^52. 1958) found 
considerable \ariation in the number of wedges seen in cross- 
sections of spines from Strongylocentrotus spp.. but primar) spines 
taken from the mterambulaeral areas above the ambitus of L. 
variegatus showed very little variabilis throughout the geographic 
range of the species I he number of wedges per spine ma\ thus be 
considered a valid taxonomic character for this genus, providing 
that only primar) mterambulaeral spines from above the ambitus 
are used and the wedges are counted in the proximal half of the 
spine. The spines afford a field character for separation of speci- 
mens of /.. variegatus and /.. williamsi; spmes of the former ap- 
pear smooth and glistening and o\ the latter distinctl) ridged. 

ABBREVIATIONS 

I lie following abbreviations are used in this paper: III), hori- 
zontal diameter of the test; I A. mtcrambulacrum; Ln, the natural 
logarithm; S.I)., standard deviation; C.V., coefficient ot variation; 
r. coefficient of correlation; S^ x standard error of Y on the \ axis 
(Croxton 1959). In the formulas. X HD in mm and Y the 
taxonomic features of the test. M.C.Z.. Museum of Comparative 
Zoology; U.S.N. M. I nited States National Museum. 

ACKNOWLEDGMENTS 

The species is named after L. T. Williams, who provided his 
boat and hospitality for the investigation of the echinoid fauna of 
the Atlantic reefs of Panama and later found additional specimens 
for examination. I would like to thank him and his wife for their 
help. K. Waterhouse deserves special thanks for his assistance 



1968 LYTECHINUS WILLIAMSI 3 

during diving operations. Dr. I. Rubinoff and the staff of the 
Smithsonian Tropical Research Institute were of great assistance 
during this project, which was supported by travel funds from the 
Smithsonian Institution. The Mollusk Department of the M.C.Z. 
provided travel funds for Mr. Waterhouse. The analyses and 
manuscript were completed during a National Science Foundation 
Postdoctoral Fellowship at Harvard University. Statistical data 
were calculated on the 940 computer at Harvard University with 
support from a Milton Fund grant to Dr. H. B. Fell. Dr. Kenneth 
H. Boss kindly read the manuscript, which has benefited from his 
suggestions. 

Lytechinus williamsi new species 
Figures 1, 2, 3a, c-i, 4, 5; Table 1 

Types. The holotype (19mm HD, preserved in alcohol) is 
deposited in the U.S.N.M. (No. El 0855), as are 14 paratypes 
(U.S.N.M. No. El 0856). Eight paratypes are deposited in the 
M.C.Z. (Nos. 8 194 and 8 195). 

Type locality. Buena Ventura, Panama, on top of a living coral 
reef 5 to 10 meters deep, 9°32'N, 79°42'W. 

Material examined. Six specimens, 15.7 to 22 mm HD, seaward 
side of coral reef at Buena Ventura cove, Panama, 9°32'N, 
79°42'W, 5 to 10 meters, Sept. 4, 1967, Chesher Coll. Seventeen 
specimens, 16 to 27.9 mm HD, same locality, Dec. 1967, Williams 
Coll. 

Diagnosis. Number of wedges per primary IA spine from above 
the ambitus less than 15, milled ring of spines poorly developed; 
naked median areas near apical system in ambulacra and IA; 
globiferous pedicellariae purple; adults with less than 14 I A plates 
per series. 

Description. Generic characters conform to those given by Mor- 
tensen (1943). All specimens have well-developed genital pores 
and are less than 30 mm HD. 

Apical system. Each genital plate has one large primary tubercle 
and spine; secondary tuberculation poorly developed; all oculars 
exsert. The relative size of the apical system measured from the 
outer edge of ocular V to the outer edge of genital plate 2 is given 
in Table 1 . 

Ambulacra. Primary tubercles form a regular series just medial 
of the pore-pairs from the peristome to the oculars. Secondary 
tuberculation is poorly developed above the ambitus, resulting in a 
naked median zone from this point to the oculars. The number of 



4 BR] \ [OR \ No. 305 

ambulacra! plates per plate-series is indicated in Table I and 
Figure 5. Tube-feet end with a well-developed, calcitic sucking 

disc, as is normal for species of Lytechinus. Spicules of the tube- 
leet are ('-shaped ( Pig. 3h ). 

Inter ambulacra. Primary tubercles form a regular series from 
the peristome to the apical system in each row o{ plates, and pri- 
ma r\ spines are well developed up to and including the genital 
[Mates. Secondary tubercles are poorl) developed above the am- 
bitus, resulting in a naked median area i Fig. I ). I he number of 
I A plates is given in Table I and Figure 5. 

Spines. The sharpK pointed primal) spines arc about 40 to 50 
per cent of the HI) in length, with the exception oi the small spines 
adjacent to the peristome. The milled ring is poorl) developed and 
the base straight sided ( I ig. 3i I I he number of wedges seen in a 
cross-section of the proximal portion of primal") l\ spines is about 
12 I I ig. 3a). Near the peristome, the number ma\ decrease and 
the small oral spines ma\ have as few as 9 wedges. The number oi 
wedges was counted from 25<> I \ piuiiaiA spines taken from be- 
tween the ambitus and the apical system and broken in half about 
2 nun from the milled ring. About 10 spines were taken from each 
specimen. I he number ol wedges varied from II to 15. with a 
mean o\ 12.34 wedges (SI) 0.7683, ( V. 6.21). 

Pedicellariae. Globiferous, tridentate, ophicephalous, and tri- 

phyllous pedicellariae are present but not diagnostic (Fig. }u-\). 
The pedicellariae are purple, and the color is retained after clean- 
ing with bleach. Spicules o\ the globiferous pedicellariae are 
dumbbell-shaped (Fig. 3g). Large, white stalk glands are present 
on main globiferous pedicellariae 

Coloration. I he spines are USUall) deep, rich green, although 
two of the specimens have white spines. I he pedicellariae are 
purple, and tlie test is beige with a purple-brown stripe covering 
the median longitudinal suture o\ the ambulacra ami intcrambu- 
lacra. After cleaning with bleach, the test is mottled green-grey. 

Habitat. I he specimens were found living in niches on the top 
of a living, shallow coral reel at Buena Ventura, Panama. Most oi 
the specimens were associated with the leaf coral Agaricia agari- 
< itei ( Fig. 4). I he gut of one specimen was filled with small grey- 
green spheres, most o\ which were o\ unrecognizable origin. These 
urchins probably graze on algae and encrusting organisms of the 
reef. There was no evidence that the) were feeding on coral. Un- 
like L. variegatus (and most other toxopneustid echinoids), /.. 
willianiM docs not cover itself with bottom debris. 



1968 LYTECHINUS WILLIAMSI 5 

The specimens collected in December began to spawn when 
placed in alcohol, which might indicate a winter spawning season. 
Moore et al. ( 1963), however, found that L. variegatus may spawn 
to some degree during the entire year. 

Relationships. L. williamsi lives in the same depth range as L. 
variegatus and in a portion of its geographic distribution. Speci- 
mens of L. variegatus can be found in the grass areas at Buena 
Ventura. Although occasionally found on rocky areas or on soft 
sediments, L. variegatus normally occurs in grass areas (Moore 
et al. 1963; personal observations), while L. williamsi appears to 
be distributed only on living reef areas. 

The new species differs from L. variegatus in several features, 
the most obvious of which is the number of wedges per primary 
interambulacral spine, which averages 12.34 in the former and 
24.14 in the latter (Fig. 3b, Tables 1 and 2). The relative size of 
the apical system and peristome is greater in the new species than 
in L. variegatus. The number of plates per ambulacral or IA 
series is much greater in L. variegatus than in L. williamsi (Fig. 
5). Jackson (1912, 1914), after examining 1,443 specimens of 
L. variegatus, found none with all oculars exsert, whereas all of 
the specimens of the new species have every ocular exsert. 

L. williamsi can be separated from the two remaining, relatively 
deep-water, Atlantic species on the basis of the number of plates 
per series, the tuberculation, the coloring, and the spicules of the 
globiferous pedicellariae. L. euerces and L. callipeplus do not 
have naked areas in the median ambulacral and interambulacral 
area; the secondary tubercles are irregularly scattered on the test 
and are almost the same size as the poorly-developed primary 
tubercles. Whereas the spicules of the globiferous pedicellariae are 
dumbbell-shaped in L. williamsi and L. callipeplus, they are C- 
shaped in L. euerces. The naked test of L. callipeplus has red 
blotches on the dorsal side, which the new species lacks. 

L. williamsi is most closely allied to L. panamensis of the Bay of 
Panama. It differs from that species, however, in having fewer IA 
and ambulacral plates (Fig. 5). In addition, the pedicellariae of 
L. panamensis are brown, whereas they are purple in L. williamsi. 
Ocular I is normally insert in L. panamensis and exsert in the new 
species, and the milled ring of the primary spines is well developed 
in L. panamensis and almost absent in the new species. The other 
Pacific species are separated from the new species in the following 
key. 



6 BREVIORA No. 305 

KEY TO THE SPECIES OF THE GENUS 
LYTIC "7/ IN US 

1. Secondarj tuberculation poorl) developed, leaving a naked median area 

in the aboral portion of the ambulacra and interambulacra; primarj 
tubercles large 3 

Secondar\ tuberculation scattered irregularl) over the test, no naked 
areas, primarj tubercles small 2 

2. 1 arge, red blotches dorsall) on the cleaned test, spicules of globiferous 

pedicellariae dumbbell shaped (Caribbean, Gulf of Guinea, deep 
water) /.. callipeplus 

Test and spines white, spicules (shaped <( aiibbean. deep Water) .... 

/ . Cllt I i ( \ 

3. Spicules of globiferous pedicellariae ( -shaped (< alifornia) .../.. pictus 

Spicules dumbbell shaped 4 

4. Primar) I \ spines from above ambitus with less than 16 wedges .5 
Spines with more than 16 wedges 6 

5. Adults with less than 14 |\ plates pei senes. pedicellai iae purple (At- 

lantic coast of I'. m. una I L. williamsi 

VdultS with more than 14 I A plates pel series, pedicellariae brown 
i Ba) of Panama » / . panamensis 

6. Ambital spines much longer than other primaries, producing a fringe: 

test low. mottled with purple (California) /.. anamensus 

Spines more or less uniform in size 7 

7. Naked I A area granular, with a iow of small tubercles along the hori- 

zontal sutures, pedicellai iae purple, covering naked area (Galapagos, 
N.W. coast of South America ) /.. semituberculatus 

Naked I A area not pronounced, smooth, pedicellariae white, confined 
to ruberculated areas 8 

S. Test green, mottled with white, spines green or tinged with purple 
i Brazil, West Indies i /.. variegatus typicus 

I est not greenish or whitish 9 

l ». Buccal membrane not densel) plated outside buccal plates K ape Verde 

Islands. Africa ) /-. variegatus pallidas 

Membrane heavily plated 10 

10. Spines slender, dark purple (Bermuda) /.. variegatus atlanticus 

Spines reddish, stout ( U.S. east coast ) L. variegatus carolinus 



1968 



LYTECHINUS WILLIAMSI 



7 



TABLE 1 

L. williamsi: equations for regression lines from plots of taxonomic 
features of the test. 

Apical system ( V-2 ax's, in mm ) : 

LnY = 0.8162LnX - 0.7688 S v .x = 0.056 r = 0.89243 

Peristome (V-2 axis, in mm): 

LnY = 0.7306LnX + 0.0646 Sv.x = 0.025 r = 0.97049 

Interambulacral plates per series: 

LnY = 0.3832LnX+ 1.3117 Sv x = 0.039 r = 0.80026 

Ambulacral plates per series: 

LnY = 0.7321LnX + 0.5368 Sv x = 0.038 r = 0.93362 

Number of wedges per IA spine from above the ambitus (N = 250) : 
Mean = 12.34. S.D. = 0.7683, C.V. = 6.21 



TABLE 2 

L. variegatus: equations for regression lines from plots of taxo- 
nomic features of the test. 



Apical system ( V-2 axis, in mm ) : 
LnY = 0.7245LnX - 0.7219 

Peristome (V-2 axis, in mm) : 

I nY = 0.824;'Lnx - 0.3291 

Interambulacral plates per series: 
LnY - 0.3916LnX + 1.6275 

Ambulacral plates per series: 

LnY = 0.5397LnX + 1.4525 



Svx = 0.081 
Sv x = 0.043 
Svx = 0.042 
Svx = 0.037 



r = 0.97798 
r = 0.99107 

r = 0.97057 



0.99248 



Number of wedges per IA spine from above the ambitus (N = 500) 
Mean = 24". 14, S.D. = 1.7924, C.V. = 7.4 



8 BREVIORA No. 305 

TABLE 3 
Material of /-. variegatus examined. 
\4.CZ. No. I o< ality 
8179 Near Canal, Panama 

Buena \ entura, Panama 
1252 Bahia, Brazil 
3346 Bahia. Brazil 
7412 Pigeon Pi., [bbago 
8042 Bancs Bay, ( uba 
5070 lamaica 

Santa Marta, c olumbia 

7939 ( edai Key, N.W. I lorida 
4444 I oi tugas, I loi ida 
I 28 1 Beaufoi i. Not ih ( arolina 
3340 Bermuda 



Size i///> in mm) 


N, 


umber 


21 to 39.6 




3 


:: to 27 




2 


10 to 41.5 




5 


41.5 




1 


30 




1 


15 to ;; 




2 


26.5 




1 


34.3 




1 


16 to 30.2 




5 


5 5 to 63 




5 


21.5 




1 


26 




1 



LITERATURE CITED 

C III Mil K. R II 

1966. Report on the I chinoidea collected b) k \ I'll I SBURY in 
the Gulf of Guinea. Stud ["rop Oceanogi Miami I (1):209- 

223. 

( K(.\ k.n I I 

1959. I lementan Statistics with Applications in Medicine and the Bio 
logical Sciences. Ne\» York, Dover, 375 pp. 

J \( kvis. K.I. 

1^12. Phylogen) of the Echini with a revision ol Palaeozoic species. 

Mem. Boston Soc. Nat. Hist 7: 1-491. 
1914. Studies of Jamaica Echini. ( arnegie Inst. Wash. Pub. 182: 

139-162. 
\I\YR. E. 

1954. Geographic speciation in tropical echinoids. Evolution 8(1): 
1-18. 



1968 



LYTECHINUS WILLIAMSI 



Moore, H. B., T. Jutare, J. C. Bauer, J. A. Jones 

1963. The biology of Lytechinus variegatus. Bull. Mar. Sci. Gulf and 
Caribb. 13 (1 ):21-53. 

MORTENSEN, T. 

1943. A Monograph of the Echinoidea. 3(2). Copenhagen, C. A. 
Reitzel, 553 pp. 

Swan, E. 

1952. Regeneration of spines by sea urchins of the genus Strongylo- 
centrotus. Growth 16: 27-35. 

1958. Growth and variation in sea urchins of York, Maine. Jour. 
Mar. Res. 17: 505-522. 

(Received 21 March 1968.) 




y-.i f'\ ' \>|ff*^ s vim V M^VJjTi It 




9 / 

■*"• r 



•5T - 



Figure l. Dorsal view of L. williamsi (Paratype, 23 mm HD). 



10 



BR] V IORA 



No. 305 


















.V'y 



'X 






• •. V' 



* •«,* 



-.« 



sf**tf. 



»• 









T ' -St.  ' 



Figure2. Lateral ;irul ventral view of /. wiMiamsi (Paratype, 23 mm 

HDl. 



1968 



LYTECHINUS WILLIAMSI 



11 




Figure 3. Spines, spicules, and pedicellariae of L. williamsi'. 

A, cross-section of a primary interambulacral spine 0.7 mm in diameter; 

B, the same, from L. variegatus 0.65 mm in diameter; C, valve of a triden- 
tate pedicellaria 1.09 mm long; D, valve of globiferous pedicellaria 0.8 mm 
long; E, valve of triphyl ous pedicellaria 0.155 mm long; F, valve of ophice- 
phalous pedicellaria 0.55 mm long; G, spicules of globiferous pedicellaria 
0.04 mm long; H, spicules of the tube feet 0.04 to 0.047 mm long: I. base 
of a primary spine, 0.8 mm in diameter. 



12 



imi \ iora 



No. 305 




Figure 4. Underwater photograph of /.. wiiiwmsi at the base of some 
le .t coral [Agarh ia agarit ih 



1968 



LYTECHINUS WILLIAMSI 



13 



30- 



20- 



10- 



20- 



cv 



■10- 



% 



t 



•I 



o o 
oo 



o 



o ooo 



• = L. variegatus 
§=L. panamensis 
° L. will iamsi 



V • 

-9. 



§ . 



i * 



»° 



O 



10 20 30 

Horizontal Test Diameter 



-i r 

40 mm 



Figure 5. The number of plates in an ambulacral plate-series and an 
interambulacral plate-series for L. williamsi, L. variegatus, and L. pana- 
mensis. 



BREVIOIA 

Museum of Comnparatlve Zoology 

Cambridge, Mass. 31 December, 1968 Number 306 

A NEW SPECIES OF ELEUTHERODACTYLUS 

(AMPHIBIA, SALIENTIA) FROM THE GUAYANA REGION, 

EDO. BOLIVAR, VENEZUELA 

Juan A. Rivero 

Abstract. A new species of Eleutherodactylus, E. pulvinatus, is de- 
scribed from the wet region of La Escalera, Estado Bolivar, Venezuela. 
The species is characterized by its small but distinct tympanum, poorly 
defined or absent vomerine teeth, large, transversely oval disks which are 
larger than the tympanum, a W-shaped marking on the occiput and non- 
spiny nuptial pads. The possible relationship between this frog and the 
genera Eupsophus, Syrrhophus, Niceforonia, Trachyphrinus and Phryno- 
pus is discussed and evaluated. 



Among the specimens recently collected by the author in the 
region of La Escalera, Estado Bolivar, Venezuela, there are seven 
specimens of a small and peculiar leptodactylid whose generic allo- 
cation has been somewhat difficult and problematical. In this 
species, the vomerine odontoids are very indistinct; in some indi- 
viduals they appear to be absent altogether. As in Eleutherodac- 
tylus, the disks are well developed and the terminal phalanges are 
T-shaped, but there are two well-developed glandular pads on the 
inner side of the first digit, a character which does not seem to have 
been described in that genus. 

It has been customary to allocate all Eleutherodactylus-like. 
forms without vomerine teeth to the genus Syrrhophus (type local- 
ity, San Antonio, Texas) and all those with vomerine teeth but 
without disks to Eupsophus (type locality, Chile). In 1932, Parker 
violated convention by describing Eupsophus wettsteini (type local- 
ity, Pongo, Peru, 2000 m) as "the first species that has been de- 
scribed as belonging to the genus Eupsophus in which vomerine 
teeth are lacking ,, (p. 344). 

Even a superficial examination reveals that Andean species of 
Syrrhophus are not closely related to the animals that are referred 



2 BREVIORA No. 306 

to that genus in Central America and the United States. Whether or 
not they have vomerine teeth, most of the Andean animals seem to 
be closely related and should perhaps be called Eupsophus until a 
more thorough study can be undertaken and their relationships 
established. It is evident that the three groups of Eupsophus 
(Thoropa, wettsteini-peruanus, and the Chilean forms) mentioned 
by Cei ( I960. 1962) constitute different evolutionary lines, one of 
which {wettsteini-peruanus) appears to continue along the Andean 
Cordillera to Venezuela, where Eupsophus ginesi is known from 
glacial streams in the paramos oi the Merida Andes. It is now 
USUall) recognized that Thoropa is a separate genus, and perhaps 
the wettsteini-peruanus-ginesi group should be recognized as an- 
other, in which case. Oreobates Jimenez de la Espada, 1872. might 
appear to be the appropriate name. Yet. although Oreobates was 
described from Quijos (whence the name quixensis), and this 
name has been interpreted to mean Quito (Peters. 1955: 347; 
Gorham 1966: 117). Espada himself states: "It occurs in the 
Oriental province, formerly Quijos. in Ecuador, at the foot of 
the Cordillera" ( \ crtcbrados del \ iajc del Pacifico, 1875: 104). 
Specimens from the Napo-Pastaza region in the M.C.Z. correspond 
to the description o\' quixensis, which looks more like a Thoropa 
than a Eupsophus. 

Two new genera, each based on a single individual from the high 
elevations of the Colombian Andes, were described by Coin and 
Cochran in 1963. The first. Nicejoronia, has distinct vomerine 
teeth, no disks and no external tympanic disk; the other. Trachy- 
phrinus, has no vomerine teeth but has small disks (and T- 
shaped phalanges) and a well-defined tympanum. In general 
physiognomy, both of these genera resemble the group of frogs 
here considered, and Trachyphrinus does not look at all like Syr- 
rhophus, the genus in which it would be included if its classifica- 
tion were to be based on conventional morphological characters. 

But all kinds of combinations of characters have been described 
in the high elevation species from Peru to Venezuela. Thus, in 
1873. Peters described Phrynopus peruanus (now called Eupso- 
phus, fide Gorham. 1966) with vomerine teeth and tympanum, but 
apparently without disks. The same combination is found in 
Borborocoetes columbianus Werner (now Eupsophus, fide Gor- 
ham. 1966). Paludicola festae Peracca (now Syrrhophus, fide 
Gorham. 1966). and Eupsophus ginesi Rivero. However, in Pal- 
udicola simonsii Blgr. (now Syrrhophus, fide Gorham, 1966). Syr- 
rhopus montium Shreve and Syrrhopus juninensis Shreve (both 
now Syrrhophus, fide Gorham. 1966) there are no vomerine teeth. 



1968 NEW SPECIES OF ELEUTHERODACTYLUS 3 

external tympanic disk, or disks. In Eleutherodactylus whym- 
peri (Blgr.) and E. surdus (Blgr.), on the other hand, 
vomerine teeth and disks are present but the tympanum is not 
apparent externally, while in Eupsophus wettsteini there are no 
vomerine teeth or disks but there is a small tympanic disk. Tra- 
chyphrinus myersi and Syrrhophus areolatus Blgr. are alike 
in having disks and external tympana (small and ill defined in 
S. areolatus) but no vomerine teeth. 

It is not the purpose of this paper to enter into the many prob- 
lems pertaining to this interesting group (or groups?) of frogs. 
But these digressions have been necessary because of the diffi- 
culty of allocating the new species described in this paper. If this 
group of Andean frogs is found to constitute a natural assemblage, 
perhaps the name Phrynopus can be used for it. It is also 
possible that it may constitute two or more evolutionary lines, in 
which case the genera suggested by Goin and Cochran may be 
found useful. It appears to me, however, that this group, or most 
of it, constitutes a latitudinal extension of Chilean and Argen- 
tinian Eupsophus along belts of similar climatic conditions in the 
Andes. If, as has been reported, some of the species are limited to 
paramos in the Andean Cordillera, their present diversification is 
to be expected. In a group where the tympanum-vomerine odon- 
toids-disks can occur in so many combinations, separation into 
various genera is hardly convenient and may actually be very 
confusing. 

A number of "Syrrhophus" have been described from the low- 
lands or at most, from elevations below 800 m in Ecuador. Such 
is the case, for example, with S. chalceus (Peters), S. calcaratus 
Andersson, and S. coeruleus Andersson. S. chalceus has also been 
reported by Lutz and Kloss, 1952, from Iuarte, along the Rio 
Uaupes. As the largest of the species described was 30 mm (5. 
coeruleus), all the others ranging from 18 to 29 mm, it is not im- 
probable that the lack of vomerine teeth can be attributed to 
immaturity in some of them. It is possible, however, that they 
represent another line of Eleutherodactylus without vomerine teeth 
(since in this genus disappearance of the vomerine teeth does not 
seem to be a difficult process) or that they are in some way con- 
nected with the Eupsophus group from Chile and/or the more 
northern Andes. 

Eleutherodactylus pulvinatus, the species described here, does 
not have the rotund physiognomy of the Eupsophus group, and its 
distribution does not suggest relationship. Nuptial pads have been 
described for some members of the Andean group (Parker, 1932: 



4 BR! MORA No. 306 

363), hut most of the species do not seem to have them. In the 
Chilean species (Cei, l l >o2). on the other hand, and in Thoropa 
(Cochran. 1955: 91), thumb and sometimes breast and axillar 
spines have been described for those species in which males arc 
known. Similar modifications have been described for Crossodac- 
tylodes Cochran (type localit) Macae. Rio de Janeiro), a inono- 
typic genus with hidden tympanum and the vomerine teeth 
represented onh b\ a roughened ridge. In view o\' the foregoing 
considerations, it has been found preferable to describe the new 
leptodactylid in the genus Eleutherodactylus, pending a more 
complete studv o\ its anatoim and relationships. 

I I I l llll RODA< H I. US Pill VINATUS sp. n. 

Type. M.C.Z. 64741, ■' , Paso del Danto, Region de la Esca- 

lera around 1400 m above San [sidro, road from II Dorado to 
Sta. Elena de Uairen. Estado Bolivar, Venezuela. Coll. J. A. 
Riveroand J. Pulido, :<> March 1968. 

Diagnosis. A small Eleutherodactylus with small but distinct 
tympanum, vomerine teeth poorl) defined or absent; two non-spin) 
nuptial pads on the inner side o\ the first digit o\ males; large trans- 
versel) oval disks, larger than the tympanum; first linger shorter 
than second; adpressed heel extending to between eye and nostril: 
and a W -shaped marking behind the occiput. 

Description. Mead as long as broad; snout short, subtriangular; 
tongue free and slightl) nicked behind: vomerine odontoids faintl) 
indicated as two irregular elevations well behind ami between the 
small choanae; eye diameter slightk greater than distance between 
eye and nostril but shorter than snout: interorbital space as 
broad as an upper eyelid; canthus rostralis rather indistinct. 
curved; loreal region onl) slightK sloping, concave; tympanum dis- 
tinct. ' .* the eve diameter; a Hat. indistinct supratympanic fold; a 
line of small, whitish tubercles from elbow to wrist; two distinct. 
oval, metacarpal tubercles; a glandular pad at the base o\' the 
inner side of the first finger and another one distal and slightK 
dorsad to this; first finger shorter than second, its disk much 
broader than the phalanx; all fingers tree: larger disks larger than 
the tympanum; a small. Hat. whitish tubercle at the distal portion 
o\ the tibial segment: a distinct and prominent inner and an indis- 
tinct outer metatarsal tubercle: toes free, with prominent subarticu- 
lar tubercles: heel of the adpressed hind limb extending to between 
eye and nostril. Skin above, shagreened. more coarsely tubercular 
in the tympanic area and anterior flanks. Below, finely granular on 



1968 NEW SPECIES OF ELEUTHERODACTYLUS 5 

throat and anterior part of the belly; more distinctly granular on 
posterior part of the belly and thighs. A very large subgular vocal 
sac with a triangular anterior end; a space (about 2 mm) between 
tip of the jaw and anterior edge of sac. A "frenulum" in the an- 
terior tip of the sac continues anteriorly almost to the tip of the jaw. 

Color. Above, brownish gray, with a dark brown interorbital 
bar, a black supratympanic streak, a W-shaped marking on the 
anterior part of the back, and some large blotches and small spots 
on the posterior two-thirds of the back; on the flanks the blotches 
continue ventrally, forming oblique, wavy bars that are separated 
by light gray areas; anterior and posterior part of the thighs uniform 
brown, this color extending as bars across the gray color of the 
upper part of the thighs; limbs and fingers with cross-bars and 
blotches; upper lip with triangular blotches on a gray background; 
no canthal streak. Ventral surfaces dirty white, darker on the 
throat and distal portion of the limbs. 

Measurements (in mm). Snout-vent 26.1; head breadth 9.8 
(between tympana); head length 10; femur 13; tibia 14.1; foot 1 1. 

Paratypes. There are six paratypes (U.P.R.-M. 1 02214-02219), 
all coming from the type locality. All are males, and the size 
varies from 26.1 mm to 23 mm. In some of the smaller specimens 
the canthus seems to be less defined than in larger specimens, and 
a cross-section of the snout would be more or less rounded if it 
were not for the slight concavities of the loreal areas. The heel of 
the adpressed hind limb extends to the nostril in only one speci- 
men, but the skin of the dorsal surfaces is more granular than in 
the type, at least in two examples. The belly and thighs are uni- 
formly granular, the throat more finely so in most specimens. 
The vocal sac does not form the anteriorly triangular pouch 
that it forms in the type, but it is possible that it has been dis- 
placed, as a similar condition can be noticed in several specimens. 
The W-shaped marking is present in all specimens, but the oblique 
bands of the flanks are less distinct in some, although a trace of 
them can be noticed in all. Vomerine odontoids cannot always be 
seen, but they can be felt with a needle. They seem to be irregu- 
lar in both size and position, and in some specimens seem to 
be almost totally absent. 

Habits and relationships. All the specimens were collected at 
night, calling from low bushes ( 1 to 3 ft. from the ground) along 
the rocky bank of the road at an elevation of approximately 1400 
m. At dusk, their calls seemed to come from under the rocks 

1 University of Puerto Rico — Mayagiiez. 



6 BREVIORA No. 306 

lining the almost vertical road bank, but none could be collected at 
the tune. Later at night, their calls were traced and recorded and 
the various specimens collected. In the dark the Erogs are yellow 
or cream, but they turn dark gray in the daytime. Their voices 
were not heard much below the elevation where all the specimens 
were obtained. No other species of Eleutherodactylus was col- 
lected or heard in this area. 

Only scattered low bushes, some soil bromeliads and. occasion- 
ally, thickets of lilaceous plants occur in the rather open areas 
where E. pulvinatm was collected. There were permanent and 
temporary streams everywhere, as the region is probablv one ol 
the rainiest in Venezuela. 1 The forest. 50 to 60 meters from the 
road, is a rather open montane forest, usuallv with man) epiphytic 
bromeliads. orchids, and araeeans. Most of the soil is a white 
quartziferous sand, which is covered in the forest by a layer of 
dead leaves and other organic matter. 

Although mam Eleutherodactylus have W-shaped markings on 
the back, the distinct markings ol E. pulvinatUS remind one oi the 
condition o{ E. W-nigrum (Boettger), which, however, is a species 
from more than 3000 m in the Amies o\ Ecuador. Other species. 
from relativel) adjacent regions, are: /•.'. inguinale Parker (New 
River, Guyana), E. beebei Cochran (Kartabo), E. pliciferus 
(Blgr.) and E. ramagii ( Blgr. ) (Iguarasu, Pernambuco, Brasil), 
/ crepitans Bockermann (San Vicente, Matto Grosso), E, car- 
valhoi B. Lutz (Itacoai R.. NW Brasil). E. peruvianus (Melin) 
(Roque, Peru). E. vilarsi (Melin) (Taracua, Rio Uaupes), / 
rosmelinus Gorham (Rio Uaupes). /.. melini Bokermann (Roque, 
Peru). /:". altamazonicus Barbour and Dunn (upper Amazon) 
and /■.. marmoratus (Blgr.) (Mt. Roraima). In all these species 
there are well-defined vomerine teeth, and nuptial pads have not 
been described for am. E. inguinale has on obtusely angular 
canthus, a broad interorbital space, much broader than the upper 
eyelid, and a black bordered, inguinal ocellus; E. beebei Cochran 
is a large species 85.5 mm in length; in E. pliciferus and E. ramagii 
the first linger is longer than the second; E. crepitans, from drv 
forest, is similar to /.. conspicillatus, and its tympanum is % the 
eve diameter; E. carvalhoi does not have a tympanum; E. peru- 
vianus and E. vilarsi are 50 and 39 to 41 mm. respectively, and 
may be synonyms of E. conspicillatus; E. rosmelinus has a 
depressed head, disks VS the size of the tympanum and first finger 

1 According to a report in a local newspaper, engineers working on the 
road from El Dorado to Sta. Elena could only work 60 to 80 days of the 
year, and even those were not always sunny days uithout any rainfall. 



1968 NEW SPECIES OF ELEUTHERODACTYLUS 7 

longer than second; E. melini has an indistinct tympanum and 
small disks; E. altamazonicus has a depressed and oval snout and a 
barely distinguishable tympanum; and E. marmoratus has a 
rounded snout as long as the orbital diameter, dark-edged marb- 
lings on the head, and a small size (only topotypical specimen: 
19 mm). 1 

From the lowland "Syrrhophus" mentioned before, E. pulvinatus 
can apparently be distinguished by the nuptial pads it has in the 
inner side of the first finger. Other differences are as follows: In 
S. chalceus (Peters), from Pastaza River, the first finger is longer 
than the second; in S. coeruleus Andersson, from eastern Ecuador, 
the canthus is distinct and there are two pairs of dorsolateral folds; 
and in S. calcaratus Andersson,- from near Archidona, 800 m, 
there is a canthal streak, and the disk of the first finger is only 
slightly broader than the phalanx behind it. 

Acknowledgments. The author wishes to acknowledge all the 
courtesies and attentions extended to him by the Instituto Vene- 
zolano de Investigaciones Cientificas of Caracas, and most espe- 
cially by Dr. Gilberto Rodriguez, Head of the Hydrobiology 
Department, who went out of his way to foster the investigations 
resulting in this and other papers. During the course of this study 
the author was employed by and received all kinds of assistance 
from the I VIC. 

For the trip to La Escalera, the author relied on an invaluable 
assistant, Mr. Juan Pulido, always faithful and always willing to 
help during the difficult situations that often arise during such 
trips. Major Luis Alfonso Godoy, Commander, Engineering 
Corps at sector A of the road from El Dorado to Sta. Elena, was 
extremely helpful in providing facilities and assistance. 

As always, the author is greatly indebted to Dr. Ernest Williams, 
not only for permitting the use of the facilities of the Museum of 
Comparative Zoology at Harvard, but also for reading the manu- 
script critically and for discussing various taxonomic and evolu- 
tionary problems. Mr. Benjamin Shreve has also read the manu- 
script and suggested changes. The author, however, assumes full 
responsibility for the shortcomings of this paper. 

1 Frogs collected in Mt. Marahuaca by the author (Rivero, 1961: 80) 
and pronounced conspecific with E. marmoratus by Parker, do not repre- 
sent E. pulvinatus. However, further comparisons should be made when 
more specimens of E. marmoratus become available. 

- Among the group of frogs here discussed, the species name calcaratus 
was first used by Giinther, 1881, for Cacotus calcaratus, now considered a 
synonym of Eupsophus roseus. 



8 bki vior \ No. 306 

REFERENCES 

VNDl RSSON, 1 \KS (i \HKII 1 

1945. Batrachians from Easl Ecuador collected 1937. 1938, b) Wm. 
Clarke-Macintyre and Kolt" Blomberg. Arkiv Zool. 37<2): 
1-88. 

B.XRUOl R. I IIOM \s. \ND I R. l)l NN 

1921. Herpetological novelties Proc Biol Soc. Wash. 34: 157-162. 

linkl KM \NN. \\ I RM R 

1958. A preoccupied name of a neotropical frog, genus Eleuthero- 

dactylus. Herpetologica I 1. pt. 2: 95. 
1965. Tres novos batraquios da regiao central de Matto Grosso, Brasil 

(Amphibia. Salientia). Rev Brasil. Biol. 25(3): 257-264 

BOI l I NG1 R, Cil I iRGl \l HI Rl 

1882. Catalogue ol the Batrachia Salientia in the Collection of the 

British Museum, ed. 2. I ondon, Kvi -♦ 503 pp. 
1888. On some reptiles and batrachians from Iguarasse, Pernambuco. 

\.m. Mag. Nat. Hist. 6(2): 40-43. 
1898 \n account of the reptiles and batrachians collected b) Mi 

W. I ll Rosenberg in western Ecuador. Proc. Zool. Soc 

l ondon 1898: 107-126. 
1900. Report on a collection made h\ Messrs. I V. Mc< onuel and 

J. J. Quelch at \lt Roraima in British Guiana. lYans. Linn. 

Soc l ondon (2) 8: 51-76. 

I I i, losi M 

I960 \ surve) of the leptodactylid frogs, genus Eupsophus, in ( hile. 

Breviora, Mus. < omp. Zool., No. MX: 1-13. 
1962. Batracios de ( hile. Santiago de Chile, 128 f cviii pp. 

( I I. losi \| . \S|) 1 I Is ( MM KRI) 

I s 5 s . Biologia \ desarrolo de Eupsophus taeniatus Girard. Inv. Zool. 
( hUenas t 159-18 

DUMERIL, \SDRl' M. C, AND GaBRIEI BlBRON 

1841. Erpetologie Generate, Vol. 8, Paris, ii - 792 pp. 

( OCHRAN, Doris 

1955. I rogs of southeastern Brasil. Bull. U. S. Nat. Mus. 206: nvi 

+ 423 pp. 

1956. A new species of frog from KartabO, British Guiana Zoologica 
11. pt. I: I 1-12. 

Coi'i . 1 DVk w<i> D. 

IX 7 K. A new genus of < ystignathidae from Texas. Amer. Natur. 12: 
252-253. 

ElUmmrsm (Newspaper) 

1968. I April 6) En junio sc enla/aran los ingenieros militares de la 
carretera El Dorado-Sta. Elena. Caracas. Venezuela. 



1968 NEW SPECIES OF ELEUTHERODACTYLUS 9 

Gallardo, Jose 

1965. A proposito de los Leptodactylidae (Amphibia Anura). Papeis 
Avulsos, Sec. Agric, Sao Paulo, 17, art. 8: 77-87. 

Goin, Coleman, and Doris Cochran 

1963. Two new genera of leptodactylid frogs from Colombia. Proc. 
Calif. Acad. Sci. 31(17): 499-505. 

Gorham, Stanley W. 

1966. Das Tierreich. Liste der rezenten Amphibien und Reptilien. 
Lief. 85, xvi + 222 pp. 

Grandison, Alice 

1961. Chilean species of the genus Eupsophus (Anura: Leptodacty- 
lidae). Bull. Brit. Mus. Nat. Hist. 8(3): 111-148. 

Gunther, Albert 

1881. Account of the reptiles, batrachians and fishes collected during 
the survey of H.M.S. "Albert" in the Straits of Magellan and 
on the coast of Patagonia. Proc. Zool. Soc. London 1881: 
18-22. 

Jimenez de la Espada, Marcos 

1872. Nuevos batracios americanos. An. Soc. Espanola Hist. Natur. 

1: 85-88. 
1875. Vertebrados del Viaje al Pacifico, Batracios. Madrid, 208 pp. 

Lutz, Bertha, and Gertrude R. Kloss 

1952. Anfibios anuros do Alto Solimoes e Rio Negro. Apontamentos 
sobre algunas formas e suas vicariantes. Mem. Inst. Osw. 
Cruz 50: 624-678. 
Melin, Douglas 

1941. Contribution to the knowledge of Amphibia of South America. 
Goteborgs Kungl. Vetensk. Vitterh.-Sam. Handl., Ser. B.. 1(4): 
1-71. 
Miranda Ribeiro, Alipio 

1926. Notas para servirem ao estudo dos gymnobatrachios (Anura) 
brasileiros. Archiv. Mus. Nac. Rio de Janeiro 27: 1-227. 

Parker, H. VV. 

1932. The systematic status of some frogs in the Vienna Museum. 
Ann. Mag. Nat. Hist. (10) 10: 341-343. 

Peracca, Mario Giacinto 

1914. Reptiles et batraciens de Colombie. Mem. Soc. Sci. Neuchatel 
5: 96-111. 

Peters, Wilhelm 

1873a. Hr. W. Peters las fiber zwei Giftschlangen sus Afrika und fiber 
oder weniger bekannte Gattungen und Arten von Batrachiern. 
Monatsb. Akad. Wissensch. Berlin, Mai: 471-478. 

1873b. Hr. W. Peters las fiber eine newe Schildkrotenart Cinosternon 
Effeldtii und einige andere neue oder weniger bekannte Amphi- 
bien. Monatsb. Akad. Wissensch. Berlin, Sept. Oct.: 603-618. 



10 BREVIORA No. 306 

Pi I I Rs. I Wll S 

1955. Herpetologica] t\pe localities m Ecuador. Rev. Ecuador Ent. 
Par. 2(3-4): 335-352. 

Kl\l Rd, J LAN A. 

I9M Salientia of Venezuela. Bull. Mus. Comp. Zool. I2i><l): 

1-208. 
|yh4. Salientios en la coleccion de la Sociedad de Ciencias Naturales 

La Salle de Venezuela Carib. J. Sci l( 1 ): 297-305. 

SHRI \ 1 . Ml M \MIN 

I y 3 s . \ new Liolaemus and i\m> new Syrrhopus from Peru. J. Wash- 
ington Vcad. Sci 28< i n 404-407 

\\ I KM K. I K\N/ 

IS 1 * 1 ' Webei Reptilien und Batrachier aus Colombien und rrinidad 
\ erh. Zool-bol (.cs Wien 19: 4 _ <i-4S4 

(Received 26 June 1968 I 



968 



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BREVIORA 

Mm senium of Comparative Zoology 

Cambridge, Mass. 31 December, 1968 Number 307 



A NEW SPECIES OF HYLA (AMPHIBIA, SALIENTIA) 
FROM THE VENEZUELAN GUAYANA 

Juan A. Rivero 



Abstract. A small species of Hyla, H. rodriguezi, is described from 
the wet region of La Escalera, Estado Bolivar, Venezuela. The new form 
is characterized by having a large, well defined tympanum, about % the 
size of the eye; nearly webless fingers; 3 A webbed toes; speckled throat and 
orange-red areas in the thighs. It may be related to the Hyla leprieurii 
group but it is also similar to Hyla grandisonae Goin, from which it differs 
in coloration and in other details. 

Among the new frogs collected by the author in the region of 
La Escalera, in southeastern Venezuela, there is a small species of 
a bromelicolous form that appears to be new to science. The new 
species is named Hyla rodriguezi in honor of Gilberto Rodriguez, 
of the Instituto Venezolano de Investigaciones Cientificas, who 
made possible the expedition during which this and many other 
interesting species were collected. Dr. Rodriguez spared no effort 
in facilitating the author's investigations of the frogs of the Vene- 
zuelan Republic. 

During this and other investigations pertaining to the Vene- 
zuelan frog fauna, the author has utilized the facilities and collec- 
tions of the Museum of Comparative Zoology at Harvard. He is 
most thankful to the Curator of Herpetology, Dr. Ernest Williams, 
for his cooperation and unfailing willingness to be interrupted and 
consulted at any time. 

Hyla rodriguezi sp. n. 

Type. M.C.Z. 64740, 6 , Paso del Danto, Region de La 
Escalera, ca. 1400 m above San Tsidro, road from El Dorado to 
Sta. Elena de Uairen, Estado Bolivar, Venezuela. Coll. Juan A. 
Rivero and Juan Pulido, 26 March 1968. 



2 BREVIORA No. 307 

Diagnosis. A small Hyla with bony head; strong, straight can- 
thus rostralis; almost vertical, concave lorcal region: anteriorly 
converging vomerine teeth; large tympanum, about 7 « the size of 
the eve; tubercular dorsal surfaces (males?); red (alive) or 
white (preserved) areas on the thighs and speckled throat. 

Description. Head slightly broader than long; snout short, 
rounded, concave above; nostrils forming a truncate tip; canthus 
rostralis strong and well defined; loreal region vertical, concave; 
interorbital space as broad as an upper eyelid; vomerine odontoids 
anteriorly converging, extending from level of palatines to about 
middle of inner margin o\' choanae; choanae moderate, oblique; 
tympanum large, distinct, about 7 * the size of the eye; larger disks 
much smaller than tympanum; eye diameter as long as distance 
between eye and nostril; a line o{ whitish tubercles from elbow to 
wrist; metacarpal tubercles not well differentiated; subarticular 
tubercles well defined; fingers almost free, the first swollen at the 
base and with an elongated brown pad which covers : i of its 
inner side; first linger not extending much beyond first subarticular 
tubercle oi second; second finder shorter than last; a small tubercle 
at the heel; no tarsal fold; an elongated inner and a flattened, 
rounded, and less distinct outer metatarsal tubercle; subarticular 
tubercles o\ toes prominent; toes about % webbed (taken in order 
from first to fifth, the following phalanges are free of web; 2; Wi; 
IV6; 2V6; I 1 »); heel o\ the adpressed hind limb extending to the 
anterior corner oi the eye. Skin above distinctly granular, most 
o\' the granules having a corneus tip; on the posterior half of the 
back, the anterior loreal region, the arms, and tibial segments, the 
granules become larger tubercles. Below, smooth on the chin and 
chest; finely granular on the throat; coarsel) granular on the belly 
and postero-ventral aspect of thighs; one almost imperceptible 
furrow on each side oi the throat. 

Color. Above, dark brown, with two somewhat lighter and ill- 
defined bands from behind the eyes and over the level of the shoul- 
ders to near the sacral region; a dark, ill-defined streak along the 
flanks below the light brown band; other longitudinal, light brown 
spots can be noticed on the dorsum; a whitish, ill-defined spot or 
line under the eye. in front of the tympanum; disk of first finger 
white, of second finger half white and half brown; posterior portion 
of arm and axillar membrane white; flanks below dark brown 
streak white, this color extending dorsally to the illea on the pos- 
terior end; thighs pink, speckled and spotted with brown; rest of 
hind limbs and fore limb with transverse blotches or bars; web 



1968 NEW SPECIES OF HYLA 3 

pink. Below, creamish white, uniform, except for the sides of the 
throat and lower lip, which are freckled with purplish brown. 

The living animal appeared to be of a solid brown color with 
orange areas on the thighs, white forearms (posterior part) and 
axilla, creamish venter with purplish specks on the throat, rosy 
hind limbs (underneath), and with cream points along the forearm 
and at the sides of the anus. 

Measurements (in mm). Snout-vent 34.5; head length 11.02; 
head breadth 11.2; femur 16; tibia 17.7; foot 12.4. 

Paratypes. The paratypes are U.P.R.-M. 1 02207-02012, all 
from the same locality and with the same data as the type. They 
vary from 21.6 to 35.2 mm in snout-vent length. In some speci- 
mens the heel of the adpressed hind limb may extend to between 
eye and nostril; in a few the web is less extensive than in the type, 
and the feet may perhaps be said to be not more than % webbed. 
The skin may be almost smooth in some of the smaller specimens. 

There is some variation in coloration, although the pattern is 
rather similar in most specimens. In a few examples the longi- 
tudinal bands or spots are darker, not lighter than the background 
color, and in the smallest individual the background color is gray 
and there are many short longitudinal streaks on the dorsum. In 
all specimens, there is a light-colored spot or line in front of the 
tympanum, but in some it is very diffused, and in one it extends 
anteriorly along the upper lip. In preserved material, the ventral 
color is as described for the type, except for the fact that the 
freckles of the throat may not be limited to the margins and may 
also extend to the central area. In living animals the color is more 
variable, and the throat may be yellow with purplish freckles while 
the rest of the venter is cream or lighter yellow. Sometimes there 
is almost no orange on the thighs, while at others the orange be- 
comes red and may extend, on the ventral side, to the tibial and 
metatarsal segments and the foot. The lower side of the forearm 
may also be orange or red 

Habitat and relationships. The place where H. rodriguezi was 
collected is probably one of the rainiest in Venezuela, and when it 
is not raining it is usually foggy and misty. The vegetation can 
probably be classified as the wet montane type, and there is a 
great abundance of epiphytes, the Aracean Philodendron nobile 
being particularly evident. 

Hyla rodriguezi was collected in ground bromeliads (apparently 
of the genus Vriesea) that grow along the road bank at an elevation 

1 University of Puerto Rico — Mayagiiez. 



4 BREVIORA No. 307 

of about 1400 in. There was never more than one specimen in 
any one bromeliad, and onl) once was there another species of frog 
(Hyla minuta) in the same plant. All were collected at dusk. At 
night, no sound was heard in the area except that of a new species 
of Eleutherodactylm (Rivero, in press), and occasionally, the 
whistling call of Leptodactylm rugosus. 

This species is well characterized, and if 1 were to relate it to 
an) other known species, it would probabK be to the Hyla lep- 
rieurii group. However, Hyla rodriguezi is a smaller species with 
a shorter snout, less webbing between the toes, and inconspicuous 
vocal sacs. It is also somewhat similar to Hyla alboguttata, but the 
head of Hyla rodriguezi is more bom. its canthus straighter and 
better defined, its feet less webbed, and its color different. The 
speckled coloration of the throat of //. rodriguezi seems to be \ei\ 
characteristic of this species. 

Hyla grandisonae Goin (type locality, Ma/arum Forests, 
Guyana) agrees in main characters with //. rodriguezi, but on 
closer examination it is found that the two species ma\ not even 
be too closet) related. In //. grandisonae, the heel of the adpressed 
limb reaches almost to the nostril, the skin above is smooth, the 
ventral color is Lira\. and the species is said to be related to Hyla 

parviceps Boulenger, //. rondoniae Bokermann and //. bokermanni 

( loin, three species that do not seem to be close to Hyla rodriguezi. 

LITERATURE CITED 

H<'KI KM \NN. \\ I KM K 

1963. Duas novas espeeies de "Hyla" de Rondonia, Brasil (Amphibia, 
Saliemia). Rev Brasil Biol. 23 1 \)\ 247 250. 

1965. lies no\os batrachios de regiao central de Mato Grosso, Brasil. 
Rev. Brasil Biol. 251 3 > 257-264. 

( rOIN, ( hi i \i \s 

I960. Description of ;i new frog of the genus Hyla from northwestern 
Brazil. \nn Mag Nal Hist 13(2): 721-724. 

1966. \ new frog from the genus Hyla from Biitish Guiana. Quart. 
lour \ la. Acad. Sci. 29(1): v '-42. 

Rivero, J. \ 

1968. A new species o/ Eleutherodactylm (Amphibia, Salientia) from 
the Guayana Region. Edo. Bolivar. Venezuela Breviora, \lus 
Comp. Zool., No. 306: I -I I. 

(Received 26 hme 1968.) 



1968 



NEW SPECIES OF HYLA 



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BREVIORA 



Museum of Comparative Zoology 

Cambridge, Mass. 31 December, 1968 Number 308 

THE MOLLUSCAN FAUNA OF AN UNUSUAL 

BERMUDIAN POND: A NATURAL EXPERIMENT 

IN FORM AND COMPOSITION 

Stephen Jay Gould 

Abstract. Twenty-four molluscan species inhabit a landlocked marine 
pond of normal salinity in Hamilton Parish, Bermuda. Walsingham Pond, 
unusual because of its isolation and the ecologic effects thereof, can be 
viewed as a natural experiment. The form and composition of its mol- 
luscan fauna are compared with "control" samples from open marine 
environments. Composition of the fauna is similar to two modern and 
three fossil samples from localities similar in ecology to Walsingham Pond 
but directly exposed to the open sea. Isolation does not seem to affect the 
presence and absence of species. Walsingham Pond molluscs are, however, 
unique in their shell form. As demonstrated by plots of weight vs. size, the 
four major Pond gastropods have thinner shells than those of the same 
species from control samples. 

It is often said that natural history is science of a lower order 
than experimental biology. If description and explanation are 
viewed as ascending orders of sophistication, this charge is valid 
when natural history remains at the level of "plain story"; for the 
very undertaking of an experiment implies a search for explanation. 
Yet there are a variety of natural situations that possess the essen- 
tial character of experiments, even though no human manipulation 
of material is involved. The evolutionary phenomena of parallelism 
and convergence, for example, serve the same function as experi- 
mental replication; both provide the repeated occurrences upon 
which inductive generalizations are based (Harris and Morren, 
1966). I am concerned here with another kind of "natural ex- 
periment" — the unusual situation amidst a large number of "nor- 
mal" occurrences that act as controls. 

Walsingham Pond in Hamilton Parish, Bermuda, is an unusual 
situation. Walsingham is a landlocked marine pond of nearly nor- 
mal salinity maintained by tidal interchange with Castle Harbor 
through underground caves. Limestone walls and mangrove roots 



2 breviora No. 308 

form the borders of the Pond; the bottom is covered with a soft 
organic mud composed primarily of decaying material, faecal pel- 
lets, and shell fragments. Dominant elements of the bottom biota 
are the alga Chaetomorpha crassa, a profusion of sponges, and. 
unfortunately for collectors, the jellyfish Cassiopeia; there are no 
hard corals in the Pond. Maximum water depth is 22 feet; the 
bottom mud layer may be several feet thick. Helz (unpublished 
manuscript) studied the water chemistry of Walsingham Pond. 
Oxygen increases steadily from top to bottom; values are con- 
sistently 1 ml 1 less than those of the normal marine waters of 
Castle Harbor. Nitrate and phosphate show no trend with depth; 
values are two to three times those of Castle Harbor. This abun- 
dance is presumably related to the continuing concentration of 
organic matter in Walsingham Pond. After several rainless days. 
Pond salinity was 100-170 ppm higher than that of Castle Harbor. 
Surface waters were 10-50 ppm above bottom waters; mixing mav 
occur by sinking of 'he denser surface water. Ten hours after a 
heavy rain, surlaee waters of the Pond were less saline than those 
o\' Castle 1 [arbor b\ 1 500 ppm. 

Two aspects of the Pond fauna — composition and form — may 
he studied to find and explain differences between this unusual 
situation and the normal open marine habitats o\' Bermuda. I 
have confined m\ attention to the Mollusca. due both to personal 
preference and to the availability of abundant comparative ma- 
terial in the magnificent collection o\' Mr. Arthur Haycock (Ber- 
muda Museum). Since Walsingham Pond's landlocked status is 
its most unique attribute, I shall concentrate on questions related 
to this point. We shall want to know if the Pond's isolation, of 
itself, influences the form and composition of its molluscan fauna. 
Failing such influence, we may still ask if a secondary result of 
isolation an unusual condition of water chemistry attributable to 
curtailed mixing of Pond with ocean, for example — can be cor- 
related with differences between the Pond fauna and "control" 
faunas from normal marine environments. 

COMPOSITION OF THE MOLLUSCAN FAUNA 

Nine thousand two hundred fifty years ago, the sea stood approxi- 
mately 78 feet below its present level (A. C. Neumann, personal 
communication, determined this figure by dating a peat bed cored 
at the bottom of Harrington Sound). With a maximum depth of 
22 feet, Walsingham Pond cannot be more than a few thousand 
years old. Entrance to the Pond can be obtained only through 
the channels of underground caves that connect Walsincham to 



1968 UNUSUAL BERMUDIAN POND 3 

Castle Harbor; I suspect that the molluscan populations now liv- 
ing in the Pond owe their origin to the infrequent transport of 
larvae through these channels. If this isolation exerts a major 
control on composition, I would expect a disharmonious as- 
semblage of molluscan species with a predominance of forms hav- 
ing long and highly mobile larval stages. If ecology is the major 
control, then neither the young age of the Pond nor its isolation 
should affect the distribution of species. In this case, I would ex- 
pect a molluscan fauna composed of those species that characterize 
similar environments in open marine areas around Bermuda. This 
finding would carry the implication that most molluscan species 
had ample access to the Pond and that the composition is a func- 
tion of the normal environmental preferences of species rather than 
of chance introductions. 

I have endeavored to compile a complete listing of the molluscs 
in Walsingham Pond. Shells were obtained both by visual collect- 
ing and by bulk sampling with a Van Veen grab at various depths; 
composition does not seem to change with depth. With the excep- 
tion of Chama macerophylla, which occur infrequently, cemented 
to the limestone walls, all species inhabit the muddy bottom. Some 
burrow into the mud (Vermicularia spirata), others crawl along 
the surface (Nassarius ambiguus), and still others (Modulus 
modulus) commonly live among the algal colonies of Chaetomor- 
pha crassa. 

Class Bivalvia 
Family Mytilidae 

Brachidontes exustus (very few specimens) 
Condylocardiidae 

Carditopsis smithii (one specimen) 
Lucinidae 

Codakia orbiculata (most common bivalve in Pond) 
Chamidae 

Chama macerophylla (infrequently on limestone walls) 
Leptonidae 

Lasaea bermudensis (very common) 
Cardiidae 

Laevicardium laevigatum (one immature specimen) 
Veneridae 

Gouldia cerina (very common) 
Tellinidae 

Tellina mera (one specimen) 
Sanguinolariidae 

Tagelus divisus (rare) 



4 bri-viora No. 308 

Class Gastropoda 
Family Rissoidae 

Alvania platycephala ( very common) 

Rissoina cancellata (one specimen) 
Vitrinellidae 

Vitrinella helicoidea I moderately common ) 
Turritellidae 

Vermicularia spirata (most common mollusc in Pond) 
Caecide 

Caecum delicatulum (a lew specimens) 

Caecum termes ( very common) 
Modulidae 

Modulus modulus ( very common) 
Ccrithiidae 

Alahinu cerithioides < quite common ) 

Cerithium variabile (ver\ common) 
Nassariidac 

Nassarius ambiguus (quite common) 
Marginellidac 

Hyalina avena ( rare ) 
Bullidae 

Bulla striata ( rare) 
Atyidae 

Haminoea antillarum ( rare ) 
Retusidae 

Retusa candei < moderately common ) 
Pvramidellidae 

Odostomia didyma (moderateK common) 

I was able to obtain samples of the molluscan fauna from five 
additional localities of similar environment (mangrove roots and 
muddy substrates), but situated in areas having free access to the 
open sea. These "controls" should serve to 'test the hypothesis 
that composition of the Walsingham Pond fauna is unusual for a 
mangrove-mud area because of the Pond's isolation and young age. 

1. Ely's Harbour, Sandys Parish (grab sample collected at a 
depth of 10 feet by Robert F. Schmalz). 

2. Mangrove root area on north coast of Ferry Reach, 50 yards 
east of Bermuda Biological station (personally collected; three 
Van Veen grab samples at depths of 5-7 feet). 

Fortunately, three small samples of fossil shells were available 
to provide some temporal control on the stability of community 
composition during the past 10.000 years. Dried mud and man- 
grove root fragments were present in all samples. 



968 UNUSUAL BERMUDIAN POND 





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6 breviora No. 308 

3. Pembroke Marsh, Pembroke Parish (collected by Charles 
Schuchert of Yale University in the 1920's). Of the five controls, 
only this sample cannot be identified as having been in open contact 
with the sea. 

4. Longbird Bridge boring. 71.8 feet below present sea level. 

5. Longbird Bridge boring. 85.5 feet below present sea level. 
Samples 4 and 5. kindly supplied by Walter S. Newman, were 

obtained from the test boring made prior to construction of Long- 
bird Bridge, St. Gcoriic's Parish. 

Table 1 lists all the mud-dwelling macromolluscs of Walsingham 
and these five localities in order of their abundance (only the rock- 
cementing Chama macerophylla is eliminated by this constraint on 
environment). I do not know how representative the small fossil 
samples are of actual diversity and abundances. The micromol- 
luscs are not included, because several oi the samples contained 
only the larger shells, with insufficient matrix for the screening of 
small species. Yet strong similarities almost surely exist between 
micromolluscs of Walsingham and the five controls. Several speci- 
mens oi Retusa candei, Caecum termes, and Odostomia didyma, 
for example, were obtained from dried mud caking the apertures 
of Vermicular ia spirata specimens in the Longbird 4 sample; all 
three of these micromolluscs are common in Walsingham Pond. 

The similarities between Walsingham and the controls are strik- 
ing. Five macromolluscs strongly dominate the Walsingham fauna 
— Vermicular ia spirant. Cerithium variabile, Codakia orbiculata, 
Modulus modulus, and Nassarius ambiguus, in that order. Except- 
ing the absence of Codakia in the Ely's Harbour sample, all of 
these species are present in all oi the five controls. Considering the 
small si/e of several samples (20 shells in Longbird 5. for ex- 
ample), this correspondence seems highly significant. Bulla .striata 
and Tagelus divisus, rare but consistently present in Walsingham. 
are found in three and two of the controls respectively. Of Brachi- 
dontes exustus and Tellina mera, the Walsingham "trace ele- 
ments." only the first is found at one other locality. 

A few unexplained discrepancies arise from species present in 
control samples but not in Walsingham Pond. The venerid clam 
Chione cancellata is present in three of the controls, dominant in 
one of these (Longbird 4) but absent from Ely's Harbour and 
Longbird 5 as well as from Walsingham. Batillaria minima, a 
homcomorph of Cerithium variabile, commonly occurs with the 
latter in rocky and sandy intertidal environments of the open sea 
(they are so associated in Castle Harbour, barely 100 yards from 
Walsingham Pond). Batillaria is absent from all but one of the 



1968 UNUSUAL BERMUDIAN POND 7 

mangrove-mud environments, but dominates among the mangrove 
roots east of the Biostation. This might argue for atypicality of the 
Biostation locality, but not for a lack of correspondence between 
Walsingham and the other controls. While it is not especially rele- 
vant to this study, the ecologic relationship of the homeomorphs 
Cerithium and Bat Maria is a fascinating problem. 1 

Thus, the association of Vermicularia spirata, Cerithium varia- 
bile, Codakia orbiculata, Modulus modulus, and Nassarius am- 
biguus in Walsingham Pond is no product of accidental access, but 
an expression of the normal molluscan community in mangrove- 
mud areas. Vermicularia is not often thought of as an inhabitant of 
muddy areas, for it usually lives cemented to hard objects in clear, 
open water. Olsson (1951), however, found abundant specimens 
on the mud flats in Tampa Bay, Florida. He described them as 
Vermicularia fargoi, but I have synonymized this species with V . 
spirata elsewhere (Gould, in press). Tagelus divisus prefers 
muddy environments (Fraser, 1967). In short, the unique isola- 
tion of Walsingham Pond is not reflected in the composition of 
its molluscan fauna. Characteristic species of the molluscan com- 
munity that inhabits mangrove-mud areas throughout Bermuda 
have all become established in Walsingham Pond despite the bar- 
riers to distribution. 

FORM OF THE MOLLUSCAN FAUNA 

Is the form of molluscan shells in Walsingham Pond different 
from that of the same species in open marine habitats? If such 
differences are found, we may again pose the question: Is the 
uniqueness of Walsingham molluscs attributable to the Pond's iso- 
lation per se or to some secondary ecologic result of that isolation? 
In the first instance, isolation of a small founding population main- 
tains the distinguishing gene frequencies present, by chance, in the 
founding individuals or attained, without regard to adaptive sig- 
nificance, by genetic drift. In the second case, an unusual condi- 
tion of water chemistry or bottom ecology works its influence upon 
the form of shells by direct phenotypic modification. A mixed case 
is also likely: unusual conditions establish strong selective pres- 
sures, while isolation favors the stabilization of new genotypes by 



1 Where they occur together, distribution is not random. Patches of 20-30 
individuals of one species are often found in large tracts dominated by 
the other. This phenomenon is particularly well seen in Coot Pond, St. 
George's Parish. 



8 BREVIORA No. 308 

removing the diluting effect o\' gene influx from normal popu- 
lations. 

When a single speeies differs from its 'control'" counterparts m 
a unique way, it may be difficult to distinguish among the three 
explanations. Walsingham specimens of Modulus modulus are 
often relatively higher than is normal for the species in Bermuda. 
The most spectacular modification, however, is found in Vermicu- 
laria spirata. This gastropod customaril) begins life as a normally 
coiled turritellid, but later cements to a hard object and uncoils its 
shell. In the absence of any hard substrate on the floor of Wal- 
singham Pond, more than 95 per cent o\' the Vermicular ia shells 
do not uncoil at all; those that do unwind slowly and regularly 
without a trace oi the normal attachment scar. We have here the 
curious case o\' an animal reverting, functionally, to the form and 
habitat oi its ancestor: Walsingham Vermicularia construct and 
inhabit burrows identical to those o\ its coiled ancestor Turritella. 
Since the Walsingham Vermicularia uncoil normally when pro- 
vided with hard coral substrate in a laboratory tank, this striking 
modification may have no genetic base other than the great pheno- 
ls pic plasticity permitted b\ the normal genotype. I have docu- 
mented this ease in some detail elsewhere (Could, in press). 

When many species differ from their "controls" in a similar 
way. some aspect o{ the Pond's unique ecolog) is probably involved 
in the explanation — either directly by phenotypic modification 01 
indirect!) via selection pressures. There is one general modifica- 
tion that affects all the major Walsingham species: Pond shells 
appear to be thinner than those o\ the same species in open marine 
habitats. I tested this obser\ation by comparing plots oi combined 
linear dimensions vs. weight for the four most common Pond 
gastropods (Fig. 1) with similar plots for the same species in 
normal marine habitats (Haycock collection. Bermuda Museum ). 
Use of a combined linear dimension (length plus width) provides a 
better measure o\ overall size than length alone, fin Modulus 
modulus, for example. Pond shells of a given length are generally 
narrower — hence lighter — than are control specimens of the 
same length. The lower weight length ratios of Pond Modulus at a 
given length might be due not to shell thinness [the hypothesis to 
be tested] but to differences in shape between Walsingham and 
control shells. | 

In the ideal case of isometric growth, shell weight varies directh 
with shell volume, and shell volume increases as the cube of a 
single linear dimension or an added combination of such measures 
(using length plus width instead o\ length alone will modify the 



1968 



UNUSUAL BERMUDIAN POND 




mm 






Fig. 1. The four dominant gastropods of Walsingham Pond (natural 
size). Upper left: Cerithium variabile. Upper right: V ermicularia spirata. 
Lower left: Modulus modulus. Lower right: Nassarius ambiguus. 



y-intercept but not the slope of this ideal weight vs. linear dimen- 
sion curve ) : 

weight = b (length plus width) 3 
Such a curve will plot as a straight line on double logarithmic paper 
with a slope of 3 and a y-value of b at x = 1. This situation is 
closely approached in the eight plots of Figure 2: slopes vary from 
2.57 to 3.53. In three of the four pairs of plots, slopes are similar 
for Pond and control samples (Table 2). The high slope of Wal- 
singham Modulus may be an artifact of a small and incomplete 
size range — see Bohlken, 1962:560. Pond b-values are lower 
than the control parameters in each case (Table 2), but this is not 
significant in itself, since x = 1 mm is too distant an extrapolation 
from the data size range to be biologically meaningful. The im- 
portant point (Fig. 2) is that for each of the four pairs of plots, the 
curve for Pond snails falls below that for control snails over the 
entire measured size range of the samples. (The intersection of the 
two Modulus curves at a size never reached by this species is a 
datum devoid of biological significance; the size range of actual data 
should always be specified in power function plots — see White 
and Gould, 1965.) At any given size for which data are available, 



10 hreviora No. 308 

Pond snails arc lighter than control snails in each of the four 
species. 

Similar modification of at least four species suggests that some 
common factor of Pond ecology is involved as direct or indirect 
cause. Due perhaps to the vast accumulation of decaying organic 
matter in Walsingham. Pond pH is a good deal lower than normal 
marine values. Six measures from various depths made in June 
1967 ranged from 7.6 to 7.7. There is some suggestion in the 
literature (Okland, 1964: 190) that low pH might inhibit shell 
formation even in the presence oi sufficient dissolved calcium in 
the water; most Bermudian waters are supersaturated with respect 
to calcite (Schmalz and Chave, 1963). The lowered pH ma\ 
make it metabolicallv more difficult to deposit available calcium; 
a given expenditure of cnerg\ may therefore produce a thinner 
shell in the Pond than in open marine habitats of normal oceanic 
pll. (We are. indeed, dealing with a deposition phenomenon 
rather than subsequent shell solution in Pond snails; only uneroded 
snails were used in these analyses. | 

TABLE 2 

Reduced Data ior Weight vs. Size Relationships in Four Gastropod 

Species: Comparison of Walsingham and Control Samples. 

Weight b (Height -\ Width ) K 



/»( /;//// ) 

.0123 
.0115 

.0356 
.0348 

.0332 
.0055 

.0303 
.0263 



Nun 


ihcr of 




Specimens 


Slope(l 


I . Cerithium variabile 






Control 


20 


3.24 


Walsingham 


10 


3.14 


2 Vermicularia spirant 






Control 


20 


2.65 


Walsingham 


15 


2.58 


3. Modulus moil ul us 






Control 


20 


3.00 


Walsingham 


13 


3.53 


4. Xassarius ambiguus 






Control 


20 


2.90 


Walsingham 


8 


2.78 



1968 



UNUSUAL BERMUDIAN POND 



11 



300 



100- 



30 



X 

O 



300 




100 



30 






10 20 

H+W 

CERITHIUM 



30 




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H+W 



30 



VERMICULARIA 



300 



100 



30 



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o 




300 



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20 
H+W 

MODULUS 



30 




NASSARIUS 



Fig. 2. Curves of weight (mg) vs. length + height (mm) for the four 
snails of Figure 1, both in Walsingham Pond and in normal marine habitats. 
Regressions marked A are for control snails, B for Pond snails. Regression 
lines extend only over the range of size and weight covered by actual 
specimens. Walsingham curves all lie below control curves, demonstrating 
that Walsingham shells of all species are lighter at any given weight. See 
Table 2 for numerical data. 



12 mrkviora No. 308 

I cannot state whether the thinner shells of these four species 
are a direct phenotypic response to unusual conditions or a result 
of parallel evolution, but I favor the former explanation, which is 
based on both the \outh of Walsingham Pond and general notions 
of simplicity. Extreme phenotypic plasticity in shell thickness has 
long been recognized in land snails. H\ adding ground chalk to 
the diet of Helix aspersa, for example. Oldham ( 1934) found that 
shell weight increased fourfold over snails deprived oi calcium. 

Thus Walsingham Pond serves as a natural experiment in the 
form and composition of moliuscan communities. The unique iso- 
lation of the Pond has no apparent effect upon the presence or 
absence of species but influences, in similar ways, the shell form 
oi its dominant mollusean populations. 

ACKNOWLEDGMENTS 

I am deepl) grateful to a number oi people: to Mr. and Mrs. 
Rutherford, owners ol Walsingham Pond, for their interest, hos- 
pitality and permission to collect on their property; to R. Jensen, 
our leading student oi Bermudian molluscs, for help in identifying 
the micromolluscs (terra incognita for me); to L. Mowbrey, di- 
rector oi the Bermuda Aquarium and Museum, for placing the 
Haycock collection at my disposal; to R. Schmalz for collecting 
the Eh's Harbour sample: to W. Newman for supplying the Long- 
bird Bridge samples; to C. MacClintOCk for loaning the Pembroke 
Marsh sample; to A. C. Neumann for supplying the date on the 
Harrington Sound peat bed; to K. J. Boss for giving me free 
run of the library and collections oi the Mollusc Department 
at the Museum oi Comparative Zoology; and to R. Houston and 
C. Jones for drafting the illustrations. This paper is contribution 
— 437 from the Bermuda Biological Station. 

LITERATURE CITED 

Bon; m n. H. 

1962. Problcme der Merkmalbewertung am Saugetierschiidel dar- 
gestellt am Bcispicl der Bos primigenius Bojanns 1827. Morph. 
Jb.. 103: 509-661. 

Kraser, T. H. 

1967. Contribution to the biology of Tagelus divisus (Tellinacea: 
IMecypoda) in Biscay ne Bay, Florida. Bull. Mar. Sci., 17: 
111-137. 



1968 UNUSUAL BERMUDIAN POND 13 

Gould, S. J. 

In press. Phenotypic reversion to ancestral form and habitus in a marine 
snail. Nature. 

Harris, M., and G. E. B. Morren 

1966. The limitation of the principle of limited possibilities. Am. 
Anthrop., 68: 122-127. 

Okland, J. 

1964. The eutrophic lake Borrevann (Norway) — an ecological study 
on shore and bottom fauna with special reference to gastropods, 
including a hydrographic survey. Folia Limn. Scan., 13: 1-337. 

Oldham, C. 

1934. Further observations on the influence of lime on the shells of 
snails. Proc. Malacol. Soc. London, 21: 131-138. 

Olsson, A. A. 

1951. New Floridan species of Ostrea and Vermicularia. Nautilus, 65: 
6-8. 

SCHMALZ, R. F., AND K. E. CHAVE 

1963. Calcium carbonate: factors affecting saturation in ocean waters 
off Bermuda. Science, 139: 1206-1207. 

White, J. F., and S. J. Gould 

1965. Interpretation of the coefficient in the allometric equation. Am. 
Nat., 99: 5-18. 

(Received 4 June 1968.) 



BREVIORA 

Museumi of Comparative Zoology 

Cambridge, Mass. 31 December, 1968 Number 309 



VARIABILITY OF RODENT INCISOR ENAMEL 

AS VIEWED IN THIN SECTION, AND THE 

MICROSTRUCTURE OF THE ENAMEL IN FOSSIL 

AND RECENT RODENT GROUPS 

John H. Wahlert 

Abstract. The microstructure in enamel of various fossil and Recent 
rodent incisors is described from thin sections. Measurements of enamel 
thickness and inclination of bands in the inner enamel layer were made from 
sagittal sections on large and small samples of Rattus norvegicus and on 
small samples of the Eocene forms Paramys copei and Knightomys depres- 
sus. Statistical analyses of these data show that a sample of size 10 yields an 
adequately close approximation to the population mean. Differences in 
enamel dimensions between upper and lower incisors and between incisors 
of different species are apparent. Thickness and inclination can be used in 
the identification of isolated incisors when data for other rodents have been 
compiled. Band width is of apparently similar utility, but small size pre- 
cludes anything more than rough measurement at 430 diameters magnifica- 
tion. Korvenkontio's "external index" and other measurements taken from 
sagittal sections are deemed unreliable. Rodent species whose enamel has 
been studied are listed according to the kind of enamel they possess. Pau- 
ciserial enamel, found only in Eocene and Oligocene forms, appears to be 
the structural predecessor of uniserial and multiserial enamels, which occur 
in all post-Oligocene rodents examined. At present, the microstructure of 
incisor enamel is useful as an additional character for determining the 
systematic position of rodent higher taxa, but it is no touchstone. 

INTRODUCTION 

That incisor enamel may be useful in the classification of ro- 
dents was first suggested by the work of Tomes (1850) and later 
by that of Korvenkontio ( 1934). Tomes examined the microstruc- 
ture of incisor enamel in a number of living rodents and found that 
within supposedly related groups the arrangement of enamel prisms 
is similar; his descriptions and figures showed that the enamel of 
modern rodents is of two basic kinds. Korvenkontio named these 
uniserial and multiserial. His extension of the research to other 



BKl \ l()k\ 



No. 3(W 









Figure 1. Sagittal sections of enamel in lower incisors: a. Rattus nor- 
vegicus; b. Metaphiomys schaubi; c. Paramys copei. Tips of incisors are to 
the right. Magnification is approximately 240 diameters. Abbreviations: o, 
outer enamel laser, i. inner enamel layer showing hands: d. dentine. 



1968 RODENT INCISOR ENAMEL 3 

modern rodents and to various fossil forms greatly enlarged Tomes 1 
work and led to the discovery of a third kind of incisor enamel 
structure, which he named pauciserial, among early fossil forms. 
In all rodents enamel covers the outer, labial side of the incisors; 
it may extend slightly on the mesial and distal sides. 

Korvenkontio published an extensive table (1934: 116-123) 
giving the dimensions of various parts of the enamel and indices 
derived from these measurements for the incisors of all the species 
he examined. These suggested the possibility that some dimen- 
sions and ratios, at least, might be characteristic of low level taxa. 
If this should prove to be true, it would be a relatively simple mat- 
ter to section, measure, and identify fossil rodent incisors, which 
are so frequently found separated from their jaws. Korvenkontio 
states (1934: 125, footnote) that in his study of 72 Recent and 
33 fossil forms he used about 520 sections. He attempted to make 
sagittal, transverse, and frontal sections of each species wherever 
possible, and thus his sample size for measurement was very small. 

The primary purpose of the work here reported has been to de- 
termine the variability of enamel and the sample size needed to 
obtain meaningful measurements. In the course of the work I 
have had occasion to section incisors of a number of rodents, both 
Recent and fossil. These are included in Table 5, which lists all 
species whose incisor microstructure is known. 

THE KINDS OF RODENT INCISOR ENAMEL 

A sagittal section of uniserial enamel, for example that of Rat t us 
norvegicus (Fig. la), reveals two layers of enamel. The inner 
layer in this aspect appears to consist of bands one enamel prism 
wide that extend outward from the dentine and upward toward the 
tip of the tooth. On close examination each band is seen to be 
divided into small units, somewhat like a string of beads. Study of 
transverse and frontal sections reveals that each band is the cross- 
sectional view of a transverse lamella of enamel prisms. The width 
of bands and prisms is the same in sagittal section; Korvenkontio's 
data show a range of 2.2 to 5.0 microns for uniserial enamels. 
The prisms in a single lamella are parallel; they do not make a 
right angle with the dentine but are at some oblique angle to the 
sagittal plane of the tooth. A band is, therefore, the cross-section 
of prisms comprising a lamella. The prisms of every other lamella 
have the same orientation. There are two kinds of differences in 
orientation of prisms of adjacent lamellae. In transverse section 
the prisms of adjacent lamellae are seen to cross each other at a 
fairly constant angle. Frontal sections reveal that the prisms are 



4 BREVIORA No. 309 

tilted to one side o\' the sagittal plane in ever) other lamella and to 
the other in the alternating set. In the outer layer of the enamel 
the prisms of all lamellae are parallel, and lamination ceases; they 
are usually inclined more steepl) toward the tip o\' the tooth than 
in the inner layer. 

A sagittal section o\' multiserial enamel, lor example that o\ 
Metaphiomys schaubi (Fig. lb), also reveals two lasers of 
enamel. The hands o\ the inner layer are many prisms wide. Kor- 
venkontio found that band widths range from 12 to 30 microns 
for multiserial enamels. Prisms are directed obliquely with respect 
to the direction oi the bands; the obliquity differs in adjoining 
bands but corresponds in alternate ones. Prisms oi the outer layer 
are parallel and more steepl) inclined toward the tip of the tooth. 
as in uniserial enamels. The three-dimensional structure is more 
complicated than in uniserial enamels, and for m\ own work 1 
have relied on Tomes' description ( 1850: 552-53), which 1 para- 
phrase closel\ here: In an oblique transverse section parallel with 
the course o\' the enamel lamellae, the inner enamel layer looks as 
though the prisms were thrown into waxes, the furrows oi which 
commence at the surface of the dentine and. proceeding obliquel) 
outwards, crop out where the prisms become parallel in the ex- 
ternal laser. The prisms pursue a serpentine course in the inner, 
lamelliform portion, where the) describe tolerablx uniform curves. 
B) altering the focus o\ the microscope it max be seen that the 
prisms of adjacent layers pursue a similar serpentine course, but 
are arranged so that the concavities and convexities point in oppo- 
site directions, thus producing a sort oi figure X. 

Pauciserial enamel, for example that oi Paramys copei < Fig. Ic). 
is also divided into two layers. The prisms oi the inner layer may 
or max not be organized into lamellae. In a sagittal section the 
bands of lamellar enamel are seen to be of somewhat variable 
width, depending on the number of prisms included, korxen- 
kontio found that band widths o\' pauciserial enamels fall between 
5 and 16 microns, between the ranges for uniserial and multiserial 
enamels, with slight overlap. The structure appears similar to 
multiserial enamel but lacks the uniformity of organization. Bands 
are usually not inclined. Prisms in the external portion are slightly 
inclined toward the tip of the tooth, though this is not visible in 
the figure. Some pauciserial enamels have fewer prisms per band, 
and thus the bands are narroxver. sugges'ing uniserial enamel in 
appearance. 

These descriptions of the kinds oi enamel seen in sections of 
rodent incisors refer only to the xicinity of the midline of the 



1968 



RODENT INCISOR ENAMEL 



tooth, where the structure is clearly seen. The terms uniserial, 
multiserial, and pauciserial apply, of course, to the inner layer of 
the enamel only. 



MEASUREMENTS AND INDICES 

Sagittal sections are the best for determining what kind of 
enamel is present in a given rodent incisor, and they are the easiest 
to make; therefore, I have considered only measurements made 
from them. 

Total enamel thickness and thicknesses of the two enamel layers 
can be measured. Korvenkontio's external index, which is the per- 
centage of total enamel thickness occupied by the outer layer, 
compares these dimensions. Total enamel thickness is defined as 
the length of a line segment normal to the enamel surface at a 
point, in the vicinity of the midline of the tooth, at which the 
enamel is thinnest. If a sagittal section contains this segment, it 
occurs where the enamel is unworn and thinnest. The geometry 
of rodent incisor enamel distribution, as seen in transverse section 
(Fig. 2a and b), makes underestimation of the total enamel thick- 
ness impossible in a reasonably good section. 




I mm 




Figure 2. Transverse sections of upper incisors: a. Rattits norvegicus; b. 
Paramys copei. Mesial side is to the right. 



6 BREVIORA No. 309 

Band width and inclination oi bands arc also measured best in 
sagittal section. Korvenkontio defined inclination as the angle 
at which bands intersect a perpendicular to the dentine in the sagit- 
tal plane. 1 When bands appear curved, a tangent to the midpoint 
o\ a band is used to determine inclination. Prisms in the external 
laser also have an inclination that can be measured; this is not 
visible in most sections, however, and I have not considered it. In 
this paper 1 use enamel thickness, external index, band width, and 
inclination in the internal enamel laser according to Korvenkon- 
tio's definitions for sagittal sections. 

MATERIALS 

[ncisors ol 48 individuals from a highl) inbred strain of albino 
Rattus norvegicus were available to me at the beginning of this 

project. Ihis is a large sample tor statistical analysis by paleon- 
tological standards. I he heads had been preserved in formalde- 
hyde, and sexes were not recorded. Incisors of Paramys (<>i>ci and 
Knightomys depressus were also available m lair quantity. I hese 
had been collected by Amherst College parties from the earl\ 
I ocene I ysite member of the Wind River Formation in the Wind 
River Basin. Wyoming. Individual incisors from a variety ol 
rodents were also sectioned; specimens were obtained from the col- 
lections of Amherst College, Albert E. Wood, the Museum of 
Comparative Zoology, the University ol lexas. ami the Yale Pea- 
bod\ Museum. 

PROCEDURE 

Standard sectioning techniques were used. All lour incisors 
from each rat were sectioned on one petrographic slide. Fossil 
incisors a\k\ individual modern incisors were sectioned singly. 
Preliminar) grinding was done on a water-bathed 600 grain dia- 
mond wheel that turned at approximate^ 100 rpm. Sections were 
hand held. Final polishing was done with wet 900 gram alundum 
on wood, and then sections were etched briefly with dilute hydro- 
chloric acid; this step makes the enamel structure visible. When 
sections of modern teeth become thin, water absorption causes 
them to buckle. This difficulty was solved by drying the section 
before it became thin and then impregnating it with mineral oil 
which acts as a waterproofing agent. 

■Tomes (1850) measured inclination as the angle made b) bands and 
the dentine surface. 



968 



RODENT INCISOR ENAMEL 



7 



Thickness measurements were made by comparing the projection 
of a thin section with a micrometer slide. The number of bands 
within a standard unit of a micrometer eyepiece divided by the 
length of the unit in millimeters yields the average band width. 
Inclination was measured on the rotating stage of a petrographic 
microscope. In each case 430 diameters magnification was used. 

Tabulation and statistical analyses of the measurements are 
given in Tables 1-4. The formulae used in computation are those 
given by Simpson, Roe, and Lewontin (1960: 84, 90, 166). Ab- 
breviations are as follows: N, sample size; OR, observed range; 



% 3 




.080 .090 .100 



140 .150 .160 



% 50 



40 



30 



20 



10 




upper 



— lower 



Figure 3. Frequency distributions for incisors of Rattus norvegicus'. a. 
total enamel thickness; b. enamel prism inclination. 



8 BREVIORA No. 30 l ) 

X, arithmetic mean: Conf. Int.. confidence interval of the mean; 
SD. one standard deviation unit; V, coefficient of variation. In all 
thickness measurements there is an error in estimation of ±.001 
mm; this error is inherent in the mean. Frequency distributions for 
enamel thickness and inclination of bands in the large sample of 
albino laboratory rats appear in Figure 3a and b. 

Although all four incisors of forty-eight rats were sectioned, not 
all individual sections were usable, and thus the sample size varies 
slightly. Inclination was not measured in Paramys copei and 
Knightomys depressus because prism bands are not well defined 
and are nearly perpendicular to the dentine surface. The sample 
M/e in both cases was too small for frequenc\ diagrams to be 
constructed. Measurements of band widths were, at best, crude, 
because boundaries are not well defined and magnification was low 
for such small objects. Consequently, I present only average width 
and range under the appropriate headings. 

TABLE 1 
Data for albino laboratory rat incisors. 



total enamel thickness (in mm) 
















959! 






Ineisoi N 


OK 


\ 


< onf. Int. 


SD 


V 


Upper 9ft 


.080-. 126 


ii')-* 


±.002 


.009 


X.7 


Lower 93 


.100-. 155 


.129 


±.003 


.012 


9.1 


Inclination of enamel prisms 


( in decrees ) 










Uppei 95 


23-42 


36 


±.6 


3 


8.3 


Lower 93 


J2-51 


46 


±.6 


3 


6.1 



DESCRIPTION OF SAMPLES 

Rutins norvegicus (laboratory strain). The incisor enamel is 
uniserial (Fig. la), and the enamel thickness is greater in the 
lower than in the upper incisors o\ each individual rat. although the 
ranges of total enamel thickness for upper and lower incisors over- 
lap appreciably (Table 1 ). In only one specimen were the meas- 
urements within the error o\ estimation of each other, the enamel 
thickness of the upper right incisor being .126 mm and of the 
lower left. .127 mm. The overlap of ranges shows that it is unsafe 
to decide whether upper or lower incisors have the thicker enamel 
on the basis of only one or two specimens. The frequency distri- 
bution is graphed in Figure 3a. 



1968 RODENT INCISOR ENAMEL 9 

The 95 per cent confidence interval for the mean is quite small 
in relation to the total thickness. The standard deviations are 
greater for lower incisors than for uppers. The coefficients of varia- 
tion are high for a Recent species (Simpson, Roe, and Lewontin. 
1960: 92). This may be accounted for by deviation of the section 
from the sagittal plane and also by natural variation, such as effects 
of age, sex differences, and inbreeding, none of which can be ex- 
cluded as a factor contributing to a high V. The distribution of 
enamel on the front of an upper incisor as seen in transverse sec- 
tion (Fig. 2a) is uneven; thus, deviation of sections from the 
sagittal plane could be the cause of considerable variation in thick- 
ness measurements. The enamel distribution is more uniform in 
lowers. Separate analysis of left and right incisors yielded the 
same mean in lowers and a difference of means in uppers that is 
within the estimated range of observational error. 

The number of incisors of a single extinct species collected from 
one locality is usually small, on the order of ten or so. To test the 
reliability of enamel thickness measurements for small samples, I 
considered the data for forty-eight rats as representing an entire 
population with a known mean and took from it ten random 
samples of ten upper and of ten lower incisors. The results of the 
analyses are presented in Table 2. Although the means of these 
samples may be as much as five thousandths of a millimeter from 
the mean of the population, the ninety-five per cent confidence in- 
tervals of the means for all but one sample (upper incisors, sample 
8) include it. This same sample has both the highest mean thick- 
ness and the least variability. With any two samples representing 
upper and lower incisors, the relative difference in enamel thick- 
ness is apparent; the smallest difference in mean enamel thickness 
obtainable from these results amounts to .020 mm and the largest 
is .039 mm. 

Attempts to measure the thickness of either layer of the enamel 
separately were unsatisfactory, because there is no clear line of 
demarcation between the two. Figure 1 shows how bands of the 
inner layer project into the outer; it is impossible to establish a 
definite line that is consistently placed in each thin section. In the - 
sample the outer enamel layer occupies approximately 30 per cent 
of the mean total thickness in uppers and 19 per cent in lowers. 
The observed ranges are 20 to 39 per cent for uppers and 14 to 27 
per cent for lowers. In two individuals the external index of an 
upper and a lower incisor is the same, but in no instance in an 
individual is that of the lower greater. A regression analysis of 
these measurements of outer layer thickness versus total enamel 



10 



HKI-.VIORA 



No. 31W 



TABLE 2 

Variability of total enamel thickness (in mm) in ten samplings, 
each of ten incisors selected at random from data tor albino labora- 
tory rats. 



Upper incisors 




95 






S.uiip. 


OK 


\ 


( onf. Int. 


SI) 


V 


1 


085-.120 


101 


±.007 


.010 




: 


080 126 


.099 


±.009 


01 i 


13.1 


) 


085 .110 


.097 


±.006 


008 


8.2 


4 


OS5-.IK) 


099 


±.005 


.007 


7.1 


5 


092-.109 


.099 


±.004 


,006 


6 I 


'• 


094-.106 


100 


±.003 


.004 


4.0 


7 


080-.1 12 


095 


±.006 


.009 


9 s 


8 


[00 .107 


.104 


±.001 


.002 


1.9 


9 


085-.126 


.102 


±.009 


.013 


12.7 


in 


092-.1 10 


101 


±.004 


.006 


5 9 


1 owei incisors 










1 


1I0-. 155 


.129 


±.01 1 


015 


1 1.6 


: 


1 I9-.153 


.134 


±.008 


.01 I 


8.2 


5 


1 10-143 


.127 


±.007 


.010 


7.9 


4 


[07- 143 


.128 


±.007 


.010 


7.S 


5 


11 1-.138 


.127 


±.007 


Din 


7.9 


6 


II 4-. 153 


.129 


±.008 


.01 1 


8.5 


7 


I1I-.I45 


.124 


±.007 


.010 


x.l 


8 


I07-.I5O 


.131 


±.01 1 


.015 


11.4 


9 


1 07-. 138 


.126 


±.008 


.01 1 


8.7 


10 


1 10-.145 


.129 


±.008 


.012 


9 ; 



thickness was carried out within upper and lower incisors of the 
entire sample. The correlation coefficient for upper incisors is .27; 
for lowers. ,18. This is extremely poor, but the gross difference 
between uppers and lowers remains. 

Band width in sagittal section appeared ncaii\ invariable at 450 
diameters magnification. It is .0033 for both upper and lower in- 
cisors. The mean inclination of enamel prisms in the inner layer 
is distinctly greater in nearly all lower incisors (Table 1. Figure 
3b). However, in one specimen the inclination of bands in ov^- 
lower incisor was less than that of the uppers; in another, an upper 
and a lower incisor yielded the same angle. Thus, two teeth, even 



I 




95% 






OR 


X 


Conf. Int. 


SD 


V 


088-.115 


.104 


±.004 


.008 


7.7 


065-.105 


.088 


±.004 


.009 


10.2 



1968 RODENT INCISOR ENAMEL 11 

from the same individual, may not indicate the usual relationship. 
The standard deviation of inclination is the same for both upper 
and lower incisors. 

TABLE 3 
Data for Paramys copei incisors. 

Total enamel thickness (in mm) 
Incisor N 

Upper 1 6 

Lower 24 

Paramys copei. The sample of incisors of Paramys copei from 
the Lysite represents a minimum of twelve individuals, all but two 
of the teeth being from the same locality. Enamel is pauciserial. 
In this species the distribution of enamel across the face of the 
tooth as seen in transverse section is fairly even (Fig. 2b). The 
error caused by the deviation of a section from the mid-plane is 
therefore not large when compared to the same situation in rat 
incisors. 

The data presented in Table 3 indicate that the mean enamel 
thickness in upper incisors is .016 greater than in lowers, the re- 
verse of the situation in rats. There is overlap in ranges, and 
examination of only a few teeth could show the opposite relation- 
ship. The standard deviations for upper and lower incisors are 
similar to the figure for upper incisors of rats. The sample is too 
small to construct a meaningful frequency distribution curve. 

The mean band width and observed range are .017 mm and 
.01 3-. 021 mm in upper incisors and .016 mm and .014-.023 mm 
in lowers. Inclination of bands is approximately zero. In any in- 
dividual tooth, bands may be slightly inclined toward or away from 
the tip. 

TABLE 4 
Data for Knightomys depressus incisors. 

Total enamel thickness (in mm) 95% 

Incisor N OR x Conf. Int. SD V 

Upper 13 .103-.150 .120 ±.009 .016 13.3 

Lower 6 .053-.095 .068 ±.005 .005 7.35 

Knightomys depressus. The sample of Knightomys depressus 
from the Lysite represents a minimum of seven individuals. Enamel 
is pauciserial. The data presented in Table 4 show that the mean 



12 BREVIORA No. 309 

enamel thickness is much greater in upper incisors and that the 
ranges for upper and lower incisors o\o not overlap. The lack of 
overlap mav simplv mean that the sample size is too small. The 
confidence interval, standard deviation, and coefficient oi variation 
tor the lower incisors seem low with regard to these figures tor the 
other samples. 1 hey are in best agreement with those tor small 
samples of rat upper incisors 

I he distribution of enamel is similar to that in Paramys copei. 
1 he mean hand width and observed range are .014 mm and .012- 
.017 mm in upper incisors and .012 mm and .009-. Old nun in 
lowers. Inclination oi bands is approximate!) zero, and the boun- 
dary between inner and outer layers of enamel is again not a 
distinct line. 

SAMPLE SIZE AND THE UTILITY OF MEASUREMENTS AND 

INDICES AS CRITERIA FOR IDENTIFICATION 

OF ISOLATED INCISORS 

A sagittal thin section of a rodent incisor is sufficient to show 
whether the enamel is pauciserial, uniserial. or multiscrial. A 
lew sections provide a rough measure of enamel thickness. A 
sample of ten or more upper and ten or more lower incisors yields 
a mean enamel thickness with a fairly small confidence interval 
and shows the relationship oi thicknesses. A sample o\ about a 
hundred individual upper and lower incisors is needed for graphing 
a frequency distribution oi thickness. The inclination oi bands in 
the inner enamel layer also vields a continuous curve when plotted 
lor a sample oi this size. I lie inclination oi prisms in the external 
layer is usually not visible. A rough measure of band width mav 
he made at a magnification oi 430 diameters, but variation is not 
apparent. 

The boundary between internal and external enamel layers is not 
a line, and thus the thickness oi either part cannot be measured 
precisely. This variability and the lack of correlation between 
thickness oi the outer layer and total enamel thickness within a 
sample suggests that a ratio of the two, Korvenkontio's external 
index, cannot be used. 

At present there are few numerical data available concerning the 
microstructure oi rodent incisor enamel. Identification of isolated 
incisors cannot be made on the basis of measurements alone. 

DISCUSSION OF INCISOR ENAMELS 

Pauciserial enamel was found by Korvenkontio to be present 
in ischyromyid. sciuravid. pseudosciurid. and some theridomyid 



1968 RODENT INCISOR ENAMEL 13 

rodents. I have sectioned incisors of some of the same Eocene 
species; these and thin sections of different ischyromyid and sciura- 
vid species reveal pauciserial enamel. Prosciurus relict us, a mid- 
Oligocene ischyromyid has uniserial enamel. Korvenkontio found 
a condition transitional between pauciserial and uniserial in the 
Oligocene theridomyid, Nesokerodon minor. 

Completely uniserial and multiserial enamels are first met with 
in incisors of early Oligocene rodents. Korvenkontio found uni- 
serial enamel in species of this age in the Theridomyidae and 
Ischyromyidae, and I have observed it in Cylindrodon fontis of the 
Cylindrodontidae. Other species having uniserial enamel are listed 
in Table 5. 

Multiserial enamel is characteristic of all caviomorph rodents so 
far examined. Wood and Patterson (1959: 292)- found it in a 
Deseadan (early Oligocene) incisor, very probably of Scotamys 
antiquus, and I have found it in Cephalomys arcidens from deposits 
of the same age. The early Oligocene African phiomyid rodents 
sectioned by me had acquired multiserial enamel; thin sections of 
incisors of Phiomys andrewsi, Metaphiomys schaubi, and a new 
species (Wood, in press) demonstrate this. The Recent Thryono- 
mys and Petromus, which Lavocat (1962) and Wood (in press) 
associate closely with the phiomyids, also prove to have multiserial 
enamel. Other species possessing multiserial enamel are listed 
in Table 5. 

Pauciserial enamel is a good structural predecessor for uniserial 
and multiserial enamels. It would appear to be the ancestral con- 
dition. As far as present evidence goes, it had probably been 
achieved by the time rodents made their first appearance in the 
record. It is known thus far in only a few post-Eocene rodents, 
and uniserial and multiserial enamels have not yet been recorded 
prior to the Oligocene. Presumably acquisition of two layers in 
which the organization of prisms is different served in some way to 
strengthen the enamel. The pauciserial structure, judging from the 
record, would appear to have been less efficient than the other two. 
There is no available evidence to suggest that one of the two 
surviving kinds is superior to the other or that one of them has ever 
evolved from the other. Seemingly, selection has favored increase 
in strengthening rather than any one method of accomplishing it. 

To the possible question: could the pauciserial condition be an 
artifact caused by diagenetic changes in materials of greater age? 



2 A lapsus in this paper may be corrected. The authors state (p. 292 n.): 
". . . the Theridomyidae are in the process of passing from a pauciserial to 
a multiserial type." For "multiserial" read "uniserial." 



14 BREVIORA NO. 309 

I would repl) in the negative. In none of the many slides of 
enamel of I ocene age that I have examined is there an\ indication 
that recrystallization has disrupted the tine structure. 

The enamels of all modern rodent incisors tall into one of the 
two major categories, uniserial and multiserial, but there are minor 
differences within each of the two as regards dimensions and orien- 
tation o\ lamellae. Stud) of the internal detail of lamellar structure 
is needed to understand the differences and to determine how the) 
ma) have evolved. 

ACKNOWLEDGMENTS 

I began this research at Amherst College and received support 
in 1964-65, through the Ceolog\ Department, from the Student 
Research Participation Program of the National Science Founda- 
tion. The work was continued in the summer o\ 1965 under Pub- 
lie Health Service (.rant 2 II (.\1 M)^-()h to the Biolog) Depart- 
ment of Amherst College, and other help was forthcoming from 
National Science Foundation Granl GB l c >77 to Professor Albert 
E. Wood. I am indebted to Professor Wood and to ProfeSSOl 
Bryan Patterson for their guidance, and to Miss Barbara Lawrence 
and Mr. Charles Mack of the Mammal Department. Museum of 
Comparative Zoology, for permitting me to section various speci- 
mens. 

TABLE 5 

I name] structure in rodent incisors. 

["he nun. ils aftei each species name .ne those of the authors who have 
described its incisoi enamel Species names are mainl) as in I Herman 
< 1441 ): familial and generic assignments are mainl) as in Simpson i 1945 I. 
Where a name given in the publications of those who have described rodent 
incisor enamels differs from the current one. I have included it in brackets. 
\ classification ol rodents above the familial level is given b) Woo. I i 1958) 

Abbreviations: B., Bohlin (1946); k. Korvenkontio (1934); I., lomes 
i I smi i ; \V . Wahlert; W. and P.. Wood and Patterson ( 1959). I OC., I ocene: 
Olig., Oligocene; Mioc, Miocene; Pleist., Pleistocene; I., early; \1 . mid; 
L., late. \f. Africa; Vs., Asia; In. Europe; N V., North America: S.A.. 
south America. (Geologic age and location given only for fossils.) 

I' \l ( isl RIAL ENAMELS 

lachyromyidae: Ischyrotomus petersoni W. L.Eoc. N.A. Knightomya de- 

pressusW., E.Eoc. N.A. Manitsha sp. W.. E.Olig. N.A. Microparamys 
lysitensis W., E.Eoc. N.A. Paramys c. copei K.. \\ I .Eoc. N.A. Paramys 



1968 RODENT INCISOR ENAMEL 15 

c. major K., W., E.Eoc. N.A. Paramys delicatior K., M.Eoc. N.A. 
Paramys excavatus K., W., E.Eoc. N.A. Thisbemys perditus W., E.Eoc. 

N.A. 

Sciuravidae: Mysops parvus W., M.Eoc. N.A. Sciuravus nitidus K., M.Eoc. 

N.A. 

Theridomyidae : Archaeomys gracilis K., L.Eoc.-M.Olig. En. Archaeomys 
major K., L.Eoc.-M.Olig. Eu. Theridomys gregarius K., L.Eoc.-M.Olig. 
Eu. Theridomys vaillanti K., Eoc. Eu. 

Pseudoseiuridae: Pseudosciurus suevicus K., E.Olig. Eu. Sciuroides quercyi 
K., L.Eoc.-M.Olig. Eu. Sciuroides sp. K., L.Eoc.-M.Olig. Eu. 

UNISERIAL ENAMELS 

Ischyromyidae: Ischyromys typus K., M.Olig. N.A. Prosciurus relictus W., 
M.Olig. N.A. Titanotheriomys veterior K., E.Olig. N.A. 

Cylindrodontidae: Cylindrodon j otitis W., E.Olig. N.A. 

Aploduntidae : Allomxs nitens K., L.Olig. N.A. Aplodontia rufa K. 

Myiagaulidae : Mesogaitlus novellus W., M.Mioc. N.A. Mylagaulus sp. W., 
L.Mioc. N.A. 

Muridae: Acomys sp. K. Arvicanthis sp. K. Conilurus [Hapalotis] albipes 
T. Hydromys chrysogaster T. M//.v. musculus K. Notomys [Hapalotis] 
longicaudatus T. Otomys sp. K. Rattus norvegicus [Mus decumanus T.. 
KJ T., K., W. /?a«HS /"M/w) ra««J K. 

Cricetidae: Cricetodon minor K., L.Mioc. Eu. Eumys elegans K., W., 
M. -L.Olig. N.A. Eumys gracilis K., Olig. N.A. Sigmodon sp. W., L.Pleist. 
N.A. Akodon arenicola K. Arvicola amphibius T. Arvicola [Microtus] 
terrestris K. Clethrionomys [Arvicola T., Evotomys K.] glareolus T., K. 
Cricetulus migratorius [phaeus] K. Cricetus cricetus [fumentarius T.] T., 
K. Ellobius talpinus K. Gerbillus sp. K. Lemmus lemmus [norvegicus] T., 
K. Meriones [Gerbillus] shawi T. Microtus [Arvicola] nivalis T. Myopus 
schisticolor K. Ondatra [Fiber] zibetluca T., K. O/'v^o'H}'^ flavescens 
[longicaudatus] K. Oxymycterus rufus K. Phyllotis [Hesperomys] darwinii 
T. Tatera sp. K. 

Spalacidae: Spalax micro phthalmus [typhlus] T., K. Tachyoryctes splen- 
dens K. 

Geomyidae: Entoptychus cavifrons K., E.Mioc. N.A. Entoptychus sp. K., 
L.Olig. N.A. Entoptychine geomyid W. M.Mioc. N.A. Heterogeomys 
hispidus K. Thomomys [Geomys] umbrinus T. 

Heteromyidae : Heteromyid W.. M.Mioc. N.A. Dipodomys sp. K. Pero- 
gnathus fasciatus K. Perognathus sp. K. Perognathus [Cricetodipus] sp. K. 



16 BREVIORA NO. 309 

Eomyidae: Adjidaumo sp. \\ '.. M.Olig. N.A. Paradjidaumo trilophus W.. 
M.Olig. N.A. 

Dipodidae: Alactaga sibirica [saliens] k. Jaculus jaculus [Dipits hirtipes] 
k. Jaculus [Jerboa T., />//>/n K.J orien talis [aegyptius] T. K. 

Zapodidae: Napaeozapus insignis k. Sicista [Sminthus] subtilis K Zapus 
hudsonius k. 

Cliridae: C/h [Myoxus] wetzleri k. L.Olig Eu. Dryomys nitedula k 
Eliomys quercinus k. G/u [Myoxus k./ g/1'5 k . W. Graphiurus ocularis 
[capensis] k Muscardinus [Myoxus 1/ avellanarius I.. k 

Seiaridae: Sciurus feignouxi k.. E.Mioc, Eu. Callosciurus [Sciurus] pre- 
vosti k. Citellus [Spermophilus] eversmanni k. Citellus parryii [Spermo- 
philus empetra] k. Citellus [Spermophilus] sp. I. ( ynomys ludovicianus 
k. Harmota caligata [Arctomys pruinosus] I Marmoia [Arctomys] 
monax [empetra I / I., k. Marmota I Arctomys k./ sp. k.. W. Petaurista 
volans [Pteromys russicus] k. Ratufa [Sciurus] macroura k. Sciurus 
niger V. Sciurus vulgaris k. lamias sibiricus [Eutamias asiaticus] k. 
Tamias striatus [lysteri 1/ I . k 

Castoridae: Steneofiber eseri K., 1 Mi<x Eu Steneofiber peninsulatus K 
1 .Olig. N.A. Cas/o/ flfter I . k. 

Eutypomyidae: Eutypomys thomsoniVf., M. Olig N V 

Theridomyidae: Archaeomys laurillardi K., Olig Eu Wesokerodon minor 
k. I Eoc M Olig. I u Sciuromys typicus k . I .Eoc.-M.Olig. Eu. 

Inomalnridae: Anomalurus fraseri k. Idiurus macrotis k. 

Ml I I ISI KIM IN \MI 1 S 

Oetodontidae: Sciamys principalis k.. I Mioc. S \ iconaemys [Schizo- 
don] fuscus I Octodon degus I Spalacopus cyanus [poeppigii] T. 

Echimyidae: tdelphomys candidus K., 1 Mioc S.A \4yocastor [Myopota- 
mus T.] coypus I k 

Ctenomyidae : ( tenomys magellanicus k 

kbrocomidae: Abrocoma [Habrocoma] bennettii I 

Capromyidae: Capromys pilorides [fournieri] T. 

Chinehillidae: Scotamys antiquus W. and P., E.Olig. S.A. Perimys pro- 
cerus K.. E.Mioc. S.A. Chinchilla laniger T. Lagostomus maximus [tri- 
chodactylus] k. 

Dasyproctidae: Cephalomys arcidens W.. E.Olig. S. \ Seoreomys australis 
k.. E.Mioc. S.A. Cuniculus [Coelogenys] paca T., K. Dasyprocta aguti 

V.. k. Myoprocta I Dasyprocta] acouchx T. 



1968 RODENT INCISOR ENAMEL 17 

Caviidae: Cavia aperea T. Cavia porcellus [cutleri] K. Dolichotis pata- 
gona K. Galea [Kerodon] ftavidens K. Galea [Kerodon] spixii K. Micro- 
cavia australis [Cavia kingii] T. 

Hydrochoeridae : Hydrochoeris hydrochaeris [capybara] T. 

Erethizontidae: Coendou [Hystrix T.] prehensilis T., K. Erethizon dorsa- 
tum K. Erethizon epixanthum K. 

Ctenodactylidae : Sayimys obliquidens B., ?Mioc. As. Tataromys cf. plici- 
dens B., ?Mioc. As. Ctenodactylus gundi K., B. 

Pedetidae: Pedetes cafer T., K. 

Hystricidae: Atherurus africanus K. Hystrix cristata T. 

I'hiomyidae: Metaphiomys schaubi W., E.Olig. Af. Phiomys andrewsi 
W., E.Olig. Af. Phiomyid W., E.Olig. Af. 

Thryonomyidae: Thryonomys sp. W. 

Petromuridae : Petromus ty picas W. 

Bathyergidae: Bathyergns suillus [maritimus] T. Cryptomys mellandi K. 
Georychus capensis K. 



REFERENCES 

BOHLIN, B. 

1946. The fossil mammals from the Tertiary deposit of Taben-buluk, 
Western Kansu. Part II. Sino-Swedish Exp., VI: 4 (Pal. Sin. 
New Ser. C. No. 8 b),259 pp. 
Ellerman, J. R. 

1940- The Families and Genera of Living Rodents. British Mus. (Nat. 

1941. Hist.), 1: 1-689; 2: 1-690. 

KORVENKONTIO, V. A. 

1934. Mikroskopische Untersuchungen an Nagerincisiven, unter Hin- 
weis auf die Schmelzstruktur der Backenzahne. Ann. Zool. Soc. 
Zool.-Bot. Fennicae Vanamo, 2: i-xiv, 1-274. 
Lavocat, R. 

1962. Reflexions sur lorigine et la structure du groupe des rongeurs. 
In Problemes actuels de paleontologie (evolution des vertebres). 
Colloq. Internat. Centr. Nat. Rech. Scient., 104: 287-299. 
Simpson, G. G. 

1945. The principles of classification and a classification of mammals. 
Bull. Amer. Mus. Nat. Hist., 85: 1-350. 
Simpson, G. G., A. Roe, and R. C. Lewontin 

1960. Quantitative Zoology. New York, Harcourt, Brace and Co., 
440 pp. 



IS BREVIORA No. 309 

loMI S, J. 

1850. On the structure of the dental tissues of the order Rodentia. 
Phil. [Vans. Roy. Soc, London, 1850: 529-567. 

W I I ID, \ I 

195S. Arc there rodent suborders? Syst. Zool., 7: 169-173. 
In press Rodentia. In: 1. I. Simons, ed., Earl) Cenozoic mammalian 
faunas. Fayum Province Egypt, United Arab Republic. Pan II. 

\\ >. \ I . \m» B. Patti rson 

1959 I he rodents of the Deseadan Oligocene of Patagonia and the 
beginnings of South American rodent evolution. Bull. Mus. 
( omp Zool . I2<> 281-428. 

I Received 1 5 August 1968 I 



BREVIORA 

MuseiMin of Comparative Zoology 

Cambridge. Mass. 31 March, 1969 Number 310 



A SHOVEL-TUSKED GOMPHOTHERE 
FROM THE MIOCENE OF KENYA 

Vincent J. Maglio 



Abstract. A tusk fragment of the first recorded occurrence in Africa of 
Platybelodon, a shovel-tusked gomphothere, is described, and its relation- 
ships to the Asiatic and North American representatives of the Amebelo- 
dontinae are discussed. It is suggested that the African form represents a 
very early member of the group, and that Africa may have been the place 
of origin of the subfamily. 

INTRODUCTION 

In 1927, E. H. Barbour described as Amebelodon fricki the 
first specimen of a rather bizarrely specialized group of probosci- 
deans, the shovel-tusked gomphotheres — Amebelodontinae. The 
geological and geographical range of this subfamily has hitherto 
been believed to extend from the late middle Miocene (Tortonian) 1 
of Asia to the late Pliocene of North America. Osborn (1936) 
proposed a subfamilial separation of the North American Amebelo- 
don from Platybelodon, which occurs in both North America and 
Asia, suggesting an independent origin of the two genera — 
Amebelodon in North America from a stock of the "Bunomasto- 
dontidae" and Platybelodon from the "Serridentidae" in Asia. 
Gregory (1945) considered the two genera to represent a more 
closely related group than did Osborn and placed them both in a 
single subfamily, the Amebelodontinae. This is the view sub- 
scribed to here. Until now, it has seemed likely that the shovel- 
tusked gomphotheres originated in Asia from a stock of the 
Gomphotheriidae sometime during the early to middle Miocene, 



1 Platybelodon danovi Borissiak, the earliest known species, occurs in the 
Chokrak beds of the Kuban region in the North Caucasus, considered to be 
of late middle Miocene age (Tortonian of western Europe) by Nalivkin 
(1960). 



2 BREVIORA No. 3 10 

migrating to North America during the early Pliocene. The speci- 
men described in the present note, however, demonstrates the 
presence in east Africa during middle Miocene time of shovel- 
tusked gomphotheres very similar to the Asiatic amebelodontines. 

The fragment (MCZ field number 3S-64K) was collected by 
Mr. James (i. Mead during the 1964 field season of the Museum 
of Comparative Zoology expeditions to east .Africa- directed by 
Professor Bryan Patterson. I he locality is 35 50' E, 02 20' N. 
approximated 4 mi. NM of the site of the abandoned town of 
Loperot and 50 mi. SSI- of I.odwar. in the Loperot area of southern 
Turkana District, northwestern Kenya (map in Joubert, 1966). 
Although none of the associated fauna has yet been studied in 
detail, the following elements have been identified: Deinotherium 
hobleyi, Dorcatherium sp. cf. D. pigotti, Chilotherium sp., 
?Brachyodm sp.. and hyracoid aff. Prohyrax. 

The specimen, weathered from the Turkana Grit, was found 
on the surface near the top of the formation, about 20 feet below 
a fine-grained olivine basalt sequence. Joubert recognized three 
successive basaltic Hows overlying the Turkana Grit in this region 
and on geological evidence placed the onset of the basal flow in 
the lower Miocene (1966: 35). I he associated fauna collected 
from the Turkana Grit is not wholl) incompatible with this view, 
in that it appears to be close to those in the Rusinga scries and. 
although less so. to that of l.otludok. both usually considered to 
be of earl) Miocene age. 

Although detailed faunal comparison is not possible at present. 
the Loperot fauna probabl) correlates with part o\' the Rusinga 
complex'; it docs not appear to be equivalent to that of Lothidok. 
Dorcatherium pigotti and Chilotherium s/>. occur on Rusinga Island. 



-The field work was supported in National Science Foundation Grant 
GP-l I ss 

:! The name Losodok as used b\ Arambourg (1933), Whitworth (1954). 
and Hooijer (1966), is replaced b) the name Lothidok on most current geo- 
logic maps of the area. as. for example. SK. 57 Kenya. I:IM (Special). 
Survej of Kens a. 1965. 

4 As Simpson has recently emphasized (I9(S7: 43). the stratigraphic and 
faunal successions at Rusinga are at present inadequate!) knovvn. as are the 
facies and age relationships of the several other earlier Miocene faunas 
known from the Kavirondo Gulf region. Evidence in support of the often 
quoted "earl) Miocene" age. or indeed of any other age. for these deposits 
is inconclusive, although the) are almost certainly older than those at Fort 
Ternan. for uhich an acceptable age (12-14 million years) is available. The 



1969 SHOVEL-TUSKED GOMPHOTHERE FROM KENYA 3 

The hyracoid is close to, but lower crowned than, Prohyrax from 
Langental, South Africa (Stromer, 1926), and may be the same 
as Meroehyrax from Rusinga (Whitworth, 1954). The Lothidok 
fauna includes Dorcatheriwn chappuisi, Megalohyrax championi 
(see Whitworth, 1954), and Aceratheriwn acutirostratum. 

The lowest basalt in the Loperot sequence has been dated at 
17.5 ± 0.9 million years by Geochron Laboratories, Inc., using 
the Potassium/ Argon method (Patterson, personal communica- 
tion). This date indicates a Vindobonian age for the basalt im- 
mediately overlying the fossil-bearing sediments. There does not 
appear to be any significant time interval between deposition of 
the upper part of the Turkana Grit and the basalt flow, and this 
age is here considered approximately correct. 

The specimen under discussion is too fragmentary for positive 
identification, but, as far as can be determined from what remains, 
the form it represents was closely related to Platybelodon. It is 
therefore tentatively placed in that genus, pending further knowl- 
edge. 

PLATYBELODON SP. 

The specimen (Fig. 1 ) is a fragment from the middle of a left 
lower incisor, and measures 56.8 mm in maximum length. The 
upper surface is broadly concave and the lower convex. The two 
surfaces are not parallel but diverge somewhat towards the sides, 
so that the tooth is thicker along the medial and lateral edges than 
at the center. The medial edge is flattened, with a broad, shallow, 
concave groove along the entire length of the fragment; the two 
lower incisors may have been closely appressed during life, at 
least in this region of the tusk. The specimen tapers in all dimen- 
sions toward one end, which is probably anterior. A similar taper- 
ing toward the tip of the tusk is seen in more complete specimens 
of other amebelodontines, e.g., the specimen of Amebelodon fricki 

age of 15.3 million years suggested by Evernden ct al. (1964) as the maxi- 
mum for the Rusinga series seems unlikely. The sample used for this de- 
termination was taken from a locality at the base of Kiahera Hill and, as 
suggested by Savage (1965) and Leakey (1967), these deposits may not be 
part of the Kiahera Series, supposedly the oldest formation on Rusinga 
Island. Until the deposits of Rusinga are better understood, correlation 
within the east African Miocene will remain uncertain. 

The relative position of the various deposits within the Miocene depends 
in part on the inclusion in or exclusion from this epoch of the Aquitanian 
and Pontian stages. 



BRt A IOR\ 



NO. 3 1 



described b\ Gregor) ( L945). The medial thickness tapers from 
14.6 mm to 11.6 mm anteriorly. The lateral edge is thinner than 
the medial and measures S.5 mm to 7.0 mm. In overall width, the 
fragment decreases from 78.5 mm to 74.5 mm. 





re 1. Platybelodon sp. Tusk fragment, NICY field number 3S-f>4K. 
from the Loperoi area, Kenya, a, Dors.il view; horizontal line indicates 
plane of section, ft, Nnterior view of cut surface. Shaded area, investment 
of laminated dentine. • 1. 



The broken ends reveal a complex internal structure of dentine 
organized into dentinal system^ or rodlike masses as in most other 
amebelodonts. although they have not weathered out as discrete 
and isolated rods, such as those described in Platybelodon grangeri 
(Osborn and Granger. 1931) and Amebelodon jricki (Barbour. 
i941 ). In thin section (Fig. 2). the dentine has a fibrous texture, 
consisting of numerous microscopic dentinal tubules. These tubules 



5 The term dentinal system was restricted by Tomes ( 1850) to that portion 
of a tooth consisting of a canal from which dentinal tubules radiate and 
around which there is no investment of enamel. 



1969 



SHOVEL-TUSKED GOMPHOTHERE FROM KENYA 




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d BRE\ [ORA No. 3 10 

appear to radiate from the center ol each dentinal system and ma\ 
curve partially around its outer edge. Near the center o! each 
dentinal system is a round to irreguiari) shaped canal, secondaril) 
filled with calcite, as are the numerous small cracks which cut 
across the tubule boundaries. I he canals probabl) correspond to 
the "small lumen" noted in the dentinal s\stems ("rod-cones") of 
Platybelodon by Osborn ( 1936: 465). The tooth is thus com- 
posed of a series of dentinal systems, each with a medullar) canal 
as m Orycteropus (Tomes, 1850). 

Some of the dentinal systems are closel) aggregated into bundles 
of two or three. A laminar substance is seen to swirl around and 
between the systems and bundles, completely tilling the spaces 
between them. Optically, this material resembles dentine; its 
structure s that of dentine and in no wa\ resembles that of cement. 
Such a condition has been described b\ I nines ( 1850) as confluent 
dentinal systems, m which "parallel dentinal systems are united to 
each other, throughout their length. b\ a thin longitudinal lamina 
of dentine.*' 

I he individual dentinal sWcnis are large, about one and one-hall 
mm in diameter, and loosel) packed: there are approximately 70 
visible on the complete cross-section of the tusk in the region 
represented b\ the fragment. I his is m strong contrast to the 

\er\ small, discrete dentinal systems found in Platybelodon gran- 
gers, in which there are about 250 in the cross-section (Osborn and 
Granger, 1931 ). In a tusk referred to Amebelodon fricki (Colo- 
rado Museum no. 1319), Osborn and Granger noted the presence 
of dentinal systems ("rod-cones") "larger and much less numerous" 
than in /'. granger i, the total number being estimated at 100 to 150. 
Dentinal s\stems have not been dcfimtcK recognized in Platybelo- 
don danovi, although Osborn (1936: 4(>2 ) interpreted the "ir- 
regular longitudinal grooves," described b\ Borissiak ( l c ->28) on 
the upper surface of the tusk, as indicating their presence. If, 
however, P. danovi was ancestral to /'. grangeri, or at least repre- 
sented a more primitive type, we might expect the internal structure 
of the tusk to be less advanced and closer to that o\ the African 
form. 

The outer surface of the tooth was sheathed in life b\ a thick 
(1.5 mm) layer o\ what appears to be strongl) laminated dentine 
with dentinal tubules oriented perpendicular to the surface. This 
superficially resembles enamel, but its internal edge is seen to merge 
into the underlying dentine, with which the tubules are continuous, 
without the sharp contact and contrasting structure characteristic 
of dentine-enamel junctions. Its microscopic structure is also not 



1969 SHOVEL-TUSKED GOMPHOTHERE FROM KENYA 7 

at all like that of cement. This sheath has been broken away from 
much of the tooth surface, but enough remains to indicate its former 
presence over the entire fragment; there is no indication of enamel 
or cement. 

In the degree of flattening, the present specimen exceeds Platy- 
belodon danovi and approaches P. grangeri, which shows the most 
extreme flattening of any species of the subfamily yet described. 
In size, the African form, as determined from available measure- 
ments, is the smallest amebelodont, with a tusk width about 28 per 
cent smaller than in P. danovi. 

DISCUSSION 

The present specimen demonstrates that by Vindobonian times 
a shovel-tusked gomphothere was present in east Africa. The 
internal modifications of the dentine into discrete dentinal systems, 
presumably for strengthening the tusk, were in a relatively rudi- 
mentary stage as compared with those of P. grangeri, being large, 
loosely compacted, and generally poorly defined. Yet if the degree 
of flattening can be used to infer the degree of adaptation to a rela- 
tively specialized ecological role, we must conclude that the present 
form had already reached a degree of specialization with respect to 
tusk shape equal to that of P. grangeri, despite the relatively primi- 
tive organization of the dentinal systems. 

It appears reasonable to suggest that the present form represents 
(with P. danovi) an early specialization toward the adaptations 
that were to characterize the later amebelodonts. Its exact relation- 
ship to the Asiatic representatives of this genus is uncertain, but in 
view of its similarity to those species, as far as can be determined 
from this specimen, a close relationship to P. danovi may be sug- 
gested, both forms being among the earliest members of the sub- 
family. 

The question arises as to the place of origin of the Amebelo- 
dontinae. The mere presence of this group in east Africa during 
the Miocene does not, of course, positively answer this question. 
However, unless the African form proves, on further evidence, to 
be an independent adaptation to this particular ecological situation, 
the existence of this group in Africa and Asia at roughly the same 
time suggests a possible African origin for the subfamily. This view 
becomes more probable when one considers the long prior evolu- 
tionary history of the Gomphotheriidae in Africa and their relatively 
late appearance in Eurasia during the early Miocene. It is to be 
hoped that further specimens will be unearthed and also that among 



8 BREVIORA No. 310 

the isolated gomphothere teeth and bones from the African Miocene 
in museum collections additional remains of this group will be 
recognized. 

ACKNOWLEDGMENTS 

I would like to express thanks to Professor Bryan Patterson for 
permission to Stud) and section the specimen, and for his comments 
on the manuscript. 

TABLE 1 

Comparative measurements of Platybelodon 
species from Asia with Platybelodon from Africa, in mm 

WIMI 26200 
P. danovi 1 P grangeri 1 M< / M< Z 

Pype Type J8-64K 2 ^ '>4k :; 

width HO 166 79.5 74.5-78.5 

medial thickness ; " ;; '6.1 11.6-14.6 

lateral thickness 15 25 10.8 07.0-08.5 



medial thickness 



width 



(i :?2 0.199 0.224 



average thickness Q2Q] () , 75 Q , 6g 

width 

1 Measurements foi P. danovi and P. grangeri taken from Osborn and 
Granger 1 1931, p. 2). 

I stimated mean measurements corrected for missing outer layer ol 
laminated dentine. 

Vctual measurements of specimen. 

LITERATURE CITED 

Aramboi rg, ( 

1933. Mammiferes Miocenes du rurkana (Afrique Orientale). \nn. 
Paleoni.. Paris. 18: 123-146. 

Barboi r, E. H. 

1927. Preliminary notice of a new proboscidean, Amebelodon fricki, 
gen. et sp. nov. Bull. Nebraska State Mus. I: 131-134. 

1941. A shovel-tusked mastodon, Amebelodon fricki, from Kansas. 
Bull. Univ. Nebraska State Mus.. 2: 37-46. 



1969 SHOVEL-TUSKED GOMPHOTHERE FROM KENYA 9 

Borissiak, A. 

1928. On a new mastodon from the Chokrak beds (middle Miocene) 
of the Kuban region, Platybelodon danovi, n. gen. n. sp., Ann. 
Soc. Paleont. Russ., 7: 105-120. 
Evernden, J. F., D. E. Savage, G. H. Curtis and G. T. James 

1964. Potassium-Argon dates and the Cenozoic mammalian chronol- 
ogy of North America. Amer. Jour. Sci., 262: 145-198. 

Gregory, J. T. 

1945. An Amebelodon jaw from the Texas Panhandle. Univ. Texas 
Pub., No. 4401: 477-484. 

Hooijer, D. A. 

1966. Miocene rhinoceroses of East Africa. Fossil mammals of 
Africa, No. 21. Bull. Brit. Mus. (Nat. Hist.), Geol., 13 (2): 
119-190. 

JOUBERT, P. 

1966. Geology of the Loperot area. Geol. Surv. Kenya, Rep. No. 
74: 1-52. 

Leakey, L. S. B. 

1967. Notes on the mammalian faunas from the Miocene and 
Pleistocene of East Africa. In: W. W. Bishop and J. D. Clark, 
eds.. Background to Evolution in Africa. Chicago and London, 
Univ. Chicago Press, pp. 7-29. 

Nalivkin, D. V. 

1960. The Geology of the U.S.S.R. — A Short Outline. London, 
Pergamon Press, 70 pp. [Translation.] 

Osborn, H. F. 

1936. Proboscidea, a Monograph of the Discovery, Evolution, Mi- 
gration and Extinction of the Mastodonts and Elephants of the 
World. Vol. 1. New York, Amer. Mus. Nat. Hist., 802 pp. 

Osborn, H. F., and W. Granger 

1931. The shovel-tuskers, Amebelodontinae, of Central Asia. Amer. 
Mus. Novit., No. 470: 1-12. 
Savage, R. J. G. 

1965. The Miocene Carnivora of East Africa. Fossil mammals of 
Africa, No. 19. Bull. Brit. Mus. (Nat. Hist.), Geol., 10 (8): 
241-316. 

Simpson, G. G. 

1967. The Tertiary lorisiform primates of Africa. Bull. Mus. Comp. 
Zool., 136 (3):39-61. 
Stromer, E. 

1926. Reste land- und susswasser-bewohnender Wirbeltiere aus den 
Diamantfeldern Deutsch-Slidwestafrikas. In: Kaiser, E., Die 
Diamantenwiiste Siidwestafrikas, 2: 107-153, Berlin. 



10 BRI MORA No. 3 10 

I OM1 s. I. 

1850. On the structure of the dental tissues of the order Rodentia. 
Phil. I rans Roy. Soc. London, Pari II: 529-567. 

Will I WOH I M. T. 

I l >54. I he hyracoids ol I ast Africa. I ossii mammals of Africa, No. 
7. London, Brit. Mus. (Nat. Hist.), 58 pp. 

i Received 25 April 1968.) 



BREVIORA 

Museum! of Contiparatlve Zoology 

Cambridge, Mass. 31 March, 1969 Number 311 



THE GENERA APENESIA AND DISSOMPHALUS 

IN ARGENTINA AND CHILE 

(HYMENOPTERA, BETHYLIDAE) 

Howard E. Evans 



Abstract. Agenesia and Dissomphalus, two of the three genera of 
Pristocerinae occurring in Argentina and Chile, have been known from 
those countries from only a few specimens representing 5 species. Recent 
use of Malaise traps has greatly increased the available material and has 
made a further study of these genera desirable. In Apenesia, 12 species are 
here reported from Argentina, 6 of them new; 2 species are reported from 
Chile, 1 of them new. In Dissomphalus, 9 species are reported from Argen- 
tina, 8 of them new; this genus has not been collected in Chile. Keys to 
species are presented and important taxonomic features are illustrated. 

This is a supplement to two papers published in the Bulletin 
of the Museum of Comparative Zoology. 1 It covers recently col- 
lected material of two of the three genera of the subfamily Pristo- 
cerinae occurring in southern South America. The remaining genus, 
Pseudisobrachium, has been treated elsewhere. 2 

Most of the material reported on here was taken by Dr. Lionel 
Stange in Malaise traps in the province of Tucuman, Argentina. 
The value of these traps as faunal samplers is shown by the fact that 
Apenesia was previously known from Argentina from two species 
represented by 7 specimens; in the present paper 12 species are 
reported, 6 of them new, these species known from over 150 speci- 
mens. Dissomphalus was previously known from Argentina from 
3 species; 9 additional species are here reported from that country, 
8 of them new. The figures for Pseudisobrachium are even more 
impressive: previously known from Argentina from only 4 species, 
each represented by only one or a few specimens, this genus is now 



i Bull. Mus. Comp. Zool., 130: 249-359 (1963). 

Bull. Mus. Comp. Zool., 132: 1-222 (1964). 
2 Acta Zoologica Lilloana (in press). 



2 BRI- VIORA No. 3 1 1 

known from 22 species, represented in all by several hundred 
specimens. Altogether, the number of known species of Pristo- 
cerinae in Argentina has been increased from 9 to 48, the number 
in Chile from to 3. Material quite recently collected by Dr. Stange 
near Tucuman and by Charles C. Porter in several localities has 
consisted wholly of these same species, suggesting that a fairly high 
percentage of the species occurring in the areas sampled is now 
known. 

A discussion o\' the terminology employed will be found in the 
papers cited earlier. The abbreviations used are listed here for 
the sake oi ready reference: 

DAO: diameter of anterior ocellus (transversely ) 

HE: height of eve ( maximum, lateral view) 

LFW : length o( fore W ing 

111: length o\' head (frontal view, including clvpeus but not 

mouthparts ) 

OOL: ocello-ocular line (minimum distance between lateral 
ocelli and nearest eye margin) 

WF: width of front (at minimum point, between eyes) 

Wll: width of head (full frontal view, including eyes) 

WOT: width of oeellar triangle ( including lateral ocelli). 

All the material discussed in this paper is in the collections of the 
[nstituto Miguel Lillo, Tucuman. Argentina, and oi the Museum 
of Comparative Zoology, Cambridge, U.S.A. All holotypes have 
been deposited in the Institute Miguel Lillo. 

GENUS A PEN ESI A WESTWOOD 

This genus has been known from 35 species in South America, 
to which 7 more species are added here. All South American 
species are presentlv known from only one sex. and this unfortunate 
situation will prevail until these insects arc reared in scries from 
their hosts or taken in copula. No females of the genus Agenesia 
are as yet recorded from Argentina or Chile. In the treatment that 
follows, arrangement of the species-groups follows my 1963 
revision. 

Key to male Apenesiu known from Argentina and Chile 

1. Pronotal disc without a transverse carina anteriorly 2 

Pronotal disc with a transverse carina anteriorly 10 

2. Eyes glabrous: clvpeus with a median lobe, not prominent on the sides 

"(Figs. 10-13) 3 



1969 APENESIA AND DISSOMPHALUS 3 

Eyes hairy; clypeus large, well developed on sides, and without a distinct 
median lobe (Figs. 7-9) {dissomphaloides group) 8 

3. Abdomen with a distinct, moderately long petiole; LFW 2.1-2.4 mm 

(laevigata group) 9. cremdata (Kieffer) 

Abdomen sessile; LFW exceeding 3 mm 4 

4. Propodeum with no evidence of a transverse carina margining the disc 

behind; pronotum very short, strongly sloping (exilis group) 

8. chilena Evans 

Propodeum with a transverse carina margining the disc behind; pro- 
notum longer and with a more or less flat dorsal surface 5 

5. Antennae with erect setae on the under surface, which stand out 

strongly above the pubescence (most noticeable on segments 4-7) 

(pilicornis group) 6 

Antennae with erect setae sparse and standing out slightly if at all above 
the prominent, bristling pubescence {columbana group) 7 

6. Transverse carina of propodeum complete, median carina also com- 

plete and reaching transverse carina; LFW 3.3-3.5 mm 

1 . angusticeps Evans 

Transverse carina of propodeum obsolescent in middle, median carina 
not nearly reaching posterior margin of disc; LFW 4.0-5.6 mm .... 
2. reducta Evans 

7. Ocelli of moderate size; OOL at least as great as WOT; head about as 

wide as high 3. flammicornis Evans 

Ocelli greatly enlarged; OOL about half WOT; head considerably wider 
than high 4. photoplula (Ogloblin) 

8. Mandibles broad, with four teeth (Fig. 23); clypeus with a sharp median 

tooth but no additional processes (Fig. 9); notauli present only on an- 
terior half of mesoscutum 7. pygmaea n. sp. 

Mandibles slender, with two or three teeth; clypeus with a small median 
tooth flanked by two rounded processes (Figs. 7, 8); notauli com- 
plete 9 

9. Mandibles bidentate (Fig. 21); punctures of front small but quite dis- 

tinct; legs testaceous 5. lilloana n. sp. 

Mandibles tridentate (Fig. 22); front obscurely punctate; coxae and 
femora medium brown 6. simulata n. sp. 

10. Abdomen sessile, first tergite broadly reaching articulation with pro- 

podeum; mandibles with three large teeth (Fig. 24); inner margin of 
volsella with only a few setae just below cuspis {mexicana group) ..11 
Abdomen short-petiolate, first tergite not reaching extreme base; man- 
dibles with 4 or 5 teeth (Figs. 25, 26); inner margin of volsella setose 
for a considerable distance (brasdiensis group) 12 

11. Clypeus broadly subangulate (as in Fig. 12); antennae elongate, the 

pubescence short and subappressed; head and thorax polished, non- 

alutaceous 10. laevicornis n. sp. 

Clypeus truncate or bidentate apically (Fig. 10); antennae shorter, with 

suberect pubescence; head and thorax somewhat alutaceous 

11. inca Evans 



4 HRI-VIORA No. 31 1 

12 Clypeus short, broadly truncate (Fig. 13); mandibles with Eour teeth 

( Fit;. 26); front polished, non-alutaceous, strongly punctate 

14. spinipes n. sp. 

C lypeus more prominent, subangulate (1 igv 1 I. 12): mandibles with five 
teeth (Fig. 25); front somewhat allltaceoUS, punctures small and 

u idel) spaced 13 

13. Ocelli enlarged, DAO .20-.22 - WF, OOL .80-.9S x WOT; mesoscutum 

evenl) alutaceous; parameres noi bighl) modified (Fig. 4> 

12. spatulata n. sp. 

Ocelli not or but slightly enlarged, DAO .16 to .20 x WF. OOL at 
least slightly exceedini; WOT; mesoscutum not or obscurely alu- 
taceous. at least medially; parameres each divided into two separate 
lobes (Fig. 5) 13. hut rata n. sp. 

1 . Apcnesm angusticcps Evans 

This species lias been known from one male from Santa Cruz, 
Bolivia. Five Argentinian males are similar to the type in color 
and sculpture. However, the antennae are somewhat shorter (seg- 
ment three 1.6-1.9  as long as wale, segment eleven about twice 
as long as wide) and the front is somewhat broader (WF 1.1-1.3 
X HE"). 1 I \\ varies from 3.2 to 3.8 mm. 

Specimens examined. — ARGENTINA: 1 '. Horco Molle, 
Tucuman. 3-10 April 1966 (L. Stange); 2  ;. Horco Molle, 
Oct.. Dec. 1967 (C C. Porter ); I t, 11 km W Las Cejas, Tucu- 
man. 1-21 Feb. 1968 (I.. Stange); 1 . San Pedro Colalao. Tucu- 
man, Nov. -Dec. 1967 ( L. Stange). 

2. Apenesia reducta Evans 

I his species has been known from a single male from Chapada, 
Brazil. Twenty-three males now before me Ice) well to this species 
if allowance is made for the I act that in about half the specimens 
the ver\ small fourth mandibular tooth is altogether absent. In 
some specimens the tip of the abdomen is weakly suffused with 
brown, while in others the last two segments are rufous as in the 
type. All these specimens are larger than the type (LFW 4.2-5.6 
mm) except lor the two from Tucuman. which are slightly smaller 
(LFW 3.6-3.9 mm). Throughout this series, the propodeum is 
broader than in the type (disc 1.2-1.4 X as wide as long), al- 
though similar in sculpturing. 

Specimens examined. — ARGENTINA: 2 6 6 , 1 1 km W Las 
Cejas, Tucuman. Jan. -Feb. 1968 (L. Stange): 5 ' . Oran, Abra 
Grande. Salta. 29 Jan.-7 Feb. 1967 (R.^Golbach); 4 $ S, Rio 
Pescado. Salta, 19-25 Nov. 1967 (C. Porter. E. Willink). PARA- 
GUAY: 12 I i . Carumbe, 1 Feb.-8 March 1966 (R. Golbach). 



1969 APENESIA AND DISSOMPHALUS 5 

3 . A penesia flammicornis Evans 

This species was described from Santa Cruz, Bolivia, and later 
reported from Jujuy, Argentina."' Nine additional males from 
northern Argentina compare favorably with the type, but some are 
smaller (LFW 4.0-5.5 mm), and there is considerable variation 
in some standard measurements (OOL 1.05-1.40 X WOT; WF 
1.05-1.20 X HE). 

Specimens examined. — ARGENTINA: 7 $ $, Oran, Abra 
Grande, Salta, 29 Jan.-7 Feb. 1967 (R. Golbach); 2 s $ , Rio 
Pescado, Salta, 19-25 Nov. 1967 (C. Porter, E. Willink). 

4. Apenesia photophila (Ogloblin) 
This species is known from a series from Loreto, Misiones, 
Argentina. It is closely related to the preceding, but has much 
larger ocelli and a reddish abdomen. I redescribed the species in 
mv 1963 revision of this genus. 

5. Apenesia lilloana new species 
Holotype. — $ , ARGENTINA: Horco Molle, San Xavier Mts., 
Tucuman, 1-7 May 1966 (L. Stange). 

Description of type. — Length 2.6 mm; LFW 2.4 mm. Head 
black; thorax and propodeum fusco-castaneous; abdomen castane- 
ous, irregularly banded with light brown, base of first segment 
piceous; mandibles in large part testaceous; first two antennal seg- 
ments testaceous, remainder of antenna dark brown; legs testace- 
ous, coxae and femora weakly suffused with darker brown; wings 
subhyaline, with dark setulae, veins and stigma brown. Head, 
thoracic dorsum, venter, and basal parts of legs clothed with dense, 
short hair. Mandibles slender, bidentate (Fig. 21). Clypeus large, 
well developed in front of antennal sockets, below which there are 
large, semicircular depressions; median carina sharply defined, nearly 
straight in profile, forming a small median tooth at its terminus, 
this tooth flanked by two small, rounded processes, so that the 
clypeus is weakly tridentate (Fig. 7). Antennae very slender, scape 
elongate, first four segments in a ratio of about 3:1:1:1, segments 
three and eleven each nearly 3 X as long as wide; flagellar pubes- 
cence erect, bristling, setulae nearly as long as width of flagellum. 
Eyes hairy, somewhat protuberant; WH .97 X LH; vertex broadly 
rounded, nearlv straight in middle; distance from eye tops to vertex 
crest somewhat less than eye height. WF 1.25 X HE; ocelli not 
notably enlarged, in a compact triangle, the front angle less than 

3 Evans, H. E., Acta Hymenopterologica, 2: 103 (1966). 



6 BREVIORA No. 31 1 

a right angle; OOL 1.3 \ WOT. Front strongly alutaceous, weakly 
shining, with an abundance of shallow but well-defined punctures. 
Thoracic dorsum wholly alutaceous. obscurely punctate; pronotum 
short, without a transverse carina: notauli linear, complete, curved 
somewhat mesad posteriorly. Propodeum of moderate length, with 
a somewhat irregular transverse carina along the top of the de- 
clivity, the median carina not quite reaching this transverse carina: 
surface strongly reticulate basally, polished apically; disc measur- 
ing 1.1 as long as wide. Mcsopleura alutaceous. without a 
well-defined callus. Pore wing with the discoidal vein strongly 
pigmented lor a distance greater than length of basal vein. Abdo- 
men verj slender at base, subpetiolate; subgenital plate broadly 
truncate; genitalia with the aedoeagUS terminating in twit simple 
lobes, the parameres with a lateral, digitiform process (Fig. 1). 

Paratypes. — 4   . same data as type except various dates 
March-July 1966. 1967 (L. Stange) 

Variation. — The paratypes varv slightly in si/e (II \V 2.3-3.2 
mm): \\ 1 1 111 \anes from .92 to >>X. \\ I III' from 1.15 to 1.25. 
OOL/WOT from 1.15 to 1.25. There is no noteworthy variation 
in color or sculpture. 

Remarks. I would assign this species to the dtssomphaloides 
species-group, previously known from Mexico and the southwestern 
United States, although it will not run to that group in my revision 
of Apenesia ( 1963 I because of the presence of a transverse carina 
on the propodeum. However, the hair\ e\es. tridentate clvpeus. 
and male genitalia all suggest that it belongs in that group. The two 
species that follow are closely related to lilloana and these remarks 
applv equall) well to them. 

6. Apenesia simuUzta ni \\ spi (ins 
Holotype. \ . ARGENTINA: Horco Molle. San Xavier Mts.. 
Tucuman. 4-12 Jan. 1966 (L. Stange). 

Description of type. — Length 3.0 mm: LFW 2.6 mm. Head, 
thorax, and base of first abdominal segment black: remainder of 
abdomen dark castaneous. irregularly banded with light brown: 
mandibles in large part testaceous; clvpeus dull ferruginous; first 
two antennal segments testaceous, remainder of antenna dark 
brown; legs medium brown except trochanters, tarsi, and femoro- 
tibial joints paler; wings subhyaline. with dark setulae. veins and 
stigma brown. Head, thoracic dorsum, venter, and basal parts of 
legs clothed with short, brownish hair. Mandibles slender, tridentate 
(Fig. 22). Clvpeus much as in lilloana, but the median tooth very 
small and flanked by two fairly large, rounded processes (Fig. 8). 



1969 APENESIA AND DISSOMPHALUS 7 

First four antennal segments in a ratio of about 16:5:6:5, seg- 
ments three and eleven each about 2.5 X as long as wide; flagellar 
pubescence erect, bristling, setulae nearly as long as width of 
flagellum. Eyes hairy, strongly protuberant; WH .96 X LH; sides 
of head roundly convergent behind eyes; occipital carina unusually 
strong, readily visible for a considerable length when head is viewed 
from front. Front broad, WF 1.4 X HE; ocelli small, in a compact 
triangle far above eye tops, OOL 1.55 X WOT. Front moderately 
alutaceous, somewhat shining, punctures small and shallow, sepa- 
rated by 2-4 X their own diameters. Thoracic dorsum alutaceous, 
obscurely punctate; notauli strong, complete. Propodeal disc 
slightly longer than wide, with a well-defined transverse carina 
behind; sculpturing similar to that of preceding species. Features 
of mesopleura and wings as in lilloana, and abdomen subpetiolate 
as in that species. Subgenital plate broadly emarginate apically; 
genitalia similar to those of lilloana but the parameres simple, the 
aedoeagus with prominent ventral rami and with elongate apical 
lobes which are serrate on their inner margins (Fig. 2). 

Paratypes. — 6 $ $ , same data as type except various dates 
Jan.-May 1966 (L. Stange). 

Variation. — LFW varies from 2.2 to 2.9 mm, WH/LH from 
.95 to .97, WF/HE from 1.30 to 1.45, OOL/WOT from 1.3 to 
1.6. In the majority of specimens the propodeal disc is about as 
long as wide. 

7. Apenesia pygmaea new species 
Holotype. — <5 , CHILE: Queb. de la Plata, Rinconada Maipu, 
Santiago Prov., 510 meters, 26 Dec. 1966 (L. Stange). 

Description of type. — Length 1.8 mm; LFW 1.6 mm. Head 
and thorax dark brown, abdomen somewhat lighter brown except 
base of first tergite black; apical half of mandibles light brown; 
antennae dark brown except second segment testaceous; legs brown, 
tarsi and apices of tibiae somewhat lighter than remainder; wings 
hyaline, with dark setulae, veins and stigma brown. Head and 
thorax dorsum with rather dense short, brownish setae. Mandibles 
with a large apical tooth and three small, sharp teeth above it 
(Fig. 23). Clypeus large, well developed in front of antennal in- 
sertions, with a median tooth but no additional irregularities; 
median carina very strong basally, then abruptly declivous to apex 
(Fig. 9). Antennae very slender, first four segments in a ratio of 
about 7:4:3:4, segments three and eleven each about 3 X as long 
as wide; flagellar pubescence suberect. bristling, setulae about as 
long as width of flagellum. Eyes small but strongly convex, hairy; 



8 BREVIORA NO. 31 1 

WH .97 X LH; vertex broadly rounded, distance from eye tops 
to vertex crest subequal to eye height. WF 1.25 X HE; ocelli 
small, widely spaced, front angle of ocellar triangle less than a 
right angle; OOL 1.3 X WOT. Front alutaceous although rather 
Strongly shining, obscurely punctate. Thoracic dorsum also alutace- 
ous, somewhat shining; pronotum unusually short, without a trans- 
verse carina; mesoscutum with a few small punctures medially, 
notauli strong on anterior half, absent behind; scutum longitudinally 
depressed on each side. Propodeum elongate, with a weak, in- 
complete carina margining the disc behind; disc 1.3 X as long as 
wide, with a long median carina and several short, radiating carinae 
basallv. Features of mesopleura and wings as in preceding species; 
subgenital plate truncate as in that species; genitalia not studied. 
Remarks. — I have seen only one specimen of this minute 
species, but since it is the second species of the genus known from 
Chile, it seems worth describing at this time. The other Chilean 
species, chilena, is much larger and has a very different clypeus, 
glabrous exes, larger ocelli, and several other prominent differences. 

8. Apenesia chilena Evans 
I described this species from one specimen from Pichinahuel, 
ArauCO, Chile.' I have seen do additional specimens, nor have T 
seen other South American species closely related to it. I assigned 
the species tentatively to the exilis group, known otherwise from 
the southern United States. 

9. Apenesia crenulata (Kieffer) 
This very distinctive species has been known only from the type. 

from Para. Brazil. Two specimens before me agree well with the 

type but are both slightly smaller (LFW 2.1-2.2 mm) and have 

the ocelli less far removed from the eyes (OOL 1.1-1.2 y WOT); 

both have propodeal sculpturing generally similar to that of the 

tvpe. but the median carina is well defined. 

Specimens examined. — ARGENTINA: 1 ". Oran. Abra 

Grande, Salta, 29 Jan.-7 Feb. 1967 (R. Golbach). BRAZIL: 1 i . 

Nova Teutonia, Santa Catarina, Feb. 1964 (F. Plaumann). 

1 0. Apenesia laevicornis new species 
Holotype.— - . ARGENTINA: 11 km W Las Cejas. Tucu- 

man, 13 ' July-2 Aug. 1967 (L. Stange). 

Description of type. — Length 3.8 mm; LFW 3.2 mm. Head 

and thorax black; abdomen dark castaneous. irregularly mottled 

4 Proc. Ent. Soc. Washington, 69: 271 (1967). 



1969 APENESIA AND DISSOMPHALUS 9 

with lighter brown; mandibles black except suffused with dull fer- 
ruginous on apical third; antennae very dark brown; legs dark 
brown except tarsi somewhat paler; wings hyaline, with pale setulae, 
veins and stigma brown. Body setae relatively sparse and long. 
Mandibles with three large apical teeth (Fig. 24). Clypeus broadly 
subangulate, median part somewhat tectiform but not carinate (as 
in Fig. 12). Antennae slender, first four segments in a ratio of 
about 14:4:9:8, segment three 3 X as long as wide, segment 
eleven 3.5 X as long as wide; flagellar pubescence short, sub- 
appressed, a few slightly longer, erect setulae standing above it. 
Eyes with scattered short setae, only slightly protuberant; vertex 
broadly rounded off well above eye tops; WH .97 X LH. Front 
broad, WF 1.47 X HE; ocelli small, widely spaced, in a right 
triangle; OOL 1.25 X WOT. Front strongly polished, non-alutace- 
ous, punctures strong, rather irregularly spaced, mostly separated 
by 5-8 X their own diameters. Thoracic dorsum also wholly non- 
alutaceous, strongly polished, sparsely punctate; pronotum with a 
transverse carina on the anterior slope; notauli and parapsidal fur- 
rows strong, nearly complete. Propodeal disc as long as wide, 
with a complete median carina and a rather irregular transverse 
carina; basal part of disc alutaceous and with strong reticulations, 
apical part smooth and shining. Mesopleura wholly polished, non- 
alutaceous, very sparsely punctate. Discoidal vein of fore wing 
very weakly developed, arising a short distance down on transverse 
median vein. Abdomen sessile; subgenital plate arcuately emar- 
ginate; aedoeagus complex although nearly parallel-sided; ventral 
arms of cuspides narrowly rounded; parameres with a rounded 
apical enlargement (Fig. 3). 

Remarks. — This unusual species is known from a single speci- 
men. It is a member of the mexicana group, similar to neotropica 
Evans in many details, but with longer and smoother antennae, a 
much broader and more polished front, and smaller ocelli. 

1 1. Apenesia inca Evans 

This distinctive species was described from Peru and Ecuador 
in 1963; in 1966 I recorded it from two localities in eastern 
Brazil. Nineteen Argentinian males have the usual tridentate 
mandibles and bidentate clypeus (Fig. 10), as well as genitalia 
resembling those of the type closely. LFW varies from 2.7 to 
3.8 mm, WF from 1.0 to 1.2 X HE; in the majority of specimens 
OOL is subequal to or even slightly less than WOT. 

Specimens examined. — ARGENTINA: 19 S $ , 1 1 km W of 



10 KREVIORA No. 3 I I 

Las Cejas, Tucuman, Sept.. Nov., Dec. Jan.. Feb.. 1967-68 
(L. Stange). 

1 2. Apenesia spatulata new species 

Holotype.- - . ARGENTINA: Horco Molle. Tucuman. 7-26 
March 1967 (L. Stange). 

Description of type. — Length 4.S mm; LFW 4.0 mm. Head, 
thorax, and abdominal petiole black; remainder of abdomen dark 
castaneous with irregular banding with lighter brown; apical hall' 
o( mandibles testaceous, teeth rufous; antennae medium brown 
except second segment light brown; legs entirely testaceous; wings 
subhvaline. with dark setulae. vein and stigma dark brown. Mandi- 
bles broad, with five strong teeth (big. 25). Clypeus broadly 
subangulate. its median line tcetiform but not carinate (Fig. 1 1 ). 
First four antenna] segments in a ratio of about 20:7:10:10, seg- 
ment three 2.5  as long as wide, segment eleven 3 X as long as 
wide; flagellar pubescence pale, suberect. majority of setulae about 
.7 as long as width of llagellum. Fycs large, somewhat protuberant, 
glabrous; w II 1.03 1. 1 1; vertex forming a broad, even arc above 
eye lops. \\ I 1.1 < HE; ocelli large, less than their diameters 
apart, front ocellus located on an imaginary line drawn between 
eye tops; DAO .22 WF; OOL .85 WOT. Front wholly 
alutaceous, weak!) shining, punctures small but sharply defined, 
separated for the most part by 3-5 their own diameters. 

Pronotal disc margined m front by a transverse carina, weakly 
alutaceous and with scattered, minute punctures: mesoscutum also 
weakly alutaceous. shining, with punctures slightly stronger than 
on pronotum; QOtauli nearly complete. Propodeum elongate, the 
disc margined behind by a series of line transverse ridges, disc 
measuring slightly longer than wide: median carina complete, disc 
also reticulate medio-basally. Mesopleurum wholly weakly alutace- 
ous. obscurely punctate anteriorly. Middle tibiae strongly spinose 
above. Discoidal cell of fore wing weakly outlined, the discoidal 
vein itself Strongl) pigmented for a distance about equal to basal 
vein. Abdomen petiolate. but the petiole very short; subgenital 
plate truncate: aedoeagus complex; ventral lobes of cuspides spatu- 
late. narrowly rounded apicallv. paramercs with a broad, sub- 
quadrate lobe which is directed mesad (Fig. 4). 

Paratxpcs. — 20 $ $ , same data as type but various dates 
Jan.-July 1966 (L. Stange). 

Variation. — In the paratvpes. LFW varies from 3.2 to 4.1 mm. 
WH from 1.03 to 1.06 LH. WF from 1.05 to 1.20 X HE. OOL 



1969 APENESIA AND DISSOMPHALUS 11 

from .80 to .95 X WOT, DAO from .20 to .22 X WF. There is 
no noteworthy variation in other characters. 

Remarks. — This species is a member of the brasiliensis group, 
apparently related to angustata (Evans). The species name refers 
to the large, spatulate lobes of the cuspis. 

13. Apenesia lacerata new species 

Holotype.— $ , ARGENTINA: 11 km. W of Las Cejas, 
Tucuman, 16-29 April 1967 (L. Stange). 

Description of type. — Length 3.3 mm; LFW 3.0 mm. Head, 
thorax, and abdominal petiole piceous; remainder of abdomen 
dark castaneous, irregularly banded with lighter brown; mandibles 
testaceous, darker at extreme base and apex; antennae testaceous 
basally, grading into medium brown beyond segment three; legs 
wholly testaceous; wings hyaline. Mandibles with five teeth (as in 
Fig. 25); clypeus broadly subangulate, tectiform medially 
(Fig. 12). First four antennal segments in a ratio of about 
18:5:10:9, segment three 3 X as long as wide, segment eleven 
3.5 X as long as wide; flagellar pubescence suberect, nearly as 
long as width of flagellum. Eyes weakly hairy, slightly protuber- 
ant; WH 1.03 X LH; vertex forming a broad, even arc above eye 
tops. WF 1.22 X HE; ocelli slightly enlarged, DAO .17 X WF; 
front ocellus slightly above a line connecting eye tops; OOL 1.1 X 
WOT. Front wholly alutaceous, somewhat shining; punctures 
small, separated by 3-5 X their own diameters. Pronotum strongly 
polished, non-alutaceous, the disc margined in front by a transverse 
carina; mesoscutum strongly polished, very weakly alutaceous 
laterad of the very strong, complete notauli. Propodeal disc as 
wide as long, sculptured exactly as in the preceding species. 
Mesopleurum shining, barely alutaceous, strongly punctate in 
front. Discoidal vein of fore wing very weakly outlined by pig- 
mented lines. Abdominal petiole very short; subgenital plate 
subtruncate; genitalia of unusual form in that the ventral arms 
of the cuspides are very long and pointed, the parameres deeply 
divided into a slender process and an enlarged mesal lobe (Fig. 5). 

Paratypes. — 17 $ $ , same data as type but various dates 
July-Nov. 1967 and Jan., Feb., May 1968 (L. Stange). 

Variation. — LFW varies from 3.0 to 3.3 mm, WH from .98 
to 1.03 X LH, WF from 1.12 to 1.30 X HE, DAO from .16 to 
.20 X WF, OOL from 1.1 to 1.2 X WOT. In the majority of 
paratypes the propodeal disc is longer than wide. In a few speci- 
mens the front coxae and femora are suffused with brown. In two 
specimens the mesoscutum is somewhat alutaceous, even medially. 



12 KREVIORA No. 31 1 

Remarks. — This species resembles the preceding closely, and 
I would be inclined to consider the two conspecific if it were not 
for the distinctive genitalia. The name laceruta refers to the greatly 
dissected parameres of the genitalia of this species. 

14. Apenesia spinipes new species 
Holotype. — 6 , ARGENTINA: 1 1 km W of Las Cejas, Tucu- 
iiuiii. 3-19 June 1966 (L. Stange). 

Description of type. — Length 4.3 mm; LFW 3.5 mm. Head 

and thorax black, abdomen dark castaneous with lighter markings 
along margin of first tcrgite as well as apically; mandibles testaceous 
except teeth rufous; antennae wholly testaceous, legs also of this 
color except coxae and femora suffused with brown; wings hyaline. 
veins and stigma dark brown. Mandibles with four large apical 
teeth (Fig. 26). Ctypeus very short, broadly truncate (Fig. 13). 
1 irst four antenna] segments in a ratio of about 4:1:2:2, seg- 
ments three and eleven each about 3 X as long as wide; flagellar 
pubescence erect, pale, setulae about one third as long as width 
of tlagellum. Lyes ver\ weakl) hairy; vertex forming a broad, 
even arc above eye tops, distance from eve tops to vertex crest 
nearl) equal to HE. WH 1.02 X LH; WF 1.2 X HE; ocelli small, 
in a compact triangle. OOL 1.4 X WOT. Front polished, non- 
alutaceous, with strong punctures which are separated by only 
1-2 X their own diameters. Thoracic dorsum polished, non- 
alutaceous; pronotal disc with a strong transverse carina in front, 
its surface covered with small punctures except along a median 
band. Mesoscutum with scattered, sharply defined punctures; 
notauli strong, complete; scutellar disc impunctate medially. Pro- 
podeal disc 1.2 X as wide as long, the transverse carina weak 
although complete, median carina also complete; disc reticulate 
medio-basallv. elsewhere strongly polished and smooth; posterior 
face shining, with a lew punctures and some medial roughening, 
but no median carina. Mesoplcurum in large part smooth and 
polished, the callus convex. Middle tibiae very strongly spinose. 
Discoidal cell of fore wing weakly outlined by pigmented lines. 
the subdiscoidal vein more heavily pigmented than the discoidal 
vein. Abdomen short-petiolate; subgenital plate truncate; genitalia 
with the aedoeagus narrowed subapically, with a pair of curved 
apical processes; ventral arms of cuspides large, rounded; para- 
meres with broad mesal lobes (Fig. 6). 

Paratypes. — 41 6 Z , same data as type except various dates 
Mav. Aug.. Oct.. Dec. 1967; Jan., Feb., 1968 (L. Stange). 



1969 APENESIA AND DISSOMPHALUS 13 

Variation. — Several specimens have the coxae and femora 
more heavily infuscated than in the type, and the apical third of 
the antennae may also be somewhat infuscated. LFW varies from 
2.7 to 4.2 mm, WH/LH from 1.01 to 1.05, WF/HE from 1.10 
to 1.25, OOL/WOT from 1.20 to 1.45. 

Remarks. — This species runs to brasiliensis (Kieffer) in my 
1963 key, although the transverse carina of the propodeum is 
not as strong. The genitalia are very different from those of 
brasiliensis and the sculpturing of the basal triangle of the 
propodeum quite different. 

GENUS DISSOMPHALUS ASHMEAD 

Seventeen species of this genus have so far been described from 
South America, and 8 more are added here. I would estimate that 
there are at least 2 to 4 times that many species on that continent, 
even though some synonymy will be discovered when the sexes are 
properly associated. The following three species are known from 
Argentina from the female sex and are not treated here: attaphila 
(Bruch), azarai (Ogloblin), and platensis (Bruch). This genus 
has not yet been found to occur in Chile 

Key to male Dissomphalus known from Argentina 

1. Second abdominal tergite with two large, pubescent pits close together 

medially and sharing a common depression (Figs. 16, 17) 2 

Second abdominal tergite not as above, containing two medium-sized to 
minute pits, marks, or processes which are separated by much more 
than their own length (Figs. 18-20) 3 

2. Pronotum with a strong transverse ridge; second tergite with four pits, 

the two middle ones very large, the tergite swollen and distorted be- 
hind them (Fig. 16) 1 . deformis n. sp. 

Pronotum without a transverse ridge; second tergite with two somewhat 
smaller pits medially, the tergite not swollen and distorted behind 
them (Fig. 17) 2. ulceratus n. sp. 

3. Notauli absent on posterior two-thirds of mesoscutum; mandibles with 

four teeth (Fig. 31) 6. teren n. sp. 

Notauli complete; mandibles with 2 or 3 teeth 4 

4. Second tergite with fairly large, round pits with raised rims (Fig. 18); 

subgenital plate with a large, V-shaped apical emargination; WF not 

exceeding HE 3. puteolus n. sp. 

Second tergite with minute pits, slits, or processes; subgenital plate with- 
out a V-shaped emargination: WF exceeding HE 5 

5. Clypeus tapering to a long, sharp median point (Fig. 15); WH only .9 X 

LH; second tergite with a pair of small, spatulate processes antero- 
medially 8. bispinulatus n. sp. 



14 BREVIORA No. 31 1 

Clypeus not so strongly pointed apically and WH subequal to HE; second 
tergite not modified as above 6 

6. Second tergite smooth, with a pair of pale, roughened spots at extreme 

anterior margin laterally, but DO Other modifications; clvpeus with a 
median process, not or very indistinctly tridentate. .9. incomptus Evans 
Second tergite with small pits, slits, or processes not nearly so close to 
anterior margin (I igs. 19, 20); clypeus Strongly tridentate (as in I ig. 
I4i 7 

7. Modifications of second tergite in the form of oblique slits (Fig, 20); 

propodeal disc short. 1.7 x as wide as long J.infissus n. sp. 

Modifications Of second tergite in the form of minute pits or hair pencils 
( 1 ig. I l >); propodeal disc 1.0-1.3 X as wide as long 8 

8. LFW l.S mm; eyes and ocelli small. WF 1.2 X HE, OOL 1.05 X WOT; 

wings pale, discoidal \cin unpigmented, basal vein subered 

4. mendicus n. sp. 

I I w :.(> mm; eyes and ocelli rathei large, WF 1.1 x HE. OOL only 
.8 > WOT; wings with dark veins and stigma, discoidal vein a short. 
hi own streak, basal vein strongly oblique 5. microstictus n. sp. 

1 . Dissomphalus deformis new species 

Holotype — 6 , ARGENTINA: Oran. Abra Grande, Salta, 
8-15 Feb. 1967 (R. Golbach). 

Description <>/ type. — -Length 2.7 mm; LFW 2.4 mm. Head 
and thorax black; abdomen dark brown, still used with lighter 
brown on parts oi basal and apical segments; mandibles largely 
testaceous; antennae testaceous, sufTused with brown on apical 
half; coxae and femora dark brown, hind tibiae medium brown, 
legs Otherwise testaceous; wings hyaline. Mandibles tridentate, 
the middle tooth the smallest (Fig. 27). Clypeus tridentate, 
median carina strong, straight in profile (Fig. 14). First four 
antenna] segments in a ratio of about 3:1:1:1, segment three 
1.5  as long as wide, segment eleven twiee as long as wide. 
Head as wide as high, the eyes large, convergent below; WF 1.1 X 
HE; ocelli widely spaced, front angle of triangle less than a right 
angle; OOL 0.9 )< WOT. Front alutaceous although somewhat 
shining, covered with shallow punctures. Thoracic dorsum strongly 
alutaceous, obscurely punctate; pronotal disc crossed anteriorly 
by a very strong carina; notauli strong, complete. Propodeal disc 
1.2 X as wide as long, disc and declivity wholly coarsely reticulate. 
Discoidal vein of fore wing barely indicated. First tergite short, 
with a strong median groove for much of its length; second tergite 
with two very large, pubescent, elliptical pits close beside the mid- 
line, the tergite strongly gibbous behind and slightly laterad of 
these pits; second tergite also with a pair of much smaller pits in 



1969 APENESIA AND DISSOMPHALUS 15 

addition, so that there are four pits in a transverse series; tergite 
with a median band of small setae behind the pits (Fig. 16). 
Subgenital plate broadly concave apically; parameres large, tri- 
angular in lateral view, subacute apically. 

Paratypes. — 3 6 £ , same data as type. 

Variation. — The paratypes are all slightly larger than the type 
(LFW 2.5-2.7 mm) and all have the propodeal disc slightly 
shorter and wider (1.25-1.35 X as wide as long). There is no 
noteworthy variation in standard measurements or in the re- 
markable configuration of the second tergite. 

2. Dissomphalus ulceratus new species 
Holotype. — s , ARGENTINA: Horco Molle, San Xavier Mts., 
Tucuman, 1-7 May 1966 (L. Stange). 

Description of type. — Length 2.0 mm; LFW 1.9 mm. Dark 
brownish-fuscous, sides of basal abdominal segments suffused with 
light brown; mandibles testaceous on apical half; mandibles dark 
brown except second segment somewhat lighter than remainder; 
coxae and femora dark brown, legs otherwise testaceous; wings 
hyaline. Mandibles tridentate, the uppermost tooth the smallest 
(Fig. 28). Clypeus with a strong median angulation and two 
small, rounded teeth beside it (much as in Fig. 7); median carina 
straight in profile. Third antennal segment 1.5 X as long as wide. 
Head as wide as high, eyes not convergent below, front broad, 
WF 1.3 X HE; ocellar triangle compact, OOL 1.2 X WOT. Front 
alutaceous, somewhat shining, covered with shallow punctures. 
Thoracic dorsum alutaceous although somewhat shining; pronotum 
short, without a transverse carina; notauli complete. Propodeum 
short, disc measuring 1.35 X as wide as long; disc mostly 
reticulate, but with a smooth area behind. Discoidal vein of fore 
wing present as a long but weakly pigmented streak. First tergite 
more rounded behind and with a shorter median groove than in 
deformis, and the second tergite not gibbous behind the pits as in 
that species; pits large, narrowly separated medially, sharing a 
common bowl-shaped depression, the tergite without other pits 
and with only a few scattered setae (Fig. 17). Subgenital plate 
broadly truncate. 

Paratype. — 1 $, same data as type except 3-10 April 1966 
(L. Stange). 

Variation. — In the paratype, LFW is 2.0 mm, WF 1.4 X HE. 
This specimen is closely similar to the type in every respect. 



16 BREVIORA No. 31 1 

3. Dissomphalus puteolus new species 
Holotype. — 6 , ARGENTINA: 1 1 km W of Las Cejas, Tucu- 

nuin. 13-27 May 1967 (L. Stange). 

Description of type. — Length 2.5 mm; LFVV 2.1 mm. Head 
and thorax black; abdomen dark brown, with paler markings 
toward base and apex; mandibles in large part testaceous; scape 
medium brown, tlagellum light brown; coxae and femora dark 
brown, remainder of legs light brown; wings hyaline. Mandibles 
with three strong apical teeth, the most basal tooth somewhat 
rounded (Jig. 29). Clypeus with a projecting median lobe which 
is tridentate (much as in Fig. 14). First four antenna] segments 
in a ratio of about 10:4:3:3, segment three only slightly longer 
than wide, segment eleven 1.5 X as long as wide. \VH 1.02 X LH; 
eyes large, strongly convergent below, WF only .93 X HE; ocelli 
widel) spaced. OOL onl\ .8 WOT. Front alutaeeous and 

Strongl) roughened b\ punctures which are separated by less than 
their own diameters. Thoracic dorsum alutaeeous; pronotum short, 
without a transverse carina: mesoscutum somewhat shining, with 
numerous small punctures, ootauli complete. Propodeal disc 
unusually short and broad. 1.7 • as wide as long, surface reticu- 
late except smooth over a small area behind. Discoidal vein of 
fore wing absent. First tergite relatively longer and second tergite 
shorter and less convex than in preceding two species; second 
tergite with a pair of fairly large, circular pits dorso-laterally, each 
pit in a depression, with a raised rim and giving rise to a small 
tuft of setulae; second tergite with numerous setae, especially 
laterad of the pits (Fig. 18). Subgenital plate with a strong. 
V-shaped apical emargination; parameres broad, tapering to an 
acute apex. 

Paratypes. — 5 6 & . same data as type except various dates 
Jan.. March, April. June, 1966-1968 (L. Stange). 

Variation, — The paratypes are very similar to the type; LFW 
varies from 1.9 to 2.1 mm; all have the basal flagellar segments 
testaceous, contrasting with the darker apical segments; in two 
the propodeum is not quite as short as in the type, the disc 
measuring 1 .6 X as wide as long. 

4. Dissomphalus mendicus new species 
Holotype. — 6 , ARGENTINA: 11 km W of Las Cejas. Tucu- 

man, 17 June- 12 July 1967 (L. Stange). 

Description of txpe. — Length 2.1 mm; LFW 1.8 mm. Head 
and thorax dark brownish-fuscous except pronotal collar medium 



1969 APENESIA AND DISSOMPHALUS 17 

brown; abdomen dark castaneous except first tergite bordered with 
light brown, second tergite light brown on extreme sides; mandi- 
bles in large part testaceous; antennae dull brown except second 
segment and adjacent parts of first and third segments testaceous; 
legs brown, tarsi and femoro-tibial joints straw-colored; wings 
hyaline, with pale setulae and pale veins and stigma. Mandibles 
tridentate, the basal two teeth small (much as in Fig. 28). Clypeus 
as described for the preceding species. First four antennal seg- 
ments in a ratio of about 5:2:2:2, segments three and eleven 
both about 1.5 X as long as wide. WH and LH subequal; eyes 
weakly converging below, WF 1.2 X HE; ocelli small, front 
angle of ocellar triangle less than a right angle, OOL 1.05 X WOT. 
Front moderately alutaceous, shining, the punctures small and 
shallow, separated by 2-4 X their own diameters. Thoracic 
dorsum alutaceous although somewhat shining; pronotum short, 
without a transverse carina; mesoscutum weakly punctate, notauli 
complete. Propodeal disc 1.25 X as wide as long, surface reticu- 
late. Discoidal vein of fore wing distinct for a short distance 
although barely pigmented. First tergite rounded behind, with a 
strong median groove; second tergite with a pair of minute lateral 
pits with raised rims, the pits occupying very shallow depressions 
and flanked by several setae (much as in Fig. 19). Subgenital 
plate broadly truncate apically. 

Remarks. — This small but distinctive species is known from 
a single specimen. 

5. Dissomphalus microstictus new species 

Holotype. — $ , ARGENTINA: 11 km W of Las Cejas, Tucu- 
man, 3-19 June 1966 (L. Stange). 

Description of type. — Length 3.0 mm; LFW 2.6 mm. Head 
black; thorax dark brownish-fuscous; abdomen dark castaneous, 
with paler markings basally and apically; mandibles in large part 
testaceous; antennae medium brown except segments two and three 
and apex of one testaceous; legs straw-colored except front and 
hind femora weakly suffused with brown; wings hyaline, with dark 
veins and stigma. Mandibles with two strong apical teeth (Fig. 
30). Clypeus with a tridentate median lobe (much as in Fig. 14). 
Third and eleventh antennal segments both about twice as long as 
wide. WH and LH subequal; eyes prominent, short-haired, inner 
orbits subparallel on lower half; WF 1.1 X HE; ocelli slightly en- 
larged, in about a right triangle, OOL only .8 X WOT. Front 
wholly strongly alutaceous, weakly shining, punctures very shallow, 



18 BREVIORA No. 31 1 

separated by 1-3 X their own diameters. Thoracic dorsum alu- 
taceous although less strongly so than front; pronotum short, with- 
out a transverse carina; mcsoscutum with scattered small punctures, 
notauli complete. Propodeal disc elongate, barely wider than long, 
disc smooth and shining behind, the median carina not reaching the 
transverse carina. Discoida] vein of fore wing lightly pigmented for 
a considerable distance. First tergite elongate, median groove con- 
fined to basal half; second tergite with a pair of widely spaced very 
small pits with raised rims, these pits in very shallow, broad depres- 
sions and flanked by a few setae (Fig. 19). Subgenital plate broadly, 
shallowly emarginate; parameres twisted mesad apically. 

Paratypes. — ARGENTINA: 53 <5 c5 , same data as type except 
various dates Nov. -Dec. 1967. Jan. -May l c )68; 1 • . San Pedro 
Colalao, Tucuman. Jan. 1968 (L. Stange). 

Variation. — LFW varies from 2.1 to 3.6 mm. The legs vary 
from wholly straw-colored to almost wholly brown. WF varies 
from 1 . 1 to 1 .3 X HE. The ocelli of some of the smaller specimens 
are only slightlv enlarged, and in these specimens OOL is only 
slightly less than WOT. There is little variation in the form of the 
second tergite in this long series. 

(•>. Dissomphalus teren new species 
Holotxpc. — | . ARGENTINA: Horco Molle. San Xavier Mis., 
Tucuman. 3-1 1 June l c K>h (L. Stange). 

Description of type. -- Length 1.8 mm; LFW 1.6 mm. Dark 
brownish-fuscous, including mandibles and antennae; legs dark 
brown except tarsi testaceous; wings h valine, \eins and stigma dark. 
Mandibles with four teeth, basal two teeth connate (Fig. 31 ). Cly- 
peus tridentate. with a strong median carina (much as in Fig. 14). 
First four antenna! segments in a ratio of about 10:4:3:3, segments 
three and eleven each about 1 .5 X as long as wide. Mead elongate, 
vertex broadly rounded off far above eye tops; WH .93 X LH; WF 
1.25 HE; front angle of ocellar triangle less than a right angle, 
OOL 1.3 X WOT. Front weakly alutaceous. moderately shining, 
punctures small and shallow. Thoracic dorsum smooth, somewhat 
shining, obscurely punctate; pronotum without a transverse carina; 
notauli present on anterior third of mcsoscutum, absent behind. 
Propodeal disc 1.3 X as wide as long, with a transverse polished 
band posteriorly. Discoidal vein of fore wing weakly pigmented. 
First tergite rounded apically. with a median groove on the basal 
.6; second tergite smooth except for scattered small setae and a pair 
of curved hair-pencils which arise from slightly elevated bases, these 
located dorsally but separated by about twice the length of one of 



1969 APENESIA AND DISSOMPHALUS 19 

the hair-pencils. Subgenital plate unusually broad, arcuately emar- 
ginate; parameres very slender apically, their tips directed mesad. 
Remarks. — This species possesses a number of unique features: 
the 4-tooth mandibles, reduced notauli, and broadly emarginate 
subgenital plate. It is known only from the type specimen. 

7. Dissomphalus infissus new species 

Holotype. — <$ , ARGENTINA: Oran, Abra Grande, Salta, 16- 
23 Feb. 1967 (R. Golbach). 

Description of type. — Length 3.2 mm; LFW 2.6 mm. Dark 
brownish-fuscous; mandibles in large part testaceous; antennae 
medium brown, dark brown beyond basal .4; front coxae, all 
femora, and hind tibiae dark brown, remaining coxae, trochanters, 
femora-tibial joints, middle and front tibiae, and all tarsi testaceous; 
wings subhyaline, with dark setulae and dark veins and stigma. 
Mandibles tridentate (much as in Fig. 28). Clypeus with a triden- 
tate median lobe (much as in Fig. 14). First four antennal seg- 
ments in a ratio of about 3:1:1:1, segments three and eleven each 
about 1.5 X as long as wide. Head about as wide as high; front 
broad, WF 1.2 X HE; vertex slightly emarginate behind ocellar tri- 
angle, the latter about a right triangle, OOL 1.15 X WOT. Front 
strongly alutaceous, covered with shallow punctures which are 
separated by only 1-2 X their own diameters. Pronotum short, 
roughened but not carinate along anterior margin of disc; meso- 
scutum somewhat shining, with scattered small punctures; notauli 
strong, complete. Propodeal disc short, 1.7 X as wide as long, 
covered with coarse reticulations. Discoidal vein of fore wing inter- 
stitial with media, pigmented for a distance exceeding basal vein. 
First tergite rounded apically, with a strong median groove on basal 
half; second tergite with a pair of dorso-lateral slits with raised rims, 
each slit flanked by a group of setae (Fig. 20). Subgenital plate 
broadly truncate. 

8. Dissomphalus bispinulatus new species 

Holotype. — $ , ARGENTINA: Oran, Abra Grande, Salta, 16- 
23 Feb. 1967 (R. Golbach). 

Description of type. — Length 1.7 mm; LFW 1.5 mm. Dark 
brownish-fuscous; mandibles testaceous apically; antennae medium 
brown except basal three segments light brown; legs straw-colored 
except front coxae, all femora, and hind tibiae medium brown; 
wings hyaline, with dark setulae and dark veins and stigma. Man- 
dibles with a large apical tooth and two minute teeth above it. 
Clypeus tapering medially to a long, acute process which is at the 



20 BREVIORA No. 31 1 

terminus of a strong median ridge (Fig. 15). First four antennal seg- 
ments in a ratio of about 9:4:3:3, segments three and eleven each 
about 1.5 X as long as wide. Head elongate, WH only .90 X LH; 
eyes glabrous, inner orbits slightly diverging above and below 
middle; WF 1.17 X HE; ocelli small, in an acute triangle, OOL 
1.3 X WOT. Front shining, weakly alutaceous, obscurely punc- 
tate. Pronotum much expanded from front to rear, sides concave as 
seen from above, without a transverse carina; mesoscutum weakly 
alutaceous, obscurely punctate, notauli complete. Propodeal disc 
1 .2 X as wide as long, most of its posterior half smooth and shining, 
median carina not nearly reaching transverse carina. Discoidal vein 
of fore wing strongly pigmented for a considerable distance. First 
tergite rounded apicallv. with a median groove on basal half; second 
tergite with a pair oi small, spatulate processes close behind the 
margin of the first tergite. these separated medially by about twice 
their own length; second tergite also with a pair of roughened spots 
anterolaterally. Subgenital plate very broadly truncate apically. 

9, Dissomphalus incomptus Evans 

I described this species from eight males from Santa Catarina, 
Brazil, in im 1964 synopsis. Its presence in Paraguay and in two 
provinces of Argentina suggest that it is widely distributed and not 
uncommon in southern South America. The specimens before me 
key readily to this species in my 1966 paper 1 and agree with the 
tvpe series in most details, including the genitalia. They average 
somewhat larger I I I \\ 1.9-2.6 mm) and show minor variation in 
head measurements (WF 1.1-1.3 X HE, OOL 1.15-1.30 WOT). 
The sinuations beside the median process of the clypeus vary in 
development, such that in some specimens the clypeus may be said 
to be indistinctly tridentate. 

Specimens examined. — PARAGUAY: 1 & , Carumbe, 1 Feb. 
-8 March 1966 ( R. Golbach). ARGENTINA: 2 6 6, Oram 
Abra Grande. Salta. 1 Jan.-7 Feb. 1967 (R. Golbach); \2 $ 3 , 
Horco Molle. San Xavicr Mts., Tucuman, Jan. -June 1966 (L. 
Stange). Dec. 1967 (C. C. Porter). 

(Received 7 October 1968.) 



5 Acta Hymenopterologica. 2: 1 10. 



1969 



APENESIA AND DISSOMPHALUS 



21 









Figs. 1-6. Male genitalia of Apenesia spp., ventral aspect. Fig. 1., A. 
lilloana n. sp. Fig. 2., A. simulate! n. sp. Fig. 3., A. laevicornis n. sp. Fig. 4., 
A. spatulata n. sp. Fig. 5., A. lacerata n. sp. Fig. 6., A. spinipes n. sp. 



27 



BREVIORA 



No. 311 





23 






26 




27 




28 




29 





31 



1969 APENESIA AND DISSOMPHALUS 23 



Figs. 7-15. Clypeus of male Apenesia and Dissomphalus. Fig. 7., A. lil- 
loana n. sp. Fig. 8., A. simulata n. sp. Fig. 9., A. pygmaea n. sp. Fig. 10., 
A. inca Evans. Fig. 11., A. spatulata n. sp. Fig. 12., A. lacerata n. sp. Fig. 
13., A. spinipes n. sp. Fig. 14., D. deformis n. sp. Fig. 15., D. bispinulatus 
n. sp. 

Figs. 16-20. Base of abdomen of male Dissomphalus, dorsal view. Fig. 
16., D. deformis n. sp. Fig. 17., D. ulceratus n. sp. Fig. 18., D. piiteolus 
n. sp. Fig. 19., D. microstictus n. sp. Fig. 20., D. inftssus n. sp. 

Figs. 21-31. Mandibles of male Apenesia and Dissomphalus. Fig. 21., 
A. lilloana n. sp. Fig. 22., A. simulata n. sp. Fig. 23., A. pygmaea n. sp. 
Fig. 24., A. laevicornis n. sp. Fig. 25., A. spatulata n. sp. Fig. 26., A. spi- 
nipes n. sp. Fig. 27., D. deformis n. sp. Fig. 28., D. ulceratus n. sp. Fig. 
29., D. piiteolus n. sp. Fig. 30., D. microstictus n. sp. Fig. 31., D. teren 
n. sp. 



BREVIORA 



Museum of Coinpairgitive Zoology 

Cambridge, Mass. 31 March, 1969 Number 312 

ECOLOGICAL OBSERVATIONS ON ANOUS OCCULTUS 
WILLIAMS AND RIVERO (SAURIA, IGUANIDAE) 

T. Preston Webster 

Abstract. Recent observations indicate that the structural niche of 
Anolis occultus on Puerto Rico is the peripheral foliage of the montane 
forests from which it has been recorded; probably best considered a canopy 
species, it descends to ground level only at the few localities where trail- 
edge vegetation merges with the tree crowns. Structurally, behaviorally, and 
in coloration it is very well adapted for a life on twigs and leaves. As ex- 
pected from other studies of Anolis ecology, the spatial preferences of oc- 
cultus show little overlap with those of the other species recorded from the 
same locality. 

In 1963 the first specimen of the very distinctive species Anolis 
occultus was captured by day at Cerro La Punta in Puerto Rico's 
Cordillera Central. Subsequent collecting produced about 40 addi- 
tional specimens from all areas of higher montane forest on the 
island. All but two of these specimens were collected by Richard 
Thomas at night along forest trails where the trail-edge vegetation 
merged more or less intimately with the foliage of the canopy. 
Thomas suggested (Williams, Rivero, and Thomas, 1965) that 
occultus is a canopy species with a preference for exposed areas 
of bare twigs and vines, although after further collecting he sug- 
gested that "their habitat requirements may not be as narrow as 
my first experience with the species indicated" (Thomas, personal 
communication, March, 1968). As Thomas observed only a single 
specimen during the day, the diurnal ecology of this elusive lizard 
remained a mystery. 

Recently 15 additional specimens were obtained 13.6 kilometers 
south of Palmer, Puerto Rico, in the Sierra de Luquillo. The 
species was first sought at night; nine specimens were easily lo- 
cated the night of April 7th. Four slept on long, exposed twigs, 
two slept on twigs near leaves, and one slept across the upper sur- 
face of a broad, stiff leaf. Twigs selected for sleeping perches were 
a quarter inch in diameter or less. Thomas's observations were 



2 breviora No. 312 

confirmed; occultus clings tightly to its perch while asleep, may 
squeak loudly when handled, and can grip the twig strongly with 
its prehensile tail. Sleeping specimens are a light grey and stand 
out clearly in the beam of a headlamp. The two remaining anoles, 
asleep in the same small clump of vegetation, one across a leaf, the 
other on a bare twig, were left for observation in the morning. 

Both lizards were found asleep at 6:15 A.M. One became active 
at 6:30. moving sluggishly to a nearby perch that seemed to offer 
greater exposure to the sun. The second became active a half 
hour later, but neither moved far for at least an additional hour. 
While both were in view, a third specimen was discovered when 
it became backlighted; far more active, it seemed to drink from 
raindrops on two occasions and prowled with frequent long pauses 
over an area of largely bare and exposed twigs, eventually wander- 
ing upward and out oi sight. The remaining two also disappeared 
in the vegetation, but with the appearance of new individuals or 
the reappearance of those previously seen a total of six were ob- 
served, at least oik- staying in sisiht at all times. Movements were 
slow, cautious, and generally along twigs of small diameter, al- 
though occasionally the anoles crossed leaves or used larger 
branches to ascend or descend through the vegetation; after seem- 
ingly great deliberation, quick and agile leaps from twig to twig 
and twig to leal were undertaken. Movement for more than a few 
inches at a time was infrequent; during the intervening pauses the 
lizard generally lay Hush with a twig or leaf. Much more rapid 
and extensive changes oi perch occurred when the foliage was 
briefly disturbed by a gust of wind. At 10: 15 the three specimens 
remaining in sight were captured without difficulty. 

When the locality was revisited about noon, an occultus was 
soon located on a dead twig in a pile of cut branches. It showed 
the same alternation of pauses with slow movements over short 
distances observed earlier. After extensive wandering among the 
twigs it leaped to the upper surface of a fern frond, where it re- 
mained for perhaps a half hour; although this small brown lizard 
was difficult to distinguish on a substrate of bare twigs, on the 
frond it was always conspicuous, and the two pale orange spots 
at the base of the tail were displayed. Eventually, after a little 
maneuvering on its leaf, the lizard jumped to another tangle of 
dead twigs and disappeared. 

That evening intensive searching yielded six additional speci- 
mens in an hour and a half; three were on living twigs near leaves, 
one was on a long dead twig, one was at the tip of a very long 
descending branch, and a juvenile was on a dead fern. 



1969 ECOLOGY OF ANOLIS OCCULTUS 3 

On Puerto Rico the genus Anolis is represented by 10 species. 
All share adaptations for an active, diurnal, arboreal life, are pri- 
marily insectivorous, and in many cases are of similar size. Rand 
(1964) used the concept of a structural niche, which for arboreal 
Anolis is a combination of perch height and diameter as deter- 
mined by quantitative observation, to separate into three ecological 
groupings seven of the eight Puerto Rican Anolis recognized at the 
time. Temperature preference provided an additional niche dimen- 
sion that permitted Rand to separate the species within each group- 
ing. It is of interest to relate the structural niche of occultus to 
those of other Puerto Rican species, especially those recorded from 
the same locality. 

Anolis occultus appears to be an inhabitant of the peripheral 
foliage. Branches and bushes along trails have characterized all 
the productive collecting localities discovered to date; these some- 
what artificial conditions give the collector access to vegetation 
continuous with the more or less high canopy characteristic of 
Puerto Rico's montane forests. Thomas's (1965) hypothesis that 
occultus is normally an occupant of the canopy is probably sta- 
tistically correct, as suitable conditions near the ground occur only 
infrequently. There is no indication, however, that areas of dead 
branches and climbing plants are preferred; the specimens observed 
in this study showed a preference for fairly dense vegetation by 
day and no well-defined selection of dead or living twigs for sleep- 
ing perches. The sleeping positions selected in thin foliage or on 
dead twigs are probably optimal for early morning sunning; one of 
the two specimens observed to awaken was asleep in moderately 
dense foliage, but moved early to a more exposed position where 
it remained motionless for a long time. 

Rand (1964) has provided a detailed study of the structural 
niches for the three common Puerto Rican Anolis present at the 
La Mina area in the Sierra de Luquillo. Of the three species pres- 
ent at La Mina, A. evermanni uses almost exclusively perches of 
several inches or more diameter. A. gundlachi, which shows some- 
what greater preference for shade than evermanni, perches lower 
but also on tree trunks and branches of large diameter. A. krugi, 
while primarily found on grasses and similar plants, occasionally 
perches low in bushes on branches of moderate diameter. Al- 
though evermanni and krugi were common, gundlachi uncommon, 
and cristatellus rare at the locality of the recent occultus collection, 
the only species showing any overlap of structural niche with occul- 
tus in early April was evermanni. Juvenile evermanni were occa- 
sionally seen on twigs and small branches, although in general they 



4 breviora No. 312 

seem to prefer perches of large diameter. This overlap is prob- 
ably over-emphasized by an examination of trail-margin condi- 
tions; although, as Rand noted, studies of Anolis ecology are biased 
by the restriction of accurate observation to the lowest level oi 
the forest, the available evidence indicates that evermanni lives 
below the canopy, while occultus is largely a species of the canopy. 
If the observations of as many as six occultus in a small clump of 
vegetation can be considered indicative of the population density 
at this locality, occultus is much commoner than juvenile ever- 
manni, onl\ one of which was noticed. Thomas recorded gund- 
Icichi. cristatellus, krugi, and evermanni from localities where he 
took occultus. 

Anolis occultus seems well-adapted structurally, in coloration. 
and behaviorally for a cryptic existence in the peripheral foliage 
of trees and bushes. Long and slender, with a downward tapering 
snout, resting specimens merge easily with the outline of a twig 
and cast little or no shadow ; from the distance of only a few feet a 
human observer finds the blending of outline of lizard and twig 
very deceptive. Although its use has only been observed at night, 
a prehensile tail is presumably of some diurnal advantage to 
occultus, if only to hold it firmly Hush with a perch. Thomas 
(1965) described the coloration of live occultus in detail. Speci- 
mens observed In the present study by day were grey or brown, 
the degree of patterning \ariable; although individuals crossed 
leaves on several occasions, and one lingered for a considerable 
length oi time on a fern frond, the green phase recorded by 
Thomas was not observed. The orange spots at the base of the 
tail are conspicuous in some postures; the) may be eye spots to 
ward olf predation or possibly function in intraspecific communi- 
cation. Motion is slow and continuous over only short distances. 
The tendency for occultus to be most active when the vegetation is 
in general motion suggests that the species seeks to remain incon- 
spicuous at all times. The selection of exposed surfaces for sleeping 
may, by warming the li/ard as early as possible on cool mornings, 
maximize the number of daylight hours during which occultus is 
alert and active. 

ACKNOWLEDGMENTS 

Field work was in part supported by National Science Founda- 
tion Grant GB6944 to Ernest E. Williams. Thanks are due to 
Dr. Williams, who sponsored this collection of Puerto Rican rep- 
tiles and expressed particular interest in the occultus problem; to 



1969 ECOLOGY OF ANOL1S OCCULTUS 5 

Mr. Richard Thomas, who supplied information on collecting in 
Puerto Rico; and to Dr. Williams and Mr. Thomas Schoener, who 
read and provided valuable criticism of this paper. 

LITERATURE CiTED 

Rand, A. Stanley 

1964. Ecological distribution of anoline lizards of Puerto Rico. Ecol- 
ogy 45: 745-752. 

Thomas, Richard 

1965. Field observations of Anolis occultus Williams and Rivero. Part 
II in: E. E. Williams, J. A. Rivero, and R. Thomas, A new anole 
(Sauria, Iguanidae) from Puerto Rico. Breviora, No. 231: 10-18. 

(Received 6 November 1968.) 



BREVIORA 

Museem of Contiparative Zoology 

Cambridge, Mass. 31 March, 1969 Number 313 



LOUIS AGASSIZ'S NUMBERS FOR STEAMER BLAKE 
STATIONS OF 1877-78, 1878-79 

Myvanwy M. Dick 

Abstract. Correct Blake station numbers corresponding to Roman 
numerals used by Louis Agassiz for specimens collected by the Blake, 1877- 
1878, 1878-1879. 

Eschmeyer (1965) demonstrated that frequent errors in locality 
data for fishes taken during Blake expeditions, 1877-1879, have 
been included in several major published works on fishes and in 
the fish catalogue, U. S. National Museum. These errors were 
caused by interpreting as station numbers Roman numerals in- 
cluded with the specimens — - numerals intended for other purposes. 
For example, Goode and Bean (1896: 296-297) reported Cal- 
lidnymus himantophorus from Blake Station XXX and used data 
for that station for the station list. "XXX," however, is a number 
applied to the specimen, apparently by L. Agassiz, the correct 
station number being 274. Eschmeyer's account of these errors 
and their perpetuation is excellent and correct, but as no catalogue 
relating Agassiz's Roman numeral designation to true Blake 
Station numbers was known to exist, the resulting specific pub- 
lished errors could be corrected only with difficulty and in many 
instances not at all. 

In collating early papers and catalogues relating to the Fish 
Department of the Museum of Comparative Zoology, a special 
catalogue of deep sea fish from the Blake expedition was found. 
Judging by the handwriting, this was prepared by Louis Agassiz. 
This contains the Roman numerals, which had been incorrectly 
interpreted as Blake Station numbers, and corresponding Blake 
Stations and localities. Each number was apparently intended to 
apply to a specific fish. Occasionally, in the Blake catalogue of 
1878-1879, the same number is given in pencil as "Duplicate," 
with a different Blake Station. Presumably this would indicate 
fish of the same species but a different locality. This presents the 



2 BREVIORA No. 313 

possibility of questionable localities for specimens having just the 
Agassiz numbers, such as "Agassiz V," which would allow seven 
localities under "duplicate Agassiz V." 

In the Agassiz catalogue of the Blake expedition of 1877 1878 
the Roman numeral is not always accompanied by a Blake Station 
number; however a locality is given, except in a very few instances 
where there are entries with "no locality." 

Several specimens from the Blake collections were sent to 
Steindachner in Vienna for examination. These were assigned 
numbers from the Agassi/ catalogue with their locality, but the 
Blake Station numbers were omitted. Hence material described by 
Dr. Steindachner would have the Agassiz number and correct 
locality but would not include a Blake Station number. 

In general, however, Agassiz's holograph catalogue can be used 
to relate his Roman numeral designations to Blake Station numbers 
and hence to the correct data as published b\ Pierce and Patterson 
( 1870), Agassi/ (1881), and Smith ( I 889). Agassiz's catalogue 
is reproduced below. 

LITERATURE CITED 

Agassiz. A. 

I SKI. 1 ist o\~ dredging stations occupied (.luring the year 1 880 by the 
U.S. Coast Survey Steamer "Blake." Bull. Mus. Comp. Zool.. 
8(4): 95-98. 

ESCHMI HR.W. N. 

1965. Station data errors for fishes collected hv the Steamer Make. 
Copeia.2: 236-238. 

Goode. G. B.. and I H. Bl vn 

1896. Oceanic ichthyology. Mem. Mus. Comp. Zool., 22: 1-553. 

J'ii R( i . B.. and C. P. Patterson 

1879. 1 ist of dredging stations occupied by the United States Coast 
Survey steamers "Corwin." "Bibb," H.issler," and "Blake," 
from 1867 to 1879. Bull. Mus. Comp. Zool.. 6 ( I ): 1-15. 

Smith, S. 

1889. lists of the dredging stations of the U. S. Fish Commission. 
Ann. Rept. Comm. Fish Fish. (1886): 873-1017. 

(Received 6 December 1968.) 



1969 



BLAKE STATION NUMBERS 



CAT. DEEP SEA FISH BLAKE EXP. 1878-79 



Present 
Number 

I 

II 
III 
IV 



V 



VI 

VII 

VIII 



IV dup. 

Vdup. 
Vdup. 
Vdup. 
Vdup. 
Vdup. 
Vdup. 
Vdup. 



IX 



VIII dup. 

VIII dup. 

IX dup. 
IX dup. 
IX dup. 

X 

Xdup. 
XI 
XII 
XIII 
XIV 

XIV dup. 

XIV dup. 

XIV dup. 
XV 
XVI 
XVII 
XVIII 

XVIII dup. 
XIX 
XX 
XXI 
XXII 
XXIII 
XXIV 
XXV 



Dredge 
Number 

276 

278 

230 

274 

147 

240 

264 

153 

222 

258 

260 

150 

185 

290 

148 

262 

116 

290 

281 

274 

275 

148 

290 

297 

210 

142 

248 

176 

227 

285 

236 

148 

281 

275 

274 

274 

297 

142 

167 

147 

147 

191 

293 



Depth 

fms. 

94 

69 
464 
209 
250 
164 
416 
303 
422 
159 
291 
375 
333 

73 
208 

92 
150 

73 
288 
209 
218 
208 

78 
123 
191 

27 
161 
391 
573 
159 
1591 
208 
288 
218 
209 
209 
123 

27 

175 

250 

250 

108 to 250 

82.. _ 



Localities 



Number of 
Specimens 



off Barbados 

off Barbados 

off St. Vincent 

off Barbados 

off St. Kitts 

off Grenadines 

off Grenada 

off Montserrat 

off St. Lucia 

off Grenada 

off Grenada 

off Nevis 

off Dominica 

off Barbados 

off St. Kitts 

off Grenada 

off Jamaica 

off Barbados 

off Barbados 

off Barbados 

off Barbados 

off St. Kitts 

off Barbados 

off Barbados 

off Martinique 

off Flannegans Passage 

off Grenada 

off Dominica 

off St. Vincent 

off Grenada 

off Bequia 

off St. Kitts 

off Barbados 

off Barbados 

off Barbados 

off Barbados 

off Barbados 

Flannegans Passage 

off Guadeloupe 

off St. Kitts 

off St. Kitts 

off Dominica 

off Barbados 



4 




BREVIORA 


NO. 3 1 3 


Present 


Dredge 


Depth 


Localities Number of 


Number 


Number 


fms. 


Specimens 


XXVI 


220 


116 


off St. Lucia 1 


XXVI! 


231 


95 


off St. Vincent 1 


XXVIII 


293 


82 


off Barbados 1 


XXIX 


293 


82 


off Barbados 1 


XXX 


274 


209 


off Barbados 1 


XXX dup. 


281 


288 


off Barbados 1 


XXX dup. 


274 


209 


off Barbados 


XXX dup. 


258 


159 


off Grenada 1 


XXX dup. 


291 


200 


off Barbados 3 


XXXI 


273 


103 


off Barbados 1 


XXX 11 


261 


340 


olTGrenad.i 1 


XXXIII 


132 


115 


off Santa Cruz 1 


XXXIV 


253 


92 


off Grenada 1 


XXXV 


136 


508 


off Santa Cruz 1 


XXXV dup. 


221 


423 


off St. Lucia 1 


XXXV dup. 


284 


347 


off Barbados 1 


XXXVI 


280 


221 


off Barbados 1 


XXXVII 


151 


365 


off Nevis 1 


XXXVIII 


2 1 2 


317 


off Martinique 

with parasite 1 


XXXIX 


121 


2393 


off Santa Cruz 1 


XL 


200 


472 


off Martinique 1 


XL] 


185 


333 


off Dominique 1 


XLII 


21 1 


357 


off Martinique 1 


XLII dup. 


222 


422 


off St. Lucia 3 


XLII dup. 


212 


317 


off Martinique 1 


XLII dup. 


151 


365 


off Nevis 3 


XL 11 


288 


399 


off Barbados 1 


\I II dup. 


260 


291 


off Grenada 2 


XLII dup. 


221 


423 


off St. Lucia 1 


XLII dup 


260 


291 


off Grenada 4 


XLII dup. 


150 


375 


off Nevis 3 


XLII dup. 


261 


340 


off Grenada I 


XL1II 


266 


461 


off Grenada 1 


XLIV 


248 


161 


off Grenada 1 


XLV 


104 


500 


Old Bahama Channel 3 


XLVdup. 


104 


500 


Old Bahama Channel 10 ± 


XLVI 


271 


458 


off Bequia 1 


XLVII 


188 


372 


off Dominica 3 


XLVIII 


227 


573 


off St. Vincent 1 


XLIX 


221 


423 


off St. Lucia 1 


XLIX dup. 


221 


423 


off St. Lucia 1 


XLIX dup. 


190 


524 


off Dominica 


L 


227 


573 


off St. Vincent 1 


LI 


148 


208 


off St. Kitts 1 



1969 



BLAKE STATION NUMBERS 



LII 275 218 

Llldup. 275 218 

LII dup. 147 250 

Llldup. 281 288 

LIU 295 180 

LUIdup. 295 180 

LUIdup. 281 288 

LUIdup. 291 200 

LIII dup. 274 209 

LUIdup. 275 218 

LIV 283 237 

LV 262 92 

LVI 231 95 

LVII 132 115 

LVIIdup. 132 115 

LVIII 129 314 

LIX 281 288 

LX 297 123 

LXI 249 262 

LXII 274 209 

LXIII 274 209 

LXIIIdup. 275 218 

LXIII dup. 274 209 

LXIIIdup. 172 62 to 108 

LXIV 297 123 

LXV 297 123 

LXV dup. 297 123 

LXVI 235 1507 

LXVII 222 422 

LXVII 222 422 

LXVIII 160 393 

LXVIII dup. 137 625 

LXVIII dup. 151 365 

LX1X 161 583 

LXX 163 769 

LXXdup. 163 769 

LXXI 163 769 

LXXII 230 464 

LXXIII 200 472 

LXXIV 295 180 

LXXIVdup. 291 200 

LXXV 281 288 

LXXVdup. 281 288 

LXXV dup. 260 291 

LXXVI 265 576 

LXXVI dup. 265 576 

LXXVI dup. 288 399 



off Barbados 
off Barbados 
off St. Kitts 
off Barbados 
off Barbados 
off Barbados 
off Barbados 
off Barbados 
off Barbados 
off Barbados 
off Barbados 
off Grenada 
off St. Vincent 
off Santa Cruz 
off Santa Cruz 
off Santa Cruz 
off Barbados 
off Barbados 
off Grenada 
off Barbados 
off Barbados 
off Barbados 
off Barbados 
off Guadeloupe 
off Barbados 
off Barbados 
off Barbados 
off Bequia 
off St. Lucia 
off St. Lucia 
off Guadeloupe 
off Santa Cruz 
off Nevis 
off Guadeloupe 
off Guadeloupe 
off Guadeloupe 
off Guadeloupe 
off St. Vincent 
off Martinique 
off Barbados 
off Barbados 
off Barbados 
off Barbados 
off Grenada 
off Grenada 
off Grenada 
off Barbados 



1 
2 
1 
1 
2 
20 
14 
3 
4 
2 



6 




BRliVIORA 




No. 313 


P rescn i 


Dredge 


Depth 


Localities 


Number of 


Number 


Number 


litis. 




Specimens 


LXXV11 


151 


365 


off Nevis 




LXXVIIdup 


151 


365 


off Nevis 




LXXVIIdup 


222 


422 


off St. Lucia 




LXXVIIdup 


260 


291 


off Grenada 




LXXVIIdup 


150 


375 


off Nevis 




1 \\\ 11 dup 


135 


450 


off Santa Cruz 


2 


I XXVII dup 


185 


333 


oil Dominica 




LXXV11I 


291 


200 


off Barbados 




1 XXVIII du] 


D. 2 l M 


200 


oil' Barbados 




LXXIX 


274 


209 


oil Barbados 




1 \\l\ dup. 


274 


209 


off Barbados 




LXXIX dup. 


291 


200 


Off Barbados 




LXXIX dup. 


193 


169 


mi Martinique 




LXXIX dup. 


L49 


60 to ISO 


Off Sl. Kills 




LXXIX dup. 


[46 


24.^ 


off St. Kitts 




LXXX 


200 


4'2 


off Martinique 




1 \X.\ dup. 


2()() 


4^2 


oil Martinique 




LXXXdup. 


175 


MS 


oil Dominica 




LXXX dup. 


227 


573 


off St. Vincent 




LXXXdup. 


135 


450 


off Santa Cruz 




LXXX dup. 


163 


769 


oil ( iuadeloupe 




LXXX dup. 


227 


573 


off St. Vincent 




LXXX1 


151 


J65 


off Nevis 




1 VXXI dup. 


151 


365 


oil Nevis 




LXXXII 


205 


334 


off Martinique 




LXXX 11 dup 


205 


334 


otl Martinique 




LXXXII dup 


130 


451 


off Santa Cruz 




LXXXII dup 


186 


333 


off Dominica 




LXXXII dup 


222 


422 


off St. Lucia 




LXXXIII 


154 


164 


off Grenada 




LXXXIV 


182 


1131 


off Dominica 




LXXXV 


151 


365 


olf Nevis 




LXXX\ 1 


180 


982 


olT Dominica 




LXXXYI du 


p. 180 


982 


off Dominica 




LXXXVII 


196 


1030 


off Martinique 




LXXXVIII 


214 


476 


off Martinique 




LXXX\ III I 


lup. 200 


472 


off Martinique 




LXXXVIII ( 


tup. 163 


769 


off Guadeloupe 




LXXXIX 


205 


334 


off Martinique 




XC 


258 


159 


off Grenada 




XC dup. 


283 


237 


off Barbados 




XCdup. 


275 


218 


off Barbados 


3 


XCI 


180 


982 


off Dominica 


1 



1969 



BLAKE STATION NUMBERS 



XCII 

XCII dup. 

XCII dup. 

XCII dup. 

XCII dup. 

XCII dup. 

XCII dup. 

XCII dup. 

XCII dup. 

XCII dup. 
XCIII 
XCIV 

xcv 

XCVI 

XCVI dup. 

XCVI dup. 
XCVII 

XCVII dup. 
XCVIII 

XCVIII dup. 

XCVIII dup. 

XCVIII dup. 
XCIX 

XCIX dup. 
C 
CI 
CH 

cm 
civ 

CI dup. 

CI dup. 
CV 
CVI 
CVII 
CVIII 
CIX 

ex 

CXI 
CXII 
CXIII 
CXIV 

cxv 

CXVI 
CXVII 
CXVIII 
CXIX 

exx 



265 


576 


off Grenada 




161 


583 


off Guadeloupe 




135 


450 


off Santa Cruz 




185 


333 


off Dominica 


2 


200 


472 


off Martinique 




227 


573 


off St. Vincent 




130 


451 


off Santa Cruz 




195 


501 


off Martinique 




188 


399 


off Barbados 




239 


338 


off Grenadines 




260 


291 


off Grenada 




100 


250-400 


off Moro Castle, Cuba 




185 


333 


off Dominica 




163 


769 


off Guadeloupe 




162 


734 


off Guadeloupe 


2 


227 


573 


off St. Vincent 


4 


274 


209 


off Barbados 


2 


274 


209 


off Barbados 


31 


227 


573 


off St. Vincent 




227 


573 


off St. Vincent 


3 


143 


298 


off Montserrat 




257 


553 


off Grenada 




262 


90 


off Grenada 


3 


262 


90 


off Grenada 


30 


132 


115 


off Santa Cruz 




231 


95 


off St. Vincent 




142 


27 


Flannegans Passage 




142 


27 


Flannegans Passage 




295 


180 


off Barbados 




291 


200 


off Barbados 


2 


297 


123 


off Barbados 




150 


375 


off Nevis 




190 


524 


off Dominica 




291 


200 


off Barbados 




148 


208 


off St. Kitts 




188 


372 


off Dominica 




258 


159 


off Grenada 




300 


82 


off Barbados 




243 


82 


off Barbados 




151 


365 


off Nevis 




290 


73 


off Barbados 




243 


82 


off Barbados 




276 


94 


off Barbados 




276 


94 


off Barbados 




290 


78 


off Barbados 




243 


82 


off Barbados 




253 


92 


off Grenada 





8 




HREVIORA 


N« 


3. 3 1 3 


Present 


Dredge 


Depth 


Localities Number of 


Number 


Number 


fms. 


Specimens 


CXXI 


128 


180 


off Santa Cruz 




CXXII 


142 


27 


Flannegans Passage 




CXXIII 


142 


27 


1 l.mnegans Passage 




CXXIV 


142 


27 


1 lanne^ans Passage 




cxxv 


271 


458 


off Bequia 




CXXVI 


278 


69 


off Barbados 




< WVII 


243 


82 


off Barbados 




CXXVIll 


300 


82 


off Barbados 




CXXIX 


132 


115 


off Santa Cruz 




1 \\x 


220 


1 \b 


oil St. Lucia 




( \\\l 


276 


94 


off Barbados 




1 WXII 


104 


500 


Old Bahama Channel 




( XXXIII 


241 


163 


off Cariacou 




c xxxrv 


293 


82 


off Barbados 




I XXXV 


104 


500 


Old Bahama Channel 




CXXXN dup. 


104 


500 


Old Bahama Channel 




( WXVI 


104 


500 


Old Bahama Channel 




CXXXV11 


104 


500 


Old Bahama Channel 




CXXXVIdup. 


104 


500 


Old Bahama Channel 




( WW III 


104 


500 


( )ld Bahama Channel 




< XXXIX 


109 


1554 


off Caya de Moa 




CXL 


HI 


1200 


la!. 19°07'N, 
Ion. 74 52'W 




( XI 1 


150 


375 


off Nevis 




CXI. II 


288 


399 


off Barbados 




( XI III 


150 


375 


off Nevis 




CXI l\ 


256 


370 


off Grenada 




CXLV 


284 


347 


off Barbados 




CXLV1 


283 


237 


off Barbados 




CXLVII 


205 


surface 


off Martinique 


2 


CXLVIII 


201 


565 


off Martinique 


2 


CXLVIIIdup. 


201 


565 


off Martinique 


3 


CXLIX 


212 


317 


off Martinique 


2 


CL 


204 


476 


off Martinique 


2 


( 1 1 


217 


398 


off St. Lucia 




CLII 


217 


398 


off St. Lucia 




CLIII 


222 


422 


off St. Lucia 




CL1V 


124 


580 


off Santa Cruz 




( 1 V 


207 


826 


off Martinique 




CLVI 


182 


1131 


off Dominique 




CLVII 


190 


524 


off Dominique 




CLV1II 


190 


524 


off Dominique 


2 


CLIX 


185 


333 


off Dominique 




CLIX dup. 


185 


333 


off Dominique 




CLIX dup. 


188 


372 


off Dominique 


4 



1969 



BLAKE STATION NUMBERS 



CLX 


188 


372 


off Dominique 1 


CLXI 


236 


1591 


off Bequia 1 


CLXII 


180 


982 


off Dominica 1 


CLXIII 


173 


734 


off Guadaloupe 2 


CLXIV 




769 


off Guadaloupe 1 


Blake Exp. 1877-78 








CLXV 


29 


955 


lat.24°36'n,lon.84°05'w 1 


CLXVI 


29 


955 


lat.24°36'n,lon.84°05'w 1 


CLXVII 


12 


36 


lat.24°46'n,lon.83°16'w 1 


CLXVIII 


36 


84 


lat.23°13'n,lon.89°10'w 1 


CLXIX 


12 


36 


lat.24°46'n,lon.83°16'w 1 


CLXX 


29 


955 


lat.24°36'n,lon.84°05'w 1 


CLXXI 


11 


37 


lat.24°43'n,lon.83°25'w 5 


CLXXII 


29 


955 


lat.24°36'n,lon.84°05'w 2 


CLXXIII 


29 


955 


lat.24°36'n,lon.84°05'w 1 


CLXXIV 


31 


1850 


lat.24°33'n,lon.84°23'w 1 


CLXXV 


186 


88 


off Alligator Keys 1 


CLXXVI 


37 


35 


Alacran Shoals, Yucatan 9 


CLXXVII 


37 


35 


Alacran Shoals, Yucatan 1 


CLXXIX 


47 


321 


lat.28°42'nlon.88°40'w 4 


CLXXX 


6 


137 


lat.24°17.5'n 

Ion. 82°09'w 1 


CLXXXI 


47 


321 


lat.28°42'nlon.88°40'w 1 


CLXXXII 




135 


off Key West 1 


CLXXXIII 




270 


Santaren Channel 1 


CLXXXIV 




128 to 240 


Two miles off Havana 1 


CLXXXV 


45 


101 


lat.25°33'nlon.84°21'w 1 


CLXXXVI 


32 


95 


lat.24°52'nlon.88°05'w 1 


CLXXXVII 




177 


off Havana 1 


CLXXXVIII 




40 


off Cary's Fort Reef 1 


CLXXXIX 


29 


955 


lat.44°36'nlon.84°05'w 5 


CXC 


46 


888 


lat. 25°43'nlon. 

84°47'30" 1 


CXCI 


44 


539 


lat.25°33'nlon.84°35'w 1 


CXCII 


36 


84 


lat. 23°13'nlong. 

84°16'w 25 


CXCIII 


36 


84 


lat. 23°13'nlong. 

84°16'w 25 


CXCIV 




175 


off Havana 1 


cxcv 




40 


off West Reef I 


CXCVI 




98 


"off Panfii Reef" 1 


CXCVII 




42 


"W. of Tortugas" 1 


CXCVIII 




117 


Illegible 1 


CXCIX 




37 


W. of Tortugas 1 


cc 




306 


W. of Tortugas 1 


CCI 




68 


W. of Tortugas 1 



10 



BRFA'IORA 



NO. 3 1 3 



P rescn I 
Number 

ecu 

CCill 
CCIV 

ccv 

CCVI 
CCV11 



Dredge Depth Localities V 'umber of 

Number jms. Specimens 

37 W of Tortugas 

118 off Alligator Reef 

31 1920 hit. 24 33' Ion. 84 23' 

43 339 lat. 24°08'lon. 82 51' 

35 804 lat. 23 54-46" 

Ion. 88°58' 
37 35 4 miles w. of whale rock 

w. of Alacran Reels 



CCV11I 


29 


955 


lat. 24 36' Ion. 84 05' 


CCIV 


11 


37 


lat. 24 43'lon. 83 c 25' 


( ( \ 




45 


off French Reef 


CCX1 




35 


off Cargoforl Reef 


( ( XII 




320 


off Cargofort Reef 


( ( XIII 




413 


off ( ape San \ntonio 


( ( \l\ 




96 


loc? 


( ( XV 




35 


W. of Tortugas 


( ( XVI 




1 19 


lat. 26 31'lon.85°03' 


( ( \\ II 


13 


"42 


3V4 N. W. off Havana 


1 ( \\ III 


47 


321 


lat 28 42' Ion. 88 40' 


( ( XIX 


30 


968 


lat 24 33' Ion. 84 05' 


< [ XX 


2" 


955 


lat. 24°36' Ion. 84 05' 


* 1 XXI 


29 


955 


lat. 24 36' Ion. 84 05' 


( ( XXII 


47 


J21 


lat. 2S 42' Ion. 88 40' 


( 1 XXIII 




84 


lat 23 13' Ion. 89 10' 


( ( XXIV 


11 


37 


lat. 24 43' Ion. 83 25' 


( ( \\\ 


47 


321 


lat. 28 42' Ion. 88 '40' 


( ( \\\ 1 


12 


J6 


hit. 24 34' Ion. 83 16' 


CCX.W II 


12 


36 


lat. 24 34' Ion. 83 16' 


( 1 \\\ 111 


12 


36 


lat. 24 34' Ion. 83 16' 


( ( XXIX 


14 


850 


lat. 23 18' Ion. 8221' 


( ( XXX 


36 


84 


lat. 2r 13' Ion. 89° 10' 


( ( XXXI 


29 


955 


lat. 24°36' Ion. 84 05' 


( ( XXXII 


37 


35 


Alacran Shoals, Yucatan 


1 ( XXIII 


29 


955 


lat. 24 36'lon. 84°05' 


CCXXXIV 


10 


37 


lat. 24°44'lon. 83°00' 


CCXXXV 


3 


243 to 450 


off Havana 


CCXXXVI 


36 


84 


lat 23 13' Ion. 89 10' 


( ( XXXVII 


12 


36 


lat. 24°34'lon. 83° 16' 


CCXXXII1 


29 


955 


lat. 24 36'lon. 84 05' 


CCXXXIX 


10 


37 


lat. 24°44'lon. 83°00' 


CCXL 


29 


955 


lat. 24'36'lon. 8405' 


CCXLI 


36 


84 


lat. 23 13'lon. 89 10' 


CCXLII 


36 


84 


lat. 23 13' Ion. 89 10' 


CCXLIII 


11 


37 


lat. 24"43'lon. 83*25' 


CCXL1V 


36 


84 


lat. 23° 13' Ion. 89 10' 



1969 



BLAKE STATION NUMBERS 



11 



CCXLV 

CCXLVI 

CCXLVII 

CCXLVIII 

CCXLIX 

CCL 

CCLI 

CCLII 

CCLIII 

CCLIV 

CCLV 

CCLVI 

CCLVII 

CCLVIII 

CCLIX 

CCLX 

CCLXI 

CCLXII 

CCLXIII 

CCLXIV 

CCLXV 

CCLXVI 

CCLXV1I 

CCLXVIII 

CCLXIX 

CCLXX 

CCLXXI 

CCLXXII 

CCLXXIII 

CCLXXIV 

CCLXXV 

CCLXXVI 



3 


450 to 243 


off Havana 


1 


29 


955 


lat. 24°36'lon. 84°05' 


1 


12 


36 


lat. 24°34'lon. 83° 16' 


9 


36 


84 


lat. 23°13'lon. 89°10' 


1 


11 


37 


lat. 24°43'lon. 83°25' 


1 


11 


37 


lat. 24°43'lon.83°25' 


1 


37 


35 


Alacran Shoals, Yucatan 


5 


37 


35 


Alacran Shoals, Yucatan 


2 


11 


37 


lat. 24°43'lon. 83°25' 


1 


37 


35 


Alacran Shoals, Yucatan 


1 


36 


84 


lat. 23° 13' Ion. 89° 10' 


9 


47 


321 


lat. 28°42'lon. 88°40' 


1 


36 


84 


lat. 23°13'lon. 89°10' 


1 


11 


37 


lat. 24°43'lon. 83°25' 


1 


36 


84 


lat. 23°13'lon. 89°10' 


2 


36 


84 


lat. 23°13'lon. 89°10' 


3 


36 


84 


lat. 23° 13' Ion. 89° 10' 


1 


14 




lat. 23°18'lon. 82°21' 


2 


11 


37 


lat. 24°43'lon. 83°25' 


1 


36 


84 


lat. 23°13'lon. 89°10' 


2 


29 


955 


lat. 24°36'lon. 84°05' 


1 


29 


955 


lat. 24°36'lon. 84°05' 


1 


29 


955 


lat. 24°36'lon, 84°05' 


1 


29 


955 


lat. 24°36'lon. 84°05' 


1 






no loc. 


1 


31 


1850 


lat. 24°33'lon. 84°23' 


1 






no loc. 


1 






no loc. 


1 






no loc. 


1 



BREVIORA 

Museumi of Comparative Zoology 

Cambridge, Mass. 31 March, 1969 Number 314 



THE CRANIAL ANATOMY OF THE 
PERMIAN AMPHIBIAN PANTYLUS 

Alfred Sherwood Romer 



Abstract. A detailed description is given of the skull structure of the 
early Permian tetrapod Pantylus, using both whole specimens and a skull 
serial sectioned and reconstructed in wax plates. This study definitely proves 
that Pantylus, often thought to be a reptile, is a microsaurian amphibian, 
and furnishes further proof that microsaurs cannot be considered reptilian 
ancestors. 

INTRODUCTION 

Pantylus, known in the past mainly from a heavily-built, heart- 
shaped skull carrying a powerful dentition, is an early Permian 
tetrapod from the Texas red beds whose systematic position has 
been open to considerable doubt. Cope, who obtained the first 
materials of this animal, originally considered it to be a batrachian 
(1881), but later classed it among the Reptilia (1892, 1896), as 
did a long series of later workers, including Case (1911, 1912, 
1915), Huene (1913), Broom (1913), Williston (1916a, 1916b, 
1925), and Wilson (1951). Two decades or so ago, however, in 
considering the nature of the Paleozoic microsaurs, it became obvi- 
ous to me that it was, as far as known, basically similar in structure 
to members of the microsaurian family Gymnarthridae, and hence 
should be placed in the order Microsauria (Romer, 1950). My 
conclusion has, I think, been generally accepted by recent workers 
(cf., for example, Dechaseaux, 1955; Gregory, Peabody, and Price, 
1956; Tatarinov, 1964). However, it seems important to add as 
much as possible to our present inadequate knowledge of its struc- 
ture in order to attempt to fix its systematic position. 

But the importance of such study goes further. Most microsaurs 
are very poorly known anatomically. If (as I hope to demonstrate) 
Pantylus is a true microsaur, a study of its cranial structure may 
prove of value in fixing the phylogenetic position of that group as a 
whole. Microsaurs have often been claimed to be related to the 



2 brhviora No. 314 

captorhinomorph cotylosaurs and to be possibly ancestral to the 
reptiles, or to at least part of that class (ef.. for example, Williston. 
1908; Westoll. 1942a, 1942b; Olson, 1947; Huene, 1948; Vaughn, 
I960, 1962; Brough and Brough, L967). 

Another possibility that has been suggested is that the microsaurs 
may be ancestral to modern amphibians, particularly the urodeles 
(Romer, 1950, etc.), although this suggestion is far from generally 
accepted (ct\, for example. Parsons and Williams. 1963). The 
pedigree of the microsaurs is in doubt. I have, since 1933, some- 
what arbitraril) combined the microsaurs with the aistopods and 
nectrideans in the Lepospondyli, in an expanded use of that term. 
But the interrelationships of the three groups are far from certain. 
And while the derivation of the l.abvrinthodontia from the rhipi- 
distian crossoptervgians seems clear, there has been little evidence 
to tie any of the three lepospondyl orders into this evolutionary 

series. 

Previous studies; materials. Cope's original materials consisted 
o( two imperfect skulls; from these he was able to describe much of 
the roof pattern and part oi the palate. later studies of these speci- 
mens by Case (1911) and lluene (1913) added little; however, 
here, as frequently, Broom (1913) was able to give greater depth to 
analysis of the specimens available to him. Mehl. in 19 12. described 
a fragmentary skull collected for the University o\' Chicago in 1908 
by Paul C. Miller, but added little o\ interest. The latter collector, 
in expeditions of 1913 to 1916. obtained a series of specimens, in- 
cluding several skulls and some postcranial materials, from the 
Mitchell Creek locality; Williston ( 19 Km) described much of this 
material, giving an adequate account of the skull roof (except the 
temporal region), some further data on the lower jaw and palate. 
and a few scattered details regarding the braincase and postcranial 

skeleton. 

No further materials were found until, in 1939. Mr. A. Witte of 
Henrietta, in charge of a WPA group under the auspices of the 
Texas Bureau of Economic Geology, came upon a "nest" of speci- 
mens near Windthorst. Archer County. Texas; this suite, now en- 
tered as No. 40.001 (1-8) in the Texas Memorial Museum. Austin, 
includes several skulls and considerable postcranial materials. In 
Mr. Wittc's possession is a further small skull from this locality. 
Post-war trips to Texas from the Harvard Museum of Comparative 
Zoology (abbreviated as MCZ below) yielded further specimens — 
two skulls from the Archer City bonebed. a partial skull from 
Montaeue County, and two incomplete crania from southern 
Archer County. 



1969 CRANIAL ANATOMY OF PANTYLUS 3 

Study of this newer material, together with re-examination of 
older Chicago specimens, now makes it possible to give a nearly 
complete account of the skeletal anatomy of Pantylus. The post- 
cranial skeleton is now being studied by Dr. Robert L. Carroll of 
McGill University. Both Dr. Carroll and I are much indebted to 
Drs. E. C. Olson and R. Zangerl of Chicago, Drs. John A. Wilson 
and Wann Langston, Jr., of Austin, and Mr. Witte for the loan of 
materials. Also, I gratefully acknowledge financial aid for the 
preparation of material from National Science Foundation Grant 
GB 500. 

For cranial anatomy, some of the older Chicago and American 
Museum specimens were restudied. Valuable were, as noted, Har- 
vard skulls MCZ 2040 and 3302. The latter is incomplete and 
broken, but the parts preserved show excellent detail, including 
some braincase structure. MCZ 2040 is complete, except that the 
roof is broken off posteriorly on one side. It shows surface struc- 
tures perfectly, and the break is valuable in that it reveals much of 
the braincase. Several of the Texas Memorial Museum specimens 
show aspects of skull anatomy. Most important was No. 40,001 
( 1 ), which, with the consent of Prof. John A. Wilson, was serially 
sectioned and reconstructed, enlarged X 6, in wax plates. The sur- 
faces of each section were preserved by the cellulose "peel" method, 
and at each grinding two peels were taken. Although this sectioned 
specimen was extremely valuable in revealing internal structure, and 
is the main "document" on which Figures 3 - 12 are based, it proved 
disappointing in two regards. ( 1 ) Subsequent to burial, it was in- 
vaded by calcite, obviously crystallized out of ground water. This 
crystallization caused multiple small fractures in the skull, which 
have rendered details of sutures in the dermal bones difficult to 
determine. (2) Before burial, the skull received one or more vio- 
lent blows, which, apart from shattering a fraction of the roof (a 
matter of small moment), jarred loose the vomers, the epiptery- 
goids, the basisphenoid plus parasphenoid, and most of the otico- 
occipital elements of the braincase. For the most part, these ele- 
ments are still present in the specimen, lying, disarticulated and 
more or less battered, between the lower jaws. Their restoration and 
proper positioning in the braincase and palate presented obvious 
difficulties. Fortunately, most needed data on these disarticulated 
or demolished elements are available in MCZ 2040 and 3302. 

Stratigraphy and systematics. Cope's original specimens were 
collected by Jacob Boll in 1 880 from the "Big Wichita." In his first 
expedition, two years earlier, Boll had explored no higher in the 
Texas beds than exposures in the Belle Plains Formation of the 



4 breviora No. 314 

Wichita Group, between the site of the modern Diversion Dam on 
the Big Wichita River and the exposures along the Little Wichita 
south of Dundee. In 1880, however, he went somewhat further 
west, and there is reason to suppose that he collected at the Mitchell 
Creek locality from which Paul C. Miller later collected many valu- 
able amphibian specimens for S. W. Williston (Romer, 1928, 1935). 
This locality lies high in the Clyde Formation, very generally re- 
garded as the uppermost formation of the Wichita Group except for 
the overlying Lenders limestones. Miller's 1908 specimen, de- 
scribed by Mehl, is entered merely as from the "Big Wichita" but is 
probablv from Mitchell Creek. His later materials, described by 
\\ illistOll m 1916, are all definitely from Mitchell Creek. 

All the early described specimens, thus, are probably from a 
single locality and horizon near the summit of the Wichita Group. 
I he newer materials extend the range stratigraphically and topo- 
graphically. W ith two exceptions, all are from the Putnam Forma- 
tion, considerably lower in the Wichita Group (and in early days 
thought by many to be part of the Cisco Group of the Carboni- 
ferous). The Texas Museum specimens, and the individual later 
collected b\ Mr. Witte, were found in an exposure in Section 55, 
Block 3. of the Clark and Plumb Subdivision, about 4 miles north 
of Windthorst, Archer County; the horizon is about the middle of 
the Putnam Formation. One Harvard specimen, MCZ 2041, is 
from the J. J. Prather Survey, about 5 miles northwest of Stone- 
burg. Montague County; this is at about the same horizon as that of 
the Texas Museum specimens. Harvard skulls MCZ 2040 and 3302 
are from a bonebed about a mile southwest of Archer City, Archer 
County, which is close to the summit of the Putnam Formation. 
Two final specimens, MCZ 1913 and 3610. are a poorly preserved 
skull and a snout fragment from Section 1834, Texas Emigration 
and Land Co.. north of the West Fork of the Trinity River in south- 
central Archer County and just below the presumed boundary be- 
tween the Moran Formation and the overlying Putnam. It now 
appears, hence, that Pantylus was in existence for at least most of 
Wichita time. 

Cope's original specimens were given the name Pantylus cordatus; 
no other species properly pertaining to the genus has been described. 
Cope in 1 896 described as Pantylus co'icodus a fragmentary speci- 
men, mainly a tooth row, from Coffee Creek in the Arroyo Forma- 
tion of the Clear Fork Group. However, these teeth are bulbous, 
pointed structures, quite unlike those of other Pantylus specimens. 
As various later writers have noted, they do not appear to belong to 
Pantylus (Case, 191 1; Gregory et al., 1956), and the specimen is 



1969 CRANIAL ANATOMY OF PANTYLUS 5 

perhaps a caseid. A jaw fragment from Coffee Creek to which Cope 
gave the manuscript name Pantylus tryptichus is probably a captor- 
hinid (Cummins, 1908; Case, 1911; Gregory et al, 1956). Wil- 
liston in 1913 gave the name Ostodolepis brevispinatus to a series 
of small vertebrae from Coffee Creek. In 1916 he noted the simil- 
arity of these vertebrae to those of Pantylus. "Ostodolepis," as de- 
scribed by Case (1929), is a gymnarthrid microsaur of a different 
nature, but as noted by Carroll and Baird (1968), there is no 
guarantee that Case's specimen is related to the type. 

It is possible that the stratigraphically lower Putnam and Moran 
materials are specifically distinct from the earlier described speci- 
mens from the Clyde. There is some variation in skull size, in 
general amounting to about 20 per cent, most skulls ranging in 
measured or estimated length from 72 to 90 mm; but this variation 
may be related to age or sex. The Witte specimen has an estimated 
length of about 36 mm, and one Chicago specimen from Mitchell 
Creek is but 39 mm long; but these are surely young individuals, 
presumably cospecific with the larger individuals found with them. 
There are dental differences, in the description and figures of earlier 
writers, regarding the number of teeth in the marginal rows, 
noted later, and the pattern of toothplate teeth. But one may 
strongly suspect that the patterns figured are in some cases diagram- 
matic and not entirely trustworthy. Quite surely age (and sex?) 
factors enter into variations in toothplate dentition. There may be 
individual differences, as well. For example, the skull sectioned by 
me has a large central tooth in both upper and lower toothplates, 
but Williston mentions a specimen in which such a tooth was pres- 
ent above but not below, and figures (how accurate I do not know) 
of other specimens show no such enlarged tooth in either upper or 
lower dentition. 

DESCRIPTION 

Skull roof (Figs. 1, 2, 7, 12). The skull of Pantylus is very mas- 
sively built, and very broad and low — its width, which, from a 
rounded snout, spreads out greatly in the cheek region, is nearly as 
great as the length. There is no otic notch. The cheek in the jugal 
region extends downward so as to cover externally a considerable 
area of the lower jaw. Skull height, even allowing for the depth of 
the overhanging cheek region, is little more than a third of the 
length. The broad roof is nearly flat, although bending sharply 
downward at the margins of the "table," and there is a somewhat 
overhanging snout. The roof bears, as in most early or primitive 



BRFVIORA 



No. 314 



amphibians, patterns of ridges and intervening valleys radiating 
from the ossification centers of the dermal elements. There is no 
evidence of grooving for lateral line canals. The small skull in Mr. 
Wine's collection shows aearly straight sutures between roofing 




Fig. 1. Dorsal surface of skull, based on \l( / 2040; right posterior area 
restored.  1*4. Abbreviations for tins and following figures: ( ;. angular; 
ar. articular; a\p. anterior splenial; ba, basal articulation of braincase with 
palatal structures; ho. basioceipital: bs, basisphenoid; c, coronoid; ch, choana. 
</. dentary; c epipterygoid; en, external nans; ,<>. exoccipital; /. frontal: jo. 
fenestra ovalis; fv, foramen for transverse vein in pituitary region; /', jugal; 
/, lacrimal; in. maxilla: n, nasal: op, opisthotic; p, parietal; pal, palatine; 
pur. prearticular; pf. postfrontal; pm. premaxilla; po, postorbital; pp. post- 
parietal; pr, prootic; prf, prefrontal: ps, parasphenoid; psp, posterior splenial; 
pt. pterygoid; </. quadrate; qj, quadrato jugal; .«/, surangular; se, sphen- 
ethmoid; sm, septomaxilla; soc, supraoccipital; sq, squamosal; st. supra- 
temporal; v, vomer; /. //, V, X, foramina for cranial nerves. 



1969 CRANIAL ANATOMY OF PANTYLUS 7 

elements. With growth, however, the superficial portions of the 
roofing bones tend to interosculate, producing, as generally in Paleo- 
zoic amphibians, a series of zig-zag sutures. For the most part, the 
pattern is a bilaterally symmetrical one, although (as generally) 
four-square meetings of bones are not to be found. However, in the 
skull figured, the right frontal has invaded and taken over an area 
properly belonging to the right parietal. 

The premaxilla is thick externo-internally, but short; it covers 
mainly the "overhang" anteriorly leading down to the tooth row be- 
tween the external nares. It extends backward laterally below the 
naris to include the region of the base of the lateral premaxillary 
teeth. The nasals are likewise short, but broad, extending down 
laterally to the upper border of the nares. Frontals and parietals 
are long. There is no parietal foramen. 

The external naris, facing laterally, is large and subcircular in 
outline. The premaxilla forms most of its anterior border, the 
maxilla the lower margin, the nasal the upper boundary, while the 
lacrimal extends broadly forward to a point close to the posterior 
margin. However, the lacrimal and, to a lesser extent, the maxilla 
and nasal are excluded from the narial margin by a highly-developed 
septomaxilla. This forms a narrow external rim along the posterior 
half of the narial opening and then turns inward to form a posterior 
wall and partial floor and medial wall to the nasal passage inside the 
narial opening. Evidently the air passage turned sharply upward 
inside the external narial opening to reach a dorsally placed nasal 
capsule. The anterior opening of the lacrimal duct pierces the 
septomaxilla just internal to the narial opening. 

The lacrimal is well developed, extending from the orbit to the 
narial region; ventrally it is overlapped to some degree by the max- 
illa, but dorsally it extends high up on the side of the face, and pos- 
teriorly it extends, in a slender process, well back to a point below 
the orbit. Sections (Fig. 3) show that the lacrimal is essentially a 
cylindrical mass of bone surrounding the lacrimal duct and is, so to 
speak, cradled below by the maxilla. The thickness of the lacrimal 
bone produces a swelling of the generally smooth internal surface 
of the dermal roof anterior to the orbit; this swelling is continuous 
with a thickened anterior margin of the orbit which extends upward 
and backward along the prefrontal. The posterior opening of the 
lacrimal duct lies just inside the orbital margin. 

The orbit, facing essentially laterally, is a large and somewhat 
oval opening, its highest point being considerably posterior to the 
lowest point in its ventral margin. The prefrontal is well developed, 
occupying a large portion of the extended dorsal and anterodorsal 



8 



BREVIORA 



NO. 314 



margins of the orbit; dorsally it extends forward between frontal 
and lacrimal to reach the nasal. The postfrontal. abutting pos- 
teriorly on the prefrontal, is large and extends far backward lateral 
to the frontal and parietal; it has. however, only a very short pos- 
terodorsal boundary on the rim of the orbit. The postorbital is 
likewise highly developed and elongate, the postfrontal and post- 
orbital between them occupying much of the area which in labyrin- 
thodonts is occupied by the ■•temporal" elements. The arrangement 
along the orbital margin is of interest. The postfrontal (as in 




i 2. I aterial vie* of skull and jaw, based on MCZ 2040, x 1V4, 



Tuditanm | pushes downward posteriori) to take over a portion of 
the orbital margin which should properly be occupied by the post- 
orbital; the latter bone, in turn, extends downward along the orbital 
margin to encroach upon the area properly pertaining to the jugal. 
A thickened ridge runs downward and forward inside the posterior 
orbital margin along the postorbital and jugal. 

The jugal is a very large bone. It borders the posterior portion of 
the lower orbital margin, and sends a narrow extension far forward 

C ... 

between lacrimal above and maxilla below. Back of the orbit it 
occupies a broad expanse of cheek, and forms most of a prominent 
flange extending downward, external to the posterior end of the 
tooth row and concealing the upper part of the lower jaw posteriorly. 



1969 



CRANIAL ANATOMY OF PANTYLUS 





Fig. 3. A series of sections, in a horizontal plane, of the region anterior 
to the left orbit, in dorsoventral order, to show the relations of the lacrimal 
duct to the surrounding bones. From the sectioned skull, X 3. The lacrimal 
duct is hatched. The duct is seen leaving the orbit in section A; by E it 
enters the narial area through a foramen in the septomaxilla, which forms 
posterior and medial walls to the naris. From B downward, the maxilla is 
seen to bound the lacrimal medially as well as laterally. 



10 



BRLVIORA 



No. 314 



The maxilla is stout mcdiolaterally, to contain the roots of the 
marginal teeth, but has only a limited facial exposure, except an- 
teriorly above the large "canine.*' It is partly excluded from the 
naris by the septomaxilla, rimming it only for a short distance be- 
tween premaxilla and lacrimal. Posteriorly, at the end of the lateral 
tooth row, the maxilla is covered externally by the jugal. 

The posterior rim of the skull roof is formed by the paired post- 
parietals. "supratemporals," squamosals, and quadratojugals. This 
series, as noted below, extends broadly on to the occipital surface. 
The postparietals are of modest size and are somewhat triangular 
in shape, in dorsal view, narrowing anteriorly. The large bone that 
may be provisionally termed the "supratemporal" is bounded by the 
postparietal and parietal medially, the squamosal and postorbital 
laterally, thus occupying essentially the position of the tabular and 




i 4. Posterior view of skull, based mainly on MC7. 2040, x l'/i. 



supratemporal of labyrinthodonts; it is in contact anteriorly with 
the postfrontal. The squamosal is likewise of considerable size in 
external exposure. The quadratojugal extends well forward along 
the lower margin of the cheek flange formed more anteriorly by the 
jugal; externally, it is narrow dorsoventrally. but the sectioned skull 
indicates that it underlaps the squamosal to a considerable degree 
internally. As noted below, it turns inward posteriorly to take part 
in the jaw joint. 

Palate (Figs. 4, 5; 8-11 in part). Part of the palate could be 
made out in earlier described specimens. In the sectioned speci- 
men, nearly the entire palatal structure can be accurately deter- 
mined. The epipterygoids and vomers were disarticulated post- 
mortem; however, these structures were found preserved between 



1969 



CRANIAL ANATOMY OF PANTYLUS 



11 



the lower jaws and could be readily restored to position. 

The pterygoids are always long and highly developed in primitive 
tetrapods; here, the development is exceptional, for they extend al- 
most the entire length of the skull. Anterior to the area of the basal 
articulation there is a narrow interpterygoid vacuity; this, however, 
extends forward little more than half the distance from basal articu- 
lation to naris. Anterior to this point, the two pterygoids meet in a 
median suture; this suture is of some thickness between the anterior 
halves of the palatines, but thins between the nares. Anterior to the 
region of the basal articulation, the pterygoids, as seen in ventral 
view, appear as a pair of rounded ridges, which bear two or three 
irregular rows of small teeth and are separated by a deep groove 
from the palatines lateral to them. Anteriorly, these ridges gradu- 
ally narrow but continue to bear teeth to a point between the vomers, 
rather forward of the level of the posterior margins of the nares. 




Fig. 5. Palatal view of skull, based on the sectioned specimen, X IVi. 



12 BRBVIORA No. 314 

The tooth-bearing surface of this most anterior region was broken 
off, together with the vomers, in the sectioned skull; but as pre- 
served, the specimen shows a further forward continuation of the 
pterygoids dorsally as a pair of narrow but obviously stout little 
rods which reach nearly to the level of the anterior margins of the 
nares. For most of the broad palate, the pterygoids are covered 
ventrally, lateral to their median ridges, bv the palatines; in dorsal 
view, however, the pterygoids are seen to extend laterally above 
(and to strengthen | the palatines, to gain contact laterally with the 
jugals. I have found no sutural evidence for the presence of an 
ectopterygoid. although such a bone ma\ have persisted in an area 
internal to the posterior rim of the orbit. 

The bony choanal openings are very large. They are separated 
from one another only b\ the narrow vomers, with, above them, the 
anterior tips of the pterygoids. Anteriorly and laterally they reach 
close to the bases of the premaxillary and anterior maxillary teeth, 
and are bounded posteriorly by the curved anterior margins of the 
palatines. Only the posterior parts of these openings, however, are 
truly choanal, for (as shown by the sectioned specimen) the large 
anterior teeth of the lower jaw extended upward into the anterior 
part of these openings when the jaws were closed. 

The vomers were displaced ill the sectioned skull, but other speci- 
mens show them in proper position. They are narrow tooth-bearing 
elements bounding the choanae medially. They are apprcssed medi- 
all\ to the slender anterior ends of the pterygoids, except that an- 
teriorly the two vomers are in contact ventrally below the tips of the 
pterygoids. Anteriorly, the vomers articulate with the premaxillae; 
this articulation, however, is well above the level of the rim of the 
upper jaw; stout paired processes of the premaxillae rise dorsopos- 
teriorly to make this contact (Figs. 8-11). 

The palatines, in correlation with the powerful dentition they 
bear, are greatly expanded as essentially quadrilateral plates. Medi- 
ally the palatine is separated from the pterygoid on the ventral sur- 
face by a longitudinal groove, deep anteriorly but fading posteriorly. 
The anterior, choanal, border is concave in outline. Medially the 
anterior margin of the bone is somewhat convex ventrally in trans- 
verse section, forming a ridge continued anteriorly by the vomer. 
Lateral to this, the surface is somewhat concave ventrally at the 
choanal margin, the concavity fading out posteriorly. Posteriorly 
the margin is concave in outline, the medial margin extending well 
posteriorly, underlain dorsallv by the pterygoid. There is no devel- 
opment of the transverse pterygoid flange characteristic of all 
primitive reptiles. 



1969 



CRANIAL ANATOMY OF PANTYLUS 



13 



Approaching, posteriorly, the region of the basal articulation, 
the pterygoid, as in all primitive tetrapods, narrows in transverse 
width and twists to a vertical plane, to form the quadrate ramus of 
the bone. Just in front of the articular region it sends upward a 
sharp spur, which aids in the support of the epipterygoid. Behind 
this spur the upper edge of the pterygoid is gently concave on its 
medial aspect, to form the lower part of the articular surface for the 
basipterygoid process. The posterior, quadrate ramus of the ptery- 
goid rapidly develops into an extensive plate, thin from side to side 




Fig. 6. Skull roof diagrammatically removed to show palate from above. 
Dermal roof sectioned horizontally through level of middle height of orbit; 
sphenethmoid and ascending ramus of epipterygoid sectioned at same level, 
but dorsal flange of sphenoid complete. Postparietals and tabular removed; 
supraoccipital shown as complete except for removal of dorsolateral flange 
on right; based on sectioned specimen, X l'/2. 



14 



HRI-VIORA 



No. 314 



but of considerable height. It extends ventrally to about the level of 
the bottom of the check flange of the jugal and quadratojugal, and 
runs back nearly to the region of the jaw articulation. Above this 
posteroventral point, it contracts somewhat in posterior extent; 
meanwhile, however, the dorsal margin of the ramus has risen 
rapidly, and the upper blade of the ramus extends far upward, back- 
ward, and laterally, to terminate high up within the temporal region, 
close to, but not in direct contact with, the squamosal. 

The epipterygoids had been jarred loose in the sectioned speci- 
men and had fallen down between the jaws, where they lodged with 
but little damage. They can be replaced without dilliculty in ap- 
proximated natural position, although with some slight doubt as to 
dorsal orientation. Below, the bone is expanded into a Hat plate 




Fig. 7. First of a scries of "dissections" in lateral view, based on the sec- 
tioned skull. I his figure comparable to Fig. 2. The course of the lacrimal 
duct is shown in broken line. X 1 Vi . 



applied closely to the lateral surface of the quadrate ramus of the 
pterygoid over a considerable area, mainly below and posterior to 
the basal articular area and extending only modestly anterior to that 
point. At the level of the upper margin of the pterygoid, the epip- 
terygoid thickens considerably; a pair of ridges extends medially to 
form a socket for much of the basipterygoid process. The anterior 
margin of the bone here is notched to receive the supporting spur of 
the pterygoid mentioned above. Dorsally. the bone slims out to a 
slender blade, which reaches far toward the skull roof. 

The quadrate is well developed. The articular surface for the 
lower jaw is broad mediolaterally, relatively narrow anteropos- 
terior^', and but slightly convex. The quadratojugal turns medially 
in a flange which is of considerable thickness ventrally; on its lower 



1969 CRANIAL ANATOMY OF PANTYLUS 15 

surface this forms approximately a third of the articular area. 
Above the articulation, the quadrate is thickened but gradually thins 
dorsally. Medial to the articulation, the quadrate is very stoutly 
developed and extends ventrally and posteriorly beyond the ventral 
and posterior limits of the "roofing" bones. There is a postero- 
medial swelling just above the articular region, presumably for a 
muscular or tendinous attachment. Anteriorly, the quadrate forms 
a very extensive plate applied closely to the outer surface of the 
pterygoid and extending far forward toward the epipterygoid. Dor- 
sally, the plate, with the accompanying pterygoid, reaches far 
dorsally and posteriorly beneath the squamosal. 

Occipital surface (Fig. 4). In the absence of an otic notch, the 
skull roof exhibits in posterior view an even, essentially semi- 
circular, curved margin. In contrast to conditions in most lower 
tetrapods is the great extension onto the occipital surface of the 
posterior elements of the dermal roof. The postparietals send a 
flange of modest size down over part of the supraoccipital on either 
side. The "supratemporal" sends a broad sheet of bone downward 
and medially; in the fashion of a labyrinthodont tabular, there is an 
articulation of the supratemporal with the otic region of the brain- 
case. The attachment is a broad one, meeting both prootic and 
opisthotic above the fenestra ovalis, and is far more ventrally placed 
than that of a typical labyrinthodont. There is a normal posttem- 
poral fenestra, bounded mainly by "supratemporal" and post- 
parietal, but ventrally and ventromedially by the otic elements and, 
to a slight degree medioventrally, the supraoccipital. 

Much in the fashion of the "supratemporal," the squamosal sends 
a broad, thin sheet of bone downward and medially over a consider- 
able lateral area of the occipital surface. Dorsally, this sheet under- 
lies the occipital flange of the "supratemporal" to some extent; 
ventrally it sheathes the occipital surface of the quadratojugal. It 
ends freely ventromedially. 

Centrally, on the occipital surface is seen the posterior aspect of 
the braincase. The usual ring of occipital elements — supra-, basi-, 
and small, paired exoccipitals — surrounds the foramen magnum. 
Above the foramen magnum the supraoccipital is deeply excavated 
by a pit, presumably for ligamentous connection with the column. 
The otic capsule is seen to extend outward far laterally (somewhat 
as in Seymour ia) rather than dorsolaterally, as in the paroccipital 
process of typical labyrinthodonts. Beneath the lateral part of this 
process the fenestra ovalis is visible; more medially, between opis- 
thotic and a lateral extension of the exoccipital, is the jugular for- 
amen for the vagus nerve (as well as, presumably, nerve IX and the 



16 



BR1ZVIORA 



No. 314 



"jugular" vein). A hypoglossal foramen is present ventrally. and 
possibly a second hypoglossal opening may be present more dor- 
sally. The occipital condyle is a broad, somewhat concave, and 
essentially tripartite structure comparable to that of such rhachi- 
tomes as Eryops. 




Fig. S. As Fig. 7. hut dermal bone of left sale removed (except for pre- 
m.ixill.ii. to show the palatal structures in lateral view. Sutures with bones 

removed indicated b) hatched areas. \t anterior end of sphenethmoid is 
seen the lateral process of that bone, which is tight!) fused to prefrontal; 
below are shown the surfaces of palatine and pterygoid, which join the 
lacrimal and jugal. Posteriori) is seen the area of contact between quad- 
ratojugal and quadrate. 




I ig. 9. As Fig. 8. but palatal structures of left side and left premaxilla 
removed to show lateral view of braincase. Below the anterior part of the 
sphenethmoid and to the front are seen the medial contacts between the 
ptervgoids and vomers and the ascending processes of the premaxillae. which 
meet the vomers. The rugose area on the otic bones above the fenestra 
ovalis is the area of contact with the "supratcmporal." 



1969 



CRANIAL ANATOMY OF PANTYLUS 



17 



soc 




Fig. 10. As Fig. 9, but sectioned through middle of braincase, to show 
internal structure. Inner surface in otic region restored with doubt. 




Fig. 1 1. As Fig. 10, but braincase removed, showing inner surface of right 
palatal complex. Anterodorsal to orbit is seen the area of contact of the 
prefrontal with the lateral flange at the front end of the sphenethmoid. A 
rugose area posteriorly is the sutural surface of the "supratemporal" articu- 
lating with the otic elements of the braincase. 



18 



BRIVIORA 



No. 314 




Fig. 12. As Fig. 11. hut palatal elements removed, showing the inner 
surface of (tic dermal roofing elements of the right side. PosteroventnilU 
is seen the area of contact of the quadratojugal with the quadrate. 



Braincase ( Figs. 4-6. 9. 10). The nasal capsule was unossilied. 
As noted earlier, the external uarial cavity is bounded posteriorly 
and mediall) bv the septomaxilla. Behind this bone, the anterior 
part ol the choanal opening: was oeeupied by the large anterior 
lower teeth; it is hence obvious thai the air passage ascended dor- 
sally from the external naris and that the nasal capsules were dor- 
sails placed, anterolateral to the front end of the sphenethmoid. 
Beginning anteriorl) at about the level of the posterior margins of 
the choanae, and extending back to about the level of the posterior 
margins of the orbits, there is a well-developed sphenethmoid. The 
entire structure is a single ossification, essentially Y-shaped in cross 
section. I he ventral edge of the bone is keeled, and in life lay for 
most of its length in a groove on the upper surface of the para- 
sphenoid rostrum: there is here a stout union between the two thin 
ascending Banges. These diverge from one another at a modest 
angle, and then approach each other to a slight degree dorsally. 
The cavity of the sphenethmoid is widely open both anteriorly and 
posteriorly. Laterally, opposite the orbit, there is on either side a 
foramen of considerable size through which the olfactory nerve pre- 
sumably passed; a second large foramen, presumably for nerve II, 
lies toward the back end of the bone. Possibly the sphenethmoid 
may have been penetrated bv small foramina for one or more of the 
eye-muscle nerves, but I have not been able to detect them in the 
sectioned specimen, and they may have emerged posterior to the 
sphenethmoid. The sphenethmoidal plates appear to have extended 
upward to the under side of the roofing elements, but I have not 



1969 CRANIAL ANATOMY OF PANTYLUS 19 

been able in the sections to determine the exact relations here (Wil- 
liston, 1916a, fig. 25, shows sections in this region). On either side 
in front of the orbital rim the prefrontal sends in a powerful medial 
flange, broadening dorsally, which articulates strongly with the 
outer surface of the sphenethmoid near its anterior end. 

The parasphenoid-sphenoid complex in the sectioned specimen 
was jolted out of position before burial; present between the lower 
jaws, it has undergone little damage except for the posterior mar- 
gins of the parasphenoid (which, however, are preserved in other 
specimens). It can be replaced fairly accurately in its proper place 
in the figures because of the known position of the basal articula- 
tions and because it can be seen in place in lateral view in MCZ 
2040. The anterior portion of the complex is a long parasphenoidal 
rostrum; a dorsal groove obviously received the ventral keel of the 
sphenethmoid in normal fashion. Posteriorly, the keel expands gra- 
dually and then abruptly produces ventrolaterally a pair of well- 
developed basipterygoid processes with stoutly rounded heads that 
fit into the sockets formed jointly by the epipterygoids and ptery- 
goids. 

In this region, the complex includes, inside the superficial para- 
sphenoidal dermal component, spongy endochondral bone repre- 
senting the basisphenoid; endochondral bone is included in the 
structure of the basipterygoid process. There is a distinct rounded 
medial pocket in the basisphenoid opposite the basipterygoid pro- 
cesses, obviously lodging the pituitary. A distinct crossridge behind 
the cavity represents the dorsum sellae; behind this point endo- 
chondral bone is absent. As in early tetrapods generally, it seems, 
there was a ventral gap in endochondral braincase ossification be- 
tween the sphenoid and otico-occipital braincase elements. A pair 
of small foramina, presumably for internal carotid arteries, pene- 
trates the bone from the under side opposite the posterior margin of 
the pituitary fossa. A remarkable feature is the presence of exten- 
sive wings extending far upward from the basisphenoid and con- 
tinuous with it on either side; the type of ossification is such as to 
indicate their endochondral nature. They appear to represent the 
pila antotica of the embryonic braincase. Near the base of either 
flange is a good-sized foramen that presumably carried the trans- 
verse pituitary vein, as in the basisphenoid of various lower tetra- 
pods. Although, as said above, the parasphenoid-basisphenoid can 
be replaced in a position close to that which it must have occupied 
in life, there is a degree of uncertainty as to the exact posture of 
these dorsal flanges. They certainly stopped short dorsally well 
below the level of the skull roof. In the sectioned specimen the 



20 brlviora No. 314 

sphenoid had broken loose from the sphenethmoid, but the two are 
seen in natural relationship in lateral view in MCZ 2040. Here 
there is seen a short area of connection between the ascending 
flanges and the sphenethmoid part way up the side, leaving a gap 
dorsally and a rounded foramen ventrally. Possibly small eye- 
muscle nerves emerged above; below is perhaps the point of exit of 
the profundus branch of nerve V, the more posterior rami of this 
nerve and nerve VII making their exits behind the dorsal flange of 
the sphenoid. 

1 1 was hoped that the sectioned specimen would show the otico- 
occipital elements in place. This was not the case. The supra- 
occipital remained essentially in position, but the other elements 
had been jarred loose before burial (suggesting, although we ap- 
pear to be dealing with an adult, rather loose suturing of elements). 
Fortunately, much oi the structure could be made out from MCZ 
2040 and MCZ 3302. Dorsoposteriorly. the supraoccipital is firmly 
fused to the overlying postparietals; in addition, the latter bones, as 
noted earlier, partially sheath the supraoccipital posteriorly. A 
narrow process extends dorsolateral!) toward the "supratemporal." 
On either side, the supraoccipital descends and expands to gain con- 
tact with the two Otic elements and. posteriorly, with the exoccipital. 
The sections show that, as commonly, the supraoccipital encloses 
the upper part of both vertical canals of the internal ear, as well as 
the cms commune. Dorsallv. the supraoccipital sends forward an 
anterior process. One would have expected this to run forward 
dorsally beneath the skull roof, as. for example, in many reptiles. 
This is not the case; it slants strongly downward, to terminate an- 
teriorly at about the level of the pituitary rather below half the dis- 
tance from the skull roof to the braincase floor and between the 
two ascending flanges of the sphenoid. Uniquely among known 
early tetrapods, this anterior extension is a dual structure, consisting 
of two closely apposed narrow rods. As is known from the develop- 
mental story in lower tetrapods, supraoccipital ossification takes 
place in a cartilaginous area formed by dorsal fusion of extensions 
upward of the two otic capsules; the situation here suggests that the 
fusion of the two was imperfect. 

The exoccipitals are, as usual in early tetrapods. small elements, 
hung below the supraoccipital on either side. Between opisthotic 
and exoccipital is a typical jugular foramen. Ventrally. the exocci- 
pitals form the lateral portion of the broad occipital condyle. The 
basioccipital is a wedge-shaped element, forming the central part of 
the condyle and tapering anteriorly on the braincase floor. 



1969 CRANIAL ANATOMY OF PANTYLUS 21 

The pro- and opisthotics jointly form a pronounced lateral exten- 
sion of the otic region (Fig. 5), tipped at its end by a broad area of 
articulation with the "supratemporal," somewhat after the fashion 
of the typical paroccipital process of many early tetrapods (which, 
however, articulates with the tabular) . Below this articular area is 
the good-sized fenestra ovalis, bounded posteriorly by the opis- 
thotic, anteriorly by the parasphenoid. This latter dermal element 
covers most of the surface of the prootic, except dorsally, and like- 
wise sheathes the opisthotic ventrally. Parasphenoid and otic ele- 
ments were apparently not bound closely together in life, since they 
have cleanly separated in the sectioned specimen. Dorsally the two 
otic structures are articulated with the supraoccipital. On the dorsal 
surface of the lateral otic process the suture between pro- and opis- 
thotics lies in a deep groove; obviously the suture between the two 
was a comparatively loose one (as confirmed by their separation in 
the sectioned specimen) . 

In the sectioned skull, three isolated otic elements are clearly 
seen, but I must confess myself baffled in attempts to interpret them 
and orient them in a reconstruction. Two of the three, obviously a 
pair, have, crudely, the shape of a clam shell; the curved outer sur- 
face is smooth except for a ridge indicating the division between a 
free surface and that sheathed by the parasphenoid. The inner as- 
pect, presumably housing part of the internal ear, shows some slight 
development of ossification. Attached to one of the two is a small 
area of ossified tissue which presumably represents an exoccipital; 
in consequence, I interpret the two as opisthotics. By elimination, 
the third element preserved is a prootic. Here again, the presumed 
external surface is featureless except for a ridge which may have 
separated a free dorsal surface from that sheathed by the para- 
sphenoid. On the inner surface there is a considerable degree of 
ossification in a complex pattern which, however, I fail to interpret. 
As a result, I can give no description of the internal structure of the 
otic region except what can be seen rather imperfectly from the 
sectioned braincase of MCZ 3302. Apparently the internal ear 
region was widely open to the brain cavity opposite the two otic 
elements, although partial anterior and posterior ossified walls are 
formed by them. 

Lower jaw (Figs. 13-15). Like the skull, this is a very stoutly 
built structure and is relatively broad, but shallow. As can be seen 
in the cross-sections figured by Williston ( 1 9 1 6a ) , both external and 
internal walls of the jaw ramus are thick. In section, the jaw is 
somewhat triangular. The upper surface is broadened for the coro- 
noid toothplate; the inner surface is essentially vertical. On the 



?? 



BRLVIORA 



No. 314 



outer surface, a longitudinal, shallow shelf is present in the region 
opposite the adductor fossa; this corresponds to the lower boundary 
o\ the jugal flange of the skull. The lower part of the outer surface 
curves gradually medially, to compensate for the dorsal width, so 
that most of the splenial-angular area and the lower part of the 
dentary are seen in a ventral rather than a lateral view. Much of the 
outer surface of the jaw is sculptured in a fashion similar to the 
skull root, hut m the posterodorsal portion oi the external surface. 




Fig. 13. Ventrolateral view of right jaw ramus, ventral margin at the 

top. • l 1 :. 




I 14 Dorsal view of left jaw ramus, x l 1 ^- 




psp 
Fig. 15. Internal view of right jaw ramus, X l'/i. 



1969 CRANIAL ANATOMY OF PANTYLUS 23 

from the coronoid process down to the longitudinal shelf already 
mentioned, the jaw (here covered by the jugal flange) is smooth- 
surfaced. The ventromedial margin of the lateral surface also lacks 
sculpture, presumably due to the fact that here, between the jaw 
rami, the skin was lifted free from the bone. Rather unexpectedly 
in such a strongly built structure, the symphysis, although stoutly 
fused, is of modest extent — a triangular area with a vertical pos- 
terior border and a curved anteroventral external surface. The 
opening of the adductor fossa is large, extending about half the 
length of the jaw back from the curved posterior border of the 
coronoid plate to the articular area. A large cavity of subcircular 
shape, corresponding to the Meckelian canal of reptiles, extends 
forward without interruption from the adductor fossa nearly to the 
symphysial region. Presumably this carried in its floor the anterior 
part of the Meckelian cartilage. 

Lateral to the opening of the adductor fossa there is an elongate 
coronoid process, which rises fairly rapidly from a point near the 
posteroexternal border of the coronoid toothplate and decreases 
gradually in height posteriorly. It slants gently toward the medial 
side above. Opposite the coronoid process there is a smaller, thin, 
process extending upward from the medial margin of the adductor 
fossa. As in the upper jaw, the articular surface is broad medio- 
laterally, narrow anteroposteriorly. It is gently concave in antero- 
posterior section. The broad posterior margin of the jaw slants 
nearly directly downward for a short distance below the articular 
surface, but then turns strongly backward to form a pronounced and 
broad retroarticular process, rather more developed medially than 
laterally. On the inner surface of the jaw there is a typical Mec- 
kelian foramen opening into the adductor fossa; this lies close to the 
ventral jaw margin, about a third of the jaw length forward from its 
posterior end. Williston (1916a) described a second, smaller, 
Meckelian foramen much further forward along the inner jaw sur- 
face; I have not been able to identify this small opening in the avail- 
able material. Presumably a fossa for the chorda tympani was 
present, but I have not observed this opening. 

As commonly, the dentary is the largest element of the jaw. An- 
teriorly, it forms most of the jaw symphysis, behind which it is ex- 
posed on the inner surface for a short distance only. Externally it 
occupies a large part of the surface of the ramus. Anteriorly, where 
it is bounded ventrally by the splenials, it occupies the greater 
part of this outer surface; more posteriorly, adjacent to the angular 
and surangular, its lower boundary retreats gradually upward. At 
the level of the longitudinal shelf mentioned above, the surangular 



24 brhviora No. 314 

extends forward in V-shaped fashion to overlap the dentary to 
some degree; above this, on the eoronoid process, however, the 
dentar\ succeeds in extending well backward, and tonus much of 
the outer surface of the process, although not reaching its summit. 
Anteriorly, the medial boundary oi the dentary descends along the 
outer surface of the eoronoid process to extend forward along the 
medial margin oi the lateral tooth row, where the suture lies in a 
marked groove. 

Of the infra-dentary scries, the two splenials are found in then- 
usual position externally, occupying a modest area near the ventro- 
medial termination of the morphologically lateral surface. The pos- 
terior Splenial extends around the lower jaw margin to be exposed 
along a narrow ventral strip on the medial surface. I he anterior 
splenial is more extensive in medial exposure. I have some doubts 
as to its boundaries, but the anterior splenial appears, as Williston 
figures it. to extend upward as a thin sheet over the anterior end of 
the prearticular to border the anterior end of the eoronoid plate and 
gam contact with the dentarv behind the anterior tooth pair; it 
forms the ventral third of the symphysis. The angular has its normal 
exposure, of considerable extent, over the posteroventral region of 
the outer surface; posteriorly it forms the outer part oi' the retro- 
artieular process. Internal I \ it is exposed ventrally, running for- 
ward to partially bound the Meekelian foramen. I he surangular is 
well developed externally, where it forms mueh oi the region of the 
eoronoid process, excepting its highest and most anterior portion. 
Posteriori) it participates in the formation oi the retroarticular 
proeess. Paralleling its upper jaw analogue, the quadratojugal. the 
surangular turns medialk at the posterior end of the ramus and 
forms the lateral third n\ the articular condvle. 

It is possible that the presumably primitive series of three coro- 
noids was present in Pantylus, but I have not been able to dis- 
tinguish sutures in available material. The eoronoid area forms a 
large plate bounded laterally by the dentary, superficially on the 
medial side by the prearticular. and anteriorly by the anterior 
splenial. Medially, beneath the last-named elements, a thick flange 
of the eoronoid. at right angles to the exposed portion, extends far 
ventrally, giving stout support to the toothplate, as figured by Broom 
(1913. fig. 4) and Williston (1916a, fig. 27). Posteriorly, behind 
the termination of the toothplate. the eoronoid complex takes part 
in the formation of the eoronoid process; it forms the inner surface 
of the high anterior portion of the process and is also exposed ex- 
ternally at its summit. 



1969 CRANIAL ANATOMY OF PANTYLUS 25 

The prearticular is extensively developed. For the most part, it 
covers the surface of the coronoid area along the inner aspect of the 
jaw as a thin plate of bone; anteriorly, it appears to be covered in 
turn by the anterior splenial. In the middle section of its extent, it 
is bounded ventrally by the splenials. It forms much of the boun- 
dary of the Meckelian foramen, posterior to which it is constricted 
in ventral extent by the angular. Back of the coronoid plate it forms 
by itself the whole of the medial wall of the adductor fossa to the 
summit of the medial dorsal process which was mentioned earlier. 
Behind this point it is constricted dorsally by the articular; it ex- 
tends to the posteromedial corner of the jaw as a restricted strip of 
bone between the articular above and the angular below. 

The articular forms the medial two-thirds of the jaw articulation. 
Below, it descends to bound the posterior end of the adductor fossa 
and form part of the substance of the retroarticular process; as far 
as I could determine, however, it does not appear on the surface of 
that process. Medially it turns forward above and internal to the 
prearticular to form a portion of the inner wall of the adductor fossa. 

Lying within the floor of the Meckelian canal for much of its 
extent is a thin sheet of bone, shown in the sections of Mehl (1912, 
fig. 4). It is somewhat broken and discontinuous in my sections, 
but is obviously not part of any one of the dermal ossifications lying 
external to it. It seems surely to be a Meckelian ossification in the 
lower surface of the cartilage which probably filled much of the 
canal. 

Dentition (Figs. 2, 5, 7-12, 14, 15). Pantylus was endowed with 
a powerful dentition of stout but blunt teeth, both on the jaw mar- 
gins and internally on broad areas of the palate and the coronoid 
area of the lower jaw. In section there is no trace of labyrinthine 
infolding of the enamel; nor is there any trace of a division of teeth 
into basal and distal portions, as might be hoped for in a member 
of a group possibly related to the modern amphibian orders (cf. 
Parsons and Williams, 1963). Evidence of tooth replacement was 
looked for, but not found. It seems clear that Pantylus subsisted 
on some type of durophagous material. A molluscan diet has been 
suggested by various authors. This is not unreasonable; but it 
is to be noted that little trace of molluscs has been seen in the beds 
from which Pantylus specimens have been collected, and probably 
some hard plant material formed the food supply. The massive 
construction of skull and jaw is reasonably to be correlated with 
strong masticatory movements, and great strength of the jaw 
muscles is indicated by the great size of the subtemporal fossae, 



26 brf.viora No. 314 

through which the temporal muscles descended — the breadth of 
these cavities being correlated with the unusual width of the tem- 
poral region of the skull. 

The upper marginal teeth, particularly the more anterior ones, 
are somewhat tilted inward at their tips (a reverse outward slant 
is present in the lower anterior teeth). Three teeth are generally 
present on each premaxilla. The first is large and powerful and 
is, next to the "canine," the largest tooth in the upper jaw . Lateral 
to this are usually two further "incisors," somewhat smaller, but 
nevertheless rather longer than any of the maxillary teeth except 
the "canine"; the two are crowded close together and are somewhat 
compressed from side to side. One small and presumably young 
specimen shows but one tooth in this position. Immediate!) behind 
the lateral "incisor" is the powerful "canine." most anterior mem- 
ber of the maxillary row. Behind the "canine*' is a series of cheek 
teeth. The first of these is little more than half the length of the 
"canine," and the first few following teeth are oi about the same 
length. More posteriorly there is a gradual diminution in size. 
The specimen sectioned is somewhat imperfect posteriorly along 
the jaw margins, so that there is some question as to the exact 
number in the scries; there are. in this specimen, certainly eight 
and possibly nine post-"eanine" teeth. Some specimens show a 
larger number of cheek teeth. Williston (1916a), for example, 
states that there are 15 or 16 maxillary teeth (but figures only 16 
for premaxilla and maxilla combined). Discrepancies in tooth 
counts ma\ be due to individual variation or. quite possibly, to 
posterior additions to the tooth row during life. 

In the marginal row oi the lower jaw. there arc two anteriorly- 
placed "incisors." The more medial of the two is the smaller, 
but this is, nevertheless, more powerful than any but the most 
anterior of the lower check teeth. This tooth is somewhat com- 
pressed from side to side at its base. The lateral "incisor" is of 
considerable size and strength, equalled only by the upper "canine." 
As noted earlier, this tooth is so highly developed that when the 
jaws were closed, it extended well up into the anterior part of 
the bony choanal opening (as did the median incisor to a lesser 
degree). Bevond the lateral "incisor" there is a slight dip in the 
height of the jaw rim and a gap, allowing for the passage of the 
upper "canine." Following this short diastema there is a series of 
marginal teeth, which mainly match in size the more anterior post- 
"canine" teeth in the maxilla. I have found but seven teeth in the 
post-"incisor" series in the lower jaw of the sectioned specimen, 



1969 CRANIAL ANATOMY OF PANTYLUS 27 

in contrast with the higher number above, and there is relatively 
little diminution in size toward the back of the series. 

Teeth are highly developed on the palate. I have seen no 
evidence of parasphenoidal teeth (found occasionally in early 
tetrapods). Beginning a short distance anterior to the basal articu- 
lation, the anterior ridge of the pterygoid is studded with small 
teeth. These are rather irregularly placed, but for most of the 
length of this long ridge they are essentially arrayed in two irregu- 
lar rows. The vomers, lateral to the anterior ends of the pterygoids, 
bear a series of teeth of somewhat larger size. 

The major palatal tooth assemblage is found on the extensive 
palatine bones. There is some irregularity in the arrangement, 
even between the two sides of a single individual (as that sectioned 
and figured), but there is, in the few palates I have seen, a uniform 
basal plan. Somewhat to the lateral side of the center is a very 
large but quite blunt tooth with an oval base and a crown slanting 
anteromedially. Just behind and medial to this tooth is a bare 
area of the palate into which, in the closed jaw position, a similarly 
large lower tooth is lodged. A longitudinal row of rather large 
teeth runs along the lateral margin of the bone; the lower marginal 
teeth in closure lay between this row and the marginal upper teeth. 
Anteromedially, a row of teeth of modest size continue backward 
from the vomerine series. A few teeth of a size comparable to 
those of the marginal row are present in front of the large central 
tooth, and good-sized teeth are present medial to it. About a score 
of teeth are present in the posterior portion of the palatal plate. 
They are of variable size; many are moderately large, but there 
is a decrease toward the angular posteromedial boundary of the 
palatine. 

The coronoid region of the broadened upper surface of the 
lower jaw bears a battery of teeth, somewhat over a score in num- 
ber, which correspond roughly to the palatal assemblage above. 
There are, however, no lower teeth to oppose those on the ptery- 
goids or (naturally) those on the vomers. Further, the lower tooth 
plate is much narrower than that of the palatine, and the teeth 
are more closely crowded. As in the case of the palatine, there 
is a large, broad-based tooth centrally situated on the coronoid 
plate. Despite the crowding of the lower plate teeth, there re- 
mains, in the sectioned specimen, a small bare area just anterior 
to this tooth, in which was received the summit of the crown of 
the major upper plate tooth. As in the palatine plate, there is a 
row of stout teeth along the lateral margin of the plate, and well- 
developed teeth are present anterior and posterior to the major 



28 



BRHVIORA 



No. 314 



tooth. Teeth forming a medial row are of smaller size. 

Individual variations in the number and position of teeth in the 
palatine and coronoid toothplates are to be expected, and it was 
mentioned earlier that in some cases the large, centrally-placed 
teeth in these plates may be absent. In the sectioned specimen, 
upper and lower jaws were in occlusion at death, and it could be 
seen that there was a perfect coordination between upper and 
lower toothplate sets. There was never any direct opposition be- 
tween an upper and lower tooth; each upper tooth fitted into a 
gap between lower teeth, and vice versa. 

Visceral arch elements (Fig. 16). As attested by the presence 
of a pebblv sheet of dermal ossicles similar to those seen in 
Tuditanus and "Ricnodon" limnophyes, the skin between the jaw 




I ig 16. I. Postcranial structures preserved behind left side of occiput, 
from reconstruction of the sectioned specimen. Outlines of the skull ele- 
ments shown for orientation; cb, ceratobranchials. eb, epibranchials, r, rih 
fragments, sta, stapes. /?, Stapes from anterior side, as oriented. C, Ventral 
\ieu of main shaft of stapes. All x 3. 



rami was still present when the sectioned specimen underwent 
burial, and various bones were preserved in this "pocket" for 
fossilization. We have already mentioned the presence here of 
detached vomers, epipterygoids, and otico-occipital elements. Fur- 
ther, the first three vertebrae had been pushed forward into this 



1969 CRANIAL ANATOMY OF PANTYLUS 29 

area, and ventrally a fraction of the dermal shoulder girdle is 
included also in the block. On the right, the sectioned block 
terminates directly back of the occiput and jaw articulation; on 
the left, however, the block extended a short distance farther back, 
so that there are present part of the cervical rib series and — much 
more interesting — part of the visceral arch series of the left side. 

Fairly high up in the block, centered about behind the left otic 
region, are three elements which appear rather certainly to be 
epibranchials. These are thin-walled, incompletely ossified in- 
ternally, with a spongy structure. They are essentially leaf-shaped, 
each with a broad distal expansion, thin anteroposteriorly; the 
three are closely appressed. Dorsally, each narrows somewhat to 
a "stem"; however, the one seen farthest to the left in the figure 
has a rather broad expansion at its "head." 

Separate from the presumed epibranchials and running trans- 
versely lower down in the block are three further structures, which 
appear to be ceratobranchials. Each consists of two contrasting 
segments. At the left, and slanting ventromedially, are "head" 
portions which resemble the epibranchials in being thin-walled, 
spongy internally, broad in the transverse plane, but thin and rather 
closely appressed, one to the other, anteroposteriorly. At their 
lower ends, each turns directly medially and assumes the form of 
a tube, thick-walled but hollow centrally. These extend for a (re- 
latively) considerable distance to the right. They terminate some- 
what irregularly; this may be due either to post-mortem damage 
or to the presence of cartilaginous terminations. At their distal 
ends each of the three narrows dorsoventrally but broadens antero- 
posteriorly, like flattened cornucopias. 

A further visceral structure is present in the sectioned specimen, 
to the right of the presumed epibranchials. This is a stout element 
which, as preserved, begins dorsally high up just behind the lower 
inner portion of the occipital plate of the left supratemporal. For 
a short distance there are two semi-distinct "heads." but these 
shortly unite to form a single, rather large structure, of rhomboidal 
shape in section, relatively thin-walled, and with a spongy interior. 
A short distance down there is a bifurcation. One branch slants 
sharply to the right and somewhat posteriorly; distally this ramus 
becomes shattered, so that nothing can be said of its extent or 
distal terminus in life. The other branch runs nearly directly 
ventrally. For a short distance it contracts, to have a relatively 
small diameter and good walls; beyond, it expands to a large 
diameter, with a spongy interior and with thin walls except toward 
the right and anteriorly. Anteriorly there develops a well-formed 



30 brlviora No. 314 

projection, thin at its base but then expanding toward each side, 
giving a T-shaped section. 

It seems certain that this element is a stapes, and rather surely 
that of the left side. In its disarticulated state there is no certainty 
as to its interpretation and position in life. At first thought, one 
would tend to interpret it in terms of the pattern common in early 
tetrapods and crossoptcr\gians (c\\. for example. Romer, 1956, 
fig. 197). On this assumption, the upper end as figured would 
be that associated with an ear drum (if present ). the shorter branch 
to the right a dorsal process, the vertical processes the main shaft, 
with the footplate at the bottom. But various considerations sug- 
gest an alternative hypothesis — that the upper terminus is a broad 
footplate, that the complex lower end oi the main shaft was asso- 
ciated with an eardrum or other auditory structure, and that the 
branch to the right braced the structure laterally in some fashion 
on the occipital aspect o\ the cheek (somewhat analogous to the 
crossoptcrvgian opercular process). On this assumption, the 
posterior surface o\ the stapes is that seen in Figure 1613. The 
somewhat bifurcate nature of the presumed footplate may be 
due to close apposition of shaft and dorsal process, but is also 
slightly suggestive of initiation o\' the lissainphibian division of the 
apparatus into stapes plus operculum. 

PANTYLUS AS A MICROSAUR 

Two facts seem immediate!*, clear: ( 1 ) PantylllS is closely re- 
lated to microsaurs of the gymnarthrid group, and (2) neither 
Pantylus nor its microsaur relatives are reptiles or ancestral to 
reptiles. 

The description above has. I think, made it obvious that 
Pantylus is a microsaur. It is specialized for a durophagous diet 
in its massive skull build, accommodation for powerful temporal 
muscles, and powerful dentition. Otherwise it compares readily 
in many regards with the contemporaneous gymnarthrid micro- 
saurs; dentitions of intermediate type are known among Carboni- 
ferous microsaurs in Canada and Bohemia. Pantylus exhibits 
the diagnostic cranial patterns of the group, such as those which I 
specified in 1950 and the additional features noted by Carroll 
(1966) and by Carroll and Baird (1968). The most obvious 
feature is, of course, the presence between parietal and squamosal 
of a large roofing element generally termed "supratemporal." In 
all labyrinthodonts we find here a row of elements, including a 
tabular posteriorly and more anteriorly a distinct supratemporal 



1969 CRANIAL ANATOMY OF PANTYLUS 31 

and, in many primitive forms, an intertemporal as well. Supra- 
temporal and tabular are present as distinct elements in primitive 
captorhinomorph reptiles. Never, however, is there any growth of 
a large element such as the microsaur "supratemporal." In special- 
ized captorhinomorphs tabular and supratemporal are reduced, 
so that — in strong contrast to microsaurs — squamosal and 
parietal come in direct contact for most of their length. 

In all early tetrapods there is a trend for the development from 
the posterior elements of the dermal roof of flanges sheathing the 
occipital surface. Such flanges are highly developed in Pantylus, 
particularly that from the "supratemporal." The occiput is poorly 
known in most microsaurs; however, a similar occipital sheathing 
is known in a few other cases, and "Ostodolepis," as figured by 
Case (1929), shows an even greater development of such dermal 
structures than that in Pantylus. It is highly probable that the 
presence of extensive dermal flanges in the occiput is a diagnostic 
microsaur feature. 

Typical of microsaurs (and early amphibians in general) is the 
rounded posterior contour of the palatal plate, in contrast with 
the development of the stout pterygoid flange seen in early reptiles. 
In various primitive amphibians we find narrow interpterygoid 
vacuities and a movable basal articulation of braincase and palate, 
in contrast to the "open" palate and fused basal articulations seen 
in most temnospondyls and modern anurans and urodeles. The 
primitive condition persists here in microsaurs in which the palate 
is known; Pantylus is in agreement with this condition. 

The braincase is poorly known in most microsaurs. It is possible, 
although of course not certain, that the structures described above 
in Pantylus are typical of microsaurs in general. We have seen in 
Pantylus a well-developed sphenethmoid of a type widespread 
among primitive tetrapods and presumably typical of microsaurs 
in general. The basisphenoid region, again, in the presence of 
well-developed basipterygoid processes and a good pituitary fossa, 
with adjacent openings for the internal carotids and a transverse 
pituitary vein, is of a primitive and generalized nature. Quite 
different, however, from other known primitive types is the struc- 
ture of the more posterior and posterodorsal regions of the brain- 
case. The sharp downward inclination of the anterior process of 
the supraoccipital is a feature not seen or expected in any other 
early tetrapod group. Case (1929), in "Ostodolepis," described 
such a structure. Until I discovered it in Pantylus, I was inclined 
to consider its position in Case's specimen as due to post-mortem 
injury. Its presence in both these forms strongly suggests that it 



32 breviora No. 314 

was characteristic and definitive of microsaurs. Again, the broad 
sheet of bone extending independently upward from the basi- 
sphenoid in Pantylus finds no homologue in other ancient groups. 
It is difficult to imagine a brain configuration fittimz this braincase 
structure. The downward tilt of the supraoccipital eliminates the 
possibility of the midbrain having extended forward into the area 
between these dorsal extensions oi the sphenoid. If brain tissues 
were shielded by them, the only possibility is that the cerebral 
hemispheres were tilted strongly upward and backward here above 
the diencephalon. 

The strong lateral projection oi the otic bones and the near- 
terminal ventral position of the fenestra ovalis are unusual features, 
somewhat reminiscent o<i the Seunouriamorpha. The occipital 
condyles are known in several other microsaurs; where seen, they 
appear to be essentially similar to the tripartite. Eryo/W-like condyle 
oi Pantylus. 

Since visceral arches are seldom reported in the older Amphibia, 
the epi- and ceratobranchials described above in Pantylus are at 
present of little significance in problems oi early amphibian rela- 
tionships. Mv identification oi a Y-shaped element as the stapes 
is supported by information given me by Dr. Carroll that an 
identical structure, likewise a presumed stapes, is present in 
Ostodolepis, and by Dr. Olson that such a structure is also present 
in an as ye! undescribed gunnarthrid from the Permian of Okla- 
homa. The whole question of stapedial elements in microsaurs 
is still an unsettled one. Since main modern amphibians have two 
auditory ossicles, operculum as well as stapes, there has been a 
tendency to look for a pair of ossicles in microsaurs, as possible 
ancestors oi the modern Lissamphibia (cf., for example. Gregory, 
Peabody. and Price. 1956). Shell-shaped structures seen in the 
ear region i^i gvmnarthrids have been interpreted as stapedial 
elements, and in some cases other fragmentary materials seen in this 
region have been thought to be accessory auditory ossicles. I rather 
doubt both interpretations. As described above, the otic bones 
in Pantylus are essentially shell-shaped and separate readily from 
the parasphenoid and other braincase elements. The shell-shaped 
presumed ossicles are probably disarticulated otic bones; the 
fragmentary materials mentioned are not improbably remains of 
the adjacent hyoid apparatus. 

MICROSAUR RELATIONSHIPS WITH OTHER AMPHIBIANS 

In 1933 I provisionally united the various amphibian types 
possessing "holospondylous" vertebrae in a single group of Lepo- 
spondyli, using this Zittel term in an expanded sense and including 



1969 CRANIAL ANATOMY OF PANTYLUS 33 

as Paleozoic components the three groups established by the Miall 
committee in 1875 — Nectridea, Aistopoda, and Microsauria. 
Study of the Pantylus skull neither adds to nor subtracts from the 
rather tenuous bonds which may possibly unite these three very 
diverse groups. 

In addition to the more typical microsaurs, such as the gymnar- 
thrids, Tuditanus, and Microbrachis, a number of other Paleozoic 
lepospondyls, such as Lysorophus , Molgophis, and the "adelo- 
gyrinids," are often united, although with some doubt, with the 
microsaurs. Recent and current studies have done nothing to 
strengthen this supposed association. It is, in fact, weakened by the 
results of "modern" studies which show, despite my fond hopes 
to the contrary, that extreme reduction of the cheek elements, 
beyond that of the microsaurs, had already taken place in the 
"adelogyrinids" of fairly early Mississippian times. These forms 
had departed at the dawn, so to speak, of amphibian evolution 
very far from the skull pattern of the Labyrinthodontia, while more 
typical microsaurs, as suggested below, could have been readily 
derived, as regards most cranial structures, from labyrinthodonts 
during Carboniferous times. 

If we compare the skull of such a microsaur as Pantylus with 
that of early labyrinthodonts, we find that the two agree in many 
features. Apart from the diagnostic change in the temporal region 
and greater overlap of roof elements onto the occiput, the dermal 
roof pattern and that of the jaw are very close to the early 
labyrinthodont plan. The palate, too, is similar to that of the 
more primitive labyrinthodonts, and even the broadening of the 
concave occipital condyle parallels the trend in temnospondyls. 
The only really distinctive skull features, apart from the "supra- 
temporal" region, are in the dorsal development of the sphenoid, 
the down-turned anterior process of the supraoccipital, and the 
lateral extension of the otic region. As far as skull structure is 
concerned, there is no difficulty in deriving typical microsaurs 
from labyrinthodonts; and, disregarding vertebral structure, the 
changes necessary need not, one would think, necessitate more 
than a fraction of Carboniferous time for their accomplishment. 
Typical microsaurs, considering skull structure alone, could have 
evolved from Lower Carboniferous — Mississippian — labyrintho- 
donts. 

I have previously (1950, etc.) suggested that microsaurs may 
have given rise to part, at least, of the modern amphibian groups. 
Parsons and Williams (1963) are inclined to doubt this, citing 
such features as the long and slender body and feeble limbs of 



34 breviora No. 314 

many microsaurs as tending to throw them out of consideration 
as ancestors of the Lissamphibia. But as Carroll and Baird ( 1968) 
point out, such forms as Pantylus and Tuditanus show more sturdy 
body proportions and stouter limbs than most earlier described 
types. There is no positive proof, but I continue to believe that 
microsaurs are still worthy of consideration as possible ancestors 
of the modern orders. From time to time, most recently by Estes 
(1965). the rhachitomes, particularly the Dissorophidae, have 
been suggested as possible ancestors of part or all of the lissamphib- 
ian groups. But although Estes is inclined to consider the balance 
slightly tipped in favor of the rhachitomes, there seems to be little 
in the way of conclusive evidence. 

MICROSAURS AND REPTILES 

Certainly typical microsaurs resemble early reptiles in various 
ways, implying a considerable degree of parallelism if not true 
relationship, notably in such characters as the tarsus and the 
attainment o\' a "holospondylous" construction of the vertebral 
column. But I am quite unable to understand how anyone who 
looks even superficiall) at the skull can believe that any reptiles, 
such as captorhinomorphs, can be oi microsaur descent. The pat- 
tern oi the temporal region has evolved in a fashion very different 
from that leading to reptiles; the "supratemporal" (or tabular), 
reduced and soon lost in reptiles, has enlarged in spectacular 
fashion in microsaurs. And even in such features as the tripartite 
condyle, the microsaurs have gone in a direction quite at variance 
with that proper to reptile ancestors, which have retained and 
emphasized the primitive single condyle of the ancestral amphib- 
ians. It seems to me high time that this impossible theory of 
reptile descent from microsaurs should be given permanent burial. 

I suggested at one time that possibly microsaurs, although not 
ancestral to reptiles, might, on the contrary, be an offshoot of 
primitive captorhinomorphs. I have never considered this as too 
probable, and now believe this to be no more worthy of belief 
than the opposite relationship. Sudden great expansion of a 
"supratemporal." already much reduced in even the most primitive 
captorhinomorphs. a radical shift in structure in the posterior part 
of the braincase away from the reptilian pattern seen in capto- 
rhinomorphs. a paralleling of temnospondyls and modern amphib- 
ians in a "last-minute" expansion of the occipital condyle — such 
shifts seem highly improbable indeed. 



1969 CRANIAL ANATOMY OF PANTYLUS 35 

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\niei -.. 16: 1597-1658. 
1950. The nature and relationships of the Paleozoic microsaurs. 

Amer. Jour. Sci., 218: 628-654. 
1956. Osteology of the Reptiles. Chicago, 772 pp. 

Tatarinov, L. P. 

1964. Microsauria. In: J. A. Orlov, ed.. Osnovy Paleontologii. V. 
Amphibians, reptiles, and birds, pp. 164-171. 

Vaughn, P. P. 

1960. On the possible polyphyletic origin of reptiles. Evolution, 14: 

274-276. 
1962. The Paleozoic microsaurs as close relatives of reptiles, again. 

Amer. Midland Natur., 67: 79-84. 



1969 CRANIAL ANATOMY OF PANTYLUS 37 

WESTOLL, T. S. 

1942a. Ancestry of captorhinomorph reptiles. Nature (London), 149: 

667. 
1942b. Relationships of some primitive tetrapods. Nature (London), 

150: 121. 

WlLLISTON, S. W. 

1908. The oldest known reptile, lsodectes punctulatus Cope. Jour. 

Geol., 16: 395-400. 
1913. Ostodolepis brevispinatus, a new reptile from the Permian of 

Texas. Jour. Geol., 21: 363-366. 
1916a. The osteology of some American Permian vertebrates. II. 

Contr. Walker Mus., 1: 165-192. 
1916b. Synopsis of the American Permo-carboniferous Tetrapoda. 

Contr. Walker Mus., 1: 193-236. 
1925. The Osteology of Reptiles. Cambridge, Mass., 300 pp. 

Wilson, J. A. 

1951. Taxonomic position of Pantylus. Bull. Geol. Soc. Amer., 62: 
1490-1491 (abs.). 

(Received 17 January 1969.) 



BREVIOR-A 

Museiuii of Comparative Zoology 

Cambridge. Mass. 31 March, 1969 Number 315 



STELUCOLA DENTIFER N. SP. (COPEPODA, CYCLOPOIDA) 
ASSOCIATED WITH A STARFISH IN JAMAICA 

Arthur G. Humes' 



Abstract. Stellicola dentijer n. sp., a Iichomolgid copepod, is described 
from Jamaica, where it is associated with the asteroid Liiidia clathrata 
(Gray). 

INTRODUCTION 

The genus Stellicola, containing twenty species, has been reported 
from Europe, West Africa, the Indian Ocean, and the western Pa- 
cific Ocean. The new species described here extends the range to 
Jamaica, and brings to fifteen the number of species in the genus 
known to occur on asteroids. 

The host of the new copepod, Luidia clathrata (Gray), is a com- 
mon starfish in shallow water from North Carolina through the 
West Indies. 

The figures have been prepared with the aid of a camera lucida. 
The letter after the explanation of each figure refers to the scale 
at which it was drawn. The abbreviations used are: A! = first an- 
tenna, A 2 = second antenna, MXPD = maxilliped, and P a =leg 1. 

ACKNOWLEDGEMENTS 

The copepods were collected by the author and Dr. Richard U. 
Gooding during field work in 1959 supported by a grant (G-8628) 
from the National Science Foundation of the United States. The 
study of the specimens has been aided by another grant (G-5838) 
from NSF. 

I wish to thank Dr. Elisabeth Deichmann, Museum of Compara- 
tive Zoology, for the identification of the host starfish. 

'Boston University, Boston, Massachusetts; Associate in Marine Inverte- 
brates, Museum of Comparative Zoology. 



2 BREVIORA No. 315 

Family L1CHOMOLGIDAE Kossmann, 1877 

Genus Stellicoi \ Kossmann, 1S77 

Stellicola den I II IK ' n. sp 

Figures 1-30 

Type material. - - 27 . ?, 17 . and 4 copepodids from 13 

Luidia clathrata (Gray) in a depth of 2 in. near mangroves, cast 
of the small boat channel leading from the hurricane anchorage, 
Port Royal, Jamaica. Collected September 10, 1 959. Holotype 
allotype, and 32 paratypes (21 .11 | | ) deposited in the 

United States National Museum, Washington, and the remaining 
paratypes in the author's collection. 

Female. -The bod) (Fig. I) has a moderatel) wide and only 
slightl) depressed prosome. I he length (not including the setae on 
the caudal ramus) is 1.39 mm (1.3 1 -1. 47 mm) and the neatest 

C 

w dth is 0.48 mm ( 0.43-0.5 I mm ). based on 10 specimens in lactic 
acid. I he ratio oi the length to the width o\' the prosome is 1.47: 1. 
The segment of leg 1 is incompletel) separated from the head by 
laterodorsal furrows and bears a transverse band i>l line striae on 
its dorsal surface. The epimeral areas o\' the metasomal segments 
are more or less rounded, except lor that of the segment of let: 4. 
which is truncated. 

The segment o\ leg 5 < I g. 2) is 104 • 161 ,». Between this seg- 
ment and the genital segment there is a distinct ventral interseg- 
mental sclerite (Fig. 3). The genital segment is 180 /< long, in 
dorsal view divided by lateral constrictions into a broad anterior 
hall' (width 187 ") with convex lateral margins and a much nar- 
rower and unexpanded posterior half (width 120 // ) with almost 
straight margins. In lateral view (Fig. 3) the thicker anterior half 
is separated from the posterior half by an abrupt transverse dorsal 
constriction (indicated also m Fig. 2). The areas o\' attachment 
of the egg s.ics are located laterally (Fig. 3). each area ( Figs. 4 
and 5 ) bearing two naked setae 20 u and I 1 <> in length and a spini- 
form process about 8 t i long. I he three postgenital segments are 
81 X 97 ft, 55 X 83 ,.. and 83  78 u from anterior to posterior. 
I he posteroventral margin of the anal segment is smooth. 

The caudal ramus (big. 6 ) is elongated. 148/< in length, its great- 
esl width proximally 31 ». and its least width distally 19 ,,. The 



1 The specific name dentifer, from Latin dens = a tooth, and fero = to 
Hear, refers to the toothlike process on leg 5 in the female. 



1969 COPEPOD FROM A JAMAICAN STARFISH 3 

ratio of length to width (greatest dimensions) is 4.77: 1. The outer 
lateral seta is 78 fi and the dorsal pedicellate seta 29 /*, both of them 
naked. The outermost terminal seta is 81 /x, finely barbed in its 
distal half. The innermost terminal seta is 86 /x, with a few minute 
barbules near its tip. The two median terminal setae are 176 [i 
(outer) and 330 p (inner), both naked and inserted a little ven- 
trally. There is a minute setule 6 /x long on the outer proximal area 
of the ramus. The ornamentation consists of a few small hairs. 

The dorsal surface of the prosome and both dorsal and ventral 
surfaces of the urosome bear numerous hairs (sensilla) and re- 
fractile points. The ratio of the length of the prosome to that of the 
urosome is 1.1:1. 

The egg sac (Figs. 1 and 7) is an elongated oval, about 420 X 
200 fi, extends as far as the anal segment, and contains 7-13 eggs, 
each approximately 100 /x in diameter, but somewhat variable in 
shape. 

The rostrum (Fig. 8) has a broad and rather truncated postero- 
ventral margin. 

The first antenna (Fig. 9) is 7-segmented, but a sclerotization on 
the ventral surface of the third segment (Fig. 10) suggests an in- 
tercalary segment. The lengths of the segments (measured along 
their posterior non-setiferous margins) are: 32 (48 /x along its an- 
terior margin), 75, 31, 33, 32, 20, and 18 p respectively. The for- 
mula for the armature is 4, 13, 6, 3, 4 + 1 aesthete, 2 -f- 1 aesthete, 
and 7+1 aesthete, as usual in the genus. All the setae are naked. 

The second antenna (Fig. 1 1) is 3-segmented, without a trace of 
division of the third segment. The first two segments are short, and 
each bears one seta. The longer third segment bears the usual inner 
group of three setae, is ornamented with a diagonal row of small 
spinules on its proximal posterior surface, and has distally three 
recurved claws and four setae. One claw (50 it along its axis and 
inserted terminally) is much larger than the other two claws (both 
about 19 p, one inserted on the subterminal anterior surface of the 
segment, the other subterminally near the large claw). One of the 
setae has a swollen base, the others are slender. All the elements 
are naked. 

The labrum (Fig. 12) has two divergent lobes, each with a small 
inner pointed process and a distal hyaline lamella. The mandible 
(Fig. 13) has on its concave margin a row of long slender spinules; 
its convex margin bears proximally a group of minute spinules fol- 
lowed by a large hyaline dentiform process and a row of smaller 
teeth. The long terminal flagellum has a row of spinules along the 
proximal half of its inner margin. The paragnath (Fig. 14) is a 



4 breviora No. 315 

small hairy lobe. The first maxilla (Fig. 15) bears four setae, three 
terminal and one subterminal. The second maxilla (Fig. 16) has 
an unarmed first segment. The second segment bears a surficial 
posterior seta and an inner setifonn spine. The terminal lash has 
on its convex side a large, weakly-selerotized tooth followed by a 
row of long slender spinules. and on its concave side a few barbules; 
across the base oi the lash there is a diagonal row of small spinules. 
The maxilliped ( Fig. 1 7 ) is 3-segmented. The slender first segment 
bears two minute distal inner spinules. The second segment is ex- 
panded medially and bears two naked setae and numerous spinules. 
The small third segment bears a barbed spine, a hyaline seta, and a 
minute spinule; the segment terminates in a large spiniform pro- 
longation with an attenuated tip. 

The ventral surface between the maxillipeds and leg 1 is slightly 
protuberant. A sclerotized line connects the bases of the maxil- 
lipeds (Fig. 18). 

Legs 1-4(1 igs. 19, 20. 21. and 22 ) have the following armature 
(the Roman numerals indicating spines, the Arabic numerals rep- 
resenting setae ) : 

P, coxa 0-1 
P a coxa 0-1 
IV coxa 0-1 

P, coxa 0-1 

The inner seta on the coxa oi legs 1-3 is long and plumose, but 
in leg 4 this seta is short ( 15 p ) and naked. The inner side of the 
basis in leg 1 is smooth, but in legs 2-4 bears a row of hairs. The 
endopod of leg 4 (Fig. 22) is much shorter than the exopod. the 
ratio being about 1 :2.3. The first segment is 34 X 29 p, with its 
inner seta 105 fx. The second segment is asymmetrical, 43 X 29 //. 
and armed with an inner seta 110/4. and two terminal setae 61 fi 
(outer) and 75 /«. (inner). All four cndopodal setae have very fine 
distal barbules. The outer margins of both segments bear a row of 
hairs, and the second segment has a row of very small spinules on 
its distal margin. 

Leg 5 (Fig. 23) has a prominent, toothlike, distally-directed 
process on the inner margin of the free segment. This segment is 
39 n long. 34 p in greatest width at the level of the tooth, and 46 p 



basis 


1-0 


exp 


1-0 


1-1 


III. 1. 4 






enp 


0-1 


0-1 


1. 5 


basis 


1-0 


exp 


1-0 


i-i 


III. 1. 5 






enp 


0-1 


0-2 


I. II. 3 


basis 


1-0 


exp 


I-o 


I-I 


III. I. 5 






cnp 


0-1 


0-2 


I. II. 2 


basis 


1-0 


exp 


1-0 


1-1 


II. 1. 5 






enp 


0-1 


2. 1 





1969 COPEPOD FROM A JAMAICAN STARFISH 5 

in greatest diagonal dimension. The two terminal setae are 91 p, and 
39 ^, and the seta on the body near the insertion of the free seg- 
ment is 31 ft. All three setae are naked. Near the outer side of 
the free segment there is a slight oblique dorsal ridge with a series 
of extremely small points (spinules ?) . 

Leg 6 is probably represented by the two setae near the attach- 
ment of each egg sac (Fig. 5 ) . 

The color in life in transmitted light is opaque, the eye red. 

Male. — The body (Fig. 24) has a prosome more pointed an- 
teriorly than in the female. The length (without the ramal setae) is 
1.18 mm (1.11-1.21 mm) and the greatest width is 0.35 mm (0.31- 
0.41 mm), based on 10 specimens in lactic acid. The ratio of the 
length to the width of the prosome is 1.74: 1. 

The segment of leg 5 (Fig. 25) is 55 X 1 12 ^t. Between this seg- 
ment and the genital segment there is no ventral intersegmental 
sclerite. The genital segment is 156 X 160 /*, with rather flattened 
lateral margins in dorsal view. The four postgenital segments are 
78 X 99 fx, 73 X 81 p, 62 X 68 /x, and 62 X 62 p from anterior to 
posterior. 

The caudal ramus resembles that of the female, but is smaller, 
110X 25/*. 

The rostrum, first antenna (with no aesthetes added), second 
antenna, labrum, mandible, paragnath, first maxilla, and second 
maxilla are like those of the female. The maxilliped (Fig. 26) is 
slender and 4-segmented, assuming that the proximal part of the 
claw represents the fourth segment. The second segment bears two 
setae and two rows of spinules. The strongly recurved claw, 159 fx 
along its axis, bears two very unequal proximal setae and is divided 
about midway. 

The area between the maxillipeds and leg 1 resembles that in the 
female. 

Legs 1-4 are similar to those in the female, with the same seg- 
mentation and armature, but the second segment of the endopod of 
leg 4 (Fig. 27) is more symmetrical. 

Leg 5 (Fig. 28) has a free segment 26 X 13 //., without an inner 
toothlike process. The two terminal setae are 72 p and 32 it, and 
the adjacent seta on the body is 32 p. 

Leg 6 (Fig. 29) consists of the usual posteroventral flap on the 
genital segment, bearing two naked setae 32 /x and 44 p. 

The spermatophores (Fig. 30), attached to the female in pairs, 
are each about 120 X 48 p., not including the neck. 

The color in life resembles that of the female. 



6 BREVIORA No. 315 

COMPARISON WITH RELATED SPECIES 

S. dentifer may easily be distinguished from all other species in 
the genus by the presence of the toothlike process on leg 5 in the 
female. The genital segment of the female also has a configuration 
unlike that of any other species. 

In most species o\' Stellicola the caudal ramus is not more than 
twice as long as wide. Two species, however, have a relatively long 
caudal ramus. In S. gracilis (Thompson and A. Scott, 1903). the 
ratio of length to width oi the ramus is 4: 1. and in S. affinis Humes 
and Ho. 1967. it is 6.3-7:1 (in the female) and 7.5:1 (in the 
male). I he new species is readik separated from these two species. 
both oi which have a single claw on the second antenna and. in 
addition, differ in the two points mentioned in the previous para- 
graph. 

S. dentifer seems to be most closely related to the four West Afri- 
can species described b\ Humes and Cresse) ( 1958), S. frequens, 
S. astro pectinis, S. luidiae, and S. hiatus, all of which similarly 
have three claws on the second antenna. Unlike most other species 
of Stellicola, the inner edge of the free segment of leg 5 in these four 
species is somewhat swollen or irregular; a toothlike process is ab- 
sent, however. 

The twenty-one species of Stellicola may be grouped geographi- 
cally on the basis of the number o\' claws on the second antenna. 
Those from the Indian and western Pacific oceans ( 15 species) have 
one such claw. The only known species from Europe and the Medi- 
terranean Sea. S. clausi ( Rosoll, 1 889 ). has two claws (see Bocquet 
and Stuck. 1962). The five species from the tropical Atlantic Ocean 
(four from West Africa and one from Jamaica) have three claws. 

SUMMARY 

The new species Stellicola dentifer is associated with the asteroid 
Luidia clathrata (Gray) in Jamaica. The copepod may be recog- 
nized within the genus by the prominent toothlike process on leg 5 
in the female. Four West African species of Stellicola appear to be 
related to the new Jamaican form. 

LITERATURE CITED 

Bocquet. C, and J. H. Stock 

1962. Copepodes parasites d'invertebres des cotes de la Manche. IX. 
CyclopoYdes associes a Marthasterias glacialis (L.). Arch. Zool. 
Exp. Gen., 101 I Notes et Revue) (2):79-91. 



1969 COPEPOD FROM A JAMAICAN STARFISH 7 

Humes, A. G., and R. F. Cressey 

1958. Four new species of lichomolgid copepods parasitic on West 
African starfishes. Bull. Inst. Fr. Afrique Noire, 20, ser. A (2) : 
330-341. 

Humes, A. G., and J.-S. Ho 

1967. New species of Stellicola (Copepoda, Cyclopoida) associated 
with starfishes in Madagascar, with a redescription of S. 
caeruleus (Stebbing, 1900). Bull. Brit. Mus. (Nat. Hist.), Zool., 
15(5): 201-225. 

KOSSMANN, R. 

1877. Entomostraca (1. Theil: Lichomolgidae). In: Zool. Ergeb. 
Reise KUstengeb. Rothen Meeres, erste Halfte, IV, pp. 1-24. 

Rosoll, A. 

1889. Uber zwei neue an Echinodermen lebende parasitische Cope- 
poden: Ascomyzon comatulae und Astericola clausii. S.-B. 
Math.-Naturw. CI. Ak. Wiss. Wien, 97 (Abt. I, Heft 4): 188- 
202. 

Thompson, I. C, and A. Scott 

1903. Report on the Copepoda collected by Professor Herdman, at 
Ceylon, in 1902. Rept. Govt. Ceylon Pearl Oyster Fish. Gulf 
of Manaar, part I, suppl. rept. no. 7, pp. 227-307. 

(Received 14 January 1969.) 



8 



HREV10RA 



No. 3 1 5 




Figures 1-8. Stellicola dentifer n. sp.. female: 1. dorsal (A); 2, urosome, 
dorsal (B); 3, segment of leg 5 and genital segment, lateral (B); 4. area of 
attachment of egg sac. dorsal (C): 5, area of attachment of egg sac, lateral 
iC ); 6, caudal ramus, dorsal (D); 7. egg sac, dorsal (E); 8. rostrum, ven- 
tral (F). 



1969 



COPEPOD FROM A JAMAICAN STARFISH 



9 




Figures 9-18. Stellicola dentifer n. sp., female: 9, first antenna, anterodor- 
sal (G); 10, third segment of first antenna, posteroventral (H); 11, second 
antenna, posterior (D); 12, labrum, ventral (D); 13, mandible, posterior 
(C); 14, paragnath, posterior (C); 15, first maxilla, posterior (C); 16, sec- 
ond maxilla, posterior (C); 17, maxilliped, antero-inner (C); 18, area be- 
tween maxillipeds and leg 1, ventral (G). 



10 



imrvioRA 



No. 3 1 5 









^ 



















/ 






21 






J 













22 









19 



20 








. 









Figures 19-22. Stellkola dentifer n. sp.. female: 19. leg 1 and intereoxal 
plated anterior (G); 20. leg 2. anterior (G): 21. leg 3. anterior (G): 22. leg 
4 and intereoxal plate, anterior (G). 



1969 



COPEPOD FROM A JAMAICAN STARFISH 



11 




Figures 23-30. Stellicola dentifer n. sp. Female: 23, leg 5, dorsal (C), 
Male: 24, dorsal (E); 25, urosome, dorsal (B); 26, maxilliped, anterior (D); 
27, endopod of leg 4, anterior (G); 28, leg 5, dorsal (C); 29, leg 6, ventral 
(D); 30, spermatophores, attached to female, lateral (G). 



BREVIORA 

Museum! of Cooiparsitive Zoology 

Cambridge, Mass. 30 April, 1969 Number 316 



CYTOTAXONOMIC STUDIES ON SOME UNUSUAL 

IGUANID LIZARDS ASSIGNED TO THE GENERA 

CHAMAELEOUS, POLYCHRUS, POLYCHROIDES, AND 

PHENACOSAURUS, WITH BEHAVIORAL NOTES 

G.C.Gorman 1 

R. B. Huey 2 
E. E.Williams 

INTRODUCTION 

Abstract. Chromosome data are presented for four species of iguanid 
lizards. Phenacosaurus heterodermus has a diploid number of 36, with 12 
metacentric microchromosomes and 24 microchromosomes. Chamaeleolis 
porcus also has 12 metacentric macrochromosomes. The exact number of 
microchromosomes was not ascertained, but it is quite similar to that in 
Phenacosaurus. The karyotypes found in these two species resemble the 
primitive iguanid condition and that most frequently found in alpha Anolis. 
Polyckrus fetnoralis has an apparent diploid number of 26, with 10 pairs of 
acrocentric and one pair of submetacentric macrochromosomes, and two 
pairs of microchromosomes. Polychrus peruvianas has 20 acrocentric macro- 
chromosomes and 8 microchromosomes (2n = 28). A karyotype with 
low diploid number (2n = 30 or less) and few metacentric macrochromo- 
somes is very unusual in iguanid lizards — previously reported only for 
Polychrus marmoratus. Its presence in the two presently studied Polychrus 
confirms their very close relationship inter se, but gives no clue to their 
relationship to other members of the family Iguanidae. P. peruvianas has 
previously been assigned to the monotypic genus Polychroides. However, 
osteological and cytological data show its very close relationship to Poly- 
chrus, and we formally propose the synonymy of Polychroides with Poly- 
chrus. Notes on behavior and ecology are appended for the four species. 
Although there may be a nomenclatural problem surrounding the use of the 
name P. femoralis, our Peruvian animal resembles P. femoralis from Loja, 



1 Museum of Vertebrate Zoology, University of California, Berkeley, Cali- 
fornia 94720. 

- Biological Laboratories, University of Texas, Austin, Texas. 



2 BREVIORA No. 316 

Ecuador. This is the first record of an apparent Polychrus femoralis from 

Peru. 

We have recentl) obtained several poorl) known iguanid lizards 

Oil Held trips to Peru (RBH). and Cuba and Colombia (GCG). 
Although our observations and chromosome data arc meagre, 
the) provide insight into the relationships of these lizards. Field 
and behavioral notes are appended because of the paucit) of in- 
formation on these species and because we have the first record 
of an apparent Polychrus femoralis? from Peru. 

The genus Chamaeleolis is endemic to Cuba and the adjacent 
Isle of Pmcs. [.on^ considered monotypic, Garrido and Schwartz 
(1968) have demonstrated that there are two distinct species. 

Both are arboreal, moderatc-si/cd lizards (snout-vent of adult 
males 155 mm), quite closel) related to Anolis. The) share with 
anoles extensible clew laps and adhesive lamellae on the toes but 
differ in a number of OSteological features, the most conspicuous 
of which is a parietal ercst that extends tar back over the neck 
(Etheridge, 1960), giving the animals a marked resemblance to 
true old-world chamaeleons. 

Phenacosaurus is an iguanid genus with ver) close affinities to 
Anolis. Dunn ( I W44 ) listed a series ol characters by which the 
genus was supposed to differ from Anolis (e.g. a prehensile tail, 
a heterogeneous scutcllation with large and small scales intermixed, 
a dorsal crest of enlarged scales), but Etheridge (PHiO) pointed 
out that all of these characters occur individual!) in a number of 
species of Anolis. However, because of a number of osteological 
peculiarities (low number of sternal ribs, high number of paraster- 
nal chevrons), Etheridge retained the genus. 

Phenacosaurus is known predominantly from high altitudes 
(1800-3500 m) in the Eastern Andes of Colombia. There are 
currentl) four recognized species, but richteri and paramoensis 
are undoubtedK synonyms of I', heterodermus, and there is at least 



I he single specimen reported here runs down i<> P. femoralis in the ke\ 
provided bv H. W. Parker (1935: 516). Examination of comparative ma- 
terial (all of it femoralis h\ Parkers key) indicates that there may he as 
man\ as three distinguishable populations. \ specimen in the United States 
National Museum from Guayaquil, the type locality of femoralis. has 
markedK larger scales than Parker's l.oja specimens, which closeh resemble 
the Peruvian animal. Pending revisionarv studies, we believe it to be prefer- 
able to use the name femoralis for all this material, including our Peruvian 
animal. 



1969 CYTOTAXONOMY OF IGUANIDS 3 

one undescribed species from Ecuador (J. D. Lazell, Jr., personal 
communication). 

Polychrus is a genus of arboreal lizards from Central and 
South America (including Trinidad) comprising about five distinct 
species (Parker, 1935). Etheridge (1960) compared 22 charac- 
ters in Anolis and Polychrus (excluding characters unique to 
Anolis or common to the family Iguanidae) and found that 20 
were shared by Anolis and Polychrus. He wrote, "Although there 
appears to be a close correspondence between Polychrus and the 
anoles, there are striking differences. Most of these differences, 
however, are features which are either unique to the anoles or 
unique to Polychrus and therefore tell us little more than that the 
anoles are separated from other members of the family, including 
Polychrus, by a wide morphological gap" (Etheridge, 1960: 111). 
He also pointed out that Polychrus and Chamaeleolis are highly 
arboreal, "and unusual in their habit of slow, deliberate move- 
ment. The similarities of these genera may well be parallel adapta- 
tions to a similar way of life, yet the occurrence of multiple parallel 
developments may, in itself, be an indication of relationship be- 
tween the groups in which it occurs" (Etheridge. 1960: 113). 

The monotypic genus Polychroides was erected by Noble (1924) 
when describing a new species of iguanid lizard (P. peruvianus) 
from the provinces of Cajamarca and Piura, northern Peru. Noble's 
comments on the affinities of the new genus were quite confused 
and are here quoted verbatim. On p. 109 in "Remarks" following 
the generic diagnosis he wrote: "The generic status of many of 
the slow-moving arboreal iguanids is very uncertain. The species 
described below cannot be referred to either Enyalius, Enyalioides 
or Polychrus as at present defined. It seems most closely related 
to Polychrus with which it agrees in its femoral pores, large head 
scales, subequal third and fourth toe and its sacculated lung. T 
have seen both Polychrus and Polychroides alive and have been 
struck by their great similarity in behavior. The pronounced nuchal 
crest of the latter readily distinguishes it from the former." 

On p. 11 0, however, in the "Remarks" which follow descrip- 
tion of the species he stated: "This species is closely allied to 
Enyalioides festae Peracca from which it differs in its larger head 
scales and gular sac, also in certain differences in the scutation of 
the digits and head. I would not hesitate to refer it to the genus 
Envalioides were it not that this procedure would require a consid- 
erable modification of our present conceptions of that genus." 



4 BREVIORA No. 316 

Etheridge I I960) was unable to find an) skeletal differences be- 
tween Polychrus and Polychroides in \-ra\ studies of iguanid gen- 
era, and he therefore cited peruvianus in his list of material exam- 
ined as another species of Polychrus. Onl\ the external charac- 
ter of a dorso-nuchal cresl distinguished peruvianus from other 
Polychrus. 

Chromosomes have been useful in elucidating relationships of 
iguanid lizards. I he majority of species studied have a karyotype 
consisting of six pairs of metacentric macrochromosomes and 
eleven or twelve pairs of microchromosomes (Gorman, Atkins and 
Holzinger, 1967; Gorman and Atkins, l l ><>7. 1968). One species, 
Plica plica, showed apparent centric fissioning with lour pairs of 
acrocentric macrochromosomes. I he chromosomes o\ the single 
species of Polychrus examined. /'. marmoratus, were so different 
irom all other species of iguanids ol which karyotypes are known 
that it was difficult to relate this genus to am of the others (Gor- 
man, Atkms, and Holzinger, 1967). Female /'. marmoratus have 
a diploid number of 30, consisting ol 20 acrocentric macrochromo- 
somes and 10 acrocentric microchromosomes. Males have a 
diploid number of 29, with l metacentric and 19 acrocentric 
macrochromosomes, and 9 acrocentric microchromosomes. In both 
Anolis (Gorman, l l M7) and Sceloporus (Lowe, Cole, and Patton, 
1967), species have now been found with high numbers of acro- 
centric macrochromosomes. I hese speeies. however, are charac- 
terized h\ high diploid numbers, greater than 2n JO, the formula 
which appears primitive lor the family [guanidae (Gorman, et al., 
1967 I I he probable explanation tor these high numbers is centric 
fission of the metacentric macrochromosomes (Gorman, Baptista, 
and Bury, 1969). 

MATERIALS AND METHODS 

Chromosome preparations were made b\ tissue culture of whole 
blood following methods alread) outlined; b\ culture of marrow ob- 
tained from the femur and treated in the same fashion as blood; 
and b\ direet testis preparations (see Gorman, et al.. 1967). 
The specimens used were one female Polychrus peruvianus (Mu- 
seum o\' Vertebrate Zoology, no. X2X34). one female Polychrus 
femoralis (M.V.Z. 82835), one male Chamaeleolis porcus (Mu- 
seum of Comparative Zoology, 100472). and one male Phena- 
cosaurus heterodermus (M.C.Z. 104409). 

Chromosome spreads were photographed and karyotypes ana- 
lyzed as best we could. Unfortunatelv. for Chamaeleolis. Polychrus 
peruvianus, and P. femoralis, we obtained few mitotic divisions. 



1969 CYTOTAXONOMY OF IGUANIDS 5 

and we are unable to present fully definitive karyotypes or diploid 
numbers, though the material permits interesting comparisons. 
Numerous divisions from the testis of Phenacosaurus were studied. 

RESULTS AND DISCUSSION 

Only four clearly resolved metaphase plates were found in 
P. peruvianas, all with 28 chromosomes, all acrocentric. There 
is not a sharp break in size between macro- and microchromo- 
somes; however, 20 might be termed macrochromosomes and 8 
microchromosomes (Fig. la). This is very similar to the karyo- 
type of female Polychrus marmoratus, and this is only the second 
species of iguanid reported to date that has a karyotype of all 
acrocentric chromosomes. So few cells were seen that we cannot 
safely assume that the diploid number is 28 — but it is quite clear 
that the karyotype of P. peruvianas resembles that of P. marmora- 
tus to such a great extent that considering these forms as members 
of separate genera obfuscates their close relationship. We formally 
propose the synonymy of Polychroides with Polychrus. 

Few divisions were obtained in our specimen of P. femoralis — 
but, again, there are enough data to establish both its close rela- 
tionship to the other Polychrus in the similarity of the specialized 
karyotype and its uniqueness among the species thus far examined. 
The diploid number appears to be 26. (This is based upon only 
four metaphase plates; additional spreads were seen, but, because 
of overlaps, prec : se counts could not be made.) In all cells, there 
is one pair of submetacentric chromosomes (Fig. lb). Within the 
genus Polychrus, a submetacentric chromosome had previously 
been seen only in the male of P. marmoratus, where it is quite 
clearly the Y chromosome (Gorman, et al., 1967). In P. femo- 
ralis also, the largest chromosome pair is clearly subacrocentric 
with minute short arms present. There is a total of 1 1 pairs of 
macrochromosomes and 2 pairs of microchromosomes in the cells 
that were carefully analyzed. The karyotype of P. femoralis is 
most likely derived from a marmoratus-\\ke karyotype of 2n=30. 
Centric fusions of microchromosomes to macrochromosomes 
would account for the reduction in diploid number and the appear- 
ance of submetacentric and subacrocentric chromosomes. 

Numerous metaphase plates were observed in Chamaeleolis 
porcus, but we lack definitive information on the number of micro- 
chromosomes. It is quite clear that the karyotype resembles the 
"typical" iguanid in having six pairs of metacentric macrochromo- 
somes and a series of more than 20 microchromosomes (Fig. 2). 



I!RI \ IOR \ No. 3 Id 



(! 5  1 3 1 1  « a 



/I '• •»«•»'»« 



X* 



Oil"* 

Fig. I. Karyotypes of Polychrus. a., P. peruvianus female. There are 
28 acrocentric chromosomes Rone marrow preparation, Giemsa Stain, b., 
P. femoralii female, ["here are 26 chromosomes; one pair nop row. far 
right) is submetacentric. Leukocyte culture, Giemsa stain. 



Onl\ testis preparations were made for Phenacosaurm heter- 
odermus. There were numerous meiotic and mitotic divisions. In 
meiosis we can clearly see six large macrobivalents and 12 con- 
siderably smaller microbivalents (lis:. 3). In mitosis, it was quite 
clear that the microchromosomes were metacentric. Thus, the 
diploid number is 36, with 24 microchromosomes. 



969 



CYTOTAXONOMY OF IGUANIDS 




Fig. 2. Mitotic metaphase of Chamaeleolis porcus male showing the 
"typical iguanid" condition of 12 metacentric macrochromosomes, and 
a sharp break in size between the macro- and microchromosomes. 



Among Anolis, Etheridge (1960) defined two major groups 
termed alpha and beta. Chromosome studies (Gorman, et al., 
1967) have shown that the primitive alpha karyotype is identical 
with that of the primitive iguanid condition ( 12 metacentric macro- 
chromosomes, 24 microchromosomes). In a recently proposed 
classification of iguanids on the basis of structure of the caudal 
vertebrae (Etheridge, 1967), Chamaeleolis and Phenacosaurus 
were placed in a group with the alpha anoles; chromosomes sup- 
port this classification. 

Polychrus remains a puzzle within the iguanids. Now that three 
species have been studied it is quite clear that they are close to 
each other inter se, but distinctly different from anoles and anoline 
genera such as Chamaeleolis, Phenacosaurus, and Anisolepis (Gor- 
man, et al., 1967). 

Since this paper was originally written, we have received the 
following new data. 



8 



Bkl \ IORA 



No. 3 I o 



On November 26, 1968, Dr. John Wright oi the Los Angeles 
County Museum collected a male Polychrus peruvianus at 3.5 km 
east of the junction between Bagua Grande and Bagua Chica, 15.5 
km west of Bagua Grande, Ama/onas Dept.. Peru, at an altitude 
of 2.0(H) feet. (Field catalog number P- c >^ 7 ; to be deposited in 
L.A.C.M ) 

I es;es were mineed and allowed to settle in a hypotonic citrate 

solution (as no centrifuge was available), the citrate was removed 
with an e\e-dropper. and fixative was added. I his field prepara- 
tion was then brought baek to the United Stales, where slides were 
made some two months later. 

Dr Wright has kindb consented to our examination of the 
slides and Utilization of the data. At diakinesis there are 13 bodies. 




Fig. 3. Diakinesis in Phenacosaurui heterodermus. There are six large 
bivalents, and 12 microbivalents. n = IS. Large round black area is a sperm 
head, not part of the meiotic figure. 



1969 



CYTOTAXONOMY OF IGUANIDS 



12 bivalents, and a trivalent. The male diploid number is thus 
expected to be 27. One mitotic metaphase was seen, and 27 
chromosomes were counted. Position of the centromere could 
not be ascertained. One clear metaphase II cell had 14 chromo- 
somes, all acrocentric. 

These data are consistent with the sex chromosome situation 
known in Polychrus marmoratus, in which the males have a diploid 



o 






\ 





4Mf 




Fig. 4. Diakinesis of Polychrus peruvianus male. There are 13 bodies, 12 
bivalents, and a sex trivalent ( arrow ) . 



10 BREVIORA No. 3 Id 

number one lower than the females, the Y chromosome is meta- 
centric (not demonstrated for P. peruvianus), and a sex bivalent 
is clearl) evident at meiosis (Gorman, Atkins, and Holzinger, 
L967). 

The finding of 12 bivalents and a trivalent in /'. peruvianus, with 
an expected male 2n o\ 27. gives further strength to our assump- 
tion that female 2n 28 (the latter figure having been arrived 
at with the examination of relatively lew cells). Figure 4 illustrates 
diakinesis in /'. peruvianus. 

NOTES ON POLYCHRUS 
POLYCHRUS PI Rl V IANUS 

Five specimens of Polychrus peruvianus were collected: 
MYZ S2S34 F.I Arenal. Rio Huancabamba, elevation 3000 
82388 ft.. 7 km V 50 km 1 of Olmos, Dept. Caja- 
83678 marea. Peru. 
83679 
MVZ 82413 (skin and skeleton). Imgo. Rio I leubamba. 
elevation 3000 ft., 30 km S, 41 km E of Bagua, 
Dept. Amazonas, Peru. 

I 1 \renal is on a terrace above the Rio Huancabamba. The 

hillsides are dr\ and rocky, and the terrace has a predominance 
of mesquite (Prosopis) interspersed with cultivated areas. All 
specimens were collected on mesquite branches. 

I he I ineo individual was captured b\ Dr. Carl Koford at nighl 
on a horizontal branch o\ an unidentified tree. Stomach contents 
included several Hymenoptera and a leaf fragment. I he largest 
ovarian follicles were less that 2 mm in diameter. 

This localit) is much wetter than El Arenal. Bromeliad-laden 
trees up to 40 feet are found on the steep hillsides, and dense 

second-growth vegetation covers the valle) floor. 

Body coloration in life I MVZ 82834) was medium yellow-green 
with a narrow, pale yellow band running from the eye to the groin 
and olive-drab saddles on the dorsum and tail. Small irregular 
yellow marks were spaced between the saddles. Some capacity to 
darken the body colors was observed in captivity. 

The tail of this specimen was slightly prehensile. P. femoralis 
gave no evidence of this ability. Both species emitted a nasal salt 
solution in captivity (see Templeton. 1967 ). 

P. peruvianus has a more protrusive gular flap than P. femoralis. 
In addition, only P. peruvianus possesses a row of raised scales 
forming a mid-gular crest. 



1969 CYTOTAXONOMY OF IGUANIDS 11 

POLYCHRUS FEMORALIS 

On 31 August 1967, one gravid female of this species was cap- 
tured near Molino, devation 2300 ft., 21 km E, 7 km N of Olmos, 
Dept. Lambayeque, Peru (5.9 S/79.6 W). This specimen 
(MVZ 82835) represents a considerable range extension from 
the record in Loja (Parker, 1932) and an apparent Peruvian 
record. 

She was oriented head down on a vertical branch of a plant 
about 5 ft. tall and remained motionless until capture. The 
locality is in a narrow river valley. Deciduous woodland alternates 
with open areas cleared for farming. 

She was kept alive and placed in a runway with a temperature 
gradient beginning 18 September. Water was sprinkled daily at 
the cool end. On 27 September she laid 12 eggs at the cool, moist 
end of the runway. The eggs were removed and placed in an incu- 
bator. Egg size eight hours later averaged 19.6 mm in length and 
1 1.3 mm in width. The shell texture was soft and leathery. Mold 
developed on the eggs by 4 October and they subsequently decayed. 

In captivity movement was rare. When prodded, she exhibited 
a leaping form of locomotion described by Boker (1935) as 
"bipedal leaping" for Polychrus marmoratus. Davis (1953) ob- 
served a similar method of locomotion in Corythophunes cristatus. 

in life she was light yellow-green with pronounced brown 
saddles on the dorsum. The capacity for color change in this 
specimen was less marked than in P. peruvianas. 

Behavior in Captivity 

Davis (1953) outlined a sequence of four defensive responses 
used by Corythophanes cristatus against apparent predators. These 
can be grouped into two general categories: 
I. Passive defense 

A. Behavioral and morphological camouflage 

B. Catalepsy to eliminate all movement not associated with 
a positional change and thus decrease the probability of 
detection 

H. Active defense 

A. Active postural movements to increase the apparent size 
of the animal 

B. Aggressive biting as a last resort 

Davis stated that the defensive response given was geared to 
the relative immediacy of the threat to the animal's safety. The 
more immed : ate the threat, the more active the response. 



12 BREVIORA No. 316 

Both Polychrus peruvianus and /'. femoralis exhibited this 
sequence of defensive traits in captivity hut did differ in degree 
of response. Since onl\ one specimen of each species was ob- 
served, these notes should be considered tentative. 

Behavioral and morphological camouflage is well marked in 
both species. The colors and patterns render detection oi these 
animals difficult in their arboreal habitats. Their relative im- 
mobility and capacit) for color change accentuate this conceal- 
ment. In both the held and captivity, /'. femoralis relied more 
on immobility than on other defensive responses. 

Behavior similar to the cataleps) oi walking sticks | Phasmidae) 
has been reported for several arboreal lizards (reviewed in Davis) 
and was present in these two species as well. /'. peruvianus could 
be easily induced into this behavior b\ forcibl) altering its position, 
but both used it when coining to rest after voluntary movement. 
When matched with other types o\ camouflage, Davis felt that 
this behavior aided in concealment. 

In response to a threat, main animals arc recorded to increase 
the apparent size o\ the bod\ or parts of the bod) b\ behavioral 
posturing. I his behavior is manifest in social encounters, as well. 
Postural movements common to both Polychrus species involve 
lateral compression ol the body, expansion of the throat fan. and 
head bobbing. I he plane of expansion is sagittal and presented 
broadside for both, as :s the ease in CorythophaneS. The expanded 

area is thus displayed to maximum advantage. 

/'. peruvianus would often gape when posturing. If approached 
closely, she would lunge forward, maintaining balance by the two 
hind legs and tail. She would bite vigorously if given the opportu- 
nity In contrast. /'. femoralis could rarely be induced to bite. But 
it would often attempt to escape b\ leaping. 

Of the two species o\ Polychrus under discussion, the behavior 
of /'. peruvianus was more similar to that n\ Corythophanes than 
was that of /'. femoralis. It is perhaps of significance that /'. 
femoralis, having a much less pronounced gular Map than the 
other two species, relied less on active postural movements and 
more on passive camouflage. 

NOTES ON CHAMAELEOLIS 

Although rare in collections. Chamaeleolis is widely distributed 
throughout Cuba. Through the help of Orlando Garrido. we ob- 
tained a juvenile male C. porcus from La Florida. Sabanilla 



1969 CYTOTAXONOMY OF IGUANIDS 13 

Oriente Province, and a larger (but subaduit) male from La 
Casimba, eight kilometers west of Maisi, Oriente Province. 

Wilson (1957) published a note on the behavior of a captive 
Chamaeleolis. Although he called this C. chamaeleontides (sic), 
the specimen has been subsequently re-identified by Schwartz as 
porcus. Wilson reported, "its behavior was strikingly chamaeleon- 
like and quite different from that of several species of Anolis 
(including the giant A. equestris Merrem) which the author has 
observed in captivity." He stated that it was very sluggish even 
when freshly captured, and that it could be left on a laboratory 
table without much danger of its wandering. Often, it would re- 
main in the same spot for hours or even days without changing 
its position. When confronted with food in the form of living in- 
sects, it would move deliberately and without hurrying. Experi- 
ments using a live male Anolis equestris and a cardboard model 
resembling a male Chamaeleolis failed to elicit any aggressive 
response. Only once, in the year that he had the specimen, did 
it extend its dewlap — and this was without any apparent external 
stimulus. 

Our observations are not at all in accord with this descrip- 
tion. Our specimens would actively jump toward prey (crickets, 
grasshoppers, and mealworms). They would often scamper away 
when left out in the open. Garrido and Schwartz {op. cit.) point 
out that C. porcus is "at least capable of quick and decisive action. " 
They give a detailed account of the escape behavior of one speci- 
men encountered in the field. Our Chamaeleolis were not so quick 
as Anolis, but certainly did not show the incredibly slow move- 
ments of true chamaeleons. Furthermore, they were quite aggres- 
sive. They would display to each other by extending the dewlap 
fully, opening the mouth and protruding the tongue. They also 
displayed in similar manner to A. equestris. When handled, they 
would extend the dewlap and often would bite. Unfortunately, no 
films were taken of the displays 

It appears, then, that the social behavior of Chamaeleolis has 
been misunderstood. Its reported sluggishness and immobility 
appear to be facultative rather than obligatory and may well be 
associated with its cryptic coloration. The ability to move quite 
quickly, and the fact that ano!e-like territorial responses were 
easily elicited implies that the behavior and social structure of 
Chamaeleolis may be quite similar to that of most anoles. 



14 BRE\ IORA No. 3 16 

NOTES ON PHENACOSAURUS 

Previous notes on Phenacosaurm have dealt with specimens 
in captivit) (Osorno-Mesa and Osorno-Mesa | 1946], on shedding 
of skin, incubation of eggs, feeding habits, etc., and Kastle 1 1965], 
a detailed ethological stud) with particular emphasis on social 
behav ior ). 

On Jul\ 10. 1968, Dr. Jorge Hernandez, oi the Universidad 
Nacional. Bogota, led Gorman and J. F. Lynch on a one-da) trip 
specificall) to collect Phenacosaurus in Chia (Cundinamarca), 
a semi-rural suburb of Bogata (at about 35 km distance, alt. ca. 
2700 m) and collected between II am and 2:30 pm. The da) 
was overcast, with intermittent showers, and cold. A thermometer 
was not available, but the temperature for most of the time was 
probabh not in excess of 14 ( . the mean temperature of that 
immediate area. 

I he roads were lined with shrubbery, often blackberries, which 
separated either pasture-land or homes. 

I luce Phenacosaurus were in the first blackberry bush ex- 
amined. I wo. a male and a female, were collected. In the 
course of the afternoon, 14 of these supposedly rare lizards were 
collected. Most often, more than one was taken at a time, usually 
in groups of two. sometimes of three. One pair was actually in 
bodil) contact the male's chm was touching the tail of the 
female. 

All but two of the li/ards were on blackberrs bushes, generally 
1-3 feel above the ground, and usualK on a bare branch. It is 
likelx that as main were missed as were caught — especially 
tin se in leaf) parts of the bushes. I he escape response is to 
drop down, often clinging to a branch with the prehensile tail, 
and then to slink slowl) away. I he most usual perch position 
was almost horizontal; onl) one individual was perched vertically, 
head down 

In review of bod) temperatures in reptiles. Brattstrom ( 1965) 
listed 18.0 C. as the minimum voluntar) temperature tor an 
iguanid lizard. This was l ma notata, which, in fact, has a mean 
bod) temperature of 38.6 when active, and the IX probably 
represents an unusual reading. Normally, iguanids living in 
cold temperatures bask in sunlight until warmed to a preferred 
body temperature of 33 ! 3 . An extreme example was reported 
bv Pearson (1954) for Liolaemus multiformis at high altitudes 
in Peru. He found that the intense isolation enables these species 



1969 CYTOTAXONOMY OF IGUANIDS 15 

to warm to body temperatures of 35°, with shade temperatures 
as low as 11°. 

Phenacosaurus was remarkable for an iguanid lizard because 
it was normally active at low temperatures with no opportunity 
to warm up. We consider it quite possible that its preferred body 
temperature is well below that of any other iguanid reported to date. 

ACKNOWLEDGMENTS 

We thank Dr. Leonard Atkins, Mr. Fernando Ortiz, Mr. Bruce 
Bury, Miss Ida Leone, and Mrs. Charlotte Kayavas for technical 
assistance in the chromosome work, which was supported in 
part by Children's Bureau Project No. 906 and NSF Grant 
GB-6944. Field work in Cuba was supported by Sigma Xi, the 
Graduate Department of Biochemistry, Brandeis University, and 
Academia de Ciencias of Cuba. Special thanks are due to Sr. 
Miguel Jaume and Sr. Orlando Garrido for their help in all our 
work in Cuba. Field work in Peru was supported by the Museum 
of Vertebrate Zoology. The help of Dr. C. Koford, Mr. R. B. 
Hamilton, and Mr. F. McCollum in the field, and Drs. Hans and 
Maria Koepcke and Mr. Robert Conly, our hosts in Peru, is 
greatly appreciated. Field work in Colombia was jointly spon- 
sored by the Museum of Vertebrate Zoology and NSF Grant 
GB-6944. We thank Mr. James F. Lynch for help in collecting 
and for his observations, and we are particularly indebted to Dr. 
Jorge Hernandez C, of the Universidad Nacional in Bogota. He 
brought us directly to the Phenacosaurus locality and provided 
facilities at the University for making chromosome preparations. 
Comparative material permitting provisional identification of the 
Peruvian Polychrus femoralis was loaned by Miss A. G. C. Grandi- 
son (British Museum, Natural History) and Dr. James Peters 
(United States National Museum). We are grateful for their help. 

LITERATURE CITED 

Borer, Hans 

1935. Einfuhrung in die vergleichende hiologische Anatomie der 
Wirheltiere. I. Jena, G. Fischer, xi + 228 pp. 

Brattstrom, Bayard H. 

1965. Body temperatures of reptiles. Amer. Midi. Nat., 73: 376-422. 

Davis, D. D. 

1953. Behavior of the lizard Corythophanes cristatus. Fieldiana: 
Zool., 35: 3-8. 



16 brfviora No. 316 

Dunn, E. R 

1944. [Tie lizard genus Phenacosaurus. ( aldasia, 3: 57-62. 

Etui riix.i . R 

1960. The relationships oi the anoles (Reptilia, Sauna, Iguanidae): 
An interpretation based on skeletal morphology. Ph.D. Thesis. 
Dept. of Zoology, Univ. of Michigan (Univ. Microfilms, Ine.. 
Ann \i boi . Mich. ). 

1967. Lizard caudal vertebrae Copeia, 1967:699-721. 

GARRIDO, ().. AND A. SCHW \R I / 

1968. Cuban lizards ol the genus ( hamaeleolis. Quart. Jour. Florida 

tcad Sci., 30: 197-220. 

GORM \N. ( , ( 

1967 Studies on the evolution and zoogeograph) of Anolis, Ph.D. 
rhesis, Depl of Biology, Harvard University 

GORM W I i ( . \M> I . \ IklNs 

1967. The relationships of tnolis of the roquet species group (Sauria 
Iguanidae). II. Comparative chromosome cytolog) Sysl /ool.. 
16: 137-143 

1968. New karyotypic data on 16 species of tnolis from Cuba, Ja- 
maica, and the ( ayman Islands Herpetologica, 24: 13-21. 

GORM VS. I i ( I \ I kiss. \\|> I HOLZINGER 

1967. New karyotypic data on 15 genera of lizards in the family 
Iguanidae, with a discussion of taxonomk and cytologies] im- 
plications ( Oogenetics, 6: 286 299 
Gorman, G. ( .1 Baptista, and R. B Hi ki 

19h l > ( hromOSOmes and SCeloporine relationships with special refer- 
ence to the horned lizards Mammalian ( hromosome Newslet- 
ter. 10 6-10. 

k \STLE, WLRM R 

1965. /in Ethologie des \nden- tnolis Phenacosaurus richteri. Zeit- 
schnft fur rierpsychologie, 22: 751-769. 
Lowe, C. H .. C. J Cole, and J. L. Patton 

1967. Karyotype evolution and speciation in lizards (genus Scelo- 

porus) during evolution of the North American desert. Syst. 
/ool. 16: 296-300. 

Noble, G. K 

1924. New lizards from northwestern Peru. Occ. Pap. Boston Soc. 
Nat. Hist.. 5: 107-113. 

Osorno-Mi sa, Hi rnando. ind Ernesto Osorno-Mesa 

1946. Anotaciones sobre lagartos del genero Phenacosaurus. Caldasia, 

4: 123-130. 

P.KRKhR. H. W. 

1932. Some new or rare reptiles and amphibians from southern Ecua- 
dor. Ann. Mag. Nat. Hist.. (10) 9: 21-26. 



1969 CYTOTAXONOMY OF IGUANIDS 17 

1935. The frogs, lizards and snakes of British Guiana. Proc. Zool. 

Soc. London. 1935: 505-530. 
1938. The vertical distribution of some reptiles and amphibians in 

southern Ecuador. Ann. Mag. Nat. Hist., (II) 2: 438-450. 

Pearson, Oliver P. 

1954. Habits of the lizard Liolaemus multiformis multiformis at high 
altitudes in southern Peru. Copeia, 1954: 111-116. 

Templeton, J. R. 

1967. Nasal salt gland excretion and adjustment to sodium loading 
in the lizard Ctenosaura pectinata. Copeia, 1967: 136-140. 

Wilson, E. O. 

1957. Behavior of the Cuban lizard Chamaeleolis chamaeleontid.es 
(Dumeril and Bibron) in captivity. Copeia, 1957: 145. 

(Received 5 November 1968.) 



BREVIORA 

Museum of Cooipsirative Zoology 

Cambridge, Mass. 30 April, 1969 Number 317 



RELATIONSHIPS OF TWO CRETACEOUS LIZARDS 
(SAURIA, TEIIDAE) 1 

Richard Estes 



Abstract. Peneteius aquilonius, n. gen., n. sp., from the late Cretaceous 
of Montana, is a small teiid lizard with complex cheek teeth. It most closely 
resembles the modern South American teiids Dicrodon and Teius and also 
shows resemblance to the large Cretaceous teiid Polyglyphanodon from 
Utah. Chamops segnis Marsh, from the late Cretaceous of Wyoming, re- 
sembles the modern teiid Callopistes mac u kit us. Meniscognathus altmani 
Estes, from the same locality as Chamops, may be related to the modern 
teiids Cnemidophorus and Kentropyx. 

These resemblances indicate that three distinct groups of primitive teiids 
living today in South America were in existence in North America during 
late Cretaceous time. 

INTRODUCTION 

In 1964 I described teiid lizards of modern type from the late 
Cretaceous Lance Formation of Wyoming. These included: (1) 
Chamops segnis, a species suggested to resemble the Recent 
Crocodilurus-Tupinamhis line in dental adaptations (see below), 
although it differed from all other so-called "macroteiids" (Van- 
zolini and Valencia, 1965) by having a parietal foramen; and (2) 
Meniscognathus altmani, which bore resemblances to the Ken- 
tropyx-Ameiva line in both dental apparatus and externally-con- 
cave mandibles. Two other forms, (3) Leptochamops denticulatus 
and (4) Haptosphenus placodon, were less clearly related to 
modern forms. 

Polyglyphanodon sternbergi, a large late Cretaceous lizard from 
Utah, was originally described by Gilmore (1940). In 1942 he 
placed it in a separate family Polyglyphanodontidae, but stated 

1 Fossil vertebrates from the late Cretaceous Hell Creek Formation, Mon- 
tana: Contribution No. 2 (Contrib. No. 1 is Estes, 1965, Copeia, No. 1, pp. 
90-95.) 



2 breviora No. 317 

that only tooth characters separated it from the Iguanidae. Hoff- 
stetter (1955) was the first to note the resemblance of Polygly- 
phanodon to the teiids and in 1962 suggested that it be referred to 
the Teiidae. A smaller, related genus Paraulxnhanodon was also 
named by Gilmore (1940, 1943). These animals are currently 
being restudied by Mr. William MacLean, 3rd. 

A vertebrate fauna recentl) recovered from the late Cretaceous 
IIJI Creek Formation, Montana, resembles that from the Lance 
Formation (Sloan and Van Valen, 1965; Estes, Berberian. and 
Meszoely, ms.). A single dentar) from the Hell Creek Formation 
sample belongs to a teiid lizard of unusual type and is described 
here. I interpret this fossil as in some ways intermediate between 
Polyglyphanodon and the modern genera Teius and Dicrodon, It 
is probably closer to the latter genera and provides an indication 
that a third major living "maeroteiid" line was already in ex- 
istence in late Cretaceous time in North America. 

ORDER SAURIA 

SUBORDER SCINCOMORPHA 

Family Teiidae 

Peneteius aquilonius, n. gen., n. sp. 

Holotype. MCZ (Museum o\' Comparative Zoology, Harvard 
University) 3612. fragmentary right dentar) with four complete 
teeth and the bases of four others (Fig. 1 ). 

Horizon and locality. West half section 9, T 22 N, R 43 E, 
McCone County, Montana; Hell Creek Formation. Collected by 
MCZ part) in 1964. 

Etymology. Latin, pent, almost; aquilonius, northern. 

Diagnosis. Differs from Recent Teius and Dicrodon in having 
tooth crests nearly transverse, but the lateral cusp anterior to 
medial cusp rather than posterior. Differs from fossil genera 
Paraglyphanodon and Polyglyphanodon in lacking transverse ex- 
pansion of tooth and in having external crests of principal cusp 
less well developed. 

Description. Jaw fragment relatively delicate; Meckelian groove 
wide, indicating a large splenial; bony separation between Mecke- 
lian canal and more lateral canal (for vascular and nervous struc- 
tures) set far medially, reducing depth of Meckelian groove. Teeth 
subacrodont, becoming molariform posteriorlv; most posterior 
(broken) tooth evidentlv the largest; tooth bases subcircular, 
relatively thin-walled; sulcus dentaUs absent. Tooth crowns antero- 
posteriorly compressed into crests; crests essentially transverse 



1969 



CRETACEOUS TEIID RELATIONSHIPS 



and formed by two main cusps, the labial one relatively the higher; 
both cusps connected by a transverse crest. Main cusps closer 
together and difference in height less pronounced in more anterior 
teeth, but cusp axis remains transverse; crests extending anteriorly 
and posteriorly from main cusps, forming slight basins on each 
side of crown; faint depressions present lingually on each side of 
main lingual cusp. 

Discussion. The widely-open Meckelian groove in combination 
with the unusual tooth crowns and heterodonty indicate relation- 
ship of Peneteius to the Teiidae. Closest resemblances within that 
family are to Recent Dicrodon and Teius from South America 










1mm 



Fig. I. Peneteius aquilonius, n. gen., n. sp., MCZ 36 12; a, labial, b, 
lingual, c, occlusal view of fragmentary left dentary. 



KRIVIORA 



NO. 3 1 7 



and the fossil Paraglyphanodon and Polyglyphanodon from the 
Cretaceous of Utah. Paraglyphanodon is the smaller of the two 
Eossil genera. Teeth oi the two described species /'. utahensis and 
P. gazini show a morphological series that could lead to the 
strongl) transverse, crested tooth condition seen in Polygtyphano- 
don, and it is possible that Paraglyphanodon is onl) a small in- 
dividual or the young of the former. The most complex and cuspate 
teeth in an\ Recent teiid occur in Teius teyou; Figure 2 shows 
occlusal views of teeth of pertinent living and fossil species. I he 








1 Lg 2 Crown patterns of tend teeth. All arc posterior teeth of letl 
dentaries. ./. hur»di>n guttulatum, MCZ 111415; />. /). heterolepis, M( / 
12329; c, Teius teyou cyanogaster, \l< Z J9982; d, I teyou, M( Z 43351b; 
e, Peneteius aquilonius, n. gen., n. sp.. MCZ 3612; f, Paraglyphanodon uta- 
hensis, United States National Museum 15668; i\ P. gazini, USNM 16580; 
h. Polyglyphanodon rternberghi, USNM 15477. a-g X about 16, h X 
about 9. 



two species of Dicrodon are quite distinct in tooth patterns and 
in man) other characteristics as well (Schmidt. 1957); Schmidt's 
species D. hohnbergi has been synonymized with P. guttulatum 
by Fugler ( 1967). The Recent species figured here all differ from 
Peneteius aquilonius in orientation of the two major cusps and in 
lacking a faint depression on the lingual side of the crown. The 
tooth pattern of Peneteius is distinct" from the Utah Cretaceous 
genera in lacking strongly-curved, prominent labial crests that may 
(Polyglyphanodon) or'may not (Paraglyphanodon) connect with 
the secondary cusp. Resemblances are shown to the modern genera 



1969 CRETACEOUS TEIID RELATIONSHIPS 5 

in the latter character, especially to Teius teyou and Dicrodon 
heterolepis. Additional similarities with the former are the ap- 
parently small number and relatively large size of the teeth. The 
basin-crest structure on tooth crowns of Recent species and 
Pene teius recalls anterior teeth of Paraglyphanodon (Gilmore, 
1942, fig. 22), but the similarity is not great. However, resem- 
blance to the Utah Cretaceous forms is shown in the more trans- 
verse rather than oblique orientation of the tooth cusps. Restudy 
of the Utah fossil forms must precede further speculation on the 
affinities of Peneteius. 

THE RELATIONSHIPS OF CHAMOPS 

Chamops segnis Marsh (1892) is relatively common for a late 
Cretaceous lizard and is known from the Lance Formation (Wyom- 
ing), Hell Creek Formation (Montana), and Wapiti Formation 
(Alberta). Estes (1964) noted that Chamops was "quite probably 
ancestral to both Crocodilurus and Tupinambis.'" Comparison of 
Chamops and Callopistes (a genus not available to me in 1964) 
requires some modification of that statement. Callopistes maculatus 
(MCZ 2751) is close to Chamops in several ways, principally in 
the relatively deep shape of the maxilla (Fig. 3c,d), the more 
normal (less conch-like, or curved) quadrate shape, and relatively 
elongate parietal (Estes, 1964, rig. 49). The parietals and 
quadrates are referred to Chamops on the basis of both size and 
frequency, as well as on their generally teiid appearance; Chamops 
is the largest and most common teiid in the Lance Formation. 

The tooth row of Callopistes maculatus is more heterodont 
than that of Chamops and fewer teeth are tricuspid; I interpret 
both heterodonty and bicuspid teeth as specialized features. How- 
ever, the nasal in Callopistes maculatus extends further forward 
on the maxilla than in Tupinambis and is thus more like the con- 
dition in Chamops (Estes, 1964: 107). The latter, Tupinambis, 
and C. maculatus share a pointed lateral premaxillary process of 
the maxilla (Fig. 3). The dentary of Chamops is relatively deeper 
than that of Tupinambis of equal size and resembles the propor- 
tions seen in C. maculatus. Tooth number is essentially the same 
in all these genera, contrary to my statement in 1964 (p. 107), 
which was based on only a few individuals. 

Facial elongation is characteristic of many "macroteiids" and 
is most extreme in Cnemidophorus. The latter does not differ from 
Ameiva in facial elongation, although I so stated in 1964 (p. 108); 
examination of a large series shows considerable size variation in 



BRLYIORA 



No. 317 




Fig. 3. a. Crocodilurtu lacertinus, American Museum of Natural His- 
tory 46290; />. Tupinambu nigropunctatus, Los Angeles County Museum 
R-74; c, Cluunops segnis. University of California Museum of Paleontology 
46033, restored dorsally from UCMP 46094 and other specimens; d, Cal- 
lopistes maculatus, MCZ 2751; not to scale, all reduced to a common length. 



1969 CRETACEOUS TEIID RELATIONSHIPS 7 

this character and the two genera are probably synonymous, as 
indicated by recent studies (Gorman, pers. comm. and ms. 1968). 
Tupinambis nigropunctatus and Callopistes maculatus of equal 
size show the former slightly exceeding the latter in facial length; 
Crocodilurus resembles the latter. Kentropyx calcaratus resembles 
Cnemidophorus in this feature; my statement to the contrary in 
1964 (p. 108) was based on a misidentified skeleton. 

In summary, the Recent Callopistes maculatus appears to be 
the closest relative of Chamops segnis. Tupinambis nigropunctatus 
is the most primitive member of that genus and is close to Chamops 
but appears more advanced than the latter and Callopistes in a 
number of features. The maxilla of Crocodilurus is relatively less 
high than that of Chamops and the former seems to be less closely 
related to the latter than it is to Tupinambis. 

CONCLUSIONS 

Current study of "macroteiids" by Gorman, Presch, MacLean, 
and myself is in general agreement with Vanzolini and Valencia 
(1965) in separating two major subgroups: one including Callo- 
pistes, Tupinambis, Crocodilurus, and Dracaena; the other formed 
of Ameiva, Cnemidophorus, Kentropyx, Teius, and Dicrodon. The 
latter two genera possess distinctive, crested, cuspate cheek teeth. 
Peneteius aquilonius, n. gen., n. sp., from the late Cretaceous of 
Montana, has similar teeth and is probably related to the Recent 
Teius-Dicrodon line. Chamops segnis, from the late Cretaceous 
of Wyoming, Montana, and Alberta, appears to be related to the 
Recent species Callopistes maculatus. The latter two species are 
probably more primitive, on the basis of high maxilla and less 
well-developed heterodonty, than are Tupinambis, Crocodilurus, 
or Dracaena. With the possible relationship noted above of the 
fossil genus Meniscognathus to the Ameiva-Kentropyx group, it 
thus appears that three distinct groups of "macroteiids" living 
today in South America were in existence in North America during 
late Cretaceous time. 

ACKNOWLEDGMENTS 

I am grateful to Mr. William P. MacLean, 3rd., Mr. William 
Presch, and Drs. Arthur C. Echternacht and George Gorman for 
comments. Figures 1 and 2 are by Mr. Laszlo Meszoly. This re- 
search was supported in part by National Science Foundation 
Grant GB-7176. 



8 breviora No. 317 

REFERENCES CITED 

ESTFS, R. 

1964. Fossil vertebrates from the late Cretaceous Lance Formation. 
eastern Wyoming. Univ. Calif. Publ. Geol. Sci . 49: 1-180. 

FUGLER. C. 

1967. Geographic variation in Dicrodon guttulatum Dumeril and 
Bibron of the Ecuadorian and Peruvian littoral. Journ. Alabama 
Acad Sci . :i«: 322. 

Gil MORI . ( 

1940. New fossil lizards from the Upper Cretaceous of Utah. Smith- 
sonian Misc. ( oil.. 99: 1-3. 

1942. Osteology oi Polyglyphanodon, an Upper Cretaceous lizard 
from l tan. Proc. I S Natl, Mus., H<»: 11-26. 

1943. Osteolog] Of Upper Cretaceous lizards from Utah, with a 
description of a new species. Proc. U. S. Natl. Mus., 93: 2<> l >- 
214 

Hoi i mi i ii r. R 

\ i > < * Squamates de i\pe moderne. /". J. Piveteau, ed.. Traiic ; de 

Paleontologie (Paris, Masson ci ( ie.), •>: 606-662. 
19h2 Revue des recentes aquisitions concernanl l*histoire el la 

sWcmatiquc des squamates. Colloq. Internat.. Prob. Pal. no. 
104: 243-2 

M \ksh. O. 

1892. Notice of new reptiles from the I aramie Formation. Amcr. 
Journ. Sci., (3) 13: 449-453. 

S( HMII>I . k 

1 1 >57. Notes on lizards of the genus Dicrodon. Fieldiana: Zool., 39: 

65-71. 

Sloan, R . \m> I Van Vai i N 

1965. Cretaceous mammals from Montana. Science. 118: 220-227. 

Vanzolini. P., and J. Vali n< i \ 

1965. The genus Dracaena, uith a brief consideration of macroteiid 
relationships (Sauria. leiidae). Arq. Zool.. 13: 7-35. 

(Received 5 February 1969.) 



BREVIORA 

Mmsemnti of Comparative Zoology 

Cambridge, Mass. 30 April, 1969 Number 318 



LEUCOLEPIDOPA SUNDA GEN. NOV., SP. NOV. 
(DECAPODA: ALBUNEIDAE), A NEW INDO-PACIFIC 

SAND CRAB 

Ian E. Eflford 1 



Abstract. Leucolepidopa gen. nov. is described along with the only 
known species in the genus, Leucolepidopa sunda sp. nov. The single 
specimen is from the Sunda Strait between Sumatra and Java. The posi- 
tion of this new genus is discussed and the relationships of all the genera 
within the family Albuneidae are reviewed. Leucolepidopa is most closely 
related to Austrolepidopa and Lepidopa. 

The Danish Kei Island Expedition collected a single specimen of a new 
sand crab during dredging operations in the Sunda Strait between Sumatra 
and Java. This new crab is from an undescribed genus closely related to 
Austrolepidopa, recently described from Australia (Efford and Haig, 1968), 
and to an entirely American genus, Lepidopa (Efford, in ms.). In this paper 
I describe the new species and present a preliminary outline of the probable 
evolutionary relationships within the family so that the position of the new 
genus can be seen. 

Leucolepidopa gen. nov. 

The type and only known species of this genus is Leucolepidopa 
sunda. This new genus is in the family Albuneidae and is closely 
related to Austrolepidopa and Lepidopa. It can be distinguished 
from other genera in the family by ( 1 ) the antennae having three 
articles in the flagellum and (2) the eyeplates possessing long 
setae on the dorsal surface. The following characters are not 
specific to the genus but can be used to separate it from others 
within the family: (3) a pleural expansion is present on the fifth 
abdominal segment; (4) the lateral spine is on the carapace and 
above the linea anomurica; (5) the scaphocerite is short, only just 



1 Department of Zoology, University of British Columbia, Vancouver 8, 
B. C, Canada. 



2 BREVIOR \ No. 3 IS 

extending up to the base of the fourth antenna! segment; (6) the 
rostrum is smoothly convex and lacks a subrostral spine; (7) 
there are no terminal spines on the anterolateral lobes; and (N) 
the anterolateral lobes cam six or seven teeth. 

The generic name refers to the whiteness of the carapace and 
to the close relationship oi this genus with the genera Lepidopa 
and Austrolepidopa. 

1 I l ( (H I 1MDOPA SUNDA SP. NOV. 

Holotype. A male. 7mm carapace length and 8mm carapace 
width (Fig. 1. 1-8, and Fig. :. 1-6). Collected on 29 July 1922 

from 75° 6' 10" S. 105° 44' E in the Sunda Strait between 
Sumatra and Java, bv the Danish Kei Island Expedition. The 
specimen was collected b\ dredging from a sand-shell bottom 
in 40 m o\ water. Deposited m the /oologisk Museum. Koben- 
havn. 

I he specific name sunda is derived from the name o\' the geo- 
graphical location. 

Description. The carapace has a rostrum that is gently rounded 
and lacks spines or teeth. It is Hanked on either side b\ a shallow. 
gently curved ocular sinus and by rounded anterolateral lobes, 
the edges o\ which cam six or seven teeth. From the antero- 
lateral lobes the anterior edge oi the carapace curves backwards 
and outwards in a smooth sigmoid curve to the large, distinct, 
lateral spines. I here are no spines along the anterior edge of 
the carapace as there are in most other members of the family. 
I lie edge is lined with long, branched setae, which gradually be- 
come shorter toward the lateral spines. From the lateral spines 
the carapace widens sligluK back to the midline, behind which 
the edges arc straight and converge. The posterior end of the 
carapace is smoothly rounded, with a deeply rounded medial con- 
cavity. 

The carapace has an extensive pattern of setae. The basic 
pattern is similar to that of both Austrolepidopa and Lepidopa; 
the main difference is that the M-shaped pattern across the an- 
terior end of the carapace extends up to the anterior edge and 
some way back along the sides. There is a smooth area in the 
middle of this pattern on either side. The setae in this area are 
longer than those on the posterior part of the carapace. Those 
lining the lateral spine groove are very long and branched, and 
extend beyond the anterior edge of the carapace. The posterior 
half of the carapace is almost equally setose, but here the setae 



1969 NEW INDO-PACIFIC SAND CRAB 3 

are in small groups rather than in one continuous expanse. In 
contrast, the posterior edge of the carapace, round the cavity, is 
free of setae and very smooth. The groove running down the 
side of the carapace stops before it reaches the posterior edge and 
does not follow the edge round as in Lepidopa. 

The carapace is chalky white with some iridescence. 

The first abdominal somite is shorter than wide, and the exposed 
posterior part forms a raised crescent-shaped area lined along 
its anterior edge by a dense row of setae. The second somite has 
broad pleural expansions with slightly concave anterior edges. 
The edges are lined with long setae that all face forward. Those 
along the lateral and posterior edges arise from a groove that runs 
round parallel with the edge. There are no setae along the edge 
of the central part of the plate. The third, fourth, and fifth seg- 
ments are also winged, but unlike those of Lepidopa, all the 
pleural expansions are rather straight, narrowing towards the ends, 
and both the anterior and posterior edges of the expansions have 
grooves lined with forward-projecting setae. The setae near the 
outside tips of the pleural expansions of the fifth segment are 
particularly long and, in this case, project laterally, away from 
the abdomen. One noticeable character is the presence of distinct 
bumps along the posterior edge of the fourth pleural expansion. 
The sixth segment is shorter than wide and narrows towards the 
anterior edge. It has a distinct waist, and the posterior edge is 
lined with backward-projecting setae. The telson is broad and 
pear-shaped. 

The eye-plate is quite distinct, as the dorsal surface is covered 
with long; setae. In Austrolepidopa and Lepidopa the upper sur- 
face of the plate is smooth, although there may be setae around 
the edges. The plate is broadest about one third from the proximal 
end. The inner edge is fairly straight and the outer edge convex. 
From the broadest point the eyes narrow towards the rounded 
distal end. The edge of the plate is stepped, and long setae are 
attached at each step. No cornea or eye-spot is visible. 

The antennules have a broad basal segment. The second seg- 
ment is broad, partiallv compressed, and has a large, right-angled 
bulge on one side. The third segment is about as long as the 
second and widens towards the distal end. The two first antennule 
flagella of the holotype have been broken off so that their length 
is unknown; however, the lower part of the first flagellum sug- 
gests that in this species they are very long, as in Lepidopa. No 
second flagellum is present on the antennule. 



4 breviora No. 318 

The antennae are unusual because the setae are arranged 
terminally or along the inner side of the segments and they are 
not found scattered around on other parts as they are in other 
genera. This gives the antennae a rather bare appearance. The 
scaphocerite is very small and hardly overlaps the base of the 
fourth segment. It has a group oi very long setae arising from 
the outer surface. The fifth segment is shorter than the fourth; 
it is cylindrical and terminates in a tlagellum oi three long cylindri- 
cal articles. The last article is longer than the other two. All 
three articles have areas of vet) long setae on their distal ends. 
In the holotype the setae are broken oil' the terminal article. 
I iie mandible is strong, smooth!) shaped, and carries a three- 
merited palp. There is a \er\ sharp cutting edge, two distinct 
teeth at the distal end. and three similar teeth at the proximal 
end. The maxillula was not examined. The maxilla is similar to 
those of others in the family, although I did not examine the 
scaphognathite. The firs! maxilliped is Battened and leaf-like; 
the endopod consists of two distinct segments. The second maxil- 
liped is similar to others found in the family; the exopod is single 
and well developed The third maxilliped has a very well- 
developed expansion o\' the antcrodoisal aid o^i the carpus, which 
extends over the propodus and reaches the junction oi the pro- 
podus with the dactylus. The dactylus is over half as long as the 
propodus. The exopod is a single, long, cylindrical segment. 

1 he pereiopods resemble those found in other members o\' the 
family. The first is subchelate with a sickle-shaped dactylus. The 
s.des oi the dactylus are smooth, and the upper surface is covered 
with long setae that point towards the proximal end of the seg- 
ment. The propodus has a distinct cutting edge lined with very 
short, blunt setae and terminating in a large, hooked tooth. The 
side is covered with rows of long setae. 1 he second and fourth 
pereiopods have broad, blunt, sickle-shaped dactyla. The third 
pereiopod has a long, smoothlv-curved dactylus without any dis- 
tinct basal projection; the carpus is very much expanded at the 
anterior end. with the anterior half of the upper surface covered 
with a dense mat of setae. The fifth pereiopod is long and chelate. 
The genital pore is round. 

Relationships. To examine the relationships of Leucolepidopa 
with the other members of the family, I used characters that seem 
to vary little between species within a genus, although they do 
vary between genera. These were: (1) the number of articles 
in the flagellum of the antenna; (2) the presence or absence of 
pleural expansions on the fifth abdominal segment; (3) the 



1969 NEW INDO-PACIFIC SAND CRAB 5 

presence of the lateral spine above or below the linea anomurica; 
(4) the length of the scaphocerite; and (5) the shape of the 
rostral region of the carapace. When the genera are examined 
using these characters, it becomes clear that some of the charac- 
ters always occur together. The presence of a pleural expansion 
on the fifth abdominal somite is always associated with a short 
scaphocerite and with a rostrum that is quite clearly convex in 
shape. The opposite three characters are associated with the 
absence of a pleural expansion on the fifth abdominal somite. 
These two groups can be used as the basis for separating the 
family into its evolutionary groups. I should mention here that 
Blepharipoda and Lophomastix are so distinct from the other 
genera in the family Albuneidae that they form a side branch off 
the main stream, and were not included in the analysis. 

The family can be divided into three groups (Fig. 3). The 
first, the Lepidopa group, contains Lepidopa, Austrolepidopa, and 
Leucolepidopa. Leucolepidopa is most clearly related to Austro- 
lepidopa, as shown by the similar shape of the eye-plates and by 
the stiff, straight, pleural expansions on the fifth abdominal somite 
compared to the rather delicate curved ones in Lepidopa. They 
differ in the length of the antennal flagellum, the presence of setae 
on the dorsal side of the eye-plate, the absence of a second 
flagellum to the antenna, and the setal pattern on the carapace. 
It is assumed that Leucolepidopa is more specialized than Austro- 
lepidopa, as it has only three articles in the flagellum, whereas 
both of the other genera have seven or eight. Austrolepidopa is 
closely related to Lepidopa, as is shown by the similar lamellated 
eye-plates, the antennal flagellum of seven or eight articles, as 
well as the elongated carpus of the third maxilliped, short scapho- 
cerite, pleural expansion, and convex rostrum common to the 
three species. 

A second group, the Albunea group, has only two species, 
Albunea and Stemenopa. The former genus occurs throughout 
the tropical regions and is rather generalized in form. Stemenopa, 
recorded from only one location in Western Australia, is a very 
specialized form with extremely long eye-stalks. It closely relates 
to Albunea, as is shown by the absence of a pleural expansion on 
the fifth segment, the concave rostrum, and the long scaphocerite. 

Zygopa is specialized in having both fused eye-plates and a 
single article in the antennal flagellum; it occupies a place some- 
where between the other two groups. According to Holthuis 
(1960), it is more closely related to the Lepidopa group because 
of the position of the lateral tooth. Possibly more weight should 



6 bri-viora No. 318 

be put on its similarity to the Albunea group in its long scapho- 
cerite, concave rostrum, and pleural expansion of the fifth ab- 
dominal segment, rather than its similarity to the Lepidopa group 
in having the lateral spine above the linea anomuriea. Until we 
have more information, we must assume that a closer relationship 
is suggested by three characters held in common than by only 
one. which places Zygopa closer to the Albunea group than to the 
Lepidopa group. 

The final group in the family is the rather separate off-shoot of 
Blepharipoda and Lophomastix. 

Apart from the position of Zygopa, the genera fall rather easily 
into these separate groups. These relationships could be tested, 
and more information obtained to confirm or reject the tentative 
position of Zygopa, b\ comparing the larvae. At the present time. 
however, we know the larvae only of Lepidopa and Blepharipoda, 
so that we are still some \\a\ from understanding the evolution of 
the family Albuneidae. 

ACKNOWLEDGMENTS 

I would like to thank Dr. I orben Wolff for allowing me to 
describe this specimen and Dr. Ernst Mayr, Dr. Herbert Levi, 
and Miss Alice Bliss for their help during my stay at the Museum 
ol Comparative Zoology. 

LITERATURE CITED 

Ej-kord, 1. E. 

(in ms. i. The species of sand crabs in the genus Lepidopa (Decapoda: 
Albuneidae i. 

I I I < »KI». I. E., AND J. HAIG 

1968. Two new genera and three new species of sand crabs (Decapo- 
da. Anomura. Albuneidae) from Australia. Aust. J. Zool., 16: 
897-914 

HOLTHl IS. L. B. 

I960. Notes on American Albuneidae (Crustacea, Decapoda, Ano- 
mura) with the description of a new genus and species. Proc. 
K. Nederiand Akad. Wet., Ser. C, 64 (1): 21-36. 

(Received 5 November 1968.) 



1969 



NEW INDO-PACIFIC SAND CRAB 




Fig. 1. Holotype of Leucolepidopa sunda gen. nov., sp. nov., male cl. 
7 mm. 1. Dorsal view of the eye-plates and carapace with the setae pat- 
tern shown. 2. Dorsal view of the abdomen and telson. 3. Left first pereio- 
pod. 4. Right third pereiopod. 5. Left fourth pereiopod. 6. Left fifth 
pereiopod showing round genital pore. 7. Left second pereiopod. Scale 
1 mm, except in 1 and 2, where it is 0.5 cm. 



8 



BREVIORA 



NO. 3 1 S 




Fig. 2. Holotvpe of Leucolepidopa sunda gen. now. sp. nov, 1. Base of 
antennule. 2. Dorsal view of left eye-plate. 3. Left antenna. 4. Left second 
maxilliped. 5. Left third maxilliped. 6. Left first maxilliped. 7. Left maxilla 
with the scaphognathite missing. S. Left mandible. Scale 1 mm. 



1969 



NEW INDO-PACIFIC SAND CRAB 



LEUCOLEPIDOPA 
(31 



5TEM0N0PA 
171 




ZYGOPA 
III 




SPINE B E 1 W LINEA 
ANOMURICA 



SPINE ABOVE LINEA 
ANOMURICA 




SPINE ABOVE LINEA 
ANOMURICA 



NO PLEURAL EXPANSIONS ON 5IH 
ABDOMINAL SEGMENT. ROSTRUM 
CONCAVE. SCAPHOCERITE LONC. 



PLEURAL EXPANSION ON 5TH 
ABDOMINAL SEGMENT. ROSTRUM 
CONVEX. SCAPHOCERITE SHORT. 



BIEPHARIPODA GROUP 
BLEPHARIPODA 
IOPHOMASTIX 




ANCESTRAL ALBUNEIDAE 




FAMILV HIPPIDAE 
HIPPA 
EMERITA 
MASTIGOCHEIRUS 



ANCESTRAL HIPPOIDEA 



Fig. 3. Evolutionary relationships within the family Albuneidae. Num- 
ber of articles on the antennal flagellum is given for each genus. 



BREVIORA 



Museumi of Comparative Zoology 

Cambridge, Mass. 30 April, 1969 Number 319 

COMPETITIVE EXCLUSION AMONG ANOLES (SAURIA: 
IGUANIDAE ON SMALL ISLANDS IN THE WEST INDIES' 

A. Stanley Rand 2 



Abstract. Each of the Greater Antilles has been a largely independent 
center of anole radiation and there has been little exchange of species 
between them. Together they are the source areas from which the other 
islands of the Antilles have been colonized. The area and number of anole 
species known from 122 of these smaller islands on 44 banks are tabulated. 

There are two kinds of small islands in the West Indies. The islands of 
one class show a correlation between area and number of anole species, 
and the larger islands may have up to 4 or 5 species. Islands of the other 
class do not show an area effect and never have more than 1 or 2 species. 

The first class includes the islands of the Great Bahama Bank and the 
islands fringing the Greater Antilles and lying on the same bank as one or 
another of them. The second class includes the Lesser Antilles and the 
small isolated islands of the Caribbean. 

The difference between the two classes seems to be a result of their Pleisto- 
cene history and the way in which they received their anole faunas. The 
islands of the first class were all connected or very close to a source area 
during the Pleistocene sea level minima and so probably received their 
anoles overland and more or less at the same time. The area effect now seen 
was probably produced by differential extinction subsequent to a sea level 
rise and separation of the various islands. The islands of the second class 
must all have been colonized over an appreciable water gap. 

The absence of islands with more than 1 or 2 species in this second 
class cannot be due to ecology, size, distance, or species of anoles involved, 
since some of the islands are large, diverse, and no further from a suitable 
source than some fringing islands are today. The data suggest that a first 
species finds it relatively easy to colonize an island. A second species finds 
it difficult, and a third may find it impossible, even though three species may 
coexist on a very small and ecologically impoverished island if they have 
colonized it overland. The exclusion effect occurs not because of the dif- 
ficulties in reaching the island, but because of the difficulties in establish- 
ment. The latter are the result of inter-specific competition in which an 

1 Research supported in part by NSF Grant GB-6944. 

- Smithsonian Tropical Research Institute, Balboa, Canal Zone. 



2 breviora No. 319 

alread) established species can exclude an invader even though if both 
were established together the) migl I coexist, it is suggested that competitive 

exclusion ma) also explain the small number of exchanges of species among 
the Greater Antilles. 

This paper deals with the factors influencing the number of 
species of Anolis per small island in the West Indies. Darlington 
(1957) relates fauna! size and complexity in the West Indies to 
island area and to distance from the postulated source area o\ 
Central America. However true these generalizations are for the 
fauna as a whole, they must he modified for the genus Anolis: 
first, because the West Indies have been a center o\' speciation 
and have not received their modern fauna directly from Central 
America and, second, because there is a class o\' small West 
Indian islands that have only one or two species i^\' Anolis regard- 
less of area or distance from source islands. The following dis- 
cussion o\' the distribution of a single genus in one area is an ap- 
proach to the theoretical problems o\' the zoogeography of archi- 
pelagos that I hope is complementary to the more mathematical 
treatments o\' Hamilton. Barth. and Rubinoff ( 1964), MacArthur 
and Wilson ( 1967). and others. 

I here are approximately 3.000 islands in the West Indies. Many 
of the islands stand on more or less extensive banks and are sepa- 
rated from one another or from one o\' the Greater Antilles by 
onl) shallow water. When the sea level was lowered during the 
last Pleistocene Emaciation, the banks must have largely been 
exposed. At that time the islands on each bank must have been 
connected to one another by dry land, so that an exchange of 
land faunas was possible. The anole faunas o{ the islands on 
each bank arc very similar to one another. The water between 
the banks is ver) deep, and there is no possibility that islands on 
different banks were connected during the Pleistocene. 

Duruig the interglacial period that preceded the last glaciation. 
the sea level was higher than it is at present, and the maximum 
probably was high enough to submerge completely many of the 
low-lying West Indian islands and exterminate any anoles living 
there. 

Anolis is represented in the West Indies by about 100 species, 
more species than are known for any other genus of land verte- 
brates in the area. These species may be the results of as few 
as two overwater invasions, one from Central America into 
Jamaica or Cuba and the other from South America into Puerto 
Rico or perhaps Hispaniola. Here and below the conclusions of 
Etheridge (I960) and Williams (personal communication) on 
the relationships within the genus Anolis have been followed. 



1969 COMPETITIVE EXCLUSION 3 

Each of the four Greater Antilles has been a largely independent 
center of evolution for anoles. Cuba has 23 species of Anolis, 
Hispaniola 21 species, Jamaica 7 species, and Puerto Rico 10 
species. Despite the fact that several of these species have crossed 
large water gaps to live on distant small islands, only one species 
is shared by two of the large islands. Even in the more distant 
past, the number of colonizations between the Greater Antilles 
was small. Only eight such events are necessary to explain the 
relationships between living forms: four between Puerto Rico 
and Hispaniola; two between Hispaniola and Cuba; two between 
Cuba and Jamaica. On the four large islands, speciation has been 
largely within and not between islands. 

The Greater Antilles, with their older and richer anole faunas, 
are probably the source areas from which the smaller (some only 
recently habitable) West Indian islands have been colonized. An 
anole invading one of the Greater Antilles from a small island 
is probably even rarer than the invasion of one of the Greater 
Antilles by an anole from another. 

Anoles have repeatedly crossed water gaps. They have not 
only reached and spread throughout the West Indies, but one 
West Indian species has reached the coast of Central America 
and another the Bay Islands of Honduras. It has been suggested 
that anoles may be blown from island to island in hurricanes or 
float across in rafts of vegetation carried by ocean currents. Per- 
haps they travel in the masses of debris and vegetation that are 
torn loose from a hurricane-lashed shore and blown out to sea. 
The heavy rains that accompany hurricanes would reduce the 
danger from salt water during the journey. 

On the assumption that the large islands have been the source 
areas, most but not all colonizations have been from east to west, 
which is the general direction of both ocean currents and prevailing 
winds, as well as the storm track of many hurricanes (see Fig. 1 ). 

Data on area and number of Anolis present have been collected 
for 126 small islands on 44 banks. Anole distributions from 
Barbour (1937), Cochran (1934, 1941), Grant (1932a, 1932b, 
1936, 1937), Grant and Roosevelt (1932), Oliver (1948), Ruibal 
(1964), Underwood and Williams (1959) have been supple- 
mented by the collections in the Museum of Comparative Zoology 
and unpublished information from the files of their curator. Dr. 
Ernest E. Williams. Mr. J. D. Lazell, Jr., and Dr. Albert Schwartz 
generously shared the results of their extensive field experience. 
Thanks are due to all three. Information on island areas was 
taken from the Columbia Lippincott Gazetteer of the World and 



imi \ loRA 



No. 3 1 9 




1969 COMPETITIVE EXCLUSION 5 

U. S. Hydrographic Office charts. The data are given in Table 1 . 
Almost all the major islands are included; the number of smaller 
ones could be increased greatly, but there is no suggestion that 
this would change the picture. 

The smaller islands of the West Indies fall into four geographical 
categories: (1) the islands fringing the Greater Antilles; (2) the 
Lesser Antilles; (3) the isolated islands of the Antilles; and (4) 
the islands of the Bahamas. Each of these groups shows a some- 
what different pattern of anole distribution. 

( 1 ) The fringing islands of the Greater Antilles stand on a 
bank with one or another of the four Greater Antilles and are 
separated from it by only shallow water. Most are close to the 
associated large island, but some of the Virgin Islands are almost 
100 miles from Puerto Rico. These islands vary ecologically from 
very simple to moderately complex. I have data on 44 islands in 
this group, all inhabited by anoles. 

The fringing islands show a definite area effect, i.e., size of 
island and number of species are directly correlated (Fig. 2). 
Many are low, and most must have been submerged during the 
last Pleistocene sea level maximum. During the last Pleistocene 
sea level minimum, these islands must have become hills on the 
coastal plains of the Greater Antilles and been inhabited by the 
common lowland species of Anolis. After the sea level rose, the 
larger islands (more than 25 square miles) kept their original 
three to five species, but on the smaller islands there appears to 
have been a tendency for species to become extinct. The number 
that remains is related in part to island area, and the very smallest 
islands (less than 1 square mile) frequently now have only a single 
species left. On none of these islands has there been any apprecia- 
ble differentiation except for the endemic, A. roosevelti, on Culebra. 

(2) The Lesser Antilles extend in a 500 mile arc from just 
east of the Virgin Islands south almost to Trinidad and Tobago. 
Many reach altitudes of several thousand feet and show a variety 
of climatically different habitats with rich floras. I have data on 
33 islands on 16 different banks, all inhabited by anoles. 

Comparing these islands among themselves there is no apparent 
area effect. Five of the banks have islands with two species, the 
islands on the remaining 1 1 banks have only a single species 
(excluding certain introductions by man). The Lesser Antilles 
have been colonized by two different stocks. One, the bimaculatus 
species group, including wattsi, has spread throughout the Northern 
Lesser Antilles, crossing at least nine water gaps. It probably 



6 BRLVIORA No. 319 

came from the Greater Antilles, though the ancestral species has 
either disappeared or changed so much that it is unrecognizable. 
The stock has differentiated on ever) island, and taxonomists 
differ on how many populations have reached the species level. 
On several islands these anoles show marked geographical varia- 
tion within the island ( l.a/cll. 19d2. 1964). Though the bimacula- 
tus group has been in the Lesser Antilles a long time, there seem 
to have been very lew cases of double invasions. On only the 
three northernmost banks o\o two species (one of them wattsi) 
occur together. 

The second stock, the roquet group, has invaded the Lesser 
Antilles from the south, crossing at least seven water gaps, but 
it is not known to be closel) related to an) living South American 
species. There are two banks on which two species of this group 
coexist. The degree of differentiation between islands is less in 
this group than in the bimaculatm group. 

(3) The isolated islands are those islands or small groups of 
islands scattered throughout the West Indies and separated from 
all other islands b\ \cr\ deep water. Most are low and quite 
simple ecologically. Some arc \cr\ far from any source area. 
Others are quite close. I have data on I I islands, on 9 banks, all 
inhabited In anoles. ( I here are also a vcr\ lew. very small. 
isolated cays [e.g. Pedro Cays, Aves Island] that are uninhabited; 
the) are the onl) islands in the West Indies known to lack anoles.) 

These islands show no area effect. One island has two species 
ol anoles. the others a single species. Most o[ these islands were 
probabl) submerged during the Pleistocene sea level maximum 
and colonized overwater since their emergence. All of these islands 
are occupied b) species that are coiispecilic with or clearly derived 
from a species now living on one of the Greater Antilles. The one 
exception is acutus on St. ( roix, probabl) derived from a Puerto 
Rican stock but not from one of the living species. 

(4) "The Bahamas are a large group of islands stretching for 
some six hundred miles in an elongate cluster from Florida, along 
the northeast coast oi Cuba, almost to Hispaniola. They are all 
low islands with little ecological diversity and were probably all 
submerged during the last Pleistocene sea level maximum. The 
islands lie on a number of banks o\ various sizes. I have data on 
53 islands on 15 banks, all inhabited by anoles. 

The largest bank is the Great Bahama Bank. During the 
Pleistocene sea level minimum, its many islands must all have 
been part of a single very large island lying next to Cuba. The 
present islands show an area effect very like that of the fringing 



1969 COMPETITIVE EXCLUSION 7 

islands. The larger islands on this bank are inhabited by four 
species of anoles, three from Cuba (sagrei, carolinensis , and 
angusticeps) and one from Hispaniola (distichus). These species 
must have reached the bank overwater and then probably spread 
through it overland. 

The remaining Bahamas lie on a number of smaller banks. Of 
the 14 banks for which I have data, the islands on 10 are occupied 
by a single species and on 4 by two species. Many of these islands 
are much larger and at least as varied ecologically as some of those 
on the Great Bahama Bank that are occupied by four species 
of anoles. 

The Bahamas on other banks that lie close to the Great Bahama 
Bank are occupied by one or two of the anoles that occur on 
the Great Bahama Bank. The Bahamas farther southeast are oc- 
cupied by a single endemic species, scriptus. The ancestors of 
scriptus may have reached the Bahamas in a single long overseas 
crossing from Puerto Rico, where its very close relative, cristatellus , 
occurs, though it may have come from a now extinct form on 
Hispaniola. A. scriptus does not occur with any other anole, 
though it lives on West Plana Cay only ten miles from East Plana 
Cay, which is occupied by A. carolinensis. 

From this survey it appears that, on the basis of the size of 
their anole faunas, there are two kinds of small islands in the 
West Indies. The first class of island includes the fringing islands 
of the Greater Antilles and the islands of the Great Bahama Bank. 
These clearly show an area effect and only the smallest have a 
single-species fauna (Fig. 2, upper). The second class of island 
includes all of the Bahamas off the Great Bahama Bank, the 
Lesser Antilles, and the isolated islands (Fig. 2, lower). These 
show little if any area effect, and even the largest islands have 
only one or two species. 

The difference in numbers of anoles between these two types of 
islands cannot be explained on the basis of differences in their 
ecological diversity or carrying capacity. The Lesser Antilles, with 
only one anole species, are larger, much richer, and more varied 
ecologically than are many fringing islands with three or four 
species of anoles. Within the Bahamas there seems to be no dif- 
ference in the ecology between the islands on the Great Bahama 
Bank, where four species coexist, and those on other banks, where 
only one or two species occur together. The difference cannot be 
in behavioral or ecological peculiarities of the species involved, for 
the species that replace one another in some of the Bahamas occur 



8 



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1969 COMPETITIVE EXCLUSION 



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10 BR] \ IORA No. 3 I 1 ' 

together on the Great Bahama Bank, rhe width of present-da) 

water gaps is another factor that will not explain the differences in 
number of species (Fig. 3). 

I he Factors that do seem important in causing the difference are 

reiated to the Pleistocene histor) of the islands and the manner in 
which the\ were colonized, either over land or over water. The 
fringing islands could, and probabh did, receive their several 
species of anoles over land, all more or less at the same time, dur- 
ing the per. od of lower sea level. Each speeies was represented 
b) mam individuals. I lie present islands of the Great Bahama 
Hank probabl) received their faunas in the same way, though the 
hank itself must have been colonized over water. I he isolated 
islands, the Lesser Antilles, and the Bahamas off the die, it Bahama 
Bank probabl) received their anoles over water. I he species prob- 
ably arrived separatel) and onl) one or a ver) few individuals 
reached an island at one tune. 

One speeies or another iA Anolis has succeeded in colonizing 
ever) West Indian island o\ more than a square mile in area, and 
almost all of the even smaller ones have been colonized, as well. 
It can be suggested that the reason so lew o! these islands have two 
speeies and none three is that the additional species have never 
readied them. But if the difficult) oi reaching the islands were 
the explanation, then one would find that the islands that wcie 
easier to reach, because the) were down wind, down current, or 
Close to the souree. would have more species than the islands that 
were harder to reach. I his is not the case. I he position of an 
island relative to the source area has more effect on what species 
will reach it than on how many. Figure 3 shows there is no simple 
correlation between number of species and distance from source 
area. 

I he limited anole faunas must result from interactions between 
speeies. Apparent!) a species established on an island makes it 
more dillicult for the second species that reaches the island to 
establish itself. If the second species is successful, it is even more 
difficult for additional species to establish. I would suggest that, 
because oi the low probability o\ overwater colonization of an 
island, the first species reaching it has time to occupy it completer) 
before an\ other species lands. The onl) island large enough and 
close enough to the souree areas for several species to colonize it 
before any one of them occupied it completely was the Pleistocene 
island that is now the (neat Bahama Bank. Once the four species 
were established, they were able to coexist, so that the modern 
islands show an area effect like fringing islands rather than like 



1969 COMPETITIVE EXCLUSION 11 

the other Bahamas. The phenomenon of an established species of 
Anolis preventing a newly arrived species from colonizing can be 
called competitive exclusion, because it must be the result of inter- 
specific competition. 

The four species that exciude one another on the Bahamas off 
the Great Bahama Bank were studied by Oliver (1948) on an 
island on the Great Bahama Bank where they occur together. He 
reports that they occupy different microhabitats. Similar differ- 
ences in microhabitats probably serve to reduce competition be- 
tween sympatric anoles on Puerto Rico (Rand, 1964) and Jamaica 
(Rand, 1967a). In both Puerto Rico and Jamaica, differences in 
microhabitat between some species are reinforced by interspecific 
territorial defense (Rand, 1964, 1967b). Where one of these 
species is abundant and another rare, all of the available territories 
in the preferred microhabitat of the abundant species may be oc- 
cupied by large adults, so that young individuals are forced to 
establish their territories in the microhabitat characteristic of the 
second species. These observations suggest that interspecific com- 
petition can occur even between species with different micro- 
habitat preferences, particularly whenever one of the species is 
very abundant. We do know that on small islands the species 
present may be there in large numbers. One species of Anolis 
that occurs without congeners, lineatus on Curacao, occupies a 
more varied microhabitat than do many populations of Greater 
Antillean species which occur sympatrically (Rand and Rand, 
1967). 

It is possible that successful colonizations of already inhabited 
islands were successful because the coionist landed at a time and 
in a place where the population density of established species was 
temporarily low. The larger and older the island and the more 
frequent the colonization attempts, the more likely it is that a 
colonist will land at a favorable time and place. This may 
explain the minor age, area, and position effects seen. 

The difficulty of colonizing an inhabited island accounts for 
the very small number of species occurring on many of the small 
West Indian islands. This phenomenon of competitive exclusion 
may also help to account for the relatively small amount of anole 
interchange among the Greater Antilles themselves and to explain 
why their anole radiations have been largely independent of one 
another. 



12 BREVIORA No. 31 ( ^ 

LITERATURE CITED 

B tRBOl R. T. 

1937. Third list of Antillean reptiles and amphibians. Bull. Mus 
Comp. Zool., 82:377-387. 
i i i hk\n. Doris 

1934. Herpetological collections from the West Indies made by Dr. 
Paul Bartsch under the Walter Ralhhone Bacon Scholarship. 
1928-1930. Smithsonian Misc. Coll., 92(7):l-48. 
1941. I he herpctolog) of Hispaniola. Bull. U. S. Nat. Mus., 177: 
i-vii, 1-398 
Darlington, P. J.. Jr. 

1957. Zoogeography: the Geographical Distribution of Animals 
New York. John Wile) and Sons. 675 pp. 
1 mi RIDOJ . Kit H\RD 

1960. The relationships oi the anoles (Reptilia:Sauria:Iguanidae). 
An interpretation based on skeletal morphology, Doctoral 
Dissertation. Univ. Mich.. University Microfilms. Ann Arbor, 
Michigan. 
Grant. C. 

1932a. I he berpetolog) of St. John and adjacent keys, l s. Virgin 

Islands. Jour. Dept. Agr. Puerto Rico. 16:331-334. 

1932b. Herpetolog) of rortola notes on Anegada and Virgin Gorda, 

British Virgin Islands. Jour. Dept. \;ji. Puerto Rico, 16:339- 
346. 

1936. rwo neu reptilian records From lost Van Dyke, B V. I., with 
notes on distribution and squamation. Copeia, 1936:9-14. 

1937. Heipetologic.il notes with new species from the American and 
British Virgin Islands, 1936. Jour. Agr. Univ. Puerto Rico, 
2\ :503-522. 

Gran i . C . IND ( . ROOSJ \ELT 

1932. I he herpetolog) of Caja de Mucrtos Island and ( aidona Key, 
Porto Rico. Jour. Dept. Agr Puerto Rico. l6(l):47-49. 

Hamilton, T. H.. R. h. Bar in. Jr.. and l ki binofi 

1964. The environmental control of insular variation in bird species 
abundance. Proc. Nat Vcad Sci I S., 52:132-140. 

1 \/i 1 1 . J. D.. Jr. 

1962. Geographic differentiation in Anolil OCltlatus on Dominica, 
pp. 466-475. In: J. D. Lazell, Jr.. and E. E. Williams. The 
anoles of the eastern Caribbean (Sauria: Iguanidae). Paris 
IV-VI. Bull. Mus. Comp. Zool.. 127(9 ) :451-478. 

1964. The anoles (Sauria; Iguanidae) of the Guadeloupeen Archipe- 
lago. Bull. Mus. Comp. Zool.. 1 31 ( I ) : 36 1 -401. 
Levins, R.. and H. Heatwole 

1963. On the distribution of organisms on islands. Carib. Jour. Sci.. 
3:173-177. 

Ma< \rihi r. R. H.. and E. O. Wilson 

1967. The Theory of Island Biogeography. Princeton. N. J., Prince- 
ton Univ. Press, 203 pp. 



1969 



COMPETITIVE EXCLUSION 



13 



Oliver, J. A. 

1948. The anoline lizards of Bimini, Bahamas. Am. Mus. Novit., 
No. 1383:1-36. 
Rand, A. S. 

1964. Ecological distribution in anoline lizards of Puerto Rico. 

Ecology, 45:745-752. 
1967a. The ecological distribution of the anoline lizards around 

Kingston, Jamaica. Breviora, No. 272:1-18. 
1967b. Ecology and social organization in the iguanid lizard, Anolis 
lineatopus. Proc. U. S. Nat. Mus., 122(3595) : 1-79. 
Rand, A. S., and P. J. Rand 

1967. Field notes on Anolis lineatus in Curacao. Studies on the 
Faunas of Curacao and Other Caribbean Islands, 24:112-117. 

RUIBAL, R. 

1964. An annotated checklist and key to the anoline lizards of Cuba. 
Bull. Mus. Comp. Zool., 130:473-520. 

SCHOENER, T. W. 

1967. The ecological significance of sexual dimorphism in size in the 
lizard Anolis conspersus. Science, 155(3761 ) :474-477. 
Underwood, G., and E. E. Williams 

1959. The anoline lizards of Jamaica. Bull. Inst. Jamaica Sci. Ser., 
No. 9:1-48. 

( Received 7 November 1968.) 

TABLE 1 

Numbers of Species of Anolis on Various Islands in the 

West Indies 



Island (those indented under 
another lie on the same bank) 

Islands fringing the Greater Antilles; 
44 islands on 4 banks. 
Cuba 

Isla de Pihos 
Hispaniola 

Gonave 

He Tortuc 

lie Vache 

Grand Cayemite 

Beata 

Saona 

Pte. Cayemite 
Jamaica 

Bogue 

Pigeon 

Salt 



Number of 


Island area 


A nolis species 


in sq. mi. 


23 


43.036.0 


8 


1.159.0 


21 


28,242.0 


5 


254.0 


3 


70.0 


3 


20.0 


3 


17.0 


2 


16.0 


3 


4.0 


1 


0.5 


7 


4,450.0 


1 


0.25 


1 


0.125 


1 


0.125 



14 


BRKVIORA 




NO. 319 


Island ( those indented 


under 


Number of 


Island area 


another lie on the same 


bank i 


Anolis species 


in sq. mi. 


Boob\ 




1 


0.062 


c abaritta 




— 


0.062 


1 merald 




1 


0.062 


1 mie 




1 


0.062 


Puerto Rico 




10 


3.421.0 


Vieques 




3 


5 1 .0 


St. Thomas 




3 


32.0 


i ot tola 




3 


24.0 


St. John 




1 


19.0 


\ne;:ada 




: 


15.0 


Cule 




4 


1 in 


\ n gin ( lorda 




3 


iDii 


lost \ .m Dyke 




: 


s II 


Petei 




3 


J.O 


Beef 




: 


2.0 


c in m.i 




\ 


2.0 


i ija Me Muertos 




2 


1.5 


Watei 




3 


1.0 


• per 




1 


1.0 


Savana 




2 


0.75 


Salt 




1 


0.75 


ii issel 




1 


0.5 


( ulebi ita 




1 


0.5 


I ii' ,n 




2 


0.5 


1 ittle St. James 




2 


0.5 


i Hen Jerusalem 




2 


0.5 


1 ittle Saba 




2 


ii 25 






2 


0.25 


< ockroach 




1 


0.25 


Dutchman's ( >p 




1 


0.25 


Dead Man's ( hest 




1 


0.25 


Sail Ke\ 




1 


0.25 


o Santiago 




3 


0.125 






1 


0.125 


1 esser Antilles: 33 islands 


on 16 banks 






Vnguilla 




2 


35.0 


St. Martin 




2 


35.0 


St Bartheln 




1 


9 s 


Vntigua 




2 


108.0 


Barbuda 




2 


62.0 


Saba 




1 


5.0 


St. Eustatius 




2 


7.7 


St. Kitts 




2 


68.0 


Nevis 




2 


50.0 


Redonda 




1 


0.5 


Montscrrat 




I 


38.0 



1969 



COMPETITIVE EXCLUSION 



15 



Island (those indented under 
another lie on the same bank) 

Desirade 

Guadeloupe 

Les Saintes, Terre de Bas 

Terre de Haut 
Marie-Galante 
Dominica 
Martinique 
St. Lucia 

Barbados 
St. Vincent 
Grenada 

Quatres 

Carriacou 

Mustique 

Petit Mustique 

Petit Nevis 

Baliceaux 

Petit Martinique 

Frigate 

Ronde 

Caille 

Diamond 
Isolated islands; 1 1 islands on 9 banks. 
St. Croix 

Buck 
Grand Cayman 
Mona 

Cayman Brae 
Little Cayman 
Navassa 
Swan (Larger) 
Swan (Smaller) 
Desecheo 
Alta Vela 
Bahamas; 53 islands on 15 banks. 
Great Bahama Bank (15 islands) 

Andros 

Eleuthera 

Cat Island 

Long Id 

Great Exuma 

Mangrove Cay 

New Providence 

Little San Salvador 

Stocking Id 



Number of 


Island area 


A nolis species 


in sq. mi. 


1 


10.5 


1 


687.0 


1 


3.0 


1 


2.5 


1 


58.0 


1 


305.0 


1 


427.0 


1 


233.0 


(2 introduced) 




1 


166.0 


2 


133.0 


2 


120.0 


1 


0.5 


2 


8.0 


1 


2.5 


1 


0.125 


1 


0.125 


1 


0.75 


1 


1.0 


1 


0.25 


1 


1.0 


1 


0.25 


1 


0.125 


1 


82.0 


1 


0.75 


1 


71.0 


1 


21.0 


1 


13.0 


2 


9.0 


1 


1.5 


1 


1.0 


1 


0.5 


1 


0.75 


1 


0.5 


4 


500.0 


4 


164.0 


4 


160.0 


4 


130.0 


4 


108.0 


3 


85.0 


4 


58.0 


1 


4.25 


1 


4.0 



16 



» BRFVIORA 




NO. 319 


Island (those indented under 


Number of 


Island area 


another lie on the same bank ) 


Anal is species 


in sq. mi. 


South Bimini 


4 


3.0 


North Cat Caj 


: 


0.75 


Margaret Island, Ragged Island Group 




0.75 


Flamingo Cay 




0.5 


Knife Cay 




0.062 


( io.it Caj 




0.062 


\cklins 




185.0 


Crooked Id 




76.0 


Long Cay 




7.0 


Castle Kland 




1.5 


Cotton Caj 




0.5 


Atwood Caj 




10.0 


( Irand ( aicos 




40.0 


North Caicos 




40.0 


Providenciales 




30.0 


1 ast c aicos 




300 


so ith ( aicos 




8.0 


West ( laicos 




7.0 


Pine Caj 




1.0 


< otton c aj 




2.0 


Elbow i 




1.0 


Conception 




4.0 


1 ast Plana ( B) 




3.0 


Great Inagua 




560.0 


Sheep ( a) 




0.25 


1 ittle Baham i Bank 






Gr. Ahaco hi 




776.0 


1 i and Bahama 




430.0 


1 ittle Ahaco 




27.0 


Elbow 1 




2.0 


Pcnsacola ( B 




1.0 


Water ( a\ . Grand Bahama 




1.0 


Strangers ( aj 




0.75 


Moraine Caj 




0.062 


I ittle Inagua 




45.0 


Mavaguana Id 




96.0 


Booby Caj 




0.25 


Rum Cay 


2 


29.0 


Grand Turk 




8.0 


Salt Ca\ 




2.5 


Cotton Ca\ 




0.5 


I .one Cay 




0.25 


Great Sand Cay 




0.25 


Watlings Id 


2 


60.0 


West Plana Cav 


1 


3.5 



BREVIORA 



Museum of Comparative Zoology 

Cambridge, Mass. 10 June, 1969 Number 320 

TAIMANAWA, A NEW GENUS OF BRISSID ECHINOIDS 

FROM THE TERTIARY AND RECENT INDO-WEST-PACIFIC 

WITH A REVIEW OF THE RELATED GENERA 

BRISSOPATAGUS AND GILLECHINUS 

Robert A. Henderson and H. Barraclough Fell 



Abstract. Taimanawa gen. nov. is erected to include two New Zealand 
fossil species, T. pulchella sp. nov. (taken as the type species) of early 
Miocene age and T. greyi (Hutton) of Oligocene age, as well as a poorly 
known extant species from the Kei Islands, Indonesia, T. mortenseni sp. 
nov., which had hitherto been tentatively referred to Plagiobrissus Pomel. In 
spite of possessing an internal fasciole, Taimanawa is referred to the Bris- 
sidae, with which it agrees in other morphological features. The closely 
related genera Gillechinus Fell and Brissopatagus Cotteau are re-evaluated 
using type material of the type species; they are maintained as distinct, but 
most of the species previously referred to Brissopatagus are here transferred 
to Gillechinus. 

INTRODUCTION 

In the course of a preliminary survey of the New Zealand Ter- 
tiary Spatangoida by one of us (H. B. F., 1947-55), the genus 
herein named Taimanawa was recognized as new. However, the 
two fossil species referable to the genus seemed to be congeneric 
with an unnamed and poorly known form from the Kei Islands, 
west of New Guinea, which had tentatively been referred to Plagio- 
brissus by Mortensen (1951). Therefore, it was deemed advis- 
able to defer publication of the material until the fossils could be 
compared with the suspected living species. In the interim, the 
responsibility for describing the entire New Zealand collection of 
Tertiary spatangoids has been assumed by the second author. In 
consequence, this report has been prepared as a joint project. 

Through the kindness of Dr. F. Jensenius Madsen (University 
Zoological Museum, Copenhagen) it has been possible to study 
fragments of two specimens from the Kei Islands and to confirm 



2 BREVIORA No. 320 

that the) represent a living species of Taimanawa. I his species, 
together with /. pulchella (the type of the genus) and /. greyi 
(Hutton), which was described from New Zealand a centur) ago, 

are all treated here. I wo additional species occur in the New 
Zealand Tertiary, Inn as the) add little to the definition of the 
izenus. the) will he described in a monograph of New Zealand 
Tertian Spatangoida now in preparation. 

I he closest relatives o\ taimanawa are the genus Gillechinus 
Fell, 1964, based on G. cudmorei Fell, and a number of species 
which previousl) have been referred to Brissopatagm Cotteau, 
1863. Brissopatagm is based on li. caumonti Cotteau. which is 
known from a single. poorl\ -preserved specimen. Recently Phillip 
i 1966) has synonymized Gillechinus with Brissopatagus. In order 
to evaluate this contention and to determine the relationship o\ 
Brissopatagus and Gillechinus to Taimanawa, type material of li. 
caumonti and G. cudmorei have been compared. Paratypes oi the 
forme: were supplied In Mi I V Darragh (National Museum o\ 
Victoria), and the holotype o\ the latter was kindl) lent to us b) 
Professor J. Roger (Laborator) o\ Paleontology, University of 
Paris). \s .i result o\ this study, Brissopatagus and Gillechinus are 
here maintained as distinct, and most o\' the species previousl) 
referred to Brissopatagus have been regrouped in Gillechinus. Tai- 
manawa is perhaps the direct descendant of Gillechinus; it is less 
closel) related to Brissopatagus. 

I he authors wish to thank Dr. ( \ Fleming and Mr. I. \\ 
Keyes (New Zealand Geological Survey), Mr. .1. D. Campbell 
(Otago University), Mr. J. A Grant-Mackie (Auckland Univer 
sit\). Mr. D. R. Gregg (Canterbur) Museum), and Dr. P. Kier 
(Smithsonian Institution) for the loan o\ specimens under their 
charge, and Mr. R .1. hosier (Hughesdale, Victoria, Australia) for 
a gift ot specimens of Gillechinus cudmorei. The project was under- 
taken during tenure at Harvard Universit) o\ a Frank Knox 
Memorial Fellowship and a Taranaki Post-Doctoral Fellowship by 
one of us ( R. A. H.). The research was sponsored by a National 
Science Foundation grant, GB-3532. 

Repositories. The repositories iA type and other specimens are 
indicated by the following abbreviations: New Zealand Geological 
Survey ( F.C. ): Geolog) Department. Otago Universit) (OU.); 
Geology Department. Auckland University (E.): and Canterbury 
Museum (zfe. ). 

Localities. Localities of the New Zealand fossil specimens have 
been arranged in order of the sheet districts of the one mile to the 



1969 INDO-WEST-PACIFIC ECHINOIDS 3 

inch topographic map series (New Zealand Department of Lands 
and Survey, series N.Z.M.S. 1 ) on which they appear. The sheet 
districts are designated "N" and "S" for the North and South 
Islands and are arranged numerically from north to south on each 
island. Sheet Fossil Numbers of the New Zealand Fossil Record 
Form System (e.g. SI 64/496) are recorded for those localities to 
which they have been allocated. 

Stratigraphy. The sequence of New Zealand Tertiary stage di- 
visions relevant to the stratigraphic ranges of T. pulchella and T. 
greyi, listed in order from youngest to oldest, are as follows: 
Whaingaroan, Duntroonian (Oligocene); Waitakian, Otaian, Awa- 
moan (Lower Miocene). 

Order Sparangoida Claus, 1 876 

Family Brissidae Gray, 1855 

Genus TAIMANAWA nov. 

Type species. T. pulchella sp. nov. Other included species: T. 
mortenseni sp. nov.; T. greyi (Hutton). 

Diagnosis. Large, flattened forms with peripetalous, subanal, 
and internal fascioles. Frontal ambulacrum deeply depressed, cross- 
ing the ambitus in a deep, narrow frontal notch. Paired petals long, 
narrow, shallowly depressed, and with subparallel pore series. An- 
terior pair widely splayed, posterior pair less so. Primary tubercles 
crenulate, restricted to the posterior plate series of the paired inter- 
ambs within the peripetalous fasciole. Labrum very short, extend- 
ing only halfway along the first plate of the adjacent ambulacra. 

Etymology. The name Taimanawa is derived from the Maori 
words tai (= sea) and nianawa (= heart); it is here treated as a 
Latin feminine first declension noun. 

Description. Test large, flattened, cordiform in outline, and al- 
most as wide as long. Aboral surface gently convex, oral surface 
flattened, ambitus sharp. Apical system at or near the axis of maxi- 
mum height, usually slightly pre-central and ethmolytic, with four 
gonopores. Peristome crescentic, labiate, and anterior; periproct 
situated on the truncated posterior test surface. Well-developed 
peripetalous and subanal fascioles and a clear but less well-devel- 
oped internal fasciole. Neither the peripetalous nor the internal 
fasciole continuous across the frontal ambulacrum. Both primary 
and secondary tubercles perforate and crenulate. 

Aboral surface: Frontal ambulacrum apetaloid, deeply depressed, 
crossing the ambitus in a narrow, deep frontal notch; ornamented 



4 BRE\ [OR \ No. 320 

with fine, dense miliary tubercles and perforated by minute, conju- 
gate pore pairs. Paired petals long, narrow, and weakly depressed. 
with subparallel pore series. Anterior petals widely splayed, at an 
angle oi approximatel) 120 : posterior petals less so. tunning an 
angle of approximate!) ("»<> . Each pore pair weakly conjugate and 
separate.! from the next In a single row of miliary tubercles. Inter- 
ambs with a broad, low ridge extending along or near the median 
suture oi the plate series; ridges oi the paired interamhulaera sub- 
tend gently sloping, flattened surfaces posteriorly to the depression 
ol each petal. Posterior plate series of the paired interamhulaera 
bear prominent fields of primar\ tubercles located within the peri- 
petalous fasciole. Tubercles surrounded by large, shallow scrobi- 
cules; those oi each field generall) arranged in a reticulate pattern 
with discernible rows both parallel and perpendicular to the axis 
oi the adjacent petal. Median suture of the posterior interamh 
bordered on each side by a single or double row of well-spaced, 
coarse secondary tubercles. Haeh interamhulaera] plate of the two 
series bordering the frontal ambulacrum bears ;i small triangular 
field iA course secondary tubercles. I he remaining intcrambulacral 
areas of the aborul surface bear tine, dense seconders and miliary 
tubercles. 

Oral surface: Plastron amphisternous, weakl) keeled, and ex- 
tending anteriorl) almost to the peristome; it bears coarse, dense 
secondary tubercles arranged in lines radiating from the posterior 
oi the keel. Subanal fasciole broad and rcniform. enclosing ;i 
densely tuberculate area with an arcuate row ol conjugate pore 
pairs on each side Labium \er\ short, extending only halfway 
along the first plate oi the adjacent ambulacra, twice as wide as 
long, and with a few secondary tubercles. Remaining interambu- 
laeral areas bear well-spaced secondary tubercles. Ambulacra! 
areas not depressed, naked except for scattered miliary tubercles; 
those margining the plastron are conspicuous, being half as wide as 
the plastron itself. Secondary spines bear prominent longitudinal 
ribs and are akin to those oi most other spatangoids. 

Remarks. Judging from the faseioles alone. Taimanawa would 
be placed in the Loveniidae alongside Breynia, which is the onl) 
other spatangoid genus with peripetalous, subanal. and internal 
faseioles. However, its other characters show that it is unrelated to 
Breynia and that it cannot be referred to the Loveniidae in spite of 
possessing an internal fasciole. In general morphology it is very 
similar to the undisputed brissid genera Gillechinus, Plagiobrissus, 
Brissopatagus, Macropneustes, and Eupatagus. Its long, narrow 
petals with subparallel pore series, short labrum. erenulate primar\ 



1969 INDO-WEST-PACIFIC ECHINOIDS 5 

and secondary tubercles, and the absence of ampullae set it apart 
from the loveniids and necessitate its inclusion in the Brissidae. 

Taimanawa, by virtue of its internal fasciole, is unique among 
the Brissidae. Its nearest relative is Gillechinus, which also has 
primary tubercles restricted to the posterior plate series of the paired 
interambulacra within the peripetalous fasciole and which has 
similar petals. Most members of Gillechinus possess flattened in- 
terambulacral surfaces subtended posteriorly from ridges of the 
paired interambs into confluence with the depressions of the paired 
petals, as in Taimanawa. However, Taimanawa, in addition to 
possessing an internal fasciole, is distinguished by its shorter labrum 
(Fig. 1 ), its much more deeply depressed frontal ambulacrum, and 
its much stronger frontal notch. It is interesting to note that the 
earliest species of Taimanawa, from the late Eocene of New Zea- 
land (as yet undescribed), resembles contemporaneous Gillechinus 
in possessing a slightly depressed frontal ambulacrum and a rela- 
tively weak frontal notch. Such a relationship strongly suggests 
that Taimanawa was derived directly from Gillechinus. The genera 
Eupatagus, Gymnopatagus, Macropneustes, Plagiobrissus, and 
Brissopatagus all bear some resemblance to Taimanawa. Besides 
lacking an internal fasciole, they are distinguished by possessing 
primary tubercles on both plate series of at least the posterior paired 
interambs (with the exception of E. ibericus Lambert). Eupatagus, 
Plagiobrissus, and Brissopatagus possess a weak frontal groove and 
frontal notch, and even in Macropneustes and Gymnopatagus, 
where these structures are better developed, they are not as strong 
as in Taimanawa. Eupatagus and Gymnopatagus are further dis- 
tinguished by possessing spatangiform petals with curved pore 
series, Plagiobrissus by possessing lateral branches of the subanal 
fasciole, and Brissopatagus by the concave depressions of its an- 
terior paired interambulacra. 

The presence of an internal fasciole in Taimanawa. was initially 
a perplexing feature, since this structure has not previously been 
reported from any member of the Brissidae and, indeed, Morten- 
sen's (1951) diagnosis of the family excludes forms with an in- 
ternal fasciole. However, it will be recalled that a similar perplexity 
arose in reviewing New Zealand species of the spatangid genus 
Paramaretia (Fell, 1963), though in this case the fasciole proves 
to be a transitory feature that disappears in the adult. The rela- 
tionship of fascioles to environmental features and the loss or ac- 
quisition of fascioles during the life-span of certain spatangoid 
echinoids have recently been studied by R. H. Chesher (private 



(-, KK 1 \ lORA No. 320 

communication), and the whole subject has now been considerably 
clarified b) Ins recent publication ( 1968). 

Most of the New Zealand Tertiai \ specimens of Taimanawa are 
from detrital limestones and from sandstones that were deposited in 
comparatively shallow water during the early and mid- Tertian 
transgression of the New Zealand region. In contrast, the extant 
species is recorded by Mortensen ( 1951 ) from two deep-water sta- 
tions (260 and 268 meters); the sediment type is known lor onl\ 
o\w of the stations and is recorded In Mortensen ( 1 c > 2 3 ) as mud 
and shells. I he bottom temperature of the two stations is not 
known but is probabl) near 20 (' and therefore some s -lo C 
higher than the mean annual temperature of surface water oil the 
southeast coast of the South Island. New Zealand, where Taiman- 
awa was abundant in OligOCene and earliest Miocene time. Such a 
relationship is consistent with the higher temperatures interred for 
the mid-Tertiar) marine environment of the New Zealand region 
on the basis ol U'smI echmoids ( I ell. 1956), and b) Bcu ( 1966) 
and others on the basis of Other fossil groups 

KEY TO SPECIES OF TAIMANAWA 

I (2i( renulation ol the primary tubercles strongl) developed, contiguous 

with the mamelon, the platform therefore aborted ; 

2(1) Crenulation of the primar) tubercles restricted to the peripher) of the 

platform, widel) separated from the mamelon mortenseni 

) (4) 23-40 primary tubercles in each interambulacral field, 22-2^ well 

developed pore pairs in the anterioi petals K" w 

4(3) 12-16 primar] tubercles in each interambulacral field, 18-19 well 

developed pore pain in the anterior petals pulchella 

TAIMANAWA PULCHELLA sp. nov. 
Plate 2, fig. 2; Plate 3; Plate 4. Figures 1 , 2a, 2c. 

Holotype. OU. 8590, Karitane, Otago; SI 55/538. 

Paratypes. S75: E. 310. Burnt Hill. Trelissick Basin. mid-Canter- 
bury; S75 517. SI 27: EC. 405, Ngapara, south Canterbury; SI 27/ 
561. S155: OU. 8540, Puketeraki, Dunedin district. Otago'. E. 302. 
304, Waikouaiti North Head. Matanaka Beach. Otago: SI 55 540. 
S163: EC. 42h. Blackhead. Dunedin district. Otago; SI 63/495. 
OU. 4736. 8566a, b, 8567b-d, 8568a-d, Blackhead. Dunedin dis- 
trict. Otago. SI 64: EC. 437. Green Island district. Otago; SI 64/ 
496. OL I. 8585, New Zealand Cement Co. pit, Burnside. Otago. 
Locality unknown: EC. 67 I . 



969 



INDO-WEST-PACIFIC ECHINOIDS 








Figure 1 . Comparison of the peristomial regions of Gillechinus and Tai- 
manawa. a) Gillechinus alabamensis (Cooke), U.S. National Museum 
562467; x3.5. b) Gillechinus cudmorei Fell, Museum of Comparative Zo- 
ology 4147; X3.5. c) Taimanawa pulchella sp. nov., holotype, OU. 8590; 
X2.5. 



8 BREVIOR \ No. 320 

Diagnosis. Adult tests compressed, with a moderately depressed 
Frontal ambulacrum and a strong frontal notch; 18-19 well-devel- 
oped pore pairs in the anterior paired petals. 12-16 primary tubercles 

in each lateral intcrambulacrum. 

Description, lest up to 120 mm in length, cordiform in outline, 
almost as wide as long, with a sharp ambitus. Aboral surface 
shallowly arched, oral surface flattened; height breadth ratio ap- 
proximately o.27. Apical system central to slightly anterior in 
position. King on the axis of maximum height of the test and 
morphologically typical of the genus. Peristome crescentic, an- 
terior, and margined b\ a prominent lip. IVriproel lies on the 
truncated posterior surface of the test; no specimen is sufficiently 
well preserved to show its shape. Peripetalous fasciole thin, not 
indented between petals. Internal lasciole thin, of variable width, 
located near the median suture of the anterior mterambs and form- 
ing a tight loop posteriori) around the apical sWem. Neither the 
peripetalous nor internal fasciole is continuous across the frontal 
ambulacrum. Subanal fasciole well developed and reniform. 

\boial surface: Frontal ambulacrum of moderate depression; 
frontal notch prominent, as wide as deep in adult tests. Paired 
pel. ils straight, lightly depressed; anterior pair generally slightly 
longer than those posterior. Pore pairs weakly conjugate and o\ate. 
Adult tests have IN-I C > pore pairs in the anterior petals and 15-16 
in those posterior. Primary tubercles restricted to the posterior plate- 
series of lateral intcrambs within the peripetalous fasciole. Adult 
tests with 12-lh tubercles m each such interambulacral field. Other 
features ol the aboral surface typical Of the genus. 

Oral surface: Labium very short, reaching only halfway along 

the first plate of the adjacent ambulacra. I here are eight pores m 
side the subanal lasciole. arranged in two lines of lour parallel to 
the lateral portions of the fasciole. In other characters the oral 
surface is characteristic o\ the genus. 

Remarks. The most distal row { )\ primary tubercles in the inter- 
ambulacral fields, .is in the other species i)\ Taimanawa, is gen- 
erally Hush with the distal end of the adjacent petal. I he anterior 
intcrambs of OU. 8540, however, have two additional rows o\ 
tubercles nearer the ambitus than the distal tip <^ the adjacent petal. 
Ljoing an unusually high number o\ tubercles (22) compared to 
that oi more normal specimens ( 12-16). One specimen (EC. 671 ) 
has a conspicuous boring perforating the test at its anterior margin. 
evidently the result oi an attack by a large, predators gastropod. 

. Lower Miocene; Waitakian — Otaian. OU. 8585 is known 



1969 INDO-WEST-PAC1FIC ECHINOIDS 9 

to be of Waitakian age and EC. 405 has the age limits of Dun- 
troonian — Waitakian. The remaining specimens, with the excep- 
tion of E. 310, are from the Caversham Sandstone, which is largely 
of Otaian Age although Awamoan microfaunas are known from its 
uppermost horizons. The holotype is known to be of Otaian Age, 
and the other specimens from the Caversham Sandstone are most 
likely of Otaian Age also. 

TAIMANAWA GREYI (Hutton) 
Plate 1 ; Plate 2, fig. 1 . Figure 2d. 

Brissus greyi Hector, 1870: 192 {nomen nudum). 
Eupatagus greyi; Hutton, 1873: 41. 

Holotype. EC. 682, Cobden, north Westland; S44. 

Additional material. N51 : E. 296, near Waikawau Stream mouth, 
west Auckland; N5 1/677. S37: EC. 480, 481, Woodpecker Bay, 
north Westland; S37/528. S44: EC. 458, Cobden Limestone, Grey- 
mouth, north Westland; S44/465. EC. 543, Cobden Limestone 
Quarries, Greymouth, north Westland; S44/476. zfe. 286, 287, 
290, Cobden Limestone, Greymouth, north Westland. S102: EC. 
407, Kakahu River, south Canterbury; S102/1. OU. 8548b, Hang- 
ing Rock, Opihi River, south Canterbury. Sill: E. 250, Lower 
Pareora Gorge, south Canterbury; SI 11/683. OU. 8606, Gordon 
Valley, south Canterbury, zfe. 284, Pareora Gorge, south Canter- 
bury. S127: EC. 615, Pigeon Rock, Waitaki Valley, south Canter- 
bury; SI 27/37 1. EC. 473, 474, Waihao River, south Canterbury; 
SI 27/376. EC. 472, Waitaki Valley, south Canterbury; SI 27/447. 
EC. 471, Awamoko Creek, Oamaru, south Canterbury; SI 27/450. 
E. 246, 248, Taylor's Road, Oamaru, south Canterbury; S127/639. 
OU. 8546, 8547, Waitaki Limestone, Duntroon, south Canterbury. 
OU. 8542, Ngapara, south Canterbury. OU. 8598, 8601, 8602. 
Earthquakes, Duntroon, south Canterbury. S136: OU. 8545, 
Weston, south Canterbury. Locality unknown: zfe. 278, Canter- 
bury, unlocalised. 

Diagnosis. Adult tests compressed, with a deeply depressed fron- 
tal ambulacrum, 22-25 well-developed pore pairs in the anterior 
petals, and 23-40 primary tubercles in each interambulacral field. 

Description. Test up to 1 34 mm in length, cordiform in outline, 
almost as wide as long, with a sharp ambitus. Aboral surface 
shallowly arched, oral surface flattened; height/breadth ratio 
approximately 0.25. Apical system varies in position from slightly 



10 



BR F MORA 



No. 320 



anterior to slightl) posterior oi center; it lies on the axis i>t maxi- 
mum test height, and its morpholog) is typical o\ the genus. Peri- 
stome crescendo, anterior, and margined by a prominent lip. Anal 
region not preserved on any available specimens. Peripetalous 

fasciole thin, not Strongl) indented between the petals. Internal 
fastiole extending lor a short distance near the sutures of the two 
plate series of interamhs 2 and 3 and forming i tight loop pos- 
tcriork around the apical system. Neither the internal nor the 
peripetalous fasciole closed across the frontal ambulacrum. 









t 



 

ft --- 



ft*^ 



\ 




a 




j -> - 



*&%* 



'1? 














I igure 2 a) Primar) tubercles of Taimanawa pukhella sp. run., holotype, 
OU. 8590; x7. b) Primary tubercles of Taimanawa mortenseni, holotvpe: 
X7. c) Plates of ambulacrum V of Taimanawa pulchella sp. nov.. holotype, 
OU. 8590; x7. d) Apical system of Taimanawa greyi (Hutton). OU. 8598; 
X7. 



1969 INDO-WEST-PACIFIC ECHINOIDS 11 

Aboral surface: Frontal ambulacrum deeply depressed, termi- 
nating in a conspicuous frontal notch that is as wide as deep on 
adult tests. Paired petals lightly depressed; anterior pair straight 
or gently curved towards the frontal ambulacrum and generally 
slightly longer than the posterior pair, which are straight or gently 
curved away from the posterior interamb. Pore pairs ovate and 
weakly conjugate; adult tests have 23-25 well-developed pore pairs 
in each anterior petal and 20-24 in each posterior petal. Fields of 
primary tubercles restricted to posterior plate series of the lateral 
interambs inside the peripetalous fasciole; 23-40 tubercles in each 
field on adult tests. Other features of the aboral surface as de- 
scribed for the genus. 

Oral surface: As described for the genus. The anal region is 
poorly preserved on all the available specimens; the presence of a 
subanal fasciole cannot be confirmed, and the nature of the tuber- 
culation within the anal fasciole is not known. 

Remarks. The material, although abundant, offers but a single, 
indifferently preserved oral surface. The petals of different speci- 
mens are somewhat variable in length, breadth, and the degree to 
which they are depressed; the anterior petals of several specimens 
of about the same test width (1 14-119 mm) vary in length from 
37 to 42 mm and vary in width from 5.0 to 6.5 mm. The number 
of primary tubercles in the interambulacral fields is also variable. 
Three specimens from the Cobden Limestone, Greymouth (zfe. 
286, 287, 290), and a single specimen registered as from Ngapara, 
Oamaru (OU. 8542), differ from the remainder of the material in 
possessing more numerous primary tubercles (35-40 in each inter- 
ambulacral field) and in bearing up to four rows of well-spaced, 
coarse secondary tubercles situated adjacent to and running parallel 
with the ambitus of the anterior interambs. The matrix of OU. 
8542, a hard, fine-grained, muddy limestone, contrasts with the 
softer, coarse-grained, almost pure limestone that forms the matrix 
of other echinoids from Ngapara; it closely resembles the matrix 
of the Greymouth specimens. It may be that the locality of OU. 
8542 has been incorrectly recorded. 

Although the holotype has never been figured, it is known to be 
the specimen used by Hutton in the erection of the species. It is 
badly crushed and shows few of the characters of the species, but 
as it has at least 25 pore pairs in ambulacrum IV and approxi- 
mately 36 primary tubercles in interambulacrum 4, it is un- 
doubtedly conspecific with the other specimens herein referred to 
T. greyi. 



12 brf.viora No. 320 

Taimanawa greyi closely resembles /'. pulchella, to which it was 
probably ancestral. It is distinguished by its petals, especially the 
anterior pair, having more numerous pore pairs (Fig. 3). Further 
differences include the deeper frontal goove and more numerous 
primary tubercles oi T. greyi. 

Age. Oligocene — lowermost Miocene; Whiangaroan — Waita- 
kian. The exact age of the holotvpe is not known. From its local- 
ity and matrix, it can confidently be assigned to the Cobden Lime- 
stone, which ranges in age from \\ hiangaroan to Waitakian. 

TAIMANAWA MORTENSENI sp. nov. 
Plate 5, fig. 1 . Figure 2b. 

Plagiobrissus sp. ? Mortensen, 1951, p 503, pi. 39, tig. 3. 

Holotype. Fragments of a specimen from Station 32 of the 
Danish Expedition to the Kei Islands ( 1922), between Little Kei 
and Tajando Islands at a depth o\ 260 meters. Ciliated by the 
University Zoological Museum. Copenhagen. 

Paratype. Fragments o\ a second specimen are represented in 
the collection from the same station. 

Description, Known from fragments onl\. the largest of which 
comprises the anterior half o\ an ahoral surface, lest huge, and 
appears to be approximated as wide as long. Aboral surface in- 
Bated and ambitus more hroadU rounded than in other members o\ 
the genus. Peripetalous fasciole well developed, not indented be- 
tween the petals and not continuous across the frontal ambulacrum. 
Internal fasciole thin. o\ variable width, bordering the frontal am- 
bulacrum for half the distance from the broken margin of the test, 
slightb anterior of the apical system, to the peripetalous fasciole. 
Subanal fasciole well developed and. as far as can be judged. 
reniform. 

Vboral surface: Frontal ambulacrum deeply depressed, apetaloid. 
terminating in a pronounced frontal notch that is approximately as 
wide as deep. Amb plates bear a fine, dense, secondary and miliar) 
tuberculatum and are perforated by minute, conjugate pore pairs. 
interior petals slightly depressed, splayed at an angle of some 
120 . and weakk tlexuous. being concave away from the frontal 
ambulacrum proximallv and concave towards the frontal ambu- 
lacrum distally. Each bears 25 ovate, weakly conjugate pore pairs; 
adjacent pairs are separated by a single row of secondary and mili- 
ary tubercles, and the interporiferous zone bears scattered miliary 
tubercles. Anterior plate series of the frontal interambulacra tumid, 
subtending a weakly concave surface posteriorly into confluence 



1969 INDO-WEST-PACIFIC ECHINOIDS 13 

with the depressions of the frontal petals. Primary tubercles re- 
stricted to the posterior plate series of the anterior interambs lying 
within the peripetalous fasciole, surrounded by prominent scrobi- 
cules, and very weakly crenulate. The partially preserved anterior 
plate series of interamb 4 devoid of primary tubercles, suggesting 
that tuberculation of the posterior lateral interambs is akin to those 
anterior. Frontal ambulacrum margined by a zone of coarse sec- 
ondary tubercles. The remaining aboral interambulacral surfaces 
bear a dense miliary and secondary tuberculation. 

Oral surface: Nature of the plastron and labrum unknown. In- 
terambs bear well-spaced secondary tubercles; ambulacra naked 
except for scattered miliary tubercles. The area enclosed by the 
subanal fasciole bears dense secondary tubercles and is perforated 
on each side by seven conjugate pore pairs. 



150 

140 
130 

_* 120 

Ui 

~* 110 

CL 100 

90- 

80- 

70 

60 

50 



/ 
/ 
/ 
/ 
/ 
/ • 
/ 
/ 
/ 
/ 

i • 

/ 
/ 
/ 
/ 
/ 

x 

/ • 

/ 
/ 

/ 
/ 
/ 
/ • • 

/ 
/ 
/ 



T. pulchella / T. greyi 



X / 

/ 



/ 
/ 

/ 
/ 
/ 



H 15 16 17 18 19 20 21 22 23 24 25 26 

pore pairs 

Figure 3. Plot of the number of fully developed pore pairs in each plate 
series of the anterior petals against test width for Taimanawa pulchella sp.. 
nov. and T. greyi (Hutton) showing the separation of the two species. 
Ordinate values have been estimated for many points and are accurate only 
to ± 5 mm. 



14 imrvioRA No. 320 

Remarks. Mortensen (1951) also recorded fragments ot this 
species from Station 44 o\ the Danish Expedition to the Kei Islands 
( l c >22). but the material is poor (Dr. F. Jensenius Madsen, private 

communication) and has not been examined in the preparation of 
this report. /'. mortenseni is conformable in all its principal char- 
acters with the fossil T. pulchella and I . greyi but represents a 
discrete species. I he possession of an internal fasciolc and a strong 
frontal notch shows that mortenseni must be referred to Taiman- 
awa. Indeed, the \er\ close resemblance between the mid- 1 ertiar\ 
New Zealand species and the present form, which is living in the 
Kei Island area ^\ the Indonesian region, is remarkable. 

I he holotype represented b\ the most complete set of fragments 
would have measured approximately 140 mm in width. 1 he test 
height cannot be accurately estimated, but the aboral surface is 
more inflated than that of either /. pulchella 01 / greyi, and the 
ambitus is more rounded than in either o\' the fossil species. I he 
anterior petals resemble those of /. <vcw in the number o{ pore 
pairs, but the anterior interambs agree with those o\ I . pulchella 
in the number o\ primar) tubercles. I he present species is readib 
distinguished from either iA the fossil species in that its primal) 
tubercles are much less distinctl) crcnulatcd and possess a much 
wider platform ( Fig. 2a. b ). 

Although the Kei Island fragments clearly represent a new 
species, more complete material is needed before it can be ade- 

quatel) defined. 

Genus BRISSOPATAGUS Cotteau, 1863 

/ \/>( species. H. caumonti Cotteau ( 1863: 144. pi. s. figs. 3-7); 
b) subsequent designation (Cotteau 1886: 135). 

Brissopatagus Cotteau, 1863: 144; Pomel, I S K 3 : 32. pars; Duncan and 

Sladen, ISS4: 226. pars; Duncan, 188 > 250, pars; I amberl and I hiery. 

1924: 491. pars; Cooke, 1942; 57, pars; Mortensen, 1951: 453. fnus: 

( ooke, 1959: 92, pars; Phillip. 1966: I 14. pars; Fischer, 1966: 5S4. pars. 

Brissospatangus ( otteau, 1XX6: 135. pars. 

nun Brissospatangus, ( otteau, 1890: IS. 

Diagnosis, rest small, with the test height approximately halt 
the test length, bearing siibanal and peripetalous fascioles. Frontal 
ambulacrum narrow and Lightl) depressed, forming a narrow, shal- 
low frontal notch. Petals lightl) depressed with subparallel pore 
series; anterior pair widely splayed, posterior pair less so. Anterior 
interambs with conspicuous depressions confluent with depressions 
of the anterior petals. Primary tubercles present on the anterior 
plate series of the frontal interambs. 



1969 INDO-WEST-PACIFIC ECHINOIDS 15 

Remarks. Brissopatagus is a poorly defined genus. It was pro- 
posed for two incompletely known species: B. caumonti from the 
Eocene of France; and B. javanicus Cotteau (1863: 144), origi- 
nally described as Spatangus? sp. by Herklots (1854: 13, pi. 3, 
figs. 2, 2a, 2b). The latter was reported from the Eocene of Java 
by Cotteau, but Gerth (1922: 520) revised the age to middle 
Miocene. Each of the species is apparently known from a single 
specimen; that representing B. javanicus is an internal mould and 
shows no tuberculation, whereas that representing B. caumonti is 
an eroded, somewhat deformed specimen. So far as their mor- 
phology can be observed, the two species show good agreement in 
test shape and in the nature of the ambulacra. 

B. caumonti, the type species by subsequent designation, forms 
the basis of Brissopatagus and is redescribed and refigured below. 
The specimen shows traces of a peripetalous and a subanal fasciole 
which have not hitherto been recorded. Cotteau's figures of the 
specimen, as typical of fossil echinoid illustrations of that time, are 
idealized. The distribution of primary tubercles is of particular 
significance. Those shown in Cotteau's figures are fictitious; the 
original specimen shows traces of a few primaries on the anterior 
plate series of the anterior paired interambs, but the remainder have 
been removed by erosion. 

Brissopatagus, in the sense of the type species, is closely related 
to Eupatagus, with which it has been grouped as a subgenus by 
some authors (Dames, 1878; Cooke, 1959). It is also closely 
related to Macropneustes. The only character by which it can 
adequately be differentiated from these two genera is the depres- 
sions of the anterior interambs. The genus is also affiliated with 
Gillechinus Fell (based on G. cudmorei Fell, 1964, p. 213, pis. 1. 
2), which is discussed in detail below. Phillip (1966: 114) has 
proposed that Gillechinus be treated as a synonym of Brissopatagus. 
A comparison of the type species of the two genera shows at once 
that such a view is untenable. G. cudmorei lacks the interambu- 
lacral depressions of B. caumonti, an essential character of Brisso- 
patagus, as Fischer (1966) indicates. Further, G. cudmorei is 
clearly distinct from B. caumonti in that its primary tubercles are 
restricted to the posterior plate series of the paired interambs. If 
Brissopatagus is to be taken as including forms lacking interambu- 
lacral depressions and possessing primary tubercles on the anterior 
plate series of the paired interambs, then it is transitional to Eupa- 
tagus through species like E. lamberti Fourtau and E. cordiformis 
Duncan and Sladen and is indistinguishable from Macropneustes. 



16 BRJ V IORA No. 320 

Subsequent to its erection, a number of Species have been re- 
ferred to Brissopatagus. Several possess characters of both Brisso- 
patagus and Gillechinus, so that their eorreet generic assignment 
is not immediately apparent. For the reasons given in the discus- 
sion of Gillechinus, those with primary tubercles restricted to the 
posterior plate series o\ the paired interambs are referred to that 
genus. The remaining species are II. colligoni Lambert ( 1933: 37. 
pi. 4. fig. 21) from the Eocene of Madagascar and II. SUlldaicus 
Bohrn (1882: 365, pi. 2. figs. 2a. b) from Madura Island north o\ 
Java. 11. sundaicus is no older than Miocene in age. as no Lower 
lertiarv rocks are known to crop out on Madura Island (van 
Bemmelen, 1949: 106). This species, like 11. javanicus, possesses 
distinct interambulacral depressions, but both species are repre- 
sented b\ internal moulds alone, so that the nature oi their tuber- 
culatum is not known and the) cannot be referred to cither Brisso- 
patagus or Gillechinus with certainty. B. collignoni was described 
from crushed material that shows primary tubercles on both plate 
series of the paired interambulacra. and thus it agrees with Brisso- 
patagUS rather than Gillechinus. However, Us interambulacral 
depressions are \er\ wcaklv developed, and it mav prove to be best 
included in EupatagUS 01 Vf<W ropneustes, I he existence ol transi- 
tional species showing characters oi more than owe genus is, ol 
course, an inevitable consequence ol evolution, since intermediate 
stages must often persist after generic stocks have differentiated . 
While such species are valuable indications o\ the relationships and 
origins ol generic Stocks, it becomes a matter oi practical con- 
venience to tolerate these transitional mter-generic taxa. whilst 
maintaining the validit) of the formal genera which they seem to 
link. Am other course would result in the fusion o\' large groups ol 
species in ever larger generic categories, as more intermediate forms 
are discovered from intermediate horizons, and as dines are eluci- 
dated from wider geographic sampling. Under the Linnaean sys- 
tem o\ taxonomy, we are compelled to base our generic diagnoses 
upon the characters seen in the original genotypes. In the case ol 
Brissopatagus and Gillechinus, these characters are given hv the 
tvpe material, and the) confirm the opinions of Mortensen ( 1951 ). 
Fell ( 1964), and Fischer ( 1966). 

In summary, Brissopatagus as here restricted has but a single 
undoubted representative. II. caumonti from the Eocene of Biar- 
ritz. France. It is doubtfully represented in the Eocene o\' Mada- 
gascar (B.? collignoni) and may occur in the Miocene of the 
Javanese area ( B.? javanicus, B.? sundaicus ) . 



1969 INDO-WEST-PACIFIC ECHINOIDS 17 

BRISSOPATAGUS CAUMONTI Cotteau 
Plate 5, figs. 2-4. 

Brissopcitagus caumonti Cotteau, 1863: 144, pi. 8, figs. 3-7. 
Brissospatangus caumonti; Cotteau, 1886: 136, pi. 30, figs. 1-4. 

Holotype. 409- 1A- Co 1-2- b 132 (Ecole des Mines, Paris). 

Description. Test of moderate size, low vaulted, elongate; apical 
system anteriorly excentric, apex near the posterior margin. An- 
terior ambitus broadly rounded and cut by a narrow, shallow, fron- 
tal notch. Posterior ambitus somewhat tapered and more narrowly 
rounded. Apical system cannot be clearly seen but appears to 
possess four gonopores. Traces of peripetalous and subanal fas- 
cioles are present, the former not indented between the petals and 
the latter complete and reniform. Peristome small, crescentic, 
anterior; periproct vertically elongate and situated on the truncated 
posterior test surface. 

Aboral surface: Frontal ambulacrum depressed on the adambital 
half of its length; its pores not preserved. Paired petals distinctly 
depressed with ovate pores. Anterior pair short, widely splayed, 
and anteriorly concave; each has about 12 well-developed pore 
pairs. Posterior pair somewhat longer, set close together, and 
straight, with about 20 well-developed pore pairs. Interambulacra 
2 and 3 with conspicuous depressions forming concavities in the 
test continuous with those of the anterior paired petals. Ornament 
almost entirely lacking due to erosion of the specimen; a few pri- 
mary tubercles preserved on the anterior plate series of interambu- 
lacra 2 and 3. The few patches that have escaped severe erosion 
suggest that much of the adoral surface was originally ornamented 
with small, dense, secondary tubercles. 

Oral surface: Not well preserved. Plastron with a well-developed 
median keel and ornamented with dense secondary tubercles. 
Labrum appears to be long and narrow. Other interambulacral 
areas also bear dense secondaries. Ambulacra adjacent to the 
plastron appear to be narrow and, like the anterior ambulacral 
areas, naked except perhaps for miliary tubercles. 

Remarks. The specimen is badly eroded and somewhat de- 
formed. The interambulacral depressions adjacent to the anterior 
petals, although undoubtedly natural structures, have been accen- 
tuated by deformation. As far as can be deduced from Cotteau's 
descriptions, the specimen to hand is the only known representa- 
tive of B. caumonti. The illustrations given by Cotteau are strongly 
idealized, as may be suspected from the discrepancies between those 
given in the two reports. The nature of the apical system and 



IS BK1 VIORA No. 320 

arrangement of the plates on the oral surface as depleted by Cot- 
tean are fictitious. On the original specimen, the groove of the 
frontal ambulacrum does not reach as close to the apical system as 

depicted h\ Cotteau, nor are the posterior petals as divergent. 

Genus GILLECHINUS Fell, 1964 

/ ype species, a. cudmorei hell ( 1 964: 2 I 3. pis. 1 . 2);b) original 
designation. 
Gillechinus fell. 1964: 213: Phillip. 1966: 114; I ischer, 1966: ^S4. 

Diagnosis. lest of moderate size, with the test height approxi- 
mated hall' the test length, hearing peripetalous and subanal 
fascioles. Frontal ambulacrum wcaklv depressed, frontal notch 
broad and shallow. Petals lightlv depressed, narrow, with suh- 
parallel pore series; anterior petals widclv splayed, posterior pair 
less so. [nterambulacra of the aboral surface ridged at or near the 
suture of the two plate series, with ridges i)\ the paired interamhs 
subtending flattened or concave surfaces posterior to the adjacent 
ambulacra) depressions. Primar) tubercles restricted to posterior 
plate series o\ the paired intcramhulacra within the peripetalous 
fasciole. 

Remarks. As alread) pointed out. Gillechinus cannot be syn- 
onymised with Brissopatagus, as advocated bv Phillip (1966), 
because the interambulacral depressions diagnostic oi Brissopatagus 
are not shown h\ the type species o\ Gillechinus. A further dis- 
tinction is provided In the restriction oi primar) tubercles to the 
posterior plate series ol the paired interainbs o\ Gillechinus, a 
feature not shown b\ Brissopatagus. I here are. however, a number 
iA species that have previouslv been referred to Brissopatagus and 
that possess both interambulacral depressions and tuberculation of 
the Gillechinus stvle. To Classif) these forms, it is necessary to 
decide which o\ the two characters is taxonomically significant. The 
species concerned are as follows: 

G. alabamensis (Cooke) (1942: 58, pi. 4. figs. 7. 8) from 

southeast U.S.A. and possibly Cuba. B. georgianus 

Cooke ( 1942: 58. pi. 7. figs. X-'l I ) and possibly B. avilensis 

Sanchez Roig ( 1951: 45. pi. 33. tigs. 2. 3). Eocene and 

possibl) ( Higocene. 
G. beyrichi (Dames) (1878: 82, pi. II. figs. 2a. b) from 

north Italy. Eocene. 
G. humei (Fourtau) (1908: 2 IS. pi. 2. fig. 8) from north 

Africa. Eocene. 
G. lummaui (Castex) ( 1930: 82, pi. 4, fig. 1 ) from France. 

Eocene. 



1969 INDO-WEST-PACIFIC ECHINOIDS 19 

G. sindensis (Duncan and Sladen) (1884: 226, pi. 38, figs. 
19-21) from India. Eocene. 

One further species, originally referred to Macropneustes, must 
be added : 

G. mexicanus (Dickerson and Kew) (1917: 134, pi. 24, fig. 
3, pi. 25, figs, la, b) from northeast Mexico. Upper Oli- 
gocene or Lower Miocene. 

Gillechinus georgianus was synonymised with G. alabamensis 
by Cooke (1959: 92). G. avilensis is not well known, but the 
grounds for maintaining it as distinct, given by Sanchez Roig (1951) 
as its more depressed petals and less pronounced groove for the 
frontal ambulacrum, are slight and likely to be due to infraspecific 
variation. G. lummaui and G. alabamensis may prove to be syn- 
onymous; G. lummaui appears to be less inflated, and its anterior 
interambs have more primary tubercles than those posterior, while 
the reverse is true of G. alabamensis. The first distinction could be 
due to the slight crushing evident from Castex's ( 1930) figure, and 
the second could be due to infraspecific variation. G. mexicanus 
is closely related to G. cudmorei, differing mainly in its less inflated 
posterior oral surface. B. sindensis is more elongate than any of 
the other species, and B. humei is likewise readily distinguished by 
its more depressed frontal ambulacrum and deeper frontal notch. 
G. beyrichi is characterized by possessing more prominent inter- 
ambulacral depressions than any of the other species. 

The six species show considerable variation in the development 
of interambulacral depressions. In G. beyrichi they are well de- 
veloped, apparently forming actual concavities on the test surface. 
The anterior depressions of G. sindensis are recorded by Duncan 
and Sladen ( 1886) as forming concavities on the test. However, in 
the remaining species the paired interambulacra appear to possess 
flattened surfaces subtended posteriorly from a tumid zone near 
the median suture of the two plate series. The flattened surfaces 
pass into confluence with the depressions of the paired petals and 
give the impression of interambulacral depressions. Homologous 
structures are well developed in Taimanawa and Plagiobrissus, and 
can be recognized on the type species of Gillechinus, G. caumonti. 
Such structures cannot be regarded as identical with the strong 
interambulacral concavities of Brissopatagus. It is significant that 
the interambulacral depressions of all six species are developed in 
all four of the paired interambs, whereas in Brissopatagus these 
structures are restricted to the anterior paired interambs. There is 
little doubt that the six species are closely related; it must be con- 
cluded that interambulacral depressions are a variable character in 



20 UKl \ [ORA No. 320 

this species group, and the point at which they can be regarded as 
present or absent is not easil\ defined. 

The restriction of the primary tubercles to the posterior plate 
series of the paired interambs is a conspicuous feature of all six 
species and is highly unusual for members o\ the Brissidae. This 
feature not only shows the six species to be \er\ closely affiliated 
with G. cudmorei but provides a convenient diagnostic character 
for Gillechinus, separating it from all other brissid genera with the 
exception o\ Lajanaster Sanchez Roig. Lajanaster differs in pos- 
sessing an elongate, strongly flattened test, and its oral surface, like 
that of Plagiobrissus, has a narrow plastron and very narrow ambu- 
lacra! areas. In consequence, the six species here discussed are 
referred to Gillechinus. 

Brissopatagus vilanovae Cotteau (1890: 19, pi. 2. tigs. 9-13) 
is somewhat transitional between Gillechinus and Eupatagus. It 
has primary tuberculatum oi the Gillechinus Style, but its petals are 
spatangiform with curved pore series rather than with subparallel 
pore series as m Gillechinus. lor this reason the species is prob- 
abl\ best referred to Eupatagus, whereupon it becomes a homonym 
of B. vilanovae Cotteau ( 1890: 10, pi. 1. tigs. 1-4) and has been 
renamed E. ibericus by Lambert. 

Gillechinus enjoyed a wide distribution in tropical and sub- 
tropical seas during I oecne time. It may range as high as the 
Miocene, but the Oligoccnc age given for (;'. avilensis by Sanche/ 
Roig (1951) and the Upper Oligoccnc lower Miocene age 
given for G. mexicanus bj Dickerson and Kew (1917) are un- 
supported and must be treated with caution. 



1969 INDO-WEST-PACIFIC ECHINOIDS 21 



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22 BREVIORA No. 320 

LITERATURE CITED 

Bl \1\11 i » N, R. W. VAN 

1449. The Geologv of Indonesia, Vol. I A. General Geology. Govern- 
ment Printing Office, The Hague, 732 pp. 

Bl I . A. G. 

1466. Sea temperatures in New Zealand during the Cenozoic Era as 
indicated by molluscs. Trans. R. Soc. N.Z.. 4(9): 177-187. 
BOHM, V 

1882. Einige Tertiare Fossilien von der lnsel Madura nordlich von 
Java. Denk. kais. Akad. Wissen., 45(2): 359-372. pis. 1-4. 
< \m i \. L. 

1930. Revision des echinides du nummulitique du Departement des 
1 andes. Actes Soc. Linn. Bordeaux, 82: 5-69. pis. 1-4. 

( HI SHKR. R. H. 

1968. The systematic^ oi sympatric species m West Indian spaiangouK 
Stud. Trop. Oceanogr., Miami. 7: vii -f 168, 35 pis. 
COOKJ . C. W. 

1942. Cenozoic irregular echinoids oi eastern United States. J. 

Paleont., L6( I): 1-62. pis. 1-8. 
1959. Cenozoic echinoids of eastern United States. Prof. Pap. U.S. 
Geol. Surv., 321: 1-106, pis. 1-43. 
Cotteau.G. H. 

1863. Echinides fossiles des Pyr6nees. Congr. Scient. fiance. Bor- 
deaux: 1-160. pis. 1-9. 
iss> Paleontologie francaise. Terrain Tertiaire. Eocene, ser. 1. I. 

1889. 672 pp.. 200 pis. 

1890- Echinides Eocenes de la Province d'Alicante. Mem. Soc. Geol. 
1891. France, ser. 3, 5(2): 1-107, pis. 22-32. 
Dami s. W. 

1878. Die Fchiniden der Vicentinischen und Veronesischen Tertiaei 
ablagerungen. Palaeontographica, 25: 1-100, pis. 1-11. 

Die M RSON. R E., AND W. S. W. Kl W 

1917. The fauna of a medial Tertiary formation and the associated 
horizons of northeastern Mexico. Proc. California Acad. Sci., 
ser. 4. 7: 125-156. pis. 17-26. 
DuN< w P. M. 

1889. A revision of the genera and great groups of the Echinoidea. 
Linnean Soc. London Jour.. Zoology. 23: 1-3 11. 
I)l N( \n. P. \1 . \m> W. P. Sladen 

1882- Fossil Echinoidea of western Sind and the coast of Baluchistan 
1886. and of the Persian Gulf, from Tertiary formations. Mem. 
Geol. Surv. India. Palaeont. Indica. ser. 14. 1: 1-392, pis. 1-58. 
Fell. H. B. 

1956. Tertiary sea temperatures in Australia and New Zealand from 
the evidence of fossil echinoderms. Proc. XIV Intern. Cong. 
Zool. Cophenhagen, 1953: 103-4. 



1969 INDO-WEST-PACIFIC ECHINOIDS 23 

1963. The spatangid echinoids of New Zealand. Zool. Pub. Vict. Univ. 
Wellington, 32: 1-8, pis. 1-6. 

1964. New genera of Tertiary echinoids from Victoria, Australia. 
Mem. Nat. Mus. Victoria, 26: 211-215, pis. 1, 2. 

Fischer, A. G. 

1966. Asterozoa-Echinozoa. ///: R. C. Moore ed.. Treatise on Inverte- 
brate Paleontology, Pt. U, Echinodermata 3. Univ. of Kansas 
Press, pp. 543-640. 
Fourtau, M. R. 

1908. Sur quelques echinides Eocenes d'Egypte nouveaux ou pen 
connus. Bull. Inst. Egyptien, ser. 5, 1: 205-220, pis. 1, 2. 
Gerth, H. 

1922. Die Fauna von Java. Echinodermata, Echinoidea. Samml. d. 
geol. Reichs Mus. Leyden, 2(4) : 497-520, pi. 52. 

Herklots, J. A. 

1854. Description des restes fossiles d'animaux des terrains Tertiaires 
de File de Java. Quatrieme partie, Echinodermes. E. J. Brill, 
Leide, 24 pp., 5 pis. 
Hector, J. 

1870. Catalogue of the Colonial Museum, Wellington, New Zealand. 
Wellington, 235 pp. 
Hutton, F. W. 

1873. Catalogue of the Tertiary Mollusca and Echinodermata of New 
Zealand in the Collection of the Colonial Museum. Govern- 
ment Printer, Wellington, 48 pp. 
Lambert, J., and P. Thiery 

1909- Essai de Nomenclature Raisonne des Echinides. Libraire 
1925. Ferriere (Chaumont), 607 pp., 15 pis. 
Lambert, J. 

1933. Echinides de Madagascar communiques par M. H. Besaire. 
Annls. Geol. Surv. Mines Madagascar, 3: 1-49, pis. 1-4. 
Phillip, G. M. 

1966. Notes on three recently proposed Australian echinoid genera. 
Proc. Linn. Soc. New South Wales, 91(2): 114-117. 

POMEL, A. 

1883. Classification Methodique et Genera des Echinides Vivants et 
Fossiles. Adolphe Jourdan (Alger), 131 pp., 1 pi. 

MORTENSEN, T. 

1923. The Danish expedition to the Kei Islands 1922. Saertryk Vi- 
densk. Medd. Dansk. naturh. Foren., 76: 55-99, pis. 1-3. 

1951. A Monograph of the Echinoidea, 5(2), Spatangoida II. C. A. 

Reitzel, Copenhagen, 593 pp., 64 pis. 
Sanchez Roig, M. 

1951. Faunula de equinodermos fosiles del Terciaro, de termino 

Municipal de Moron, Provincia de Camaguey. Mem. Soc. 

Cubana Hist. Nat.. 20(2): 37-64, pis. 23-40. 

(Received 18 September 1968.) 



24 



HRI MORA 



NO. 320 



'<• ":v 



{ 



\ 



3 



i 



^s 




/ 





IM Ml 1 



Figs. 1. 2. Taimanawa greyi (Hutton). dorsal view (Fig. 1 >. lateral view 
(Fig. 2). OU. 8598. test length 135.5 mm: < K% ). 



1969 



INDO-WEST-PACIFIC ECHINOIDS 



25 



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>-j J 

 '■■■ 4 J f 



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L 






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PLATE 2 

Fig. 1. Taimanawa greyi (Hutton), ventral view. OU. 8598. test length 
133.5 mm; (X%). 

Fig. 2. Taimanawa pulchella sp. nov., lateral view. Holotype, OU. 8590. 
test length c 110.0 mm; (x%). 



26 



imi \ [OR \ 



No. 320 



' | ' 



_/ f 



/•"*'' 



m 





,^Aj 



>• 



PI \ I I J 

/ manawa pulchella sp. nov., dorsal view. Holotype, 01 8590, tesl 
length < 1 10.0 mm; <  1 >• 



969 



INDO-WEST-PACIFIC ECHINOIDS 



27 



w>- •; -^ 




! w -■; 



PLATE 4 

Taimanawa pulchella sp. nov., ventral view. Holotype, OU. 8590, test 
length c 1 10.0 mm; ( x 1 ). 



2S 



HKl \ [OR \ 



No. 320 






V 

4 i 

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1969 INDO-WEST-PACIFIC ECHINOIDS 29 



PLATE 5 

Fig. 1. Taimanawa mortenseni sp. nov., dorsal view. Holotype, frag- 
ments of a specimen from Station 32 of the Danish Expedition to the Kei 
Islands (1922), University Zoological Museum Copenhagen, width of 
fragment 105 mm; (Xcl). 

Figs. 2-4. Brissopatagus caumonti Cotteau, dorsal view (Fig. 2), ventral 
view (Fig. 3), lateral view (Fig. 4). Holotype, 409- 1A- Co 1-2- b 132, 
Ecole des Mines, Paris, test length 40.5 mm; (Xcl). 



BREVIGRA 



Museum of Comnpairative Zoology 

Cambridge, Mass. 10 June, 1969 Number 321 

OBSERVATIONS ON THE AGONISTIC AND BREEDING 
BEHAVIOR OF LEPTODACTYLUS PENTADACTYLUS 
AND OTHER AMPHIBIAN SPECIES IN VENEZUELA 

Juan A. Rivero 

and 

Andres Eloy Esteves 1 



Abstract. Observations were made of breeding aggregations of Lepto- 
dactylus pentadactylus, Phyllomedusa trinitatis, Engystomops pustulosis, 
Phrynohyas venulosa and Hyla minuta in northeast Venezuela. L. penta- 
dactylus makes hollows near the water's edge and produces a foam nest where 
the eggs are deposited. During amplexus and oviposition, the male moves 
the feet in alternate, coordinated movements over the cloacal openings of 
the embracing pair, apparently assuring, in this way, proper fertilization and 
distribution of the eggs, besides beating the foam into the proper consist- 
ency. During amplexus, the male shows definite agonistic behavior towards 
other members of his species or to any other object that may touch him. 
Quite often fights between males occur. These may last several minutes and 
consist of standing up and hugging the opponent with the thumb tubercles, 
pushing, toppling, and apparently embracing, and pulling the adversary 
towards the breast tubercles. 

Engystomops seems to prefer road puddles and fouled water for breeding. 
Its voice is similar to that of a puppy, and a group of this species will stop 
calling simultaneously, and for relatively long intervals, when approached 
by an observer. 

Phyllomedusa was extremely abundant in Dieffenbachia and other broad- 
leaved plants. Egg clusters were found on leaf surfaces only. It appears that 
when leaves of the right size are not found, Phyllomedusa does not need to 
make the leaf nests (funnels) described for many of the species. Besides its 
normal mating call, the male has a particular release call. It is possible that 
spent females produce a similar sound. 

Phrynohyas venulosa were occupying approximately equidistant positions 
in a shallow pool. Both this species and Hyla minuta, which were extremely 
common during the first night of observation, had completely disappeared 
the following night. 

1 Instituto Venezolano de Investigaciones Cientificas. 



BR] mora No. 321 



INTRODUCTION 



On June 25. 1968, the authors made a trip to the region of 
Guiria. l : .studo Sucre, in the Peninsula de Paria of Venezuela, and 
collected extensivel) in the vicinity of the village of Rio Salado and 
on Pica del Mango at a slightb higher elevation (100 m ) . The 
conditions near Guiria are xcrophytic to mesophytic. but higher up 
on Pica del Mango the vegetation is quite luxuriant and there is a 
great abundance oi heliconias. Araeeae and Marantaceae. Most oi 
the observations reported here were made at Pica del Mango. 

The excursion to Pica del Mango was made at night after a 
heavy rain during the afternoon. As a result oi the rain, a depres- 
sion in the forest had been filled with water to a depth of a few 
inches. The pool, of about ten by three meters, was undoubtedly 
temporary, but it ma) have been in existence prior to the rainy 
period oi that afternoon. The margins, three to live meters beyond 
the edge of the water, were more or less clear o\ an\ vegetation cx- 
eept at one end. where there were a number of broad-leaved Mar- 
antaceae and Dieffenbachia growing in a thicket. I he forest, with 
its heaw undergrowth of heliconias, CostUS and Araeeae. com- 
menced at about five meters from the edge of the pool. I he pool 
was. however, shaded b\ two or three large trees that grew near 

us margins. 

I he roar produced In frogs calling from the pool could be heard 
at a distance o\ about 200 meters. Upon approaching, the noise 
increase. 1 in intensit) and became almost deafening. Calling from 
the leaves o\ plants were ll\la miniita. sometimes three, four, or 
five H' each leaf oi Dieffenbachia. It was often possible to get three 
specimens with one sweeping movement of the hand over the leal 
surface. 

When flashlights were pointed at the water. Phrynohyas venulosa 
were seen to be occupying the pool, spaced in a pattern at a dis- 
tance ii\ about three to four feet from one another. As there was 
no vegetation inside the pool, this "'formation'" could be easil\ 
observed. 

Also contributing to the uproar coming from the pool were 
Leptodactylus pentadactylm and Phyllomedusa trinitatis in great 
abundance. Engystomops pustulosis in lesser abundance, and a few 
specimens of Leptodactylus />. petersii. 

LEPTODACTYLUS PENTADACTYLUS 

Most attention was given to /.. pentadactylus, since it was hoped 
that the following night could be dedicated to the other species 



1969 



BREEDING BEHAVIOR 



breeding in the pool. Large specimens of L. pentadactylus were 
everywhere; occasionally one would even step on them. In one 
place, eight breeding pairs were counted in a space about three 
meters by one meter, a condition reminiscent of the breeding 
rookeries of some shore birds. Nests were sometimes eight to ten 
inches apart, but more commonly they were about one foot from 
one another. As the eyes of L. pentadactylus shine red at night, it 
was not difficult to survey the area and locate and count speci- 
mens. Remaining stationary and moving the flashlight around the 
pool, we counted 60 individuals, but, if it is considered that nests 
were more abundant in the Dieffenbachia thicket, that females in 
nests were probably not seen, and that many specimens were pro- 
tected from view by other obstructions, the total number may easily 
have been double that. More specimens, especially females, were 
pouring in from the forest all the time. 

Most specimens were males, however. Apart from the mated 
specimens, which were usually quiet in their nests, unattached in- 
dividuals were moving all the time, the jumps being short and fre- 
quently changing in direction. When the butt end of a flashlight 
was presented to a free male, the animal would move away, and no 
attempt was made at amplexus ( in contrast to what a Bufo would 

8 



Q 

o 

Id 
CO 



a. 



4- 



2- 







«►* 




T 



T 



.2 
TIME 



4 .4 .6 

IN SECONDS 



.8 



Fig. 1. — Audiospectrogram of the breeding call of Leptodactylus pei 
tadactylus, Pica del Mango. Estado Sucre. Venezuela, 25 June 1968. 



4 imi mora No. 321 

do). Males wore often soon fighting, the two animals rising on their 
roar logs and attempting to puncture the adversary with their 
thumb spines b) inward embracing movements of their powerful 
forelimbs. The movements were brisk, and the light did not usually 
last more than a few seconds, one of the frogs moving away, or 
being pushed away, after the first "round." On other occasions, 
the light lastod for five or ten minutes, and there was much push- 
ing, toppling (from the raised position), and embracing before the 
fighl was over. 1 Hiese fights created considerable turmoil, and the 
noise could be heard at a distance of 15 or 20 feet without diffi- 
culty. In the plant thicket there was much movement o\' the plants 
as the wrestling animals pushed against them. 

I he male attached to a female showed agonistic behavior to- 
wards anything of the right si/e it ,i frog of a smaller species was 
brought close to the male by the observer, the male would not 
usualK release its hold but would try to push the animal away with 
one or both arms. It the "attack" wore more persistent, or it a 
bigger animal were involved, u would release its hold and fight the 
intruder in the usual manner, trying to puncture it with the Strong 
inward movements of the forearms. M the intruder wont away, or 
if it was moved away, the male it it had boon displaced — would 
give a low short hops near or around the nest, call, and then come 
back, either to the original female or to any other neighboring fe- 
male that was uostmg and was not pre-empted by another male. II 
the male had not boon displaced, he would just sit above the 
female, turn around and call repeatedly. 

It appears that a male does not have an\ strong attachment to 
an) particular female, and. it it happens to stop on something hav- 
ing the proper consistency, it gives a tew short hops, turns around 
several times, and then attempts amplexus. During this time the 
female is completely passive. 

During the period of observation, it was the authors' impression 
that the female does not respond, locally, to an individual male's 
call. Females were coming to the pool, apparently attracted by the 
males' calls, but once there they did not seem to go to calling males 
and. when the) were moving freely, the males did not pay any at- 
tention to them. In practically all eases of matings observed, the 
females were Hat on a depression apparently made by them, and 

'Similar lights have been described for dendrobatids (Test, 1954; Sexton 
I960; Duellman, is><s«s i. Rana (Brode. 1959) and Hyla (Lutz, Bertha. 
I960), hut only the latter anil Leptodactylus are reported to use their 
thumb spines as offensive weapons. 



1969 BREEDING BEHAVIOR 5 

the meeting — or rather the stepping of the male over the female 
— was apparently fortuitous, a situation that may explain the con- 
tinuous and erratic movements of unattached males. To be sure, 
males were sometimes seen calling from shallow depressions, but 
they were never seen making the holes, and no females were seen 
to be attracted to them. Mated males continue to call regularly dur- 
ing the sexual embrace, an indication that the call may constitute 
an innate mechanism to call females to the breeding site but may 
not have any local value. It might have a territorial function, 
but embracing males are often attacked, and not much respect is 
apparently given to their call. However, after a male has rejected 
an intruder, he hops around the nest and calls repeatedly, as if he 
were looking for another "fight/ 1 or as a rooster would, signalling 
its victory. 

As nests (depressions) were sometimes very close together, it 
was often possible to have several breeding pairs under observation 
at the same time. In one particular place, there were three breed- 
ing pairs about six to eight inches from one another, three un- 
attached males one to two feet away and at approximately that 
same distance from one another, and two females that arrived from 
the forest during the period of observation. No foam had been 
produced in any of the nests as yet. The embracing males were 
closely attached to the females, with the last finger over the shoulder 
and the others, including the tubercular thumb, in the axilla. Oc- 
casionally there was some shifting movement on the part of the 
males, perhaps to secure a better position, although it resembled 
the fondling movements of mammals. One of the males suddenly 
released its hold and moved away. The female stayed flat ( actually 
slightly concave, the head and posterior ends being higher than the 
center of the body) in place. After a while, another male came by, 
apparently by accident, stepped on her, and embraced her. She 
pushed the mud forward with her forearms, much as a crab would 
do with its pincers. The occasional "fondling" movements con- 
tinued. 

Another male coming from somewhere else jumped over one of 
the embracing males, dislodging it. When the attack is sudden in 
this fashion, the animal is apparently caught by surprise and, if 
physically displaced, does not usually come back or offer a fight. 
The intruding male did not, however, take "possession" of the 
vacant female and, after a while, left the place. In a second case, 
the attacking male took possession of the female and, in a third, the 
attacking animal took possession, not of the vacated female, but of 
another vacant one a few inches away. 



6 BREVIORA NO. 321 

I Ik- two incoming females wandered into the place at different 

times. I he) passed unattached calling males without incident. 
\\ hen a female was captured and offered to a male by an observer, 
no attempt at amplexus or light was made. Captured males did not 
produce a release call or vibrations that could be noticed. No at- 
tempt at mating was seen with anything not Hal on a depression, 
although one pair was found in amplexus. not in a depression, but 
about a toot from the edge of the pool. ProbabK it a male were to 
attempt amplexus with another male, the attempt would be met not 
with a release call, but with strong agonistic behavior. In many in- 
stances, males were sometimes lound fighting awa\ from nests, but 
it is possible that the lights originated in nests, as in no case were 
two unattached males seen to start a fight and neither were males 
seen fighting females. 

One of the observed incoming females settled lor a while in a 
loam nest lull of eggs Alter some shitting movements, she flattened 
there lor a lew minutes but then departed lor another depression 
apparently left b\ another individual, and settled there. Alter a 
while, she was embraced 

The other incoming female did a lot ol wandering, trying several 

sites, passing onl\ inches awa\ from calling, unattached males, and 
then disappearing for a few minutes. I his specimen was probably 
the one seen later in amplexus near the water's edge. It is not 
known if this female was grabbed by a male while passing, but this 
was the onl\ instance in which a mated pair was seen out o\ a nest 
( depression ). 

On one occasion a commotion was heard at the plant thicket and 

i male was seen trying to clasp the horizontal stem of a Dieffen- 
bachia plant. The stem came out o\ the ground vertically, but 
about 3 inches from the ground it bent and continued horizontally 
f< r about a foot before becoming vertical again. The frog was hold- 
ing the horizontal portion of the stem (which was about 1 ' z inches 
in diameter and quite fleshy) and was trying to clamp it with jerky 
inward movements o\ his arms. It is not known if the stem was 
confused with a female or with a male; it is possible that a light 
between two males had been going on and. as one retreated, the 
other continued to light with the best possible substitute: the Heshy. 
shghtk raised stem oi Dieffenbachia. During this unilateral fight, 
the male was seen to raise its hind legs, first one. then the other, as 
if attempting to scratch the antagonist with its foot. 

When a flashlight butt was pushed at an embracing animal, he 
released his hold, clasped the flashlight with his arm. moved his 
arms inward several times, making a noise as the thumb tubercles 



1969 BREEDING BEHAVIOR 7 

hit the metal, and apparently pushed the flashlight tube towards the 
breast tubercles with his clasping arms. Some "scratching"' move- 
ment of the hind limbs was also noticed, but again it is not known 
if this was done to produce harm or to be able to have all four legs 
on top of the flashlight, which was presented in an oblique position. 

The nest is a shallow depression apparently made by the female, 
who flattens out and moves mud forward with her forelimbs. She 
also makes sidewise movements, as a Phrynosoma would do to 
cover itself with sand, and may also do some pushing with the hind 
limbs, but this was not observed. The nest depression is round, only 
slightly bigger than the animal, and, since it is generally made near 
the edge, but out of the water, it usually has a little water in the 
bottom. Some foam nests were at the margin of the pool, but in- 
side the water, and one was associated with floating debris. No cen- 
tral depression or opening can be observed in the foam. In fresh 
nests some radiating and spirally arranged lines can be seen com- 
ing from the center, but these soon dissipate, perhaps because the 
wandering males do not hesitate to step on nests and other females 
may flatten up in them as if the foam were non-existent. 

In the region of La Escalera, Estado Bolivar, where several foam 
nests were observed in a shallow pool, these were usually in the 
water (not deeper than one inch) and associated with sticks, leaves, 
and other debris. A few were out of the water, but under the leaf 
litter at the edge of the pool. 

Unfortunately, the production of foam could not be observed. In 
many of the nests, foam was already produced, and in others with 
amplexing pairs no foam was produced during the period of obser- 
vation. After the foam is produced, male and female seem to be 
absorbed in their activity. Every few seconds (about Vi minute) 
the female dumps her head in the foam and raises the posterior end 
of the body. The male follows her, and for a while both heads dis- 
appear in the foam while both posterior ends are raised above it. 
Then follows a most interesting movement. The male starts a fan- 
ning movement with his feet, moving them up and down, one fol- 
lowing the other in a rhythmic, well-coordinated fashion. As the 
toes are moved over the cloacal opening of the male, they may 
carry sperm down to the cloacal opening of the female, while at the 
same time they apparently spread the fertilized eggs out into the 
foam. Foam, by the way, is flowing between the two cloacal open- 
ings during the whole procedure. While this process is occurring, 
the foam may be seen to increase in quantity. It appears that the 
fanning movement of the feet may have three functions: passing 
the sperm over the eggs, distributing the fertilized eggs in the foam. 



S BREVIORA N(). 321 

and agitating the water in an egg heater fashion so that it mixes 
with the sperm fluid and produces a foam of the right consistency. 
Substances produced with the sperm or eggs or both ma\ contribute 
to add VlSCOSit) to the liquid. 

ENGYSTOMOPS PUSTULOSUS 

Some Engystomops were observed in die forest pool described 
above, but apparently this is not their preferred kind of site, as only 
10 or 12 individuals were seen here. I he same tanning movement 
observed in Leptodactylus was seen in Engystomops pustulosus, but 
in this ease mating occurred in shallow water ami not in nests or 
depressions out of the water, Engystomops has a notable prefer- 
ence lor pools made in dirt roads In passing cars. In these muddv 
and agitated places the) seem to be at their best, calling continu- 
ously with a voice that reminds one ol a pupp\ recently separated 
from its mother. In the Andes (near la A/ulila and at Isnotu) 
/ ngystomops was collected in water discharged from latrines, where 
it was calling together with H\la crepitans and /.. poecilochilus 
(Isnotu). Approach to a pool in which Engystomops is calling 
must be made quicllv as the\ can stop calling suddenly, as it b\ 
signal, and then keep quiet tor a while. 

PHYLLOMEDUSA TRINITATIS 

Another species breeding in the forest pool together with Lepto- 
dactylus pentadactylus and Engystomops was Phyllomedusa trini- 
tatis. II the sight of breeding /.. pentadactylus was grandiose, that 
of Phyllomedusa w.is spectacular. The Marantaceae-Dieffenbachia 

plant thicket described above was literally loaded with them. Main 
leaves were bent as a result of the weight, and others had broken 
at the stem and hung down with several Phyllomedusa attached or 
walking, in monkev fashion, along the stem. Some Phyllomedusa 
were in amplexus. as well described and photographed In Kenny 
(1966), but females were relatively scarce; quite often a male 
would grab another male, onlv to release him upon the production 
ot his release call and vibrations. A specimen that hung from its 
hind limbs, dead, was 'covered" by a male for about one hour. In 
some places there were bunches of four or five individuals, all 
apparently attempting amplexus with a female. These bundles hung 
like grapes; they were never seen on top of a leaf or on the floor, 
but embracing pairs were sometimes found on broad leaves or on 
stems. 



1969 BREEDING BEHAVIOR 9 

The movements of Phyllomedusa remind one of those of a 
monkey. The limbs are moved slowly and deliberately, the fingers 
and toes grabbing the twigs or branches more or less as a monkey 
would. Obviously, its walking gait is adapted to moving along 
twigs and small branches, but, when placed on the floor, it can 
also walk, although it may jump fairly fast too. When Phyllo- 
medusa is placed in the sun, the color first becomes spotted dark 
green on a lighter green background and later, but in a matter of 
seconds, it becomes dark, bottle green. In the collecting bag in the 
shade, most were leaf green, but a few were olive brown, and this 
latter color is kept after preservation. It is stated by Kenny that 
the non-breeding color is bright green, whereas the breeding color 
is olive brown. This could not be confirmed, as most of the speci- 
mens observed, including amplexing pairs and unspent females, 
were green. Calling males from San Casimiro, 850 m, were also of 
a bright green color. 

5 



3- 



o 

z 
o 
o 

Ui 
CO 

ce 

UJ 

Q_ 



S 1 



'ML. JU  — J»".1 1 . 
I I 



.5 1.0 

TIME IN SECONDS 

Fig. 2 — Audiospectrogram of the breeding call of Phyllomedusa trinitatis, 
Monte Oscuro, near San Casimiro. Estado Aragua. Venezuela, 15 June 
1968. 



In one instance a Phyllomedusa was seen to be walking along a 
stem. Suddenly it stopped, more or less "sat," and, while holding 
the twig with the two feet and one hand, raised the right arm and 
waved it around as if looking for another twig. On another oc- 
casion the animal raised both arms and did a similar operation. It 



10 



BR] \ [OR \ NO. 32 I 



appears that the animal "•feels" the twigs and branches rather than 
sees them, hut this ma\ have been as a result of the light point- 
ing at it. This behavior looked so unfroglike that it was thought to 
be accidental until it was observed on several other occasions. 

Only three or tour bundles of eggs (of about 25 or 30). all 
attached to the upper surfaces of broad-leaved plants, were ob- 
served. This is contrar) to the situation described in other Phyllo- 
medusa { Budgett, is c ' l >: Agar. 1909; von Ihering, 1866) or for this 
species in Trinidad ( Kenny, 1966), where a funnel made from one 
or more k\i\cs is used as" a nest. A. and B. but/ ( 1939) wrote 
that a nest o\ /'. appendiculata was found in a hollow o\ a wet block 
of stone covered bv a fallen tree trunk in a mountain brook. It 
appears that when small leaves are not available, Phyllomedusa 
can adapt lo other conditions. 

How Phyllomedusa made itself heard under the roaring sound o\ 
I', venulosa and /.. pentadactylm is difficult to understand Perhaps 
it comes earlier to the pool; or perhaps the scarcity o\ females can 
be attribute,! to this factor, kemn (1966:16) reports that the 
females of Phyllomedusa are also vocal and that males have a 
mating call (■"huh. huh. huh") and a non-mating call which the) 
produce throughout the night I he authors onl) heard two calls, 
one the regular mating call, and the other, a release call (and vibra- 
tions) produced bv males when other males louch them or attempt 
amplexus with them. I his call is also produced when a male is 
grabbed by a human or when several specimens are placed together 
ui a bag. It is possible that spent females produce a similar sound, 
although this was not recorded. 

PHRYNOHYAS VENULOSA 

I ew observations were made on Phrynohym venulosa, in the 

hope o\ paying more attention to this species the next day. How- 
ever, it was observed that they occupy equidistant positions in the 
pool, that the) retain amplexus alter capture, that one was seen to 
grab a /.. pentadactylm in amplexus and ride it until they both dis- 
appeared, and that even when not calling, the two vocal sacs seem 
to act as floaters, looking as if the animal were wearing water-wing 
life preservers. 

The voice of Phrynohym venulosa has been described by a num- 
ber of authors. The call can perhaps best be described as like the 
bleating of a goat, but when many specimens are calling together. 
the noise produced can be compared to that of a low-flying heli- 
copter or to that produced bv automobiles with holes in their 
mufflers. 






969 



BREEDING BEHAVIOR 



11 



The following afternoon, the surface of the pool was seen to be 
almost completely covered with what, at first sight, appeared to be 
the pupae of giant mosquitoes; these were the eggs of P. venulosa. 
That same night both P. venulosa and Hyla minuta had disappeared 
completely. And they were neither seen nor heard in another pool 
a few kilometers away, where they were heard abundantly the 
night before. 

8 



© 
o 

Ui 
W 

id 
^2 




.4 



1.0 1.2 1.4 

TIME IN SECONDS 



1.6 



1.8 



2.0 



Fig. 3 — Audiospectrograms of breeding choruses. Left, Leptodactylus 
podicipinus petersii, El Pilar, Estado Sucre, Venezuela, 27 June 1968. 
Center, Pseudis paradoxus, Rio Salado, Estado Sucre, Venezuela, 25 June 
1968. Right, Phrynohyas venulosa, Pica del Mango, Estado Sucre, Vene- 
zuela, 25 June 1968. 



It is not known when the forest pool described above was filled 
with water. If it was during the afternoon rain preceding our visit, 
then the breeding; of Phrynohyas venulosa lasts only one night. 
During the periods of observation, no ordinary frog predators were 
seen. Leptodactylus can easily swallow Engystomops, but it was 
probably too active, breeding and fighting, to bother about eating. 
On the first night, two enormous pigs were observed grabbing frogs 
and apparently eating them (the actual eating was not seen, as the 
pigs were immediately scared off by the observers, but their insist- 
ence on coming back left little doubt about their "intentions"). 
Three of the P. venulosa seen were dead, the other came out of the 
water dying. The cause of this is not known, but it is not improb- 
able that they accidentally bumped into a male Leptodactylus, which 
can be a dangerous adversary. In one instance, Esteves' hand was 
accidentally grabbed by a male Leptodactylus and, before he could 
remove it, the animal tried to puncture it with its thumb tubercles 



12 BREVIORA NO. 321 

by jerky inward movements and did its best to push the hand against 
the breast spines. A /'. venulosa would probabl) be in serious 

trouble it' similarly treated. 

ACKNOWLEDGMENTS 

I he observations resulting in this paper were made possible b\ 
the assistance given b\ the Instituto Venezolano de [nvestigaciones 

( lenutieas. the institution b\ which both authors were employed at 
the time the work was conducted. The) are especially thankful to 
Dr. GilbertO Rodriguez for his help and encouragement in all 
phases of their work. Partial support was also given b\ NSF grant 
GB 6944 to Ernesl 1 Williams. 

bibliography 

\ GAR, \\ II I Ki n i 

1909 I lie nesting habu of the tree frog Phyllomedusa sauvagii. Proc 

Zool Sot I ondon 1909 ! pi. 84. 

Hri hi h. Chari is \t . Jr 

[925 rhe Darien Jungles Nal Hist 25(4): 325-337, 15 figs. 
1946. Amphibians and reptiles of the Rio Chucunaque Drainage, I'an- 

.1111,1. with notes on theii life histories anil habits. Bull. Amer. 

Mus Nat Hist Hf.isi J75-436, 25 figs., pis. 42-60. 

Brodi . Wn i hm t 

1959. rerritorialit) in Rana clamitans. Herpetologica 15(3): 1 40. 

Bi doi 1 1 . I . S 

1899. Notes on batrachians of the Paraguayan ( haco, with observa- 
tions upen the breeding habits and development, especially with 
regard to Phyllomedusa hypochondrialis ( ope \lso a descrip- 
tion of a new species Quart, Join Micro Sci >2 305-333, 
pis 28 J2 

D III MAN, Wll 1 [AM 1 

1956. Ih; frogs of the hylid genus Phrynohyas I itzinger, 1843. Misc. 
Publ. Mus Zool. t ni\ Mich. No. 9h 1-47, 10 figs., 4 maps. 
6 pis. 

1960. \ distributional stud) of the amphibians of the Isthmus of 
Tehuantepec, Mexico, Univ, Kansas Publ. Mus. Nat. Hist. 13: 
19-72. 

1966. Aggressive behavior in dendrobatid frogs. Herpetologica 22(3): 
217-221. 2 figs 

1 I RNANDEZ, K \l I . WD MlOl El I IKS \M>I / 

1921. Sobre la biologia v reproduccion de algunos batracios amen 
tinos. I. Cystignathidae. An. Soc. Cient. Argentina 91: 97-140. 
4 figs.. 3 pis. 



1969 BREEDING BEHAVIOR 13 

Gallardo, Jose M. 

1958. Observaciones sobre el comportamiento de algunos batracios 

argentinos. Ciencia e Investigacion 14(7): 291-302. 
1964. Consideraciones sobre Leptodactylus ocellatus (L.) (Amphibia 
Anura y especies aliadas. Physis 24(68): 374-384. 3 pis. 
Gans, Carl 

1960. Notes on a herpetological collecting trip through the south- 
eastern lowlands of Bolivia. Ann. Carneg. Mus. 35, Art. 13: 
283-314. 

Hemming, Francis (ed.) 

1958. Opinion 520. Suppression under the Plenary Powers of the 
specific name tibiatrix Laurenti, 1768. as published in the com- 
bination Hyla tibiatrix, and of the generic name Acrodytcs Fit- 
zinger, 1843, and interpretation of the same Powers of the 
nominal species Rana venulosa Laurenti, 1768 (Class Am- 
phibia). Opin. Declar. Intl. Comm. Zool. Nomen. 19: 169-200. 

Hudson, William H. 

1892. The Naturalist in La Plata. London, 394 pp. 
Ihering, Hermann von 

1886. On the oviposition in Phyllomedusa iheringii. Ann. Mag. Nat. 
Hist. 17: 461-463, 1 fig. 
Jimenez, de la Espada, Marcos 

1875. Vertebrados del viaje al Pacifico, Batracios. Madrid, 208 pp., 
6 pis. 

Kenny, Julian S. 

1966. Nest building in Phyllomedusa trinitatis Mertens. Carib. J. 
Sci. 6(1-2): 15-22, 6 figs. 

Lutz, Adolpho, and Bertha Lutz 

1939. Notes on the genus Phyllomedusa Wagler. Ann. Acad. Bras. 
Sciencias 11(3): 219-263, 8 pis. 
Lutz, Bertha 

1960. Fighting behavior and an incipient notion of territory in male 
tree frogs. Copeia 1960 ( 1 ) : 61-63, 4 figs. 
Lynch, W. Gardner 

1959. Some reptiles and amphibians from Trinidad. Herpetologica 
15(3): 1 13-117. 

Mayorga, Horacio 

1967. Informe sobre una coleccion de anfibios (Salientia) procedentes 
de Nicaragua. Carib. J. Sci. 7( 1-2) : 69-77. 

Netting, Graham M. 

1930. The systematic status and breeding habits of Eupemphix trini- 
tatis Boulenger. Ann. Carnegie Mus. 19(4): 249-254. 
Noble, Gladwin Kingsley 

1927. The value of life history data in the study of the evolution of 
the Amphibia. Ann. N. Y. Acad. Sci. 30: 31-128. 30 figs., pi. 9. 



14 BREVIORA NO. 321 

PORTI R. kl NNF.TH K 

1962 Mating calls and noteworthy collections of some Mexican 
amphibians. Herpetologica 18(3): 165-171. 

SI \ MIS, ()\\ I N 

I960. Sonic aspects of the behavior and of the territory of a dendro- 
batid frog. Prostherapis trinitatis. Ecology Hi It: 107-115. 
6 figs. 

1462. Apparent territorialism in Leptodactylus insularum Barbour. 
Herpetologica 18(3): 212-214. 

Sl I .INI (.1 K. 1 I tiNHVKD 

1402. An annotated list of batrachians and reptiles collected in the 
vicinity of La Guaira, Venezuela, \«. nh descriptions of two 
new species of snakes I'roc. U. S. Nat. Mus. 21: 179-192, 
figs i J3 

Iim. I Kl ni RICK H. 

I*>54. Social .mgiessixencss in .in amphibian. Science 120(3 108): 
140-141. 
ZWEIFEL, Ki< HVKD ( i 

1964 Life hisioi\ of Phrynohyas venulosa (Salientia: Hylidae) in 
Panama* ( opeia |,> ' ,,l ' »' 201-208. 

(Received 5 March 1464.) 



BREVIORA 

Museum of Commpsir&itlve Zoology 

Cambridge, Mass. 10 June, 1969 Number 322 



THE FOSSIL RECORD OF AMPHIUMID SALAMANDERS' 

Richard Estes 



Abstract. The first pre-Pleistocene records of amphiumid salamanders 
are described here. Proamphiwna cretacea, n. gen., n. sp., from the late 
Cretaceous Hell Creek Formation of Montana, differs from Amphiuma in 
having less well-developed vertebral muscle crests, basapophyses, and 
neural spines. The species is represented only by vertebrae. Amphiuma 
jepseni, n. sp., from the late Paleocene Polecat Bench Formation, Wyoming, 
is the earliest record of the modern genus. Its vertebrae do not differ from 
those of Recent Amphiuma, but its skull lacks the facial elongation char- 
acteristic of the modern species. It is represented by vertebrae and by a 
partial skull. 

Widespread transgressive seas of the Cretaceous, by providing extensive 
coastal plains and abundant shallow-water riparian habitat, may have been 
a major factor in the evolution of amphiumids and other elongated, aquatic 
salamanders. 

INTRODUCTION 

No previous fossil record for the salamander family Amphiu- 
midae has been described, except for two Pleistocene records, one 
reliable, the other spurious. Auffenberg and Goin (1959) and 
Brattstrom (1953) have correctly ascribed to Amphiuma speci- 
mens from Florida Pleistocene localities. An unusual zoogeographic 
record was described for the Pleistocene of Germany by Brunner 
(1956), as a new species, Amphiuma norica. This specimen may 
be the parasphenoid of a teleost; it bears no resemblance to that of 
Amphiuma, and there is no basis for the identification (Fig. 3g-h). 

Study of a North American Cretaceous floodplain herpetofauna 
(Estes, 1964) has disclosed ancestors of some lower vertebrates 
now living only on the Gulf Coastal Plain of North America. The 



1 Fossil vertebrates from the late Cretaceous Hell Creek Formation, 
Montana: Contribution No. 3. 



2 BR1VIORA No. 322 

fauna of the latter region today differs in main ways from the 
ancient floodplain fauna, hut hoth share the presence of the fishes 
Amia, Lepisosteus, Acipenser, a polyodontid, sirenid salamanders. 

crocodiles, the turtle Trionyx, and alligators. These animals toda\ 
are all relicts of forms once widely distributed on fioodplains oi 
late Cretaceous epicontinental seas, the relict drainage of which is 
the Mississippi River system. The description here of another Gulf 
Coast element in the fossil fauna <^ the late C "retaceous of Mon- 
tana and Paleocene of Wyoming is further indication of the relict 
nature and former wide distribution of the (iulf Coast forms noted 
above. 

ACKNOWLEDGMENTS 

I am grateful to Dr. Francis Rose for helpful suggestions. Dr. 
Glenn Jepsen has provided the Silver Coulee specimens, which are 
described here in advance o\ a collaborative paper on Silver Coulee 
lower vertebrates, and 1 take pleasure in naming the new species for 
him. Collection of the Princeton material was supported by the 
Scott Fund. Princeton University. This research has been sup- 
ported in part b) National Science Foundation Grant GB-7176. 
I he figures are In Mr. l.as/lo Meszoly. 

FAMILY AMPHIUMIDAE 

Proamphiuma cretacea, n. gen., n. sp. 

(Figs. 1, 2) 

Holotype. M( / 3504, vertebra. 

Paratypes. MCZ 3506-3509, 3627-3636, vertebrae: 3505. 

3637, atlantes. 

Locality. Bug Creek Anthills ( BCA), NO feet below the Tullock 
Formation in the west half oi Section 9, T 22 N, R 43 F, McCone 
County, Montana; collected bv A. D. Lewis and party. 

Horizon. Hell Creek Formation. 

Age. Fate Cretaceous. 

Diagnosis. An amphiumid with neural arch of atlas horizontal 
rather than dorsally-pointing posteriorly; vertebrae relatively nar- 
rower, more constricted medially, and with muscle crests, basa- 
pophyses, and neural spines less prominent than in Recent Am- 
phiuma of comparable size; shares with Amphiwna described below 
the relatively more constricted centrum and neural arch. 

Description. Centra amphicoelous, cotyles teardrop-shaped with 
thin internal coating of calcified cartilage; subcentral keel promi- 
nent or low, with or without channels for the segmental blood 



1969 



AMPHIUMID SALAMANDERS 



vessels; prominent or flattened anterior basapophyses usually pro- 
jecting beyond anterior centrum margin; indications of two closely- 
appresscd rib-bearers appearing on the four anterior vertebrae, the 








5mm 







Fig. 1. Proamphiuma cretacea, n. gen., n. sp.: a, left lateral, b, an- 
terior, c, posterior, d, dorsal, and e, ventral views of holotype vertebra, 
MCZ 3504; /, anterior, g, posterior, /;, dorsal, i, ventral, and /', right lateral 
views of atlas, MCZ 3505. 



HRl VIORA 



No. 322 



others with only one; ventral lamina of rib-bearer present but ap- 
parently not well developed anteriorly; zygapophyses anteroposteri- 

orly elongate, prominent, widely separated; posterior zygapophyses 
with keeled crests dorsally, extending forward to level of root of 
transverse process, these crests more medial in presumed anterior 
vertebrae and more lateral in presumed posterior vertebrae; neural 
spme relatively low, but prominent, thin and keeled, squared-off in 
lateral view; posterior border of spine slightly forked, often in three- 
pronged fashion. Atlantes with rounded anterior cotyles and prom- 
inent intercotylar process; centrum short; neural arch relatively 
high and blunt, with dorsal surface horizontal; neural spine only a 
faint ridge. 

Discussion. The vertebrae are distinctively amphiumid in the 
combination o\ anterior basapophyses. subecntral keel, and pos- 
terior zygapophyses with prominent distinctive crests on their dorsal 




Fig. 2. Proamphiuma cretacea, n. gen., n. sp.: (/. MCZ 3507. right lateral 
view of trunk vertebra; /'. MCZ 3508. righf lateral view of anterior trunk 
vertebra; c, MCZ 3506, dorsal view of trunk vertebra; </ ami <•. MCZ 3509 — 

J. ventral, and e, left lateral views of trunk vertebra, probably near an- 
terior end of column. White = broken surfaces. 



1969 AMPHIUMID SALAMANDERS 5 

surfaces that reach anteriorly to the roots of the transverse pro- 
cesses. There is close resemblance to vertebrae of Amphiuma, but 
specimens of equal size of the latter and of Proamphiwna differ in 
that the Cretaceous species has relatively narrower centra, less 
prominent zygapophyses, and weaker muscle crests. These are 
small differences but are on the order of those in closely-related 
modern salamander genera. Presence of posterior zygapophyseal 
crests implies development of the specialized IV-6 intervertebral 
muscle fibers found in Recent Amphiuma (Auffenberg, 1959). 

No skull parts are known for this species, but the Paleocene 
Amphiuma described below has a less elongated snout than Recent 
Amphiuma, and probably a similar short skull was present in Pro- 
amphiwna as well. The two genera are closely related; Proam- 
phiuma is structurally an ancestor of Amphiuma and there is no 
impediment to considering it a real ancestor, as well. It resembles 
the Paleocene Amphiuma described below in having relatively 
narrow, constricted centra. This is the earliest record of the 
Amphiumidae. 

Amphiuma jepseni, n. sp. 
(Figs. 3-5) 

Holotype. Princeton University (PU) 14666, at least fourteen 
associated but dislocated vertebrae in a single block of matrix; 
associated left quadrate and left dentary. 

Paratypes. PU 14668, partial skull and associated mandibles. 

Locality and collector. Silver Coulee Quarry, S21, T 57 N, R 
100 W, Park Co., Wyoming; collected by G. L. Jepsen and party. 

Horizon. Polecat Bench Formation, Silver Coulee beds. 

Age. Late Paleocene (Tiff anian). 

Etymology. Named for its collector, Dr. G. L. Jepsen, Princeton 
University. 

Diagnosis. Vertebrae relatively narrow as in Proamphiuma, 
crests and basapophyses well developed as in modern Amphiuma; 
snout short and blunt in contrast to modern Amphiuma; vomers 
less larval, with fewer and relatively more pointed teeth than in 
modern Amphiuma. 

Description. Vertebrae amphicoelous, cotyles suboval anteriorly, 
teardrop-shaped posteriorly; anterior basapophyses strong, ven- 
trally flattened, crested on their posterior surfaces, subcentral keel 
well defined, subcentral foramina simple, elongate; rib-bearers of 
posterior vertebrae unicipital, blunt distally, no rib articulation 
facet; anterior vertebrae with bicipital but closely approximated 



imr.vioRA 



No. 322 











l ig. 3. a - /. Amphiuma jepseni, n. sp.. PU 14666: a, labial, and b, ven- 
tral views of left dentar) associated with holotypc: <. nuht lateral. </. ven- 
tral. <■. anterior, and /. dorsal views of best-preserved vertebra from holonpe 
series, g, A. means, ventral view of parasphenoid; //, A. norica Brunnei. 
ventral view of parasphenoid (?), after Brunner; g, //. X about 4. Coarse 
stipple = matrix, white area = broken surface 






1969 



AMPHIUMID SALAMANDERS 



rib-bearers; ventral lamina of transverse process prominent in pos- 
terior vertebrae, absent or reduced in anterior vertebrae; zyga- 
pophyseal ridge little developed near midpoint of vertebrae, neural 
arch narrow, pinched in at level of transverse process; well-devel- 
oped keels on posterior zygapophyses reaching anteriorly to the 
level of transverse process; neural spine prominent, thin, squared 
off dorsally; posterior border of neural arch forked, a median sep- 
tum bisecting the fork in three-pronged fashion. 

Only anterior part of skull present, crushed and flattened, but 
restoration indicates a snout relatively wider and shorter than in 
Amphiuma; premaxilla unpaired, flattened dorsally, elongated pos- 
teriorly, a strong median septum formed, total actual tooth number 
not determinable because of breakage, but presence of a median 
tooth and six left lateral ones indicating a total of 13; ethmoid 
processes of frontals visible in section posteriorly; nasals not pre- 
served, prefrontals sculptured, elongated, a long prefrontomaxillary 





Smm 



-j — i — I 




Fig. 4. Amphiuma jepseni, PU 14668, anterior portion of skull: A, re- 
stored dorsal view of snout and dentaries; B, dorsal view of fossil as pre- 
served, right maxilla dislocated and flattened; C, restored palatal view; D, 
palatal view of fossil, elements of left side in approximately natural articu- 
lation. Abbreviations: d = dentary, f = frontal, m = maxilla, n = nasal, 
p = premaxilla, pa = parasphenoid, pf = prefrontal, v = vomer. 



8 



BREVIORA 



No. 322 









L 



5min 

I 



Fig. 5. Amphiuma jepseni, PU 14668, fragmentary dentaries originally in 
articulation with skull and removed to prepare palate: a, right lingual, b, left 
lingual, c, left ventral, and d. right ventral views. 



1969 AMPHIUMJD SALAMANDERS 9 

suture extending posteriorly to orbit; frontal broken, dislocated, a 
strong sculpture present; maxilla bluntly curved, sculptured on 
nasal process, a row of foramina present exteriorly; teeth placed in 
a deep, channelled sulcus dentalis; teeth pedicellate, sharply 
pointed, and with recurved crowns, total tooth number unknown be- 
cause of breakage; vomer expanded anteriorly, a small hollowed- 
out sulcus dentalis present near the midline, 10 or 11 teeth present, 
choanal excavation present laterally in vomer, reaching to level of 
seventh tooth from front; ethmoid visible medial to vomer, an an- 
terior ethmoid foramen visible on the left; blunt parasphenoid ex- 
tending anteriorly, covering ethmoids and separating vomers to 
anterior end of sulcus dentalis; orbitosphenoids missing, but grooves 
for them present on ventral surfaces of ethmoid; flange of vomer 
covers orbitosphenoid area ventrally. 

Dentary strongly curved with deep sulcus dentalis; teeth as in 
upper jaw; symphysis flat, prominent; tooth number unknown, but 
comparison of dentary associated with type and PU 14668 indi- 
cates about 18; prearticular robust, medially crested, reaching an- 
teriorly to about sixth tooth from symphysis; prominent fossa for 
adductor muscle attachment posterior to last tooth; blunt coronoid 
process present. 

Quadrate blunt, robust, articular surface gently concave, palato- 
quadrate and hyoid connections well separated. 

Discussion. Vertebrae of Amphiuma jepseni do not differ sig- 
nificantly from those of Recent species in general features, and are 
especially similar in degree of muscle crest development. Vertebrae 
of Proamphiuma cretacea have substantially less well-developed 
muscle crests relative to those of A. jepseni and Recent Amphiuma 
of equal size. The relatively constricted centra resemble those of 
P. cretacea, a resemblance not unlikely in view of their close geo- 
graphic and temporal association. 

The snout of Amphiuma jepseni is less elongated than that of 
Recent Amphiuma, although the specialized enclosure of olfactory 
tracts by the frontal is as well developed as in the Recent species. 
Tooth crowns of the fossil are less chisel-shaped than those of 
Recent species. Tooth number of premaxilla and dentary is within 
the range of Recent Amphiuma, but the number of vomerine teeth 
is significantly less (10-11 rather than 17-18; Rose, 1968). The 
latter condition and the greater development of vomers anterolateral 
to the tooth row indicate a lesser degree of paedomorphosis in A. 
jepseni than in Recent Amphiuma. 

These skull differences probably reflect no more than specific 
difference from the Recent species. The lack of complete skeletons 



10 breviora No. 322 

of the fossil form and the existence of only two closely-related 
modern amphiumid species for comparison makes the differences 
somewhat difficult to assess. However, some substantiation of this 
a mclusion can be drawn from the fact that Proamphiuma cretacea 
differs substantially in development of vertebral muscle crests when 
compared with Amphiuma jepseni and with Recent individuals oi 
equal size. I consider these muscle crests to be useful in defining 
generic boundaries, since their development reflects muscle differ- 
ence oi greater adaptive significance to the animal than is elonga- 
tion of the snout. 

EVOLUTION AND FAUNAL ASSOCIATES 

Other Cretaceous records of modern and extinct families oi 
elongated, aquatic salamanders have also been described (Estes, 
1964, 1969). The addition oi amphiumid salamanders to this 
record suggests that the widespread epicontinental seas of the Cre- 
taceous were an important factor in evolution of this abundant sala- 
mander fauna, by providing extensive coastal plains with slow 
drainage and abundant shallow-water, riparian habitats 

The presence of these amphiumids in Cretaceous (Maestrichtian) 
and Paleocene (Tiffanian) sediments in Wyoming and Montana 
implies an origin oi the Amphiumidae earlier than the late Cre- 
taceous. The family is thus almost equally as ancient as the Sireni- 
dae. the earliest record of which occurs m rocks of late Cretaceous 
(Campanian) age in Wyoming (unpublished record o\' Habrosaurus; 
I stes. 1964, p. 170). 

The reduction in si/e and numbers oi dinosaurs and the pres- 
ence oi plant fossils of Paleocene aspect imply warm-temperate 
rather than sub-tropical conditions for the Bug Creek locality, which 
is stratigraphically close to the arbitrary Cretaceous-Palcocene 
boundary in Montana (Sloan and Van Valen, 1965). 

The fauna at the Paleocene Silver Coulee locality is unusual for 
late Cretaceous and early Paleocene localities in that fish remains 
are lacking. However, the aquatic salamanders Scaphcrpeton and 
Opisthotriton occur at both Silver Coulee and Bug Creek localities. 

LITERATURE CITED 

\l FFENBERG. W. 

1959. The epaxial musculature of Siren. Amphiuma, and Necturus 
I \mphibia). Bull. Florida State Mus. 4:253-265. 



1969 AMPHIUMID SALAMANDERS 11 

AUFFENBERG, W., AND C. GolN 

1959. The status of the salamander genera Scapherpeton and Hemi- 
trypus of Cope. Amer. Mus. Novit. 1979: 1-12. 
Brattstrom, B. 

1953. Records of Pleistocene reptiles and amphibians from Florida. 
Journ. Florida Acad. Sci. 16:243-248. 
Brunner, G. 

1956. Nachtrag zur Kleinen Teufelshohle bei Pottstein (Ober-franken). 
Ein Ubergang von der letzten interglaziale Riss-WLirm-Warm- 
fauna zur Wurm I-Kaltfauna. Neues Jahrb. Geol. Pal., Monatsh., 
1956:75-100. 

Estes, R. 

1964. Fossil vertebrates from the late Cretaceous Lance Formation, 
eastern Wyoming. Univ. Calif. Publ. Geol. Sci. 49:1-180. 

1969. The Batrachosauroididae and Scapherpetontidae, late Cretaceous 
and early Cenozoic salamanders. Copeia, 1969, no. 2, in press. 
Rose, F. 

1968. Ontogenetic changes in the tooth number of Amphiuma tridac- 
tylum. Herpetologica, 24:182-184. 
Sloan, R., and L. Van Valen 

1965. Cretaceous mammals from Montana. Science, 148:220-227. 

(Received 24 March 1969.) 



BREVIORA 

Museum of Comparative Zoology 

Cambridge, Mass. 10 June, 1969 Number 323 

ON THE EARTHWORMS OF ASCENSION AND 
JUAN FERNANDEZ ISLANDS 1 

G. E. Gates 2 



Abstract. Recent collections enable addition of several species to lists of 
earthworm faunas of the two oceanic islands. As all species are widely dis- 
tributed anthropochores. differences in the two faunas may be due mainly 
to climatic factors. 

ASCENSION ISLAND 

Ascension is a small island in the South Atlantic with an area 
of only 38 square miles. Green Mountain, reaching a height of 
2,817 feet, is surrounded by a table land with a height varying 
from 1200-2000 feet. The island was discovered by Joao da Nova 
in 1501 but probably was first settled only in 1815 when Napoleon 
was exiled to St. Helena, some 500 miles distant, the nearest 
land. Previously the island was bare except for vegetation at the 
summit of Green Mountain. 

Only two megadrile species had been recorded (Sims, 1964) 
from the island. Through the efforts of herpetologist Arthur Love- 
ridge, who retired to St. Helena, a small collection was secured 
from Ascension. 

LUMBRICIDAE 

AllolobophoraEiscn, 1874 
Allolobophora sp. 

The aclitellate worm (Sims, 1964) was not further identified. 

1 From research financed by the National Science Foundation. 
- Zoology Dept., University of Maine, Orono, Maine. 



2 BREVIORA No. 323 

Ml (.ASC'OLECIDAE 

Pheretima Kinberg, 1866 

Pheretima calif ornica Kinberg. 1866 

Green Mountain, 30 August 1962, (3?)-l-6. John Packer per 
A. Loveridge. 

Pheretima hawayana ( Rosa. 1891 ) 

Green Mountain. 30 August. \ ( )(>2. (3?)-l-l. John Packer 
per A. Loveridge. 

Pheretima morrisi ( Beddard, 1892) 

Three oi the tour specimens of this species previously available 
(Sims. 1964) were from garden soil. 

OC \l RODRI1 ID \l 

Green Mountain. 3d August, 1962, 2-0-0. John Parker per 
\ I overidge. 

Setae, closelv paired. Gizzards, probabl) lacking. Calciferous 
glands, long, seemingl) extending through ix-\ deeply constricted h\ 

9 10. opening into the gut posteriorly in \, the anterior half more 
firm. Prostates, one pair. \erv long, external apertures unrecog- 
nizable. Spermathecae, non<: found. 

rhese worms (cfi. 1mm thick) were so softened that further 
information was unobtainable. No Other ocncroJnlc is known to 
have glands of the kind indicated above. Although reference to 
any known genus is impossible, the worms obviousl) were ocnero- 
drile. 

Remarks. The pheretimas are widely distributed anthropochores. 
I hey almost certainly were brought from the Orient to the island, 
directly or indirectly, by man. Any lumbricids on the island 
probably were brought, also by man. from England. Evidence to 
be presented elsewhere suggests that the pheretimas may be better 
adapted than the lumbricids to the climates o\' tropical islands such 
as Ascension. 

/'. californica and hawayana have been collected on St. Helena, 
the nearest land. /'. morrisi was once recorded from St. Helena but 
was not present in any of the various collections which were made 
on that island b\ Loveridge and others during the last twenty 
scars. 



1969 EARTHWORMS OF TWO OCEANIC ISLANDS 3 

REFERENCE 

Sims, R. W. 

1964. Oligochaeta from Ascension Island and Sierra Leone including 
records of Pheretima and new species of Dichogaster. Ann. Mag. 
Nat. Hist.. (13) 7: 107-113. 

THE JUAN FERNANDEZ ISLANDS 

The Juan Fernandez islands, all small, are three: Mas a tierra 
(36 square miles), Mas Afuera (33 square miles), and Santa 
Clara. The first is inhabited. The islands were discovered by the 
man for whom they are named in 1563. The first settlement to 
become permanent was in 1877. 

Prior to Dr. Y. Kondo's visit, four species had been recorded 
from Juan Fernandez: Allolobophora caliginosa (Savigny, 1826), 
Dendrobaena rubida (Savigny, 1826), Eiseniella tetraedra 
(Savigny, 1826), Eukerria saltensis (Beddard, 1895). In the ab- 
sence of any indication to the contrary, all presumably were found 
on Mas a tierra. Collections made by Dr. Kondo on Mas a tierra, 
enable the listing of three more species. 

LUMBRIC1DAE 

Allolobophora Eisen, 1874 
A llolobophora tuberculata Eisen, 1 894 

Quebrada, Portozuela East, 200m, 2.xi.l965, 4 (+25?) -5-2. 
No. 31, 11. xi. 1966, 5-0-1. 

A. tuberculata is one of at least four species long retained in a 
classical congeries known as A. caliginosa. Worms previously re- 
corded from Juan Fernandez as A . caliginosa may well have been 
A. tuberculata. 

Dendrobaena Eisen, 1874 
Dendrobaena octaedra (Savigny, 1826) 

No. 31, 11. xi. 1966, 1-0-2. 

Dendrobaena sp. 

Asondalan, 29. x. 1966, 1-0-0. 

This small juvenile is not of D. octaedra, but its condition was 
too poor to permit a positive identification, and it may be of D. 
rubida (Savigny, 1826), previously recorded from the island. 



breviora No. 323 



Octolasion Oerley, 1 885 
Octolasion tyrtaeum (Savigny, 1826) 

No. 31, 11. xi. 1966. 2-0-1. 

MON1LIGASTRIDAE 

Drawida Michaelsen, 19(H) 

Drawida bahamensis (Beddard, 1892) 

No. 31. 11. xi. 1966. 0-1-1. 

The left copulatory chamber, of one of the posterior amputees. 
is everted and the penis is protruded. Apertures of the other three 
chambers are tightly closed. 

Gizzards, in \ii-\iv ( 1 ). in xiii-xiv ( I ). Ovisacs, confined to xii. 

DISCUSSION 

Mas a tierra is only two square miles smaller than Ascension. 
Both islands are in the Southern Hemisphere. The Atlantic island 
has a known megadrile fauna of live species, and the Pacific island 
has seven: each island has one ocnerodrile. E. \altcn\i\ originated 
in southern South America, but in which part is unknown. The 
Ascension ocnerodrile could have evolved m South America or 
even in Africa. Ascension pheretimas. like the San Juan monili- 
v:astrid, originalK were from the Orient, l.umbricids. of course, 
came from Europe. 

Each species of the two islands is more or less widely distributed 
throughout the world. Each very probably was brought to the 
islands accidentally by man. Lumbricids. so far as can be indi- 
cated b\ the samples, seem to be dominant on Mas a tierra. 
Megascolecids. according to a smaller sample, seem to be dominant 
on Ascension. The Juan Fernandez Islands are well south of the 
tropics. Ascension well within them. The dilTerence in dominance 
may. then, be due to climatic factors. Nevertheless, the pheretimas. 
like the lumbricids. probably are from the Temperate /one of 
the Northern Hemisphere. 

( Received 4 March 1969.) 



BREVIORA 

Muscimol of Comparative Zoology 

Cambridge, Mass. 15 September, 1969 Number 324 



POLYMORPHISM AND EVOLUTION 
OF THE HISPANIOLAN SNAKE GENUS UROMACER 

(COLUBRIDAE) 

Henry S. Horn 1 

Abstract. Four species of the endemic genus Uromacer are recognized: 
catesbyi, fremitus, oxyrhynchus, and dorsalis. All save dorsalis show a re- 
markably discrete di- or trimorphism in a color pattern that is apparently 
cryptic. The adaptive significance of this polymorphism is discussed using an 
intuitive model that draws on the theory of Levins. 

The species are linearly ordered on the basis of body shape. Surprisingly, 
the stomachs of the stoutest species contain arboreal lizards and frogs, while 
the slimmer species take progressively more terrestrial lizards. The linear 
order suggests a pattern of geographic speciation occurring at the time when 
the northern and southern portions of Hispaniola were separate islands. Thus 
the pattern is consistent with patterns of evolution and migration proposed 
for other Hispaniolan reptiles by Williams and others. Speculations about 
the origin of dorsalis, endemic to He de la Gonave, may explain why the 
fauna of that islet is a curious mixture of species characteristic of the northern 
and southern portions of Hispaniola. 

INTRODUCTION 

The snakes of the genus Uromacer are endemic to Hispaniola 
and the surrounding islets. Some years ago A. Stanley Rand dis- 
covered a color polymorphism in several local populations of 
Uromacer oxyrhynchus. Examination of the other species has 
shown that several of them are also polymorphic. Since the same 
morphs of different species are superficially more similar than are 
different morphs of the same species, this polymorphism sometimes 
results in the misidentification of museum specimens. Conse- 
quently, the polymorphism has obscured the pattern of distribution 

'Department of Biology, Princeton University, Princeton, New Jersey. 



2 BREVIORA No. 324 

of the Forms of I 'romacer, making it difficult to relate their distribu- 
tion to that o\' other Hispaniolan reptiles. 

The recent concentration on the herpctologv of Hispaniola at 
the Museum of Comparative Zoolog) (MCZ) has provided large, 
synchronous collections of U romacer from several localities. The 
geographical coverage of even these large collections is not yet 
extensive enough to warrant a definitive discussion o\' intraspeciflc 
variation. However, the available collections allow me to assess 
the validity oi the described species of I romacer, to suggest the 
pattern of their relationships, to document their striking poly- 
morphism in some detail, and to comment on some interesting 
evolutionary problems posed by the genus. Though I have studied 
only preser\ed material. I have made inferences about the eeolog\ 
oi the species from their stomach contents. However, I must leave 
several questions unanswered because I lack field knowledge of 
Hispaniola. particularly o\ seasonal changes during the year. 

Where \1( / material was insufficient, I have examined speci- 
mens from the American Museum ol Natural Histor\ ( AMNH), 
the British Museum (Natural Histor\ ) (BMNH), the Field Mu- 
seum o\ Natural Histor\ (I \1NII). the Yale Peabod) Museum 
(YPM), the Albert Schwartz Field Series ( ASFS). and the United 
States National Museum (USNM). I am grateful to the curators 
o\ these collections for allowing me to examine specimens in 
their charge. 

I thank A. Stanle\ Kand and Ernest E. Williams tor discussion 
and criticism throughout this stud\. Dr. Rand introduced me to 
{J romacer, and his unpublished notes on their polymorphism 
were the skirting point for m\ work. Dr. Williams sponsored part 
of this stud) as m\ undergraduate diesis at Harvard College in 
1962. \lv ideas ^w polymorphism developed from discussions 
with Richard Levins and Robert II Mac Arthur. For criticisms o\' 
several manuscripts. I am indebted to I nomas H. Irazzetta. Eliza- 
beth G. Horn. Thomas W. Schoener, Albert Schwartz, and George 
(i. Simpson. Dr. Schwartz kindlv compiled the fauna! list for 
lie dc la Gonave. 

MATERIAL AND METHODS 

Cochran (1941 ) recognized six species of V romacer: catesbyi, 
scandax, frenatus, wetmorei, dorsalis, and oxyrhynchus. There is 
reason to doubt the specific distinctness o\ scandal and wetmorei, 
which Cochran recognized as respectively allied to catesbyi and 
frenatus. For reasons given in the Appendix. I consider scandax 



1969 



EVOLUTION OF UROMACER 



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4 bki \ [OR \ No. 324 

to be identical with catesbyi, and wetmorei to be a subspecies of 
frenatus. The remaining species are easily separated by the 
diagnostic characters ot Cochran's key: snout length relative to 
eye diameter, ventral scale count, and midbody scale rows. The 
ranges of snout length and ventral scale count overlap between 
species, but these characters arc usually sufficient tor tentative 
identification ot" single specimens. I he number of scale rows at 
midbody is constant in each species, but is diagnostic only for 
oxyrhynchus. The subcaudal scale count for each species has a 
laree range, and though there may be significant differences be- 
tween the means oi different populations, it is seldom useful in 
identifying a single specimen. Similarly, the presence or absence 
of an anterior loreal ma\ be used to infer relationships among 
populations, but it is useless as a diagnostic character. 

Stomach contents of the MCZ and YPM specimens were re- 
moved and identified. I he snakes had fed primarily 00 Anolis 
whose ecolog) is rclati\el\ well known, so inferences about the 
ecolog) of Uromacer ma) be made on the basis of these stomach 
contents. Schwartz (in litt. ) and Merteiis (1939) have observed 
that freshly caught Uromacer often swallow the cohabitants of a 
collecting bag. but even those stomach contents that were not 
eaten by free snakes will retleet the syntopic fauna in which the 
snakes were collected. To minimize this difficulty, those stomach 
contents that were undigested were ignored in analysis. 

Where different color morphs are described for a single species 
of Uromacer, these morphs are entirely distinct, with no inter- 
mediates except between the gra\ and the striped gray forms. 
There was never any hesitation in assigning any well-preserved 
specimen to one o\ three alternative morphs: green, green above 
and below, lighter below; gray-green, green above, mottled gray 
below; or gra) . gray above and below. A detailed description of 
each morph and its variations in each species is given in the 
Appendix. 

RESULTS AND INTERPRETATIONS 

The ranges of the species of Uromacer are mapped in Figure 1 
and listed in Table 1 . 

External metric characters are listed in Table 2, and there are- 
photographs of the several species in Figure 2. The species may 
be linearly ordered, as in the table, on the basis of characters that 
are correlated with slimming and lengthening the body. U. catesbyi 
is typically colubrid in shape, with a short, blunt snout, low 



969 



EVOLUTION OF UROMACER 



TABLE 1 
Ranges of the species of Uromacer 



Species 



Southwest and 
Barahona 
Peninsulas 



Port-au-Prince 
and vicinity 



N . and E. lie de la Gondve 
Dominican 
Republic 



cateshyi 
frenatus 
oxyrhynchus 

dorsalis 



TABLE 2 
Metric morphology of the species of Uromacer 



Snout length 
relative 
Species Number to eye Ventral Subcaudal 
examined diameter scales scales 



Scale 

rows at Anterior 
midbody loreal 



catesbyi 254 



2.0 167 ±0.7 183 ±1.0 



17 — 



frenatus 102 2.6 187 ±0.5 204 ±1.9 



oxyrhynchus 129 



3.0 200 ±0.5 199 ±1.1 



17 



19 present in 25 



dorsalis 



84 



2.9 200 ±1.2 210 ±2.2 



17 



+ 



Entries are the means for each character ± the standard error of the mean. 
The standard error of the mean snout length (±0.01) was, in all cases, less 
than the reading error of each measurement (±0.2). Numbers separated by 
* are significantly different at the .05 level by a one-tailed t test (Snedecor, 
1956), for snout length, ventral count, and subcaudal count, or a binomial 
test (Siegel, 1956), for scale rows at midbody and anterior loreal. 



HRI \I()K\ 



No. 324 








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IMIIIlit 

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IMillllt 



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1 969 EVOLUTION OF UROMACER 



Figure 2. a, Uromacer catesbyi MCZ 57764 nr. Carpintero, Dominican 
Republic; b, U. frenatus MCZ 65530 Perrine nr. Jeremie, Haiti; c, U. oxyr- 
hynchus MCZ 62713 Bombardopolis, Haiti; d, V. dorsalis MCZ 80815 
Pointe-a-Raquettes, He de la Gonave, Haiti. 



3 



imi \ I or \ 



No. 324 



ventral and subcaudal counts, and a relatively thick bod\ covered 
with rhomboidal scales that arc nearl) square. U. oxyrhynchui 

and dorsalis arc shaped more like typical tree and vine snakes. 
with long, pointed snouts, high ventral and subcaudal counts, and 
more slender bodies covered with obliquely rhomboidal scales. 
U. frenatus is intermediate between these extremes. 

The stomach contents of I romacer are listed in Tables 3 and 3a. 
In Table 3 the stomach contents of the snakes are pooled within 



TABLE 3 
Habitat of food taken by the species of I romacer 

Habitat catesbyi fremitus oxyrhynchtu dorsalis 

Trees 

I ree trunks and bushes >» 

( n.iss and open areas 
( i round and rocks 



3 


1 


2 


5 


II 


4 


) 


4 


II 


4 


4 


10 



TABLE 3a 

Stomach c intents ol the species of i 'romacer 
Habitat and species eaten <<//<s/>\; frenatus oxyrhynchus 



ilor\tili\ 



rREES 










ll\ln puh hrilineata 


1 








do/ninh ensis 


I 








I noli\ i hlorocyanui 


5 




1 


2 


( oelestinus 


1 


3 






1 Rl K TRUNKS \ Bl sill s 










A nulls cy botes 


4 


4 


8 


3 


hrc\ iros'i s 








1 


distil hits 


5 


1 


3 




(.K \Ss,V Ol'l N \RI \S 










A nulis semilineatus 




i 


3 




olssoni 






1 


11 


(.ROUND& ROCKS 










Ameiva taeniura 




1 


: 


1 


chrysolaema 








ft 


Leiocephalus melanochlorus 




i 






vinculum 








3 


schreibersii 






: 




Sphaerodactylus sp. 




1 







1969 EVOLUTION OF UROMACER 9 

common habitats of the food species; the habitat groupings were 
made by referring to Mertens (1939), Williams and Rand (1961), 
and Rand (1962). Since the groupings are only generalizations, 
and the samples are small and heterogeneous, the data do not 
fully characterize the diets of the species of Uromacer. However, 
the differences among the diets are so striking, when compared 
to the uniformity of diets within each species, that crude relative 
characterizations can be made with confidence. 

U . catesbyi takes arboreal food almost exclusively, and dorsalis 
feeds primarily on terrestrial food, whereas /. jrenatus and oxyrhyn- 
chus are intermediate and more similar to each other than either 
is to catesbyi. The tendency for the colubrid-like catesbyi to take 
arboreal prey, and for the longer-snouted species to take pro- 
gressively greater proportions of terrestrial prey, is the opposite 
of expectations based on the morphology of the species (e.g. 
Mertens, 1939, who suggest that oxyrhynchus is the most arboreal 
species of Uromacer) . 

Polymorphism. Of the four species of Uromacer, only the gray 
dorsalis is not demonstrably polymorphic in at least one locality 
(see Table 4). Green U. catesbyi occur throughout the range of 
the species, but with a pattern unlike the corresponding morph 

TABLE 4 
Distribution of the color morphs of the species of Uromacer 

Species Green Gray-green Gray Gray (striped) 

catesbyi ENTIRE RANGE Miragoane 

/. jrenatus He a Vache mainland He a Vache 

/. wetmorei Barahona Isla Beata 

oxyrhynchus Haiti & Haiti 

ENTIRE RANGE Eastern D.R. 
dorsalis ^ENTIRE RANGE 

of any other species; catesbyi is polymorphic, with a gray form, 
near Miragoane, in the middle of the southwest peninsula of Haiti. 
U. f. jrenatus is monomorphic gray on the mainland, and poly- 
morphic gray and gray-green on lie a Vache. The gray form on 
lie a Vache resembles that of dorsalis more than that of mainland 
jrenatus or oxyrhynchus. U . j. wetmorei on Barahona may be 
polymorphic, but the few poorly-preserved specimens presently 
available from scattered localities are insufficient evidence. 



10 BR] V ioka No. 324 

V. oxyrhynchus is apparently monomorphic green through much 
of the Dominican Republic, but it is dimorphic green and gray- 
green throughout the Haitian portion of its range. In scattered 
Haitian localities it is trimorphic. green, gray-green, and gray. 
U. oxyrhynchus may in fact be trimorphic throughout the Haitian 
portion of its range, perhaps in parts of the Dominican Republic 
as well. The gray morphs are rare enough to be found only in 
large, synchronous collections from Haiti; no such collections have 
been made in the Dominican Republic. 

The discreteness ol the morphs in all the polymorphic species, 
and the lack of intermediates, suggests that the mechanism con- 
trolling the polymorphism is an allelic switch at a single genetic 
locus. In the trimorphic oxyrhynchus, die distribution of the 
morphs is consistent with the interpretation that the green and the 
gray tonus are homozygotes and that the commoner gray-green 
form is the heterozygote; i.e. in all the localities from which both 
green and gra\ specimens have been taken. gra\ -green specimens 
have been taken as well. 

I he distribution ol the color forms among the localities of the 
polymorphic species and among the species of the genus suggests 
that the various tonus are cryptic tor different surroundings. U. 
catesbyi is green throughout its range, darker m the western por- 
tions ol Hispaniola than in the eastern, with some gray individuals 
on the southwest peninsula in the Miragoanc area. U. frenatUS 
is gray 0:1 the southwest peninsula and polymorphic on lie a \ ache, 
while its ecological counterpart, oxyrhynchus, has green, gray- 
green, and gra\ morphs in Haiti, but a preponderance o\ green 
morphs in the Dominican Republic (see I able 5 ). I hese distribu- 
tions correspond roughh to the pattern of humidit) and vegetation 
on Hispaniola. as described b\ Wetmorc and Swales ( 1931 : 2-7). 

TABLE 5 

Geographical distribution of the color morphs of frenatUS 

and oxyrhynchus 



S/>< i ;, \ 


1 rea 


Gret n 


( ,r,r -gn • 1, 


Gray 


/. fremitus 


lie .1 \ ache 




15 


22 


f. frenatiis 


Haitian mainland 




2 


56 


f. wetmorei 


Barahona 




4 


1 


oxyrhynchus 


Haiti 


47 


48 


4 


oxyrhynchus 


Dominican Republic 


17 


4 





1969 EVOLUTION OF UROMACER 1 1 

The grayer and darker (i.e. less bright green) forms are associated 
with the drier, western portion of the island. The presence of the 
gray-green morph of fremitus on He a Vache and the brighter green 
of the catesbyi from the islet are correlated with the presence of 
denser woodland on the islet than on the adjacent mainland. This 
correlation of the distribution of the color morphs with foliage 
density suggests cryptic coloration as a possible function of the 
different morphs. 

The two most strictly monomorphic species, catesbyi and 
dorsalis, are characterized by the most restricted diets. The green 
catesbyi takes only arboreal prey, while the gray dorsalis takes 
predominantly terrestrial prey. This additional correlation of 
color pattern with ecology suggests again that the coloration of 
Vromacer is cryptic, concealing the snake as it stalks agile prey. 

Polymorphism in a cryptic pattern of coloration poses an interest- 
ing evolutionary question. If a specific color pattern is indeed 
cryptic, why is more than one pattern present at all? The theory 
of polymorphism provides some likely answers and suggests the 
kinds of data that should be sought to choose between alternatives. 
Levins (1962-6; Levins and MacArthur, 1966) has examined 
the adaptive significance of polymorphism, using an abstract mathe- 
matical model of extreme generality. I shall develop a much 
simpler model to examine polymorphism in the special case of 
cryptic coloration, but the intellectual basis and results are the 
same as those of Levins. This model is almost absurdly simple, 
but the concepts illustrated are quite general and are not easily 
apparent without the model. 

The environment is a checkerboard with black and white 
squares and the cryptic animals are checkers. Later I shall intro- 
duce reproduction and "fitness" into the model, but first let us 
look at crypticity alone. If we randomly place black or white 
checkers on the squares of the checkerboard, roughly half the 
black checkers will fall on black squares and half the white 
checkers, on white. Thus half the population of either black or 
white checkers will be cryptic. However, if we throw gray checkers 
on the checkerboard, none of them will be cryptic, unless, of 
course, the checkers are very large and the squares of the checker- 
board are so tiny that it looks like the gray of a halftone illustra- 
tion. Thus if two alternative morphs are to be cryptic, the environ- 
ment must be a mixture of large patches of different kinds, each 
with an extreme morph that is more cryptic than an intermediate 
morph. 



12 BREN IORA No. 324 

I assume that "adaptation" is achieved through natural selection, 
which favors that individual whose offspring form the greatest 
proportion o\' breeders in the next generation. Correspondingly 
the "fitness" of an individual is defined as the Dumber o\' his 
offspring that breed in the next generation, and the individual 
with the greatest fitness in a given environment is the "'best 
adapted*' to that environment. 

We inns allow each checker to reproduce an average of X 
offspring in each generation; in each reproductive period we 
throw checkers onto the board and give each checker X+d oil- 
spring ii it is cryptic Inn only X d offspring if it is not cryptic. 
Black checkers have black offspring; white checkers have white 
offspring. We define "polymorphic" checkers to have either black 
or white offspring with a 50-50 chance, whether the phcnolvpc 
of the "polymorphic" checker is black or white. I he genetic or 
developmental basis of this polymorphism need not be specified, 
so I shall call it '"sibling polymorphism," since it requires that 
the offspring of a given morph maj include both morphs. If the 
checkerboard has an equal number of black and white squares, 
then the offspring o\ black, while, or poKmorphic checkers have 
an equal chance of being crvptic or non-cr\ptic. and the fitness 
dt each type is an average of X • d and \ d. or simpK X. (You 
can easil) s how that it the environment is an uneven mixture of 
black and white squares, the morph that is cryptic in the commoner 
square has a higher fitness than the other morph or the poly- 
morph. ) I hus, it the environment is a spatial mixture of large 
patches of two types, a monomorphic population is better adapted 
than a poKmorphic population, or at least as well adapted. 

If, instead o\ throwing the checkers onto the board randomly. 
we allow each ot the checkers to choose and reproduce in its 
appropriate habitat, then the white squares will be occupied pre- 
dominant!) b\ white checkers and the black squares by black 
checkers. On sampling a large and heterogeneous area, we then 
obtain both morphs. and the population appears to be polymorphic. 
However, the two morphs could represent two species, rather than 
a poKmorphic species. In tact, the average fitness of individuals 
of the two separate species would be higher than that of a poly- 
morphic species, unless the polymorphism continually adjusted 
the ratio of the morphs to match the proportions of the corre- 
sponding patches in the environment. Since this polymorphism 
ma\ result from sampling a heterogeneous environment and does 



1969 EVOLUTION OF UROMACER 13 

not require that a given individual be capable of having poly- 
morphic offspring, I shall call it "sampling polymorphism' 1 as dis- 
tinct from "sibling polymorphism." (Under certain restricted con- 
ditions the two kinds of polymorphism are convergent, and sam- 
pling polymorphism may evolve a genetic basis. See Levins, 1965, 
for a discussion.) 

Now instead of throwing the checkers onto a checkerboard, we 
throw them alternately onto a white board or a black board for 
each generation. That is, the environment is a temporal mixture 
of different habitats, rather than a spatial mixture. The black 
and white checkers are alternately cryptic and non-cryptic. Each 
individual in a cryptic generation has X+d offspring, each of whom 
has X— d offspring in the next generation; thus the average fitness 
of black or white checkers over two generations is (X-fd)(X— d) 
= X-— d-. On the other hand, the "polymorphic" checkers have 
an equal number of cryptic and non-cryptic offspring in each 
generation and their average fitness is V2 (X+d) + '/2 (X— d) =X 
in each generation; then the average fitness of polymorphic check- 
ers over two generations is X 2 , which is always greater than 
X 2 — d-. If the black and white environments alternate several 
times within each generation, fitness is averaged over the two 
environments for each generation, and the results are the same as 
in the case of spatial heterogeneity. However, sibling polymorph- 
ism yields a higher average fitness than monomorphism when the 
environment varies drastically with a period of the same order 
of magnitude as the time between generations. 

What does the checkerboard model predict about Uromacer! 
If the polymorphism of Uromacer is simply a sampling poly- 
morphism, then the different morphs should be associated with 
distinctly different habitats or microhabitats. The habitats might 
be different types of vegetation, in each of which one of the 
morphs is most cryptic. 

If the polymorphism of Uromacer is a sibling polymorphism, 
the alternative morphs should still be cryptic in distinctly different 
habitats, but the habitats will alternate in an inescapable temporal 
pattern, e.g. wet-dry season. Therefore, if the polymorphism is a 
genetic or developmental adaptation to a temporally heterogeneous 
environment, the different morphs will often be found in habitats 
in which they are not cryptic. 

Documenting the adaptive significance of polymorphism in 
Uromacer is a difficult, long-term project. It is first of all a field, 
not a museum project. It is necessary to define the habitats in 



14 hkiaiora No. 324 

which each of the morphs is most cryptic. Then the alternatives 
ot genetic and sampling polymorphism must he separated by deter- 
mining the distribution of the morphs among the several habitats. 
over a time equal to several generations oii Uromacer. As interest- 
ing as the problem is. it will probabl) not be quickly solved, per- 
haps not even quickly undertaken, since the required study is 
longer than the lifetime of the average research grant. 

Relationships within the genus. I here has been no previous dis- 
cussion oi the relationships oi the species oi Uromacer. 

It is at once tempting to suggest that the morphological and 
dietarv series described corresponds to an evolutionary series as 
well; that catesbyi is the most primitive of the Uromacer, oxyrhyn- 
chus and dorsalis the most recently derived, and that frenatus 
represents an intermediate step. When the present ranges o^i the 
species, the physiographic historj of Hispaniola, and the mechan- 
isms ol geographic speeiation are taken into account, this sugges- 
tion is strengthened. 

During the Pleistocene. Hispaniola was divided by an arm of 
the sea into two large islands, a northern island. Haiti north o\ 
the Cul-de-SaC Plain and the Dominican Republic exclusive of 
the Barahona peninsula, and a southern island, the southwest 
peninsula o\ Haiti and the Barahona peninsula ( Schuchcrt. 1935; 
Weyl, L966; Williams. 1961). I he ancestral colubrid ( Maglio, 
in prep., suueests something similar to Alsophis) had given rise 
to a catesbyi-like form, to be called "proto-catesbyi," and to 
"proto-frenatus." autochthons of the northern and southern is- 
lands respectively. Proto-frenatus invaded the northern island, 
and as it came into contact with the well-established proto-catesbyi. 
the characteristics that distinguished the tWO species were inten- 
sified (character divergence o\ \la\r. 1963. after Darwin). By 
its increased divergence from proto-catesbyi. the invading proto- 
Erenatus gave rise to 'proto-oxyrhynchus.'" ancestor oi oxyrhyn- 
chlU and dorsalis. 

I he proposed sequence accounts for the presence in dorsalis 
of characters similar to both frenatus and o.wrhxnehus. The pro- 
genitor of dorsalis on Gonavc was presumably proto-oxyrhynchus. 
which would have retained similarities to proto-frenatus and thus 
to frenatus. while differentiating in the direction of o.xyrhynchus. 
Alternatively, dorsalis could have been derived directly from 
proto-frenatus. and its similarity to o.xyrhynchus could represent 
parallel evolution of the species. Lack of evidence relevant to 
these alternatives is in itself sufficient reason for provisionally call- 
ing dorsalis a separate species. 



1969 EVOLUTION OF UROMACER 15 

This evolutionary sequence is only an interpretive hypothesis 
and cannot be tested, but it embodies processes that have been 
suggested to account for the present distribution of other His- 
paniolan reptiles. The postulated distinction of proto-catesbyi and 
proto-frenatus, as the respective autochthons of the northern and 
southern islands, is paralleled by the distinction of Anolis coeles- 
tinus and Anolis chlorocyanus, of Anolis aliniger and Anolis 
singularis, and perhaps of the Anolis of the monticola group and 
the subspecies of Anolis ricordii (Thomas and Schwartz, 1967; 
Williams 1965a, 1965b). The subsequent invasion of the northern 
island by proto-frenatus is paralleled by the Anolis of the setni- 
lineatus group. Williams (1961) suggests that Anolis olssoni and 
A. semilineatus are indigenous to the northern and southern islands 
respectively and that their present distributions are the result of 
the relatively recent invasion of the northern land mass by semi- 
lineatus. Similarly, Schwartz ( 1967) suggests that Ameiva taeniura 
originated on the southern island and has since invaded the 
northern land mass. 

There are probably other patterns of radiation that are con- 
sistent with the present ranges of the species of Uromacer, and 
almost as simple in the changes of morphology that they would 
require. However, several conclusions will remain unchanged 
given such alternative patterns. U. catesbyi is clearly more differ- 
ent, in morphology and diet, from the other species than they 
are from one another. It is then significant that only catesbyi is 
broadly sympatric with the other species of the genus. U. frenatus, 
oxyrhynchus , and dorsalis are similar in morphology and may 
provisionally be considered allopatric ecological replacements. 

Explaining the origin of dorsalis on Gonave as more than a 
historical accident requires bold speculation in the absence of 
critical data. This speculation is justified, however, as it also 
suggests a possible derivation for the problematical fauna of 
Gonave. 

Present knowledge of the fauna of Gonave is scanty, but it is 
already apparent that the fauna is an odd mixture of species 
characteristic of the northern and southern islands of the Pleisto- 
cene (See Table 6). During part of the Pleistocene, Gonave was 
reduced to about one fourth of its present area (Butterlin, 1960). 
When the northern and southern islands were rejoined, Gonave 
may have been connected to the mainland by a spit at least 3 km 
wide to the north and 7 km wide to the south (Fig. 1; also Wood- 
ring et al., 1924), which would have allowed the fauna of both 
regions to invade Gonave. When Gonave became insular again, 



16 



KKl \ [ORA 



No. 324 



those species whose presence had depended on continuing invasion 
would he eliminated, leaving only those best adapted to the 
restricted habitats of Gonave and those potential competitors that 
were most divergent. The geographic origin of the invading species 
would be irrelevant to their subsequent persistence. 



TABLE 6 
Reptiles of Gonave and their areas of origin 



\/'ci /< \ 



Origin 



Gonatodi i albogularis 
Sphaerodat tylus < inereus 

fteliigt r i ci hranae 
Audits brevirostris (end. subsp i 

( hlorot yanus 

i \ bott I 

olsoni 

singularis 

( m luni ( hi nnlti 

1 1 ocephalus vinculum (end subsp i 



North 

North 
South 



Reference 



Williams. 1965 a 

Williams. 1961 
Williams. 1965a 



Ameiva < (try solai ma ( end. 
taeniura 


sub* 


».) 


North 
South 2 


Schwai t/ & 

Klinikowski, 1966 
v wartz, 1967 


Diploglossus curtissi 
Amphisbaena gonavensis i 


end 


siihsp. ) 


South ' 


Ihomas. 1965 


/ yphlops capitulatus (end. 
pusillus 


sub 


sp.) 


North 


Richmond. 1964 


Eph rates angulifer 
fordi 










I ropidophis haetianus 










Dromicus parvifrom 










Hypsirhynchus ferox 










laltris dorsalii 










Uromacer i atesbyi 

dorsalii < endemic I 




North - 
North 





1 North and south under this heading refer to the northern and southern 
islands into which Hispaniola was periodical divided during the Pleistocene. 

See Fig. 1 . 

-' Island of origin of parent species, though invasion may have been from 
a population on the other island. 



1969 EVOLUTION OF UROMACER 17 

In particular, if proto-catesbyi, proto-frenatus, and proto- 
oxyrhynchus all reached peninsular Gonave, the ultimate per- 
sistence of the most divergent pair, and the elimination of the 
intermediate proto-frenatus is predictable. The dietary divergence 
of dorsalis and catesbyi confirms the role of interspecific competi- 
tion in the origin of dorsalis. U. dorsalis clearly differs from 
catesbyi in foraging habitat, if not in foraging behavior, to a 
greater extent than jrenatus and oxyrhynchus differ from catesbyi 
(Table 3). Interspecific competition may necessitate this increased 
divergence of dorsalis and catesbyi on Gonave, whereas less diver- 
gence insures the coexistence of two sympatric Uromacer on the 
more extensive and heterogeneous Haitian mainland. 

If the history above is wrong, Gonave deserves intensive study 
as an extraordinary example of an island colonized from two com- 
parable and equidistant sources. If the suggested history is cor- 
rect, then Gonave is a model of the faunal transformations of a 
continental island that when peninsular had the full complement 
of species of the adjacent mainland: while several closely related 
species could persist when immigration from the mainland was 
easy, fewer were able to persist after their populations became 
insular. Those that did persist on the island had to be strongly 
divergent ecologically. According to such a model, an interpreta- 
tion of the fauna of a continental island must emphasize the eco- 
logical compatibilities of the potentially resident species, rather 
than their relative vagilities and routes of invasion from the ad- 
jacent mainland. 

LITERATURE CITED 

BUTTERLIN, J. 

1960. Geologie generate et regionale de la Republique d'Haiti. Travaux 
et Memoires de ITnstitut des Hautes Etudes de l'Amerique 
Latine, Paris, 6, 194 pp. 
Cochran, D. 

1931. New reptiles from Beata Island, Dominican Republic. Proc. 

Biol. Soc. Washington. 44: 89-92. 
1941. The herpetology of Hispaniola. Bull. U.S. Nat. Mus.. 177: 
1-398. 
Dunn, E. R. 

1 920. On the Haitian snakes of the genera Leimadophis and Uromacer. 
Proc. New England Zool. Club, 7: 37-44. 
Levins, R. 

1962. Theory of fitness in a heterogeneous environment. I. The fitness 
set and adaptive function. Amer. Nat., 96: 361-373. 



IS hki \ IORA No. 324 

1963 rheorj ol fitness in .1 heterogeneous environment. II Develop 
menial flexibility and niche selection, \nu-i Nat., *>7: 75-40. 

1964. rheon of fitness in .1 heterogeneous environment. 111. ["he re- 
sponse to selection. J. rheoret. Biol . 7 224 240. 

I9h5 ITieory of litness in a heterogeneous environment l\ l he 
adaptive significance ol gene H iw I volution, IK: 635 638 

1966. I he strategy of model building in population biolog) \mei 

Sci . •"> I: 421-431. 

I I \ INS. K . \M> R II M \< \KI III K 

I9hh. The maintenance of genetic polymorphism in .1 spatially hetero- 
geneous environment: variations on a theme by Howard 1 evene 
\nu-i Nal . 100 585 5J 
Mayr, I 

1963 XnniKil Species and Evolution Cambridge, Mass., Belknap 
Press, Harvard I Diversity Press, 797 pp. 

Mertens, R 

193 ' Herpetologische I rgebnisse einei reise nach der Insel Hispaniola, 
Westindien. \bh. Senckenberg N.iiuit Ges., 149: 1 84 
Rand, \ S 

I l >(i2 Notes Dii Hispaniolan herpetology 5. I Ik- natural history 
ol three sympatric species of Anolis, Breviora, Mus ( omp. 
Zool . No 154 1-15 

Rll HMOND, N H 

1964 I he bliod snakes 1 / yphlopa 1 of Haiti with descriptions of three 
new species. Bre\ a Mus I omp Zool., No. 202: 1-12. 

S< HI l III Kl . ( 

1935 Historical Geology ol the ^ntillean-Caribbean Region. New 
> oik. John w iley & Sons. 81 1 pp 

S( HW VKI/. \ 

1965 I Ik- Leiocephalus il acertilia, Iguanidae) of Hispaniola. I Leio- 
cephalus meianochlorus < ope. lourn. Ohio Herp, Soc., 5: 39-48. 

1967. The Ameiva (Lacertilia, reiidae) ol Hispaniola. Ill Ameiva 
taeniura Cope. Bull. Mus. (omp Zool., 135 $45-375. 

Si HW \RI/. \ . \N|) K 1 Kl INIKOWSK1 

1966 I Ik- Ameiva il acertilia, reiidae) of Hispaniola. II. Geographic 
variation in Ameiva chrysolaema ( ope Bull. Mus (omp. Zool., 
133: 427-4$ 

Siegi 1 . S 

1956 Nonparametric Statistics for the Behavioral Sciences. New 
York, McGraw-Hill, 312 pp. 

SM Dl ( ciR. (1 \V. 

1956. Statistical Methods \pplieil to I uperiments in Agriculture and 
Biology. Vines, Iowa, Iowa state Univ. Press. 534 pp. 
Thomas. R. 

1965. Two ne\s subspecies oj Amphisbaena (Amphisbaenia, Reptilia) 

from the Barahona Peninsula of Hispaniola Breviora, Mus 
Comp. Zool.. No. 215: 1-14 



1969 EVOLUTION OF UROMACER 19 

Thomas, R., and A. Schwartz 

1967. The monticola group of the lizard genus Anolis in Hispaniola. 
Breviora, Mus. Comp. Zool.. No. 261: 1-27. 
Wetmore, A., and B. H. Swales 

1931. The birds of Haiti and the Dominican Republic. Bull. U.S. Nat. 
Mus., 155: 1-483. 
Weyl, R. 

1966. Geologie der Antillien. Beitrage zur Regionalen Geologie der 
Erde, Vol. 4, Berlin, Gebruder Borntraeger, 410 pp. 
Williams, E. E. 

1961. Notes on Hispaniolan herpetology. 3. The evolution and rela- 
tionships of the Anolis semilineatus group. Breviora, Mus. 
Comp. Zool., No. 136: 1-8. 
1965a. The species of Hispaniolan green anoles (Sauria, Iguanidae). 

Breviora, Mus. Comp. Zool., No. 227: 1-16. 
1965b. Hispaniolan giant anoles (Sauria, Iguanidae): New data and a 
new subspecies. Breviora, Mus. Comp. Zool., No. 232: 1-7. 
Williams, E. E., and A. S. Rand 

1961. Notes on Hispaniolan herpetology. 2. A review of the Anolis 
semilineatus group with a description of Anolis cochranoe, new 
species. Breviora, Mus. Comp. Zool., No. 135: 1-11. 
Woodring, W. P., J. S. Brown, and W. S. Burbank 

1924. Geology of the Republic of Haiti. Rep. of Haiti Dept. of Public 
Works, Port-au-Prince, 631 pp. 

(Received 20 January 1969.) 



Appendix 
SYSTEMATICS 

The synonomy of each species cites only the original descrip- 
tion and additions to the synonomy of Cochran ( 1941 ). The data 
for the cited geographical range are given in Figure 1 . The extreme 
measurements of the snakes examined are given for each metric 
character; the standard error for each measurement is listed in 
Table 2. A list of all specimens examined, their localities and 
color morphs, and the stomach contents listed in Table 3 are filed 
with the specimens at the Museum of Comparative Zoology. 

Uromacer catesbyi (Schlegel) 

Dendrophis catesbyi Schlegel, 1837, Essai sur la physionomie des serpens, 

2: 226 (type loc. "L'ile de St. Dominque"). 
Uromacer scandax Dunn, 1920, Proc. New England Zool. Club, 7: 43 (type 

loc. Tortuga Island). 



Jo BRKVIORA No. 324 

Range. Hispaniola: the coastal area aorth of the Cul-de-Sac 
Plain, the southwest peninsula of Haiti, upland valleys of the in- 
terior of the island, lie a Yaehe. He de la Gonave, an J lie de la 
Tortue. It is sympatric with ever) other species in the genus. 

Diagnosis. Snout 2.0 (range: 1.7-2.7) times eye diameter; scale 
rows at midbody 17; ventrals lb7 (range: 160-180); subcaudals 
183 (range: 163-205); anterior loreal absent. 254 specimens 
were examined: 14 W1NH. 7 l > ASFS, 6 BMNil. 21 I'MNH, 
126 MCZ. 8 USNM. 

Color pattern. Green morph: dark green above, light yellow- 
green below, the colors meeting on the 3rd scale row; head dark 
green above, cream below, a light line above the eye and a dark 
line from the snout to the end o\' the head i alter M< / 64800). A 
few specimens have a light lateral line on the 2nd and or 3rd scale 
row rhe dorsal dark green varies geographically, from a dark. 
brownish green in western Haiti to a \er\ light green in the 

eastern Dominican Republic. Specimens from lie a \ ache are 
somewhat lighter than those from the adjacent mainland. 

Graj morph: gray above, cream below, the colors meeting on 
the 3rd scale row; head gray-green above, cream below, a light 

line above the eve (after \i( / 66350). I his morph has been 

found in 5 specimens from the Miragoane area. 

Nomenclature. Dunn (1920) defines scandal as "like Uro- 
macer catesbyi but with 19-11 scale rows and 181 ventrals." 

Examining the holotype (USNM 59438), I count 17 scale rows 
at midbody and 180 ventrals, with \ ( > scales around the neck 
persisting to within 5 em o\ midbody. A count oi \ ( ) scale rows 
persists for some distance down the body in six o{ the 254 
catesbyi examined: there are also three typical catesbyi from 
scattered localities with ventral counts of 177. 178. and 179. 
Two catesbyi typical in scale characters (NIC/. 37607, USNM 
59920) have been taken from He de la I ortue. but larger collec- 
tions ma\ reveal an endemic subspecies oi catesbyi for which 
the name scandal would be appropriate. 

Uro\i\( ik prenatus (Gunther) 

Ahaetulla frenala Gunther, 1X65. Ann. Mag. Nat. Hist.. Scr. 3. 15: 94 (type 

'oc. "no: known" ). 
Uromacer wetmorei Co:hran. 1931. Proc. Biol. Soc. Washington. 44: 91 
( type lot Isla Beata i 
Range. Southwest peninsula of Haiti east and north to the 
Cul-de-Sac Plain, lie a \ ache. Isla Beata. and the eastern coast 



1969 EVOLUTION OF UROMACER 21 

of Barahona Peninsula. Records outside this area have been 
based on misidentifications (Cochran, 1941). It is sympatric 
with catesbyi throughout its range and with oxyrhynchus at the 
base of the southwest peninsula. 

Diagnosis. Snout 2.6 (range: 2.1-3.0) times eye diameter; 
scale rows at midbody 17; ventrals 187 (range: 177-196); sub- 
caudals 204 (range: 185-212; but AMNH 51416 has 135 and its 
tail appears intact); anterior loreal absent. 102 specimens were 
examined: 4 AMNH, 39 ASFS, 2 BMNH, 46 MCZ, 11 USNM. 

Color pattern. Gray-green morph: Gray-green frenatus from 
He a Vache are dark blue (probably green in life) above, mottled 
gray to gray-brown below, the colors separated by a white line on 
the 2nd and 3rd scale rows from four head lengths down the neck 
to the vent; head blue (probably green in life) above, cream below, 
a light line above the eye, dorsal edge of upper labials black (after 
MCZ 37630). The posterior extent of the white line is variable; 
on some specimens it fades as far anteriorly as midbody. Gray- 
green /. wetmorei from Barahona are as above, but the white line 
is on the 4th and 5th scale rows from the snout to one head length 
behind the vent (after AMNH 51421). One specimen (MCZ 
43806) is not mottled ventrally and may represent a green morph, 
but it has been so blackened by the preservative that it is impossible 
to infer its color in life. 

Gray morph: Mainland /. frenatus are finely mottled gray above 
and below; a trace of green on the top of the head, light green on 
upper labials, chin, and ventral neck (after MCZ 65530). Gray 
frenatus from He a Vache and the type specimen of /. wetmorei 
(USNM 83891, Isla Beata) are the same as above but with the 
4th and 5th scale rows very light, thus giving the appearance of a 
wide, dark gray dorsal stripe (after MCZ 37629). 

Nomenclature. Cochran (1931) described wetmorei on the 
basis of a single specimen (USNM 83891 ) from Isla Beata, noting 
its close relationship to frenatus. I have re-identified 5 specimens 
from the Barahona mainland (MCZ 43806, AMNH 51416, 
51419-21) as wetmorei. These additional specimens narrow the 
morphological distinction between frenatus and wetmorei. U. f. 
wetmorei has a significantly lower ventral count (180) than /. 
frenatus (189) (t=5.7, 45d.f.), although there is extensive over- 
lap. The color patterns of /. wetmorei are unlike that of mainland 
/. frenatus, but are quite similar to those of /. frenatus from He a 
Vache. Until enough specimens are available to characterize the 
Barahona population, I prefer conserving the name wetmorei to 
designate a subspecies of frenatus. 



22 BR] \ IORA No. 324 

Specimens of frenatus from Diquini show some characteristics 
that might be attributed to introgression hv oxyrhynchus. BMNH 
1913.7.17.29 has a ventral count of l%;'USNM'l 17282 has the 
anterior loreal scale; MCZ 8683, USNM 117282. and 117346 
differ from typical f. frenatus in coloration, being gray with a 
broad stripe oi gray-green down the back. Each oi these speci- 
mens can be detiniteK identified as an aberrant /. fremitus, but it 
seems strange that all the frenatus from Diquini are aberrant. 
Recent collections from the Diquini area include no frenatus, but 
typical frenatus and oxyrhynchus are found together in the Jacmel 
and Miragoane areas, and in Port-au-Prince, adjacent to Diquini. 
With DO evidence of interbreeding. Without a larger sample, fur- 
ther speculation i^n the significance o\ the Diquini population 
seems pointless 

URcm \< i k ox 1 ! Kin m in s Dumeril and Bibron 

Uromacei oxyrhynchtu Dumeril and Bibron, 1854. Erp6tologie Generate, 
7: ""2 (type loc. "Senegal"). 

I\an^i\ Hispaniola: coastal area north of the Cul-de-Sac Plain, 
southwest peninsula ol Haiti west to Miragoane. lie de la Tortue. 
and Isla Saona. It is sympatric with catesbyi throughout its range. 
and with frenatus at the base of the southwest peninsula. V. 
oxyrhynchus is not known to occur with f. wetmorei, though 
Mertens I 1939: 7 l >) reports seeing an oxyrhynchus and a frenatus 
in the same bush at "Camberlen" (near San Cristobal. D.R.); he 
did not catch the frenatus, however, so that il is not possible to 
be sure that the snake was not a gi"a\ oxyrhynchus. One specimen 
of oxyrhynchus (USNM 75927), collected by J.S.C. Boswell 
in 192S. is reporte.l as taken from Anse a Calets. Cionave. I \- 
ceptional localities o\ other specimens in Boswell's 1 1 >2S collection 
have been attributed to labeling errors (Schwartz, 1965). Collec- 
tions from lie de la Gonave in the last ten years contain many 
catesbyi and dor sous, but no oxyrhynchus. 

Diagnosis. Snout 3.0 (range 2.5-3.7) times eye diameter; scale 
rows at midbody 19; ventrals 200 (range: 189-212); subcaudals 
199 (range: 183-211); anterior loreal present in 25 of the 129 
specimens examined: 6 AMMI. 8 BMNH. 2 FMNH. 97 MCZ, 
15 USNM. 1 YPM. 

Color palter)}. Green morph: dark green above, light yellow- 
green below, the colors separated by a white lateral line on the 
1st and 2nd scale rows from the snout to two hcadlengths beyond 
the vent, ventral edge of the scales in the 1st row black; head 



1969 EVOLUTION OF UROMACER 23 

green above, cream below, a light line above the eye, dorsal edge 
of upper labials black (after MCZ 62701). There are variations 
in color intensity. The lateral line may be edged by black, yellow, 
both, or neither. In a few specimens, as much as the anterior half 
of the lateral line is indistinct or lacking. 

Gray-green morph: like the green morph, but with the ventral 
surface finely mottled gray to gray-brown from the neck to the 
tip of the tail (after MCZ 62703). This morph has been found 
in Haiti and in the eastern Dominican Republic. Variations are as 
in the green morph. 

Gray morph: finely mottled gray above, more coarsely mottled 
gray below, 2nd, 3rd, and 4th scale rows lighter gray, dorsal edge 
of scales in the 5th row black from midbody to vent; head as in 
the green morph (after MCZ 62702). Four specimens of this 
morph have been taken from scattered localities in Haiti. 

Uromacer dorsalis Dunn 

Uromacer dorsalis Dunn. 1920, Proc. New England Zool. Club, 7: 43 (type 
loc. Gonave Island). 

Range. lie de la Gonave. It is sympatric with catesbyi. 

Diagnosis. Snout 2.9 (range: 2.5-3.7) times eye diameter; 
scale rows at midbody 17 (19 in MCZ 93146); ventrals 200 
(range: 192-210); subcaudals 210 (range: 197-228); anterior 
loreal present (absent in MCZ 93139). 84 specimens were ex- 
amined: 7 ASFS, 65 MCZ, 12 YPM. 

Color pattern. The species is monomorphic: finely mottled gray 
above and below, the dorsal region between the 5th scale rows 
darker; head green above, light green below (after MCZ 61043). 
In very light specimens the dorsal stripe is indicated only by black 
spots on the dorsal edge of scales in the 4th row. 

Nomenclature. U. dorsalis is more similar to oxyrhynchus than 
to frenatus in general body form, snout length, ventral count, and 
the presence of the anterior loreal scale, but more similar to 
frenatus in midbody scale count, subcaudal count, and color pat- 
tern. Since dorsalis is unique in its diet and morphologically inter- 
mediate between two distinct mainland species, I retain it as a 
distinct species. Although I admit a close relationship between 
dorsalis and oxyrhynchus, the evidence is not compelling enough 
to warrant calling dorsalis a race of oxyrhynchus. 



BREVIORA 



Museum of Connparsitive Zoology 

Cambridge, Mass. 15 September, 1969 Number 325 

THE GENUS PHENACOSAURUS (SAURIA: IGUANIDAE) 

James D. Lazell, Jr. 1 



The members of this order are the animals ihe Lord of the 
Hebrews did not create to grace his Eden; they are not among 
the products of the six days' labor. These are the sports, the 
offthrows, of the universe. . . . These are the weird children 
of the lust of the spheres. 

Charles Finney 

Abstract: A complete revision of the Andean genus Phenacosaurus, us- 
ing all available museum specimens, is presented. The genus is separated 
from Anolis on the bases of pelvic osteology and digital pad structure. Of 
four previously described species, two are retained: heterodermus (including 
"richteri" and "paramoensis") and nicefori. A new species, P. orcesi, is 
described from Ecuador. 

Among the strange and varied productions of the high Andes 
is a small assemblage of grotesque, big-headed, short-legged, pre- 
hensile-tailed lizards: the genus Phenacosaurus. Since Dunn's 
(1944) work on this genus in Colombia, it has not been re-exam- 
ined as a whole. Through the efforts of E. E. Williams, Museum of 
Comparative Zoology (MCZ), it has now been possible to amass 
at one time and place the majority of specimens of all the species 
of this genus and thus attempt a complete review. Without the 
cooperation of the curators of many institutions, of course, this 
would not have been possible. I express here my gratitude to the 
curators of the following institutions : Instituto de La Salle, Bogota 
(ILS), British Museum (Natural History) (BMNH), Museum 
of Vertebrate Zoology (MVZ), University of Michigan Museum 
of Zoology (UMMZ), Stanford University Museum of Natural 
History (SU), Field Museum of Natural History (FMNH), Phila- 
delphia Academy of Natural Sciences (PANS), Fundacion La Salle 



1 Massachusetts Audubon Society, Lincoln, Mass., 01773 



2 breviora No. 325 

de Ciencias Naturales, Caracas (FCN), United States National 
Museum (USNM), American Museum of Natural History 
(AMNH). Vienna Museum (WEEN), Munich Museum (MUN), 
Senckenberg Museum (SMF). Berlin Museum (ZMB), Museum 
National d'Histoire Naturelle de Paris (MHNP). and Universidad 
National de Colombia (UNC). 

Professor Gustavo Orces-V, Esquela Politecnica Nacional, 
Quito. Ecuador, lias very kindly donated the first phenacosaur 
recorded from Ecuador, which I take great pleasure in designating 
as the type of a new species named in his honor (see below). 

Dr. Fred Medem, Cartagena. Colombia, has made available an 
excellent collection of Colombian phenacosaurs, including the 
type and paratopes of P. richteri Dunn. Those specimens actually 
collected by him are fresh alcoholics and combine color reten- 
tion with very accurate data — including elevations. Dr. George 
Gorman, Museum of Vertebrate Zoology. Berkeley, has recently 
collected series in the Bogota area. 

1 he Orsono-Mesas ( 1946) have provided an informative treat- 
ment o{ several aspects of phenacosaur ecology and behavior. 

I lor photographs of living phenacosaurs taken around Bogota 
have been provided by Fred Medem and Jose Muho/, Hans Zenke, 
and Victor I lutchison. 

In addition to the problems posed by the species-group level 
taxonomy of Phenacosaurus, it seems necessary to re-examine the 
status and validity of the genus. Anolis heterodermus Dumeril 
(1851) was included by Boulenger (1885) in his concept of 
Xiphocercus Fitzinger (1843). Xiphocercus 1 ' was then sep- 
arate,! from Anolis (and ChamaeleoUs) on the character com- 
bination of tricuspid lateral teeth plus a prehensile tail, and from 
"Norops" on the character of a raised terminal phalanx. Barbour 
(1920) separated heterodermus from the West Indian form {"Xi- 
phocercus" [~- Anolis] valencienni) on the basis of body squa- 
mation. dorsonuchal crest, elevated rim of head plates (= cir- 
cumoccipital swelling or casque of Dunn. 1944), digital dilations, 
and tail structure. "Xiphocercus" heterodermus thus became the 
type of the new genus Phenacosaurus Barbour ( 1920). However, 
Williams ( 1962) has shown "Xiphocercus" to be a synonym of 
Anolis; the validity of Phenacosaurus may be questioned also. Cer- 
tainly neither body squamation nor dorsonuchal crest is useful in 
separating Phenacosaurus from Anolis. Caudal vertebrae that lack 
autotomy septa are found in such distantly related Anolis as dar- 
lingtoni and latijrons; though no Anolis has yet been demonstrated 



1969 PHENACOSAURUS 3 

definitively to use its tail in a prehensile manner, it seems reason- 
able to infer from structure that some, like darlingtoni, do. 

The "circumoccipital swellings," or "casque," referred to is 
indeed a striking feature of adult male heterodermus. Actually 
posterolateral^ directed flanges of the parietals, this "casque" is 
but weakly developed in juveniles and females of heterodermus, 
or in most specimens of P. nicefori, regardless of sex, and is re- 
duced in the new form from Ecuador. In addition, some species 
of Anolis (e.g. richardi of the Lesser Antilles) develop similar 
parietal flanges; there is, admittedly, still a definite degree of differ- 
ence between the condition in some Phenacosaurus and that shown 
even by adult male Anolis richardi, but one would hesitate to ac- 
cept a genus wholly based on such a degree of difference. 

The digital dilations of Phenacosaurus are distinctive. The toes 
are comparatively shorter than the toes of Anolis of similar size; 
the digital pad is expanded not only beneath the second and third 
phalanges of the fourth toe, but beneath the first phalanx as well; 
proximal continuation of the expanded digital pad is the rule 
with all the toes, and produces a rather gecko-like appearance of 
the foot. Although this condition is not matched by any species of 
Anolis, the variation in structure and dilation of the digital pad in 
Anolis is sufficiently great (see Williams, 1963) so that any degree 
of difference shown by the phenacosaur condition is surely of di- 
minished importance. 

After examination of skeletal material of more than 70 species 
of Anolis in the MCZ, I have discovered that not only is the struc- 
ture of the ilium remarkably constant in that genus, but that it is 
strikingly different from the condition seen in all forms of Phena- 
cosaurus. I therefore propose a redefinition of the genus Phena- 
cosaurus and provide the following definition of Anolis for com- 
parative purposes: 

Anolis is a genus of iguanid lizards without femoral pores; sub- 
digital lamellae bear cuticular hairs; pre- and post-zygopopheses 
of trunk vertebrae not expanded to form great lateral flanges; 
angular absent; lateral teeth cusped; ilial shaft subcylindrical, 
dorsal border forming an obtuse angle with the acute or nodular, 
elongate, anterior iliac process (see Figure 6). 

Phenacosaurus Barbour 

Anolis (part), Dumeril, 1851, Cat. Meih. Rept., p. 59. 
Xiphocercus (part), Boulenger, 1885, Cat. Lizards Brit. Mus. II, p. 10. 
Phenacosaurus Barbour, 1920, Proc. New England Zool. Club 7, p. 62. 
Type species. Anolis heterodermus Dumeril, 1851. 



4 breviora No. 325 

Definition. [guanid lizards without femoral pores; subdigital 
lamellae bear cuticular hairs; pre- and post-zygopopheses of trunk 
vertebrae not expanded to form great lateral flanges; angular ab- 
sent; lateral teeth cusped; ilia! shaft compressed and bladelike, in- 
clining smoothly into a blunt, subrectangular anterior iliae process 
( see I ig. 6). 

The condition of the ilium can be readily checked without un- 
due damage to the specimen by merely lifting the skin above one 
hind limb and separating the muscle attachments from the bone. 
This has been done for all oi /'. nicefori and the new species, as 
well as many P. heterodermus. The only dry skeletons of Phena- 
cosourus examined are Il.S 55d and MCZ 17111. both lietero- 
dermus. 

Polychrus have femoral pores present in males and lack an an- 
terior iliac process. 

Anisolepis and Aptycholaemus (nearly or quite synonymous, 
tnle Etheridge, pers. eonun.) have an ilium basically similar to that 
of Phertacosaurus, but possess a "fairly large'* angular, fide Ether- 
idge (in Hit.). Anolis and Phertacosaurus lack an angular bone. 

1 he remaining anoline genera isensn Etheridge. 1959) have an 
ilium like that oi Anolis. In addition, Tropidodactylus is unique 
in lacking cuticular hairs on the lamellae; Chamaeleolis has sim- 
ple, peglike lateral teeth and a small angular; ChomaeUnoropS has 
trunk vertebrae bi/arrely modified by expansion and amalgama- 
tion of the pre- and postzygopophases to form great lateral llanges. 

The usefulness of separating Phenacosourus from Anolis may 
be questioned, and Dr. R. A. Etheridge (in lilt.) has questioned 
the separation in principle. This, certainly, is the place to make 
a case for the genus, especially as I regard it as valid. Etheridge 
notes that phenaeosaurs are an early offshoot of "alpha Anolis 
stuck in South America that have developed some peculiar be- 
havioral specializations and assorted anatomical specializations." 
In that, we concur; what alarms Etheridge. however, is that "Phen- 
acosourus heterodermus (for example) is much more closely re- 
lated to Anolis jacare than Anolis jacare is to Anolis nebulosus (for 
example).'* That is also true, but it in no way invalidates the 
genus Phenacosourus. The difficulty, and the principle at stake, 
is the old bugbear of relationship. It is perfectly true that birds 
are. in a very real sense, more closely related to crocodilians than 
crocodilians are to lizards, and far more closely than crocodilians 
are to therapsids (for example). Yet. birds are not included within 
the Reptilia; they belong to a distinct class of their own. Simpson 



1969 PHENACOSAURUS 5 

(1961, pp. 129-132 and 140-145) has given detailed con- 
sideration to this problem. Its solution lies in defining taxa in 
higher categories in a way that best expresses their evolutionary 
position and adaptive similarities. In the present case, Anolis 
jacare and A. nebulosus are members of a presumably monophy- 
letic group that have so much in common, from an adaptive and 
evolutionary point of view, that they are best regarded as con- 
generic despite obvious differences. Also, a plethora of other Anolis 
species spans the gap between them, and thus helps to unite them. 
The species of Phenacosaurus, on the other hand, are sharply dis- 
tinct, in important adaptations, from any Anolis; the gap is not 
spanned by living intermediates. Of course, intermediates must 
once have lived, and this leads us to recognize a compelling truth: 
Any taxon at the level of a higher category (genus or above) must 
— either now or in the past — contain some member more closely 
related to members of some other taxon than that member may 
be to some members of its own taxon. I draw a line by defining 
the genus Anolis: the species of Phenacosaurus are outside that 
line. 

It is only my opinion that Phenacosaurus — an apparently nat- 
ural group of species — qualifies for the rank of genus; no defi- 
nition of the category genus exists to tell me whether Phenacosaurus 
is one or not. I rest my case, therefore, on the evolutionary and 
adaptive significance of the characters that separate the groups: 
the structure of the ilium and the digital dilations. These seem to 
me to be quite as important as the distinctions, noted above, which 
separate the other anoline genera. 

Phenacosaurus heterodermus (Dumeril) 

Anolis heterodermus Dumeril, 1851, Cat. Meth. Rept., p. 59. Syntypes: 

MHNP 1664, A1664, B1664, 6798, A6798. 
Xiphocercus heterodermus, — Boulenger, 1885, Cat. Lizards Brit. Mus. II, 

p. 10. 
Phenacosaurus heterodermus, — Barbour, 1920, Proc. New England Zool. 

Club 7, 61-63. 
Phenacosaurus richteri Dunn, 1944, Caldasia 3, p. 60. Type: MCZ 69120. 
Phenacosaurus paramoensis Hellmich, 1949, Dtsch. Aquar.-u. Terrar.-Ztschr. 

2, p. 91. Type: MUN 118/37. 
Diagnosis. A Phenacosaurus with large dorsal scales: 8-19 dor- 
sals contained in the standard distance 1 (not counting interstitial 



i Defined as the distance from the center of the eye to the tip of the snout. 



6 rrf.yiora No. 325 

granules); head plates largo: interparietal sometimes as long as dis- 
tance aeross head between orbits, but often shorter; subdigital lamel- 
lae under second and third phalanges oi fourth toe 18-24. 

Lectotype. MHNP 1664, an adult male with 14 dorsals in the 
standard distance and 19 subdigital lamellae. 

Type locality. "Nouvelle Grenade" (= Colombia), here re- 
stricted to the vicinity of Bogota. Colombia. 

Discussion. The extreme variability of this species has led to 
taxonomic difficulties and confusions, and to the naming, there- 
fore, of two additional species richteri Dunn ( 1944) and 
paramoensu Hellmich (1949). 

Dunn, in his 1944 review, used the following characters to 
separate "richteri" and heterodernuis: 

Dorsal crest squamation. Dunn separated "richteri" (and iiicc- 
fori) from heterodermus on the basis o\' single, rather than double, 
dorsal crest scale row. In attempting to make sense out of the 
remarkable variation I found in this character. I assigned numbers, 
from one to ten. to the various sorts of crest scale arrangements 
noted. In Table 1 each specimen is graded according to the sort 
of crest scale variant it possesses. Where specimens showed more 
than one sort of arrangement, the two or more tirades arc indi- 
cated. The following ten of variation were noted: 

1. Undifferentiated middorsal scales; i.e., no dorsal crest at all, 
except sometimes a median scries of single, enlarged scales on the 
nape or tail. 

2. Some enlarged scales along the middorsum, but not arranged 
in a particular pattern. 

3. Enlarged, tectiform to carinate scales along the middorsum, 
but separated from each other by undifferentiated smaller scales. 

4. Single row of enlarged, tectiform to carinate scales in contact 
with each other along the middorsum. 

5. Single row of spike-like scales in contact with each other 
along the middorsum. 

6. Staggered series of tectiform to carinate scales, every other 
one separated by undifferentiated smaller dorsals, but each usually 
contacting the scale immediately adjacent at the dorsal midline. 

7. Alternating series of tectiform to carinate scales, continu- 
ously in contact with each other. 

8. Double row of spike-like scales on anterior body combined 
with alternating tectiform to carinate scales posteriorly. 

9. Alternating series of spike-like scales. 
10. Double row of spike-like scales. 




PHENACOSAURUS 



L» 






CT> 



o 

.9 

—  

03 

c 
03 

"a, 
x 



X 



00 




CO 




oo 



s 

o 

«J 
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03 



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8 breviora No. 325 

In Figure 1 each of these ten variants is pictured. I do not find 
the condition of the dorsal crest squamation diagnostic for any 
of the named forms. 

Cranial ridges. Dunn (1944) differentiated both of his new 
species from heterodermus on the basis of the structure of the 
circumoccipital and postocular ridges. P. heterodermus was sup- 
posed to have swollen scales making up these ridges, whereas 
"riehteri" and nicefori were said to have swollen bone structure 
underlying the scales: the scales themselves were not supposed to 
be swollen in the latter two forms. Of "riehteri" he states: "nor are 
they [these scales] enlarged" except slightly in old males (p. 60). 
In all the specimens I have seen, including the entire tvpe series 
of "riehteri" and the svntvpes of heterodermus, the scales of the 
circumoccipital and postocular ridges are greatly enlarged, tecti- 
form to carinate, and. in addition, lie on bony ridges of the skull. 
This character, therefore, fails to separate taxa. 

Chin shields. Dunn (1944) separated "riehteri" from hetero- 
dermus on the basis of the extent of contact between the enlarged 
chin shields and the infralabials. Anteriorly, the chin shields and 
infralabials are always in contact, posteriorly they arc separated 
by small sublabials. The point at which separation begins is very 
variable, and the condition supposedl) representative of hetero- 
dermus is to be found in the type series of "riehteri." I could not 
correlate variation in this character with any of the others men- 
tioned by Dunn. 

Contact of dorsal scales. As with the preceding characters, 
Dunn recognized two conditions: enlarged dorsal scales either 
contacting each other, or separated. Both conditions do, ad- 
mittedly, occur, but very often on the same li/ard. I could not 
correlate this condition, even when clear-cut. with any of the 
other characters. 

Leg length. As noted by Aleman ( 1953). leg length is variable 
and will not separate taxa unequivocally. In general, however, 
nicefori do tend to have much shorter legs in the southern part of 
their range (e.g., Pamplona); heterodermus appear to have longer 
legs; in the northern part of their range (e.g., Saota, Paz del Rio). 
Therefore, though leg length cannot be used to diagnose the 
forms, it may indicate character divergence between heterodermus 
and nicefori, and is discussed further under the latter species. Leg 
length cannot be used to divide heterodermus as here defined into 
taxa, and "riehteri" is. therefore, not separable from heterodermus 
on any character or combination of characters discernible by me. 



1969 PHENACOSAURUS 9 

Scale size. As may be noted from the above diagnosis of hetero- 
dermus, I do regard scale size as useful in separating forms. I 
have used two indices to scale size: the size of the head plates is 
here indicated by the length of the interparietal plate compared to 
the distance between the orbits across the top of the head. As 
noted, heterodermus has rather large head plates; the interparietal 
plate may be as long as the distance between the orbits, though it 
is usually somewhat shorter. The size of the dorsal scales, indi- 
cated by the number contained in the standard distance just lateral 
to the midline at midbody, separates nicefori and heterodermus. 
The new form from Ecuador combines scale size measured by 
both indices in a way that is radically different from either of the 
previously named forms. Both nicefori and the new species are 
discussed fully below. Dorsals in the standard distance just lateral 
to the midline at midbody are tabulated for heterodermus and 
nicefori in Table 1. 

Coloration and pattern. Although not utilized taxonomically by 
Dunn (1944), coloration and pattern are obviously subject to 
some striking variation, at least in heterodermus as here defined, 
and must be considered more fully. Hellmich (1949) described 
a single specimen from Paramo de Sumapaz, 3750 meters, as a 
new species, P. paramoensis. The coloration and pattern of this 
individual are striking (see Figs. 2 and 3). Two other specimens 
I have seen match this condition: the smaller of ZMB 4256, a 
female (as is MUN 118/37, the type of "paramoensis"), and 
AMNH 91752, a male. Numerous other specimens approach this 
condition. 

Two specimens, MUN 9/49 and MVZ 68686, from Tambo, 
Cauca, and San Antonio, Valle, respectively (the most south- 
western localities for P. heterodermus), are pale ash-grey with 
very broad, brown bands across the dorsum. Excellent color 
transparencies taken at Bogota by Hutchison show that the colors 
noted occur in living individuals at the type locality. Hutchison 
further notes (pers. comm.) that individuals are capable of chang- 
ing their patterns as well as their colors. 

Range. Phenacosaurus heterodermus, as may be seen in Fig- 
ure 4, occurs in both the Cordillera Oriental and the Cordillera 
Central. The species occurs at least in the extreme southern 
Cordillera Occidental near the head of the Cauca River, at San 
Antonio, Valle. Because the Cauca River, which separates the 
Cordillera Occidental from the Cordillera Central, is nowhere as 
great a barrier as is the Magdelena, which separates the latter 



10 



BRLVIORA 



No. 325 







X 



=: 

O 

=: 

5 



u 



Z 

s 

c 



o 

to 



! 

- 

I 

- 

-=: 

a. 



I 



1969 



PHENACOSAURUS 



11 




N.STRZK . 



Figure 3. Phenacosaurus heterodermus, top of head. Specimen MUN 
1 18/37, the top of " paramoensis" Hellmich. 



12 breviora No. 325 

from the Cordillera Oriental, and because all three Cordilleras 
merge in southern Colombia (e.g., vicinity of Tambo, Cauca), I 
expect the total distribution of the species, at least in the western 
range, to be greater than is presently known. Dunn (1944) gives 
the altitudinai range as from 1800 m to 3500 m. Hellmich (1949) 
cites the elevation for "paramoensis" (= heterodermus) as 
3750 m. This is the highest recorded elevation for the species. 

Size. The largest specimen measured is MCZ 78531, from Paz 
del Rio, Boyaca. It is 86 mm snout to vent. Remarkably, it is 
a female. In the more southern populations of heterodermus 
males seem to average larger than females, and reach 83 mm snout 
to vent (male of ZMB 521 1 ) in the Bogota area; females in the 
Bogota area reach 80 mm (MHNP 1923.55). I therefore expect 
some males from the northern part of the range to be larger than 
any specimen here recorded. Si/e is discussed comparatively 
under the following species: 

Phlnacosaurus nicefori Dunn 

Phenacosauriu nicefori Dunn, 1944. Caldaaia 3, p. - s ^. Type: ILS 64. 

Diagnosis. A Phenacosaurus with small dorsal scales: 21-34 
contained in the standard distance (not counting interstitial gran- 
ules): head plates small, interparietal much shorter than the dis- 
tance across head between orbits; subdigital lamellae 16-20. 

Type. ILS 64, an adult female with 24 dorsals in the standard 
distance and 17 subdigital lamellae. 

Type locality. Pamplona, Norte de Santander, Colombia. 

Discussion. On the basis of the eight specimens I have seen, 
this species is quite closely allied to heterodermus. The only defin- 
itive difference I have been able to discern is dorsal scale size. 
With respect to this character, it must be noted that for both nice- 
fori and heterodermus the range of variation within the species, 
respectively, is greater by far than the gap which separates them. 
I have indicated in my discussion of heterodermus that I believe 
there is some evidence for character divergence between these two 
forms as they approach the area of potential sympatry. Though 
heterodermus and nicefori have not yet been demonstrated to be 
sym- or parapatric at any point (see Fig. 4), what evidence for 
character divergence we do have should be considered in full. 
Specimens of both forms have been arranged in Table 1 in a north- 
south sequence from Cerro Tetari (northernmost locality for nice- 
fori, and the genus) to Tambo (the southernmost locality for 



1969 PHENACOSAURUS 13 

heterodermus) . Thereafter, in Table 1, the sequence is northward 
through the more western heterodermus localities in the Cordil- 
leras Occidental and Central. Characteristics of dorsal scale size, 
dorsal crest type, and subdigital lamellae are tabulated; these, 
with the exception of dorsal crest type (which I do not regard as 
indicative of either taxon or character displacement), are included 
in the discussion below: 

Dorsal scale size. Although the numbers of specimens in the 
samples do not permit any meaningful statistical analysis, nicefori 
and heterodermus do seem to become more different as they ap- 
proach one another. The smallest dorsals in heterodermus ( 1 9 in 
the standard distance) are in FMNH 69673, from San Antonio, 
Huila, one of the southernmost examples. Similarly, the largest 
dorsals in nicefori (only 21 in the standard distance) are in FCN 
664, from Cerro Tetari, Venezuela — the northernmost repre- 
sentative of that form. The differences between the more nearly 
adjacent populations — Pamplona (with one specimen of nicefori 
with 22) and Saota (heterodermus with 14) — are not spectacu- 
lar, but with more evidence might prove meaningful. 

Subdigital lamellae. In this case the situation is not indicative; 
assuredly, the Cerro Tetari nicefori has the highest number of 
lamellae found in that species, but the highest counts in hetero- 
dermus come from the southern part of that range. The more 
nearly adjacent populations hardly differ with respect to this char- 
acter. 

Leg length. As mentioned previously, leg length does seem to 
provide an example of character divergence. However, females of 
both forms seem to be shorter-legged than males, and I can find 
little difference between them. Fortunately, the three southern- 
most heterodermus localities (San Antonio, Huila; Tambo; and 
San Antonio, Valle) are represented by males. In all of these 
the appressed hind limb barely reaches the axilla. In most males 
from around Bogota, the one male from Paz del Rio, and in the 
Saota specimen, the appressed hind limb extends beyond the 
axilla. In Pamplona males of nicefori (2) the hind limb is, as in 
females, shorter than the distance to the axilla, whereas in the 
Cerro Tetari male, as noted by Aleman (1953), the hind limb is 
quite as long as this distance. Even in a case like this, however, 
where the evidence seems clear cut on the face of it, the number 
of individuals involved is much too small for any sort of surety. 

Size. Here again there is but little evidence. The largest hetero- 
dermus examined, as noted, was from a northern locality, one of 



14 breviora No. 325 

the two closest to a nicefori population. Similarly, the largest 
nicefori, FCN 664, is 86 mm snout to vent, and from the northern- 
most locality, farthest removed from the known range of hetero- 
dermus. Because all phenacosaurs begin life small, and because 
I have such a paucity of specimens and information about living 
animals, I cannot claim that this is necessarily meaningful. 

Coloration and pattern. Most of the specimens of nicefori are 
formalin specimens, and show nothing in the way of coloration or 
pattern. FCN 664. however, is well preserved and appears to 
have been rather uniform green with a white stripe from the 
supralabials to the shoulder, as in many heterodermus. There is 
nothing about any of the other specimens of nicefori that could 
contradict the assumption that this is how they all look. 

In the foregoing discussion I have tacitly assumed that FCN 664 
actually represents a nicefori. If it does, it is by far the largest 
representative seen; those from around Pamplona are all less than 
65 mm snout to vent Also, it has the largest dorsals, the highest 
number of siibdigital lamellae, and the longest legs of any nicefori 
seen. I have suggested that these differences may be attributable to 
character divergence in two species. On the basis of present infor- 
mation I see no alternative; if FCN 664 is not a nicefori, then a 
great deal of additional material will be required to demonstrate its 
true relationships. Certainly heterodermus and nicefori. whether 
they are distinct species or geographic representatives of the same 
species, are closely related. 

Extreme with respect to major characters, the relationships of 
the new form from Ecuador are in no way so clear. I describe 
it as: 

Phenacosaurus orcesi sp. nov. 

Type. MCZ 38937, collected iv. 1957 by Jorge Olalla. 

Type locality. Mt. Sumaco. Napo Pastaza Province, Ecuador. 

Paratype. USNM 16533, collected by James Peters between 
L'Alegria and La Bonita, Ecuador. 

Diagnosis. A Phenacosaurus with very small dorsals, 38-48 
contained in the standard distance (no interstitial granules pres- 
ent); head plates very large, interparietal much longer than the 
distance across the head between the orbits; subdigital lamellae 
under second and third phalanges of fourth toe 16-18. 

Description of the type. MCZ 38937 is a female measuring 
58 mm snout to vent, with a 61 mm tail. Both the foot structure 
and the ilium (dissected on left side) are typically phenacosauran. 



1969 



PHENACOSAURUS 



15 



cerJo \ VENEZUELA 




'A Tetari -f { 



i/\ Pamplona 

^ San Pedro ^ ±*T* 
-~ f\\ . -+- Paz di 

-> ' -r- Sonson ' 



del Rio 



}~j$- Bogota 

4- Paramo de Sumapaz 
San Antonio, Valle-f-/,' "" 

-|- San Antonio, Huila 
-j- Tambo 

COLOMBIA 





+ heterodermus 
A nicefori 
(g> orcesi 



Figure 4. Localities for three species of Phenacosaurus. Names are 
not cited here for localities in the immediate vicinity of Bogota but are 
tabulated in north-south order in Table 1. 



16 breviora No. 325 

There are three large plates across the snout at the level of the 
second canthals. Two scales, the right one of which appears to 
be two smaller scales anastomosed, border the rostral. The nasal 
is entire, in contact with the first supralabial. and separated from 
the rostral by a single, subrectangular prenasal. 

The supraorbital semicircles are composed of large plates, three 
of which are broadly in contact at the midline. The supraocular 
disks are composed of several large plates, two or three of which 
border directly on the supraorbital semicircles, that decrease in 
size laterally. There are no supraeiliarv scales distinguished from 
the orbital granules. 

There are two rows of loreals. The interparietal is very large, 
ca. 1.3 times as long as the distance between the orbits across the 
top of the head, and is in contact with the supraorbital semicircles. 
I he circumoccipita] ridges are reduced to the point of absence. 
The suboculars are in contact with the supralabials. The mental 
is sutured, bordered posteriorly by the infralabials, sublabials, and 
four gulai granules — eight scales in all. 

A single series of sharply leetiform crest scales, separated by 
undifferentiated dorsal granules, begins on the nape and continues 
to the rump; this is a '.V' type crest scale pattern. The dorsal 
scales are granular and of rather uniform si/c; there are 38 con- 
tained in the standard distance at midbody. The ventrals are 
smooth, slightly swollen, and subimbricate medially. 

There are 18 subdigital lamellae under the second and third 
phalanges of the fourth toe. The digital dilations are very broad 
and become only gradually narrower proximally. 

The tail is very slightly compressed; there is a single row of 
slightly enlarged, tcctiform scales middoisally beginning some- 
what posterior to the level of the vent. There are no scale whorls 
or other indications o( autotomy septa. 

There is apparently not a functional throat fan in this female. 
This is a formalin specimen; it has been rendered a dark, disa- 
greeable, mud-brown all over; no trace of coloration or pattern 
remains. 

The type is illustrated in Figure 5. 

The paratype. The only other specimen of P. orcesi currently 
available is a male 60 mm, snout to vent, with a 67 mm tail. The 
digital pads of both fourth toes on the hind feet taper more abruptly 
proximally than is usual in Phenacosaurus, but are broader under 
the first phalanx than in Anolis of similar size. The ilium (dis- 
sected on the left) is typical of the genus: blade-like, with a blunt, 
obtuse anterior prominence. 



1969 



PHENACOSAURUS 



17 




Figure 5. Phenacosaurus orcesi sp. nov., type, MCZ 38937, from Mt. 
Sumaco, Ecuador. 



18 breviora No. 325 

The paratype is similar to the type in all details described above 
except the following: There are four large plates across the head 
between the second canthals. and three scales border the rostral 
posteriorly. Two plates of the supraorbital semi-circles are in 
contact at the midline. The circumoccipital ridges, though low, 
are quite evident. Because one row of gular granules fails to reach 
the mental, only seven scales border it posteriorly. There are no 
enlarged crest scales on nape or back; this is a "1" type crest scale 
pattern. The dorsals are granular but variable in size; counts of 
46 to 48 can be made in the standard distance at midbody. There 
are 16 or 17 lamellae under the second and third phalanges of the 
fourth toe. The tail bears a scries of enlarged, tectiform scales, 
separated by smaller granules, beginning just posterior to the level 
ol the vent; this is a "3" type crest scale pattern. The throat fan 
appears about as well developed as voung males of P. nicejori or 
P. heterodermus: not apparently greatly extensible. 

Dr. James Peters reports (in lilt.): "The specimen was picked 
up by me personally. ... It was found on the trail between L'Ale- 
gria and La Bonita. in Napo Province, Ecuador, high above the 
river that forms the boundary between Ecuador and Colombia, on 
the eastern slopes, on June 26, 1962. It was found dead on the 
trail, in a horse's hoofprint. ... It was rather badly beaten up . . . 
— in the state you see it now. unfortunately. The elevation at 
which it was collected is approximately 5800 feet." 

Discussion. While Phenacosaurus heterodermus and P. nicejori 
seem quite closely allied forms. P. orcesi appears remote. The dor- 
sal squamation lacks the large scales interspersed with granules 
that both other species exhibit. The dorsals are smaller than in 
nicejori, though the head plates are larger than in heterodermus. 
The general appearance is very much like some Anolis (e.g., soli- 
tarius); the small scales augment this effect. The pelvic girdle, 
however, is distinctly phenacosauran; P. orcesi, in fact, differs 
from Anolis with respect to the ilium in the same ways that hetero- 
dermus and nicejori do, but to an even greater extent; the ilial 
prominence is very obtuse and weakly developed in both speci- 
mens available. 

All considered, we know so little about the phenacosaurs in 
general that any attempt at a reasonable assessment of relation- 
ships is pointless. Until series of specimens can be collected from 
the many crucial areas presumably included within the range of 
the genus, and until comparisons can be made utilizing such 



1969 PHENACOSAURUS 19 

characteristics as coloration in life and behavior in the field — 
features demonstrably of great importance in the systematics of 
anoline lizards — our knowledge of Phenacosaurus will remain 
reprehensibly elementary. 

ACKNOWLEDGMENTS 

Figures 2 and 3 were prepared by N. Strekalovsky. Dr. Richard 
Etheridge has provided data on anoline osteology. 

This work was supported by National Science Foundation 
Grants G- 16066 and GB6944, held by Ernest E. Williams. 

LITERATURE CITED 

Aleman, C. 

1953. Contribucion al estudio de los reptiles y batracios de la Sierra 
de Perija. Mem. Soc. Cien. Nat. La Salle (Venezuela) 13: 
205-225. 
Barbour, T. 

1920. A note on Xiphocercus. Proc. New England Zool. Club 7: 
61-63. 

BOULENGER, G. A. 

1885. Catalogue of the Lizards in the British Museum, II. London, 
xiii -+- 497 pp. 
Dumeril, A. A. 

1851. Catalogue methodique de la collection des reptiles. Museum 
National d'Histoire Naturelle, Paris, 224 pp. 
Dunn, E. R. 

1944. The lizard genus Phenacosaurus. Caldasia 3 (11 ) : 57-62. 
Etheridge, R. 

1959. The relationships of the anoles (Reptilia: Sauria: Iguanidae): 
an interpretation based on skeletal morphology. Univ. Micro- 
films 60-2529, Ann Arbor, Mich., xiii + 236 pp. 
Fitzinger, L. J. 

1843. Systema Reptilium. Vienna, vi -f 106 pp. 
Hellmich, W. 

1949. Auf der Jagd nach der Paramo-echse. Dtsch. Aquar.-u. Terrar.- 
Ztschr. 2 (5): 89-91. 
Orsono-Mesa, H., and E. Orsono-Mesa 

1946. Anotaciones sobre lagartos del genero Phenacosaurus. Caldasia 
4 (17): 123-130. 
Simpson, G. G. 

1961. Principles of Animal Taxonomy. Columbia University Press, 
xii + 247 pp. 



20 



BREVIORA 



No. 325 



Williams, E. E. 

1962. Notes on the herpetology of Hispaniola. 7. New material of 
two poorly known ;mo!es: Anolis monticola Shreve and Anolis 
christophei Williams. BrevioraNo. 164: 1-11. 

1963. Studies on South American ancles. Description of Anolis mirus, 
new species, from Rio San Juan, Colombia, with comment on 
digital dilation and dewlap as generic and specific characters in 
the ancles. Bull. Mus. Comp.^Zool. 129 (9): 463-480. 

(Received 29 April 1969). 






Figure 6. Ilia of two anoline lizard genera: A. Anolis (specimen 
MCZ 61027, A. richardi, Tobago, West Indies); B. Phenacosaurus (speci- 
men MCZ 17111, P. heterodennus, Guasca, Colombia). 



1969 



PHENACOSAURUS 



21 



TABLE 1: SPECIMENS EXAMINED 

Specimens of the three species of Plienacosauriis and three of their 
characters: dorsal scales counted in the standard distance, type of dorsal 
crest, and number of subdigital lamellae. See text and map (Fig. 5). The 
locality "Bogota" is of highly variable precision. 

PHENACOSAURUS NICEFORI 













Lamel- 


Specimen 


Sex 


Locality 


Dorsals 


Crest 


lae 


FCN 644 


$ 


Cerro Tetari 


21 


3 


20 


FMNH 5684 


9 


Paramo de Tana 


28 


6 


17 


FMNH 56443 


juv. 


Pamplona 


31 


3 


18 


PANS 25865 


$ 


44 


27 


3 


17 


ILS 64 


9 


44 


24 


3 


17 


ILS 64a 


9 


44 


22 


3 


16 


MCZ 6729 


$ 


44 


34 


3,4 


17 


USNM 72745 


9 


Tapata 


33 


3 


16 



PHENACOSAURUS HETERODERMUS 



ILS 63 


$ 


Saota 


14 


9 


18 


UNC 692 


9 


Paz del Rio 


10 


2,9 


21 


UNC 693 


$ 


44 44 


14 


5,7 


20 


MCZ 78520 


S 


Cogua 


11 


3,5 


20 


MCZ 78521 


$ 


(( 


12 


4 


20 


ILS 62 


S 


Subachoque 


13 


8 


18 


ILS 62a 


S 


tc 


15 


9 


19 


MCZ 78524 


£ 


Cajica 


13 


5,4 


21 


MCZ 78525 


8 


^ 


10 


5,4 


21 


MCZ 78526 


9 


t< 


9 


5 


20 


MCZ 78527 


$ 


u 


11 


5,3 


20 


MCZ 78528 


$ 


(4 


12 


5 


20 


MCZ 78529 


$ 


41 


14 


5,3 


19 


MCZ 78534 


juv. 


M 


12 


4 


20 


UNC 705 


9 


(4 


10 


5,4 


21 


MCZ 104409 


$ 


Chia 


10 


4 


19 


MCZ 110133 


4 


44 


10 


7 


10 


MCZ 110134 


9 


44 


12 


4 


18 


MCZ 110135 


juv. 


44 


17 


2 


20 


MCZ 110136 


9 


44 


8 


2 


22 


MCZ 110137 


9 


44 


14 


2 


22 


MCZ 110138 


S 


44 


11 


2 


20 


MCZ 110139 


6 


)» 


11 


7 


19 


MCZ 78522 


9 


Usaquen 


10 


8 


19 


MCZ 78523 


S 


Bosque Popular 


16 


6 


20 



22 



BREVIORA 



No. 325 



Specimen 



Sex 



Locality 







Lamel- 


Dorsals 


Crest 


lae 


12 


4,7 


19 


10 


4 


20 


9 


5.7 


19 


12 


4 


21 


10 


5 


23 


12 


8 


19 


10 


8 


20 


11 


8 


18 


9 


5 


19 


11 


5 


20 


10 


5 


19 


12 


3 


20 


8 


5 


20 


12 


5,6 


18 


12 


5 


20 


10 


5,6 


19 


10 


5 


20 


13 


5,6 


20 


10 


5.6 


20 


10 


10 


19 


1 1 


9.5 


20 


14 


4 


19 


14 


l >. 5 


21 


13 


9.6 


20 


14 


9 


19 


12 


9.5 


19 


10 


6 


21 


12 


5 


20 


1 1 


5,6 


20 


13 


9. 10 


20 


13 


9,6,4 


21 


12 


9 


20 


15 


5 


20 


14 


5 


19 


12 


1 


21 


1 1 


8 


21 


13 


1 


19 


13 


5. 6, 3 


20 


1 1 


8,6 


20 


12 


8 


21 


14 


9,5 


20 


11 


8 


21 


14 


8 


20 


13 


8 


20 



MCZ 78519 


$ 


Ten a 


MCZ 69120 


$ 


Tabio 


MCZ 69121 


9 


" 


MCZ 69122 


<5 


.. 


MCZ 69123 


$ 


.4 


PANS 25074 


juv. 


.. 


PANS 25192 


6 


.4 


PANS 25193 


& 


.. 


PANS 25194 


6 


.» 


PANS 25195 


$ 


II 


PANS 25196 


9 


II 


PANS 25197 


<J 


•• 


ILS 55 


6 


Sopo 


ILS 55a 


9 


II 


ILS 55b 


<5 


M 


ILS 55c 


9 


M 


ILS 55e 


$ 


Sopo 


ILS 55f 


9 


** 


UMMZ 56755 


9 


Guasc 


\H / 117111 


9 


m 


MCZ 117112 


6 


m 


MCZ 14164 


^ 


Bogot 


M( Z 14165 


juv. 


m 


BMNH 68.3.3.9 


6 


M 


BMNH 68.3.3.10 


juv. 


'* 


BMNH 1919.3.6.1 


i 


*' 


BMNH 1919.3.6.2 


9 


14 


BMNH 1919.3.6.3 


i 


41 


BMNH 1919.3.6.4 


9 


M 


BMNH 1919.3.6.5 


S 


H 


BMNH 1919.3.6.6 


i 


II 


SU 8275 


9 


" 


SU 8276 


9 


M 


SU 8277 


& 


M 


PANS 21058 


6 


II 


PANS 21059 


6 


it 


PANS 21060 


9 


• • 


ILS 61 


3 


U 


ILS 61a 


i 


14 


ILS 61b 


6 


M 


ILS 61d 


9 


II 


ILS 61e 


9 


it 


ILS 61g 


<J 


II 


ILS 61h 


6 


»t 



1969 



PHENACOSAURUS 



23 



PHENACOSAURUS HETERODERMUS 



Specimen 



Sex 



Locality 





Lamel 


■yrsals 


Crest 


lae 


8 


9,5 


19 


15 


9 


20 


10 


5,1 


20 


11 


5,3 


19 


9 


5,3 


20 


10 


5 


21 


15 


5,3 


21 


14 


5,1 


20 


10 


8,5 


20 


11 


8 


20 


11 


5 


20 


13 


5 


21 


11 


5 


20 


16 


9,4 


21 


14 


9,4 


21 


16 


6,9,4 


19 


15 


6,9,4 


22 


14 


9,5 


20 


15 


9,6 


20 


10 


5,7,2 


20 


10 


5,7,6 


21 


14 


9,7,6 


19 


14 


8 


20 


12 


9.4 


20 


12 


8 


21 


12 


5,2 


20 


13 


9 


19 


10 


8 


21 


10 


9 


19 


14 


5 


19 


13 


5,9,2 


19 


13 


9 


23 


18 


8,4 


22 


14 


6,3 


19 


14 


7 


20 


15 


4,7 


18 


9 


5 


19 


13 


9 


22 


9 


9 


22 


17 


5,9 


20 


13 


9,10 


20 


13 


9 


24 


13 


9, 10 


21 


13 


9 


19 



ILS 61i 


9 


ILS 61j 


S 


ILS 56 


9 


ILS 56a 


$ 


ILS 56b 


9 


ILS 56c 


9 


ILS 56d 


S 


ILS 56e 


9 


MCZ 74314 


9 


MCZ 74315 


$ 


UNC 687 


juv 


MUN 119/37 


$ 


MUN 119/37 


9 


WIEN 72 


$ 


WIEN 72 


$ 


WIEN 72 


S 


WEIN 72 


$ 


WEIN 72 


6 


WEIN 72 


$ 


WEIN 72 


$ 


WEIN 72 


S 


WEIN 72 


9 


WEIN 72 


9 


WEIN 72 


juv. 


MHNP 1923-55 


9 


ZMB 4255 


$ 


ZMB 4255 


$ 


ZMB 4256 


$ 


ZMB 4256 


9 


ZMB 5211 


$ 


ZMB 5211 


$ 


UMMZ 56258 


S 


SMF 10789 


$ 


AMNH 91752 


$ 


AMNH 91753 


$ 


AMNH 91754 


$ 


AMNH 14024 


$ 


AMNH 14023 


$ 


AMNH 27567 


9 


AMNH 7641 


$ 


AMNH 24215 


$ 


USNM 93225 


S 


USNM 95923 


$ 


USNM 92495 


$ 



Bogota 



24 



BREVIORA 



No. 325 











i 


Lamel- 


Specimen 


Sex 


Locality Dorsals 


Crest 


lae 


USNM 95922 


9 


ii 


1 1 


9 


18 


AMNH 44987 


9 


no locality 


15(?) 


8,6 


21 


AMNH 44986 


(5 


•■ 


12 


7.6 


19 


MHNP 1664 


3 


"Nouvelle Grenade" 


14 


9,7 


19 


MHNP A. 1664 


S 


II M 


13 


5.7 


20 


MHNP B.1664 


3 


II II 


12 


5.8 


19 


MHNP 6798 


9 


II l| 


10 


10,7,2 


18 


MHNP A.6798 


6 


II II 


12 


5,7 


18 


WIEN 12662 


6 


II || 


17 


5 


21 


WIEN 451 


S 


"Alto de los Cruces" 


15 


9,6 


20 


WIEN 12660 


9 


Paramo de Cruz Verde 


10 


6,4 


21 


WIEN 12661 


6 


.. 


15 


9,4 


20 


UNC 680 


3 


u 


16 


5.7 


21 


UNC 681 


9 


.. 


16 


5.7 


19 


UNC 686 


9 


.. 


1 1 


8.6 


19 


MCZ 78514 


i 


\i raeachal 


13 


5. 2 


21 


\1( / 78515 


£ 


II 


16 


5. 2 


21 


PANS 24150 


6 


Vguadeta 


1 1 


5 


19 


PANS 24151 


9 


•• 


13 


9 


22 


PANS 24152 


6 


.. 


17 


8.6 


21 


ILS 57 


9 


1 usagasuga 


1 1 


5 


19 


USNM 127099 


4 


•• 


13 


3 


19 


USNM 12710D 


6 


H 


16 


3 


20 


USNM 127101 




ii 


8 


3. 2 


20 


MUN 118/37 


. 


Paramo de Sumapaz 


10 


8 


20 


WISH 32680 


6 


Gutierrez 


12 


5.9 


19 


AMNH 32681 


9 


•• 


9 


5.9 


20 


FMNH 69673 


<J 


San Antonio. Huila 


19 


3.5 


21 


MUN 9/49 


6 


W. Tarn ho 


16 


3,5 


24 


MVZ 68686 


9 


San Antonio. V'alle 


16 


3.5 


21 


PANS 25285 


i 


1 aguneta" (?), Caldaa 


16 


5.2 


19 


SMI 10443 


6 


Rio Samana 


13 


2 


22 


I \l\l/ 65209 


9 


Sonson 


15 


5,4 


20 


AMNH 35303 


9 


•■ 


9 


5.4 


21 


AMNH 35304 


9 


.. 


14 


5 


19 


AMNH 35305 


$ 


•• 


12 


5 


20 


AMNH 35306 


9 


Sonson 


13 


3, 1 


21 


W1NH 32673 


6 


Medellin 


14 


5 


19 


ILS 58 


i 


San Pedro 


17 


5 


19 


ILS 59 


S 


.. 


9 


8,4 


21 


ILS 60 


6 


.1 


14 


8,6,4 


22 



Phenacosaurus orcesi 



MCZ 38937 


9 


Mt. Sumaco 


38 


3 


18 


USNM 166533 


$ 


btwn. L'Alegria and 
La Bonita 


48 


1 


16 



Harvard MCZ Library 




3 2044 066 302 761 




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