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HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



DO£S NOT CIRCUJLAII 



BREVIORA ^^ ^^ 



MUSEUM OF COMPARATIVE ZOOLOGY 
Harvard University 



Numbers 265-295 
1967-1968 



CAMBRIDGE, MASS. U.S.A. 
1969 



Edited 

by 

Nelda E. Wright 



CONTENTS 

BREVIORA 

Museum of Comparative Zoology 

Numbers 265-295 

1967 

No. 265. New land-locked fish species of the genus Galaxias from 
North Auckland, New Zealand. By R. M. McDowall 
1 1 pp. April 6. 

No. 266. A new species of Vesicomya from the Caribbean Sea 
(Mollusca: Bivalvia: Vesicomyidae). By Kenneth 
Jay Boss. 6 pp. April 6. 

No. 267. A new generic arrangement for some Australian scincid 
lizards. By Allen E. Greer. 19 pp. June 29. 

No. 268 The ecology and behavior of two sympatric Lyeodactylus 
geckos. By Allen E.Greer. 1 9 pp. June 29.' 

No. 269. Two new species of Aniphitarsus (Ophiuroidea) from the 
Western North Atlantic. By Amy Schoener 9 pp 
June 29. ' ^^' 

No. 270. Additions to the unionid fauna of the Gulf drainage of 
Alabama, Georgia, and Florida (Mollusca: Bivalvia) 
By Richard I. Johnson. 21 pp. June 29. 

No. 271. Chelonia depressa Garman re-investigated. By Ernest 
E. Williams, Alice G. C. Grandiso^n. and Archie F. 
Carr, Jr. 15 pp. November 17. 

No. 272. The ecological distribution of the anoline lizards around 
Kingston. Jamaica. By A. Stanley Rand 18 pp 
November 17. 

No. 273. Studies on neotropical Pompilidae (Hymenoptera). III. 
Additional notes on Epipompilus Kohl. By Howard 
E.Evans. 15 pp. November 17. 



No. 274. The genus Nops (Araneae, Caponiidae) in Panama and 
the West Indies. By Arthur M. Chickering. 19 pp. 
November 17. 

No. 275. A new scincid lizard from the northern Solomon Islands. 
By Allen E. Greer and Fred Parker. 20 pp. Novem- 
ber 17. 

No. 276. The generic relationships of the African scincid genus 
Eumecia. By Allen E. Greer. 9 pp. November 24. 

No. 277. Lestidiuni bigelowi, a new species of paralepidid fish with 
photophores. By Michael J. F. Graae. 10 pp. No- 
vember 24. 

No. 278. Notes on the emberizine sparrow Rhynchospiza stolz- 
inanni. By Raymond A. Paynter, Jr. 6 pp. Novem- 
ber 24. 

No. 279. A second skink with fragmented head scales from Bou- 
gainville, Solomon Islands. By Allen E. Greer and 
Fred Parker. 12 pp. November 24. 



1968 

No. 280. The notatus group of Sphaerodactyhis (Sauria, Gekkoni- 
dae) in Hispaniola. By Benjamin Shreve. 28 pp. 
March 19. 

No. 28 1 . Additional notes on batoid fishes from the Western Adan- 
tic. By Henry B. Bigelow and William C. Schroeder. 

23 pp. April 5. 

No. 282. The extinct baboon, Porapapio jonesi, in the early Pleisto- 
cene of northwestern Kenya. By Bryan Patterson. 
4 pp. April 5. 

No. 283. Scythian ammonoids from Timor. By Bernhard Kum- 
mel. 21 pp. April 5. 

No. 284. The relationships of Arwlis of the roquet species group 
(Sauria: Iguanidae) — III. Comparative study of 
display behavior. By George C. Gorman. 31 pp. 
April 5. 

No. 285. The genera of puffbirds (Bucconidae). By G. W. Cot- 
trell. 5 pp. April 5. 



No. 286. The status of Nesogalaxias neocaledonicus (Weber and de 
Beaufort) (Pisces. Galaxiidae). By R. M. McDowall. 
8 pp. April 5. 

No. 287. The sequence of genera in the Estrildidae (Aves). By 
Ernst Mayr. 14 pp. April 5. 

No. 288. The epidermal glands of Lygodactylus (Gekkonidae, 
Lacertilia). By P. F. A. Maderson. 35 pp. May 29. 

No. 289. The genus Miagrammopes (Araneae, Uloboridae) in 
Panama and the West Indies. By Arthur M. Chicker- 
ing. 28 pp. May 29. 

No. 290. Geographic Variation in the Hispaniolan frog Eleuthero- 
dactylus wetmorei Cochran. By Albert Schwartz. 
13 pp. May 29. 

No. 291. A new species of Tribolonotiis (Lacertilia: Scincidae) 
from Bougainville and Buka, Solomon Islands, with 
comments on the biology of the genus. By Allen E. 
Greer and Fred Parker. 23 pp. May 29. 

No. 292. Herpetogeography of Puerto Rico. V. Description of a 
new species of Sphaerodactylus from Desecheo Island. 
By Harold Heatwole. 6 pp. May 29. 

No. 293. Intra- and interspecific chromosome variation in the lizard 
Anolis cristatellus and its closest relatives. By George 
C. Gorman. Richard Thomas, and Leonard Atkins. 
13 pp. May 29. 

No. 294. Distribution and biology of the opisthoproctid fish Win- 
teria telescopa Brauer 1901. By Richard L. Haedrich 
and James E. Craddock. 1 1 pp. May 29. 

No. 295. The Chaiiares (Argentina) Triassic reptile fauna. IV. 
The dicynodont fauna. By C. Barry Cox. 27 pp. 

May 29. 



INDEX OF AUTHORS 

BREVIORA 

Museum of Comparative Zoology 
Numbers 265-295 

1967-1968 

No. 
Atkins, Leonard 293 

Bigelow, Henry B 28 1 

Boss, Kenneth Jay 266 

Carr, Archie F., Jr 271 

Chickering, Arthur M 274, 289 

Cottrell, G. W 285 

Cox, C. Barry 295 

Craddock, James E 294 

Evans, Howard E 273 

Gorman, George C 284, 293 

Graae, Michael J. F 277 

Grandison, Alice G. C 27 1 

Greer, Allen E 267, 268, 275, 276. 279. 291 

Haedrich, Richard L 294 

Heatwole, Harcld 292 

Johnson, Rich \rd 1 2 73 



No. 
KUMMEL, Bernhard 283 

Maderson, p. F. a 288 

Mayr, Ernst 287 

McDowell, R. M 265, 286 

Parker, Fred 275, 279, 291 

Patterson, Bryan 282 

Paynter, Raymond A.. Jr 278 

Rand, A. Stanley 272 

ScHOENER, Amy 269 

schroeder, william c 281 

Schwartz, Albert 290 

Shreve, Benjamin 280 

Thomas, Richard 293 

Williams, Ernest E 271 



BREVIORA 

Mmseuim of Coimparsitive Zoology 

Cambridge, Mass. 6 April, 1967 Number 265 



NEW LAND-LOCKED FISH SPECIES 

OF THE GENUS GALAXIAS 

FROM NORTH AUCKLAND, NEW ZEALAND' 

R. M. McDowall 

Museum of Comparative Zoology, Harvard University 

Abstract: Two species of Galaxias Cuvier (Galaxiidae) are described 
from small land-locked coastal dune lakes in North Auckland, New Zealand. 
They are small shoaling species similar to and apparently derived from the 
sea-going Galaxias maculatus attenuatiis (Jenyns). 

INTRODUCTION 

In the New Zealand Galaxiidae, speciation appears to be taking 
place by the land-locking of sea-going species. This is a pattern 
familiar in the Northern Hemisphere Salmonidae, and also in the 
Southern Hemisphere in the Retropinnidae (McDowall, 1965). 
Stokell (1964) described a new species of Galaxias (G. parrishi), 
which he considered to be a land-locked derivative of G. maculatus 
ignotus, and noted another land-locked form from Victoria, Aus- 
tralia. The writer (McDowall, 1966) noted a similar form col- 
lected from another Victorian lake (see Pollard, 1964), and the 
occurrence of further such derivatives from G. maculatus attenu- 
atus, in New Zealand coastal lakes. 

About 15 years ago, the New Zealand Marine Department con- 
ducted a survey of the chain of small lakes on the west coast of 
the North Island of New Zealand, to determine their ecology in 
relation to the introduction of suitable game fishes. In their report 
on this survey, Cunningham et al. (1953) described the fish faunas 
of these lakes as follows: "Various species of Galaxias, Gobio- 
morphus, and Retropinna were recorded," and they listed in a table 
the lakes from which these genera were taken. In 1963, the 
writer examined a sample of G. maculatus attenuatus-Vike fishes 
from one of these lakes, which, on close examination, appeared to 

1 Fisheries Research Publication No. 93. 



BREVIORA 



No. 265 



be distinct from G. maculatus attenuatus in a number of details. 
Because this sample was rather old, it was felt desirable to re- 
collect the fish, before description, and this was done in March 
1965. Large samples of Galaxias were collected from Lake 
Waiparera, a few miles north of Kaitaia, and from Upper Lake 
Rototuna, on the peninsula west of the northern arm of Kaipara 
Harbor (Fig. 1). Inasmuch as these lakes are land-locked, it 
was to be expected that the Galaxias populations would differ mor- 
phologically and behaviorally from Galaxias maculatus attenuatus. 




L WAIPARERA- 



Figure 1. Locality map of North Island, New Zealand, showing location 
of: 1. Lake Waiparera, 2. Lake Rototuna. 



Being land-locked, their life cycles are also necessarily modified 
from that of G. maculatus attenuatus which has marine juveniles. 
Examination of the fresh samples from these two lakes, showed 
them to be distinct from G. maculatus attenuatus and from each 
other. They do not resemble any other New Zealand species of 
Galaxias. Characters suggesting relationship to G. maculatus at- 
tenuatus, apart from their general appearance and coloration, in- 
clude shoaling behavior as adults, forked caudal fins, similar denti- 
tion, lack of pyloric caeca, and the long, many-rayed anal fin. 



1967 NEW SPECIES OF GALAXIAS 3 

The isolation of sea-going species in small lakes, as appears to 
have taken place in these instances, poses a problem in species 
delimitation. The alternatives lie between describing each isolate 
as a distinct species or subspecies, or including all the populations 
in a single, widely variable species, derived from the parent sea- 
going species. Since there has been a fundamental change in the 
life history pattern of the land-locked species, it seems likely that 
the parent and derivative species have acquired reproductive iso- 
lating mechanisms. This conclusion is supported by the fact that 
the derivative populations have developed characteristic morpho- 
logical differences from the parent population. The specific dis- 
tinctness of the two derivative populations under examination is 
another problem. Ideally, breeding experiments and behavioral 
studies of the two populations would be desirable to provide the 
answers, but these are outside the scope of the present study. On 
examining the morphological differences between the two popula- 
tions, it seemed likely that they were independently derived from 
the parent stock and therefore more closely related to the parent 
than to each other. The population in Lake Rototuna is, from both 
morphological and geological considerations, probably an earlier 
derivative of G. maculatus attenuatus than is the Lake Waiparera 
population. Lake Rototuna occurs in well stabilized rolling sand 
dunes, once covered in bush, at an altitude of 300 ft. (91 m). 
whilst Lake Waiparera is in still shifting sand dunes, about a hun- 
dred feet (36 m) above sea level. Corresponding to their apparent 
difference in age, and perhaps to differing ecological conditions in 
the two lakes, the degree to which these species have diverged 
from their common parent species differs greatly. The differences 
are most obvious in vertebral count, a character which has proved 
useful in separating Galaxias species in New Zealand. Assuming 
relationship to G. maculatus attenuatus, the Lake Rototuna popu- 
lation has traversed the full range of vertebral number in the New 
Zealand Galaxiidae, from the maximum in G. maculatus attenuatus 
of 59-64, to a minimum of 47-50. The vertebral number in the 
Lake Waiparera population is also reduced and almost completely 
disjunct from both G. maculatus attenuatus and the Lake Rototuna 
population, viz. 54-59 (Fig. 2). 

The direction of divergence from G. maculatus attenuatus, seen 
in the two derivatives, varies; sometimes the counted structures are 
fewer in number, as in vertebrae and most of the fin-ray counts 
(Fig. 3). In the case of gill rakers (Fig. 2), in contrast, G. macu- 
latus attenuatus has a total count on the first arch of 13-17, the 
Waiparera fish 15-17. and the Rototuna fish 18-23. These data 



BREVIORA 



No. 265 



C. CBACmS 



11 



Jk 



C ATTENUATUS 



iiL 



g. USITATUS 



h 



jL 



ll 



iL 



Jll 



jlll 



4 S 6 7 8 13 14 1^ 16 17 18 19 SO 21 22 23 



an ANCHrOSTECALS 



ClLl RAKERS 



47 48 49 50 SI 52 51 54 S5 S6 57 58 59 60 61 62 63 k4 
VERTEBRAE 



Figure 2. Frequency distribution for counts of branchiostegals, gill rak- 
ers, and vertebrae in Galaxias gracilis, G. maculatiis attenuatiis, and G. 
usitatus. 




DOR5 Al f I N RAYS 



ANAL FIN RAYS 



PEL\-1C FIN RAYS 



Figure 3. Frequency distribution for counts of dorsal, anal, and pelvic 
fin rays in Galaxias gracilis, G. maculatiis atteniiatHS, and G. usitatus. 



1967 NEW SPECIES OF GALAXIAS 5 

suggest that the meristic distinctions may not be simply a case of 
temperature related differences. Differences were also found to 
occur in body proportions. In some cases both the derivatives 
diverged from G. maculatus attenuatus in the same direction (Fig. 
4-2, 4-5), the measured structure being proportionately either 



(D 



A 
B 
C 



© 



8 9 10 11 12 

STANDARD LENGTH / LENGTH CAUDAL PEDUNCLI 



A 
B 
C 



(D 



1.7 1.8 1.9 2.0 2.1 

STANDARD LENGTH / PRE-PELVIC LENGTH 



A 
B 

C 



® 



2.6 2.8 3.0 3.2 3.4 3.6 3.8 4.0 4.2 4.4 4.6 

STANDARD LENGTH / PELVIC - ANAL LENGTH 



A 
B 



C 



(D 



9 10 II 12 13 

STANDARD LENGTH / LENGTH DORSAL FIN BASE 



A 
B 
C 



© 



3.8 4.0 4.2 4.4 4.6 4.8 5.0 5.2 5.4 5.6 

STANDARD LENGTH / HEAD LENGTH 



A 
B 

c 



2.3 2.4 2.S 2.6 2.7 2.8 2.9 3.0 3.1 3.2 3.3 3.4 3.5 

HEAD LENGTH / 1 NT E R - ORB IT A L V/IDTH 

Figure 4. Ranges for proportional measurements in Galaxies gracilis 
(A), C. maculatus attenuatus (B), and G. usitatus (C). 



6 BREVIORA No. 265 

larger or smaller in both derivatives than in G. maculatus attenua- 
tus. In other cases, the divergence has gone in opposite directions, 
and the range for G. maculatus attenuatus was found to lie be- 
tween the ranges for the derivative populations (Figs. 4-1, 4-4). 

These data indicate that each of the two populations should be 
regarded as a full species, distinct from G. maculatus attenuatus 
and from each other. 

SYSTEMATIC DESCRIPTION 

Galaxias gracilis^ n.sp. 

Figure 5; Table la. 

Holotype: Deposited in collection of Fisheries Research Di- 
vision, New Zealand Marine Department. 

Paratypes: Dominion Museum, New Zealand. Reg. No. 4499; 
Museum of Comparative Zoology, Harvard University, No. 45053; 
Smithsonian Institution Fish Collection No. USNM 201224. 

Type locality: Upper Lake Rototuna, Kaipara Harbour, North 
Auckland, New Zealand. 



Figure 5. Galaxias gracilis n.sp. Length to caudal fork, 59 mm. 

Description: A small, slender, shoaling species with long head, 
low vertebral count and rearward placement of the pelvic fins. 
Maximum known fork length 62.5 mm. D i-iii, 7-10; C 16; A i-iii, 
12-16; Pel 6-8; Pec 12-14; Branchiostegals 4-6; Vertebrae 47-50; 
Gill-rakers 5-7 (upper), 13-17 (lower limb of first arch), total 
18-23. 

Trunk cylindrical, slender, somewhat depressed on head, later- 
ally compressed on caudal peduncle, deeper than broad. Caudal 
peduncle slender and short. Head small, slender, a little broader 
than deep, lower jaw protruding a little, or equal in length to 
upper; lips thin, cleft of mouth slightly oblique, extending to a 

1 Named on account of its slender form, from the Latin gracilis, slender. 



1967 NEW SPECIES OF GALAXIAS 7 

line through anterior eye margin. Eye large, at upper head pro- 
file, with inter-orbital flat. 

Mandibular and premaxillary teeth without canines, lingual 
teeth strong, entopterygoidal teeth moderately developed; gill-rak- 
ers strongly developed; pyloric caeca lacking. 

Fins membranous and short, except anal which is long based. 
Distal margin of dorsal fin rounded to straight, anterior rays long- 
est. Anal origin more or less below dorsal origin, distal margin 
of fin straight, anterior rays longest, maximum fin length little 
longer than base length. Pelvic fins inserted relatively far back, 
pelvic-anal interval short, fin relatively short in pelvic-anal length. 
Pectoral fin disposed laterally, fin short. Caudal fin forked, fin tips 
rounded, depth about equal to body depth. 

Coloration: Trunk covered with a dense peppering of large 
melanophores, which intensify on the head and dorsum of the 
trunk, failing lateroventrally, and ventrally. Fresh material is silver 
in these latter areas. 

Meristic variation: Dorsal rays (segmented), 7(1), 8(21), 
9(23), 10(5); Caudal 15(4), 16(42), 17(4); Anal 12(1), 
13(5), 14(30), 15(13), 16(1); Pelvic 6(7), 7(40), 8(3); Pec- 
toral 12(8), 13(28), 14(14); Vertebrae 47(1), 48(16), 49(24), 
50(9); Branchiostegals 4(2), 5(27), 6(21); Gill-rakers (total 
count) 18(2), 19(3), 20(13), 21(7), 22(3), 23(2). 

Differs from G. maculatns attennatus (Jenyns) in having a much 
lower vertebral count, fewer dorsal rays, a somewhat lower bran- 
chiostegal count and more gill rakers; also in a longer pre-pelvic 
length, longer head, narrower inter-orbital, larger eye, short dorsal 
fin base, smaller maximum size reached, and the lack of a marine 
whitebait juvenile stage. Differs from G. usitatiis n.sp. in having 
fewer vertebrae, fewer branchiostegals, a greater number of gill 
rakers, a longer pelvic base-anal origin length, shorter dorsal fin 
base, lack of serrations on the opercular membrane and the smaller 
maximum size attained. 

Galaxias usitatus^ n.sp. 
Figure 6; Table lb. 

Holotype: Deposited in collection of Fisheries Research Divi- 
sion, New Zealand Marine Department. 



1 Named on account of its lack of highly distinctive characters, from the 
Latin iisitatiis, ordinary. 



8 BREVIORA No. 265 

Paratypes: Dominion Museum, New Zealand Reg. No. 4500; 
Museum of Comparative Zoology, Harvard University, No. 45054; 
Smithsonian Institution Fish Collection No. USNM 201223. 




Figure 6. Galaxias iisitatus n.sp. Length to caudal fork, 72 mm. 

Type locality: Lake Waiparera, North Auckland, New Zealand. 

Description: A slender, lacustrine, shoaling species, similar to 
Galaxias maculatus attenuatus, with long head, a short pelvic-anal 
interval, with many of the fishes examined having six rather than 
the more usual seven pelvic fin rays. Distinctive in the presence 
of fine but definite serrations on the free margin of the opercular 
membrane. Maximum known fork length 81.5 mm. D i-iii, 9-12; 
C 14-18; A ii-v, 12-16; Pel 5-7; Pec 11-15; Branchiostegals 5-7; 
Vertebrae 54-59; Gill-rakers 4-5, 11-13, total 15-17. 

Trunk cylindrical, slender, somewhat depressed on head, lat- 
erally compressed on caudal peduncle and somewhat deeper than 
broad. Caudal peduncle slender and short. Head long and nar- 
row, slender, about as deep as broad; jaws equal, prominent, cleft 
slightly oblique reaching to about anterior eye margin, gape rather 
narrow. Eye large, placed close to upper head profile, inter-orbital 
more or less flat, broad relative to head width. Mandibular and 
maxillary teeth without canines, lingual teeth strong, entopterygoi- 
dal teeth well developed; pyloric caeca lacking; gill rakers well 
developed; free margin of operculum finely serrate. 

Fins membranous and short, except anal which is long based. 
Dorsal fin set well back, fin with moderately short base, distal 
margin of fin somewhat rounded. Anal fin origin more or less 
below dorsal origin; anal fin long based but greatest fin length little 
greater than base length, distal margin of fin about straight 
and inclined to body axis. Pelvic fin inserted at about mid-point 
of standard length, pelvic-anal length short, pelvic fin moderate in 
pelvic-anal interval, thus short. Pectoral fin base disposed laterally, 
fin short. Caudal fin forked, fin tips rounded, fin depth about equal 
to body depth. 



1967 NEW SPECIES OF GALAXIAS 9 

Coloration: Trunk a dusky gray-brown and covered with irregu- 
lar dark blotches similar to G. maculatus attenuatus. When alive, 
the lateroventral and ventral abdomen is silvery, in preserved ma- 
terial, largely unpigmented. 

Meristic variation: Dorsal rays (segmented) 9(4), 10(28), 
11(8), 12(l);Caudal 14(1), 15(4), 16(34), 17(1), 18(1); Anal 
12(2), 13(10), 14(13), 15(15), 16(1), Pelvic 6(30), 7(11); 
Pectoral 11(1), 12(5), 13(21), 14(12), 15(2); Vertebrae 54(2), 
55(5), 56(10), 57(12), 58(9), 59(3); Branchiostegals 5(1), 
6(25), 7(15); Gill-rakers 15(2), 16(11), 17(7). 

Differs from G. maculatus attenuatus in having a lower vertebral 
count, fewer pelvic fin rays, longer head, somewhat larger eye, 
shorter pelvice-anal interval; differs from G. gracilis in details dis- 
cussed under the diagnosis of that species. It differs from both 
G. maculatus attenuatus and G. gracilis by the presence of serra- 
tions on the opercular margin. 

DISCUSSION 

The two new species described above are interpreted as land- 
locked derivatives of one of the New Zealand sea-going species, 
G. maculatus attenuatus. In Lake Okataina, thermal lakes district. 
North Island, and in one of the Kaihoka Lakes, near Cape Fare- 
well, there are two land-locked populations of the sea-going G. 
fasciatus Gray. The upland-alpine lake species G. koaro Phillipps 
and G. lynx Hutton are also envisaged, by the writer, as fresh- 
water derivatives of G. brevipinnis GUnther, a species with marine 
stages. It was suggested (McDowall, 1965). that Retropinna ab- 
breviata McDowall and R. lacustris Stokell had speciated under 
similar circumstances — in isolation, in lakes cut off from the sea 
with a migratory population captured in the impounded water 
body. Looked at from this perspective, the distribution patterns of 
some of the apparently younger species in New Zealand's fresh- 
water fish fauna make greater sense, e.g. G. lynx, present in alpine 
lakes all along the eastern side of the main divide of the South 
Island, to the west of the divide in the Nelson Lakes district, with, 
according to the arrangement of Stokell (1949) a highly disjunct 
population in Lake Waikaremoana, near East Cape in the North 
Island. This pattern appears best explained by derivation of the 
freshwater lacustrine form, probably several times, from the par- 
ent species G. brevipinnis. G. koaro probably has the same mode 
of derivation, from G. brevipinnis, one or several times, in the 
lakes of the volcanic plateau of the North Island, and further study 
may well suggest that G. lynx and G. koaro are conspecific. 



10 BREVIORA No. 265 

This view of speciation in the Galaxiidae probably suppHes the 
key to understanding much of the fauna, both phylogenetically 
and zoogeographically. 



ACKNOWLEDGEMENT 

The writer is grateful to Dr. Giles W. Mead for his advice 
and for reading the manuscript, and to Messrs. K. F. Maynard and 
W. Skrzynski for assistance in collecting the specimens. The finan- 
cial support of a New Zealand National Research fellowship dur- 
ing the study is gratefully acknowledged. 

LITERATURE CITED 

Cunningham, B. T., et al. 

1953. A survey of the western coastal dune lakes of the North Island, 
New Zealand. Aust. J. Mar. Freshw. Res. 4 (2): 343-386, 2 pis. 

McDowALL, R. M. 

1965. A new species of Retropinna from Lake Omapere, North Auck- 
land. Rec. Dom. Mus. N.Z. 5 (13): 89-91, 1 fig. 

1966. Further observations on Galaxias whitebait and their relation to 
the distribution of the Galaxiidae. Tuatara 14(1): 12-18, 2 figs. 

Pollard, D. A. 

1964. Studies on the reproductive cycle of Galaxias attemiatus. Aust. 
Soc. Limn. Newslett. 3(1): 10. 

Stokell, G. 

1949. The systematic arrangement of the New Zealand Galaxiidae. IL 

Specific classification. Trans. Roy. Soc. N.Z. 77 (4): 472-496, 

10 figs. 
1964. A new species of Galaxias from Victoria, Australia. Rec. Dom. 

Mus. N.Z. 5 (6): 45-48, 1 fig. 



(Received 10 October, 1966.) 



1967 NEW SPECIES OF GALAXIAS 11 

TABLE 1. 
Morphometric variation in Galaxias gracilis and G. usitatus. 

Ratio a. G. gracilis b. G. usitatus 

Standard length/Body depth at vent 
Standard length/Caudal peduncle length 
Caudal peduncle length/Peduncle depth 
Pre-dorsal length/Standard length 
Standard length/Dorsal fin base length 
Dorsal fin length/Fin base length 
Standard length/Anal fin base length 
Anal fin length/Fin base length 
Pre-pelvic length/Standard length 
Pectoral-pelvic length/Standard length 
Pectoral fin length/Pectoral-pelvic length 
Pelvic-anal length/Standard length 
Pelvic fin length/Pelvic anal length 

Standard length/Head length 
Head length/Head width 
Head length/Inter-orbital width 
Head length/Eye diameter 
Head length/Gape width 



8.25-9.50 


7.70-10.08 


7.46-9.65 


9.14-11.60 


1.54-2.08 


1.31-1.69 


0.73-0.78 


0.74-0.79 


10.90-13.86 


8.00-11.78 


1.38-1.71 


1.29-1.64 


6.44-7.68 


6.02-8.20 


1.11-1.33 


1.15-1.33 


0.53-0.56 


0.49-0.56 


0.28-0.33 


0.28-0.33 


0.32-0.42 


0.36-0.43 


0.20-0.24 


0.22-0.27 


0.41-0.50 


0.36-0.43 


3.92-4.39 


4.06-4.54 


1.87-2.14 


1.87-2.29 


3.01-3.50 


2.67-3.25 


3.67-4.50 


3.72-4.58 


3.29-4.00 


2.90-3.80 



BREVIORA 

Mmseimm of Comparsitive Zoology 

Cambridge, Mass. 6 April, 1967 Number 266 



A NEW SPECIES OF VESICOMYA 

FROM THE CARIBBEAN SEA 

(MOLLUSCA: BIVALVIA: VESICOMYIDAE) 



By 



Kenneth Jay Boss 

Museum of Comparative Zoology 
Harvard University 



During Cruise 104 (fail 1965) of the M /V OREGON of the 
Bureau of Commercial Fisheries Pascagoula Fishery Station. Pas- 
cagoula, Mississippi, an unusually large bivalve was trawled, alive, 
from a depth of over two hundred fathoms (366 m) off the coast 
of Colombia in the Caribbean Sea. Upon examination and com- 
parison with material in the United States National Museum, the 
specimen was found to be referable to the rare and enigmatic 
family Vesicomyidae. The unique specimen is not conspecific 
with any known species of this family and is herein described 
as new. 

Vesicomya (Callogonia) caribbea sp. nov. 
Figures 1-6 

Holotype: Division of Mollusks, U. S. National Museum, cata- 
log number 674573. 

^Type locality: M/V OREGON Station 5692, 12 = 3 1'N; 70 58'W; 
205 fathoms (375 m); Caribbean Sea, off Cabo la Vela, Peninsula 
de Guajira, Colombia; bottom temperature 55T (12.8°C); 10 
October 1965. The specimen was collected alive by a 40' shrimp 
trawl. 

Description: Shell 96 mm long, 67 mm high and 54 mm wide, 
irregularly elliptical, inequilateral, equivalve, solid, frangible, in- 
flated with both valves of equal convexity, without posterior gape, 
but with slight undulate flexure to right posteriorly. Umbos an- 
terior, conspicuous, involute, pointed, prosocline, slightly elevated 



2 BREVIORA No. 266 

and inflated; umbonal cavity deep. Anterior margin not extensive, 
very broadly rounded and weakly convex; ventral margin ex- 
tensive, weakly convex, rising gently posteriorly; anterior dorsal 
margin short, semisigmoid, concave umbonally, rounded and con- 
vex distally; posterior dorsal margin convexly arcuate and coexten- 
sive with posterior margin, forming broad blunt, oblique trunca- 
tion; outline weakly pointed posteroventrally. Sculpture closely 
set, fine concentric laminae, grouped between larger, more conspic- 
uous, irregular growth rings; radial sculpture not evident. Posterior 
ridge, semisulcated dorsally, radiating from umbo toward postero- 
ventral region, becoming weaker distally. Ligament strong, pro- 
tuberant, black, elongate, lanceolate, slightly inset basally, sup- 
ported by thickened hinge line, and forming broad elongate nymph. 
Lunule not developed or circumscribed by impressed isocardio- 
form line. Hinge dentition invested with periostracum. Right 
valve with two cardinal teeth beneath umbo; ventral tooth stronger 
and protuberant; dorsal tooth with anterior laminar portion and 
posterior thickened portion. Left valve with single central cardinal 
tooth developed, strong and protuberant; anterior laminate portion 
broken and obsolete; secondary laminate tooth beneath umbo 
broken. No true internal radial rib, but rib-like thickening present 
on posterodorsal margin of anterior adductor muscle scar. Muscle 
scars moderately impressed, large, irregularly ovate, anterior 
smaller than posterior. Pallial line weakly convex; pallial sinus 
irregular, narrow; pallial line not directly or obviously contiguous 
with posterior adductor muscle scar. Internal surface of valve 
with radial vermiculations. Periostracum dehiscent, olivaceous, 
wrinkled, heavier peripherally, also thickened over cardinal and 
anterior portions of hinge internally. Shell basically white, ex- 
ternally grayish to olivaceous, discolored centrally with russet 
band; earthy and chalky in texture; internally shining, with dark- 
ened blotches and pale salmon centrally. 

Remarks: The family Vesicomyidae (originally spelled Vesi- 
comyacidae) was established by Dall and Simpson (1901). It 
has been reviewed by Lamy ( 1920), and recently Odhner (1960) 
has listed the species referable to Vesicomya. The status of the 
family has been questioned: Thiele (1935). Odhner (1960) 
and Clarke ( 1962) included Vesicomya in the Kellyellidae. Jukes- 
Browne (1913) even mentioned it in relation to the Veneridae. 
On the other hand, Dall (1908), Thiele and Jaeckel (1931), 
Prashad (1932), Taki and Habe (1955), and Newell (1965) 
have maintained the Vesicomyidae separately. 



1967 NEW SPECIES OF VESICOMYA 3 

There is also a question as to what genera should be included 
in the Vesicomyidae. For example, Woodring (1938) placed 
Calyptogena Dall. along with his newly described genera Ectena- 
gena and Phreagena, in the Vesicomyidae. An historical survey 
of the family and a critical revision of at least the western Atlantic 
species is currently being prepared. 

No western Atlantic Vesicomya approaches the size of V . carib- 
bea. V. leeana (DalK 1889, p. 440; 1890, pi. 10, figs. 6-9) 
is the only western Atlantic species that is closely related to and 
which might be confused with V. caribbea. The latter is easily 
distinguished by being much larger and having a heavier shell with 
a distinct posterior ridge. In addition, the hinged teeth of V. carib- 
bea are stronger and more blunt. The eastern Pacific species. V. 
gigas (Dall, r896, pi. 18; 1908, pi. 16, fig. 9), attains the size of 
V. caribbea but it is usually less heavily shelled and its teeth are 
more distinct. It may be diflferentiated from V. caribbea by its 
much shorter ligament and shorter, more distinct, nymphal callos- 
ity. Further, V. gigas lacks a posterior ridge and is generally of 
longer, narrower proportions. Also, V. gigas is mesially constricted 
ventrally while V. caribbea is not. 

The two species most closely related to V. caribbea are known 
only from widely separated localities and are also rare in collec- 
tions. The most closely related is Vesico)n\a cliuni described by 
Thiele and Jaeckel (1931, p. 244, pi. 4. fig. 100) from VAL- 
DIVIA Station 63, 2 N; 8'4.3'E, from a depth of 2492 m in the 
Gulf of Guinea, eastern Atlantic Ocean. Unfortunately, the au- 
thors did not figure an internal view of the shell and no specimens 
of this species were available for comparison. The outline of the 
valves of V . caribbea and V . cliuni are quite similar; however, the 
semisulcated posterior ridge of V. caribbea appears to be a dis- 
tinguishing character. Further, according to Thiele and JaeckeFs 
description, V. chuni is relatively thin shelled, whereas V. caribbea 
is thick shelled. 

The other species closely resembling V. caribbea is from the 
East Indian area. Vesicomya winckwortJii, described by Prashad 
(1932, p. 153, pi. 5, figs. 7-8) from SIBOGA Station 17, 7°28.5'S; 
1 15'^28'E, in 1060 m depth, north of Java, is founded on a single 
right valve which, fortunately, was well illustrated externally and 
internally. In contrast to V. winckworthi, V . caribbea is apparently 
more inflated, its anterior margin more narrowly rounded, and its 
nymph stronger and thicker. 

Vesicomya caribbea is placed in the subgenus Callogonia Dall 
1889 (type-species, by monotypy, Callocardia [Vesicomya] leeana 



4 BREVIORA No. 266 

Dall 1889), as its pallial sinus is, though indistinct, somewhat 
angular. Thiele and Jaeckel (1931) placed V. chuni, the species 
most closely related to V. caribbea, in Callogonia. 

ACKNOWLEDGMENTS 

Mr. Harvey BuUis, Jr., Director of the Bureau of Commercial 
Fisheries Pascagoula Fishery Station made it possible for me 
to participate in Cruise 104 of the M/V OREGON and gener- 
ously transferred specimens into my care. Drs. J. Rosewater of 
the Division of Mollusks of the U. S. National Museum, D. Cohen 
of the Bureau of Commercial Fisheries Ichthyological Labora- 
tory, and W. J. Clench and R. D. Turner of the Museum of Com- 
parative Zoology, Harvard University, critically read the manu- 
script. The drawings were inked by Mrs. Mildred Carrington. Dur- 
ing the preparation of this report the author was on the staff of 
the Ichthyological Laboratory, Bureau of Commercial Fisheries, 
Washington, D. C. 

LITERATURE CITED 

Clarke, A. H., Jr. 

1962. Annotated list and bibliography of the abyssal marine mollusks 
of the world. Bull. Nat. Mus. Canada, No. 181, 114 pp. 

Dall, W. H. 

1889. Report on the Mollusca [BLAKE]. Part II. Gastropoda and 
Scaphopoda. Bull. Mus. Comp. Zool., 18: 1-492, pis. 1-40. 

1890. Preliminary report on the collection of Mollusca and Brachio- 
poda obtained in 1887-'88 [ALBATROSS]. Proc. U.S. Natl. 
Mus., 12(773): 219-362, pis. 5-14. 

1896. Diagnoses of new species of mollusks from the west coast of 
America. Proc. U.S. Natl. Mus., 18(1034): 7-20. 

1908 The Mollusca and Brachiopoda [ALBATROSS]. Bull. Mus. 
Comp. Zool.. 43(6): 205-487. pis. 1-22. 

Dall. W. H. and C. T. Simpson 

1901. The Mollusca of Porto Rico. Bull. U.S. Fish Comm., 20(1): 
351-524. pis. 53-58. 

Jukes-Browne, A. J. 

1913. On Collistu, Amiuntis, and Pitaria. Proc. Malac. Soc. London, 
10: 335-347. 

Lamy, E. 

1920. Revision des Cypricardiacea et des Isocardiacea vivants. Jour. 
de Conch., 64: 259-307. 



1967 NEW SPECIES OF VESICOMYA 5 

Newell, N. D. 

1965. Classification of the Bivalvia. Amer. Mus. Novit., No. 2206, 
25 pp., 3 text figs. 

Odhner, N. H. 

1960. Mollusca. Kept. Swed. Deep-Sea Exped., 2(Zool. 22): 365-400, 
2 pis., 12 text figs. 

Prashad, B. 

1932. The LamelHbranchia of the SIBOGA Expedition. Systematic 
Part, Pelecypoda 11 (exclusive of the Pectinidae). SIBOGA- 
Expeditie, Monographie 53 c, 353 pp., 9 pis. 

Taki, I., and T. Habe 

1955. [In] Kuroda, T., Vesicomyidae. Illus. Cat. Jap. Shells, 2(1): 
6, text-figs. 1-4. 

Thiele, J. 

1935. Handbuch der systematischen Weichtierkunde. Jena, Gustav 
Fischer, vol. 2, pp. 779-1022, 110 figs. 

Thiele. J., and S. Jaeckel 

1931. Muscheln der deutschen Tiefsee-Expedition. Wiss Ergeb. 
Deutsch. Tiefsee Exped., 21(1): 161-268 (1-110), pis. 6-10 
(1-5). 

WOODRING, W. P. 

1938. Lower Pliocene mollusks and echinoids from the Los Angeles 
Basin, California. U.S. Dept. Interior, Geol. Surv. Prof. Papers, 
190, 67 pp., 9 pis. 

(Received 20 October, 1966.) 



BREVIORA 



No. 266 




VesicoDiMi iCallogonia) carihbea n.sp.. Holotype. USNM 674573. 

Fig. 1. Internal view of left valve. Fig. 2. Internal view of right valve. 

Fig 3. External view of left valve. Fig. 4. External view of right valve. 

Fig. 5. Dorsal view of valves, showing posterior ridge. Fig. 6. Anterior 
view, showing involute umbos. 



BREVIORA 



MiiseiLiim of Com para tive Zoology 



Cambridge, Mass. 29 June 1967 Number 267 

A NEW GENERIC ARRANGEAAENT FOR SOME AUSTRALIAN 

SCINCID LIZARDS 

By Allen E. Greer 

In Boulenger's (1887, vol. 3) classification of skinks, the 
genus Lygosoma was by far the largest and most heterogeneous 
group that defied further spHtting on the basis of the characters 
used. Unable to recognize "natural" distinctions which would war- 
rant full generic rank, Boulenger designated 1 1 "natural" species 
groups of Lygosoma as subgenera and thereby established a rank- 
ing system that then, as now, was little used in lizard taxonomy. 

The relationships of one of Boulenger's Lygosoma subgenera, the 
subgenus Rhodona, and part of the genus Ablepharus are re- 
examined in the present paper. 

Boulenger's (1887:223) subgenus Rhodona was based on an 
unusually large suite of external characters, being defined as fol- 
lows: "Limbs short or rudimentary. Lower eyelid with a trans- 
parent disk. Ear distinct, minute. No supranasals. Praefrontals 
very small and widely separated, or absent. Frontal broader than 
the supraocular region." 

To this list could have been added the following: nasals enlarged 
so that they just fail to touch or do touch medially, 2-5 pairs of 
nuchals, and a single pair of enlarged preanals. In addition, it 
might have been noted that the group is found only in Australia 
and Tasmania. 

The widespread Old World genus Ablepharus, on the other 
hand, was based essentially on a single character, Boulenger (1887: 
344) including in this genus all those skinks lacking "movable 
eyelids" and with a "transparent disk covering the eye." Obviously, 
Boulenger in this instance put great confidence in a single character, 
the "ablepharine" eye. 

This single character approach should not be thought of as being 
too incongruous with Boulenger's general philosophy. In most 
other reptile groups with the "ablepharine" eye (non-eublepharine 
geckos, pygopodids. xantusids, and snakes), the character was 



2 BREVIORA No. 267 

then and may still be rightly regarded as the hallmark of natural 
taxa. The "ablepharine" skinks, however, are an exception. 

Boulenger's basic conception of Rhodona and Ablepharus was 
accepted by the authors of the two subsequent revisions of the 
'lygosomine" skinks (Smith, 1937; Mittleman, 1952). Malcolm 
Smith (1937) gave Rhodona full generic rank and included within 
it three southeast Asian (aiiguinoides, roulei, and gyldenstolpei) 
and one Australian (australis) species that in 1935 he had con- 
sidered the sole representatives of the genus Ophioscincus} This 
arrangement, however, was suspect on two counts. First, the species 
of Ophioscincus differ from those of Rhodona in lacking both fore 
and hind limbs and external ear openings, and in having no or only 
one pair of nuchals, and second, such an arrangement involves an 
improbable geographic distribution. 

All skinks with an immovable transparent disc covering the eye 
were retained in Ablepharus by Smith ( 1937). In this he followed 
Boulenger, although he noted in 1935 that the group was almost 
certainly polyphyletic, probably having arisen from Leiolopisma 
and Emoia stocks. 

Mittleman (1952), taking his cue from Smith's reasonable sup- 
position of the polyphyletic nature of Ablepharus, cut the Gordian 
knot by separating out all Ablepharus that had the frontoparietals 
and interparietal fused into a single scale and placing them in the 
genus Cryptoblepharus. It is interesting to note that the "able- 
pharine" eye was still sacrosanct; no "ablepharine" skink was 
lumped with a non-"ablepharine" skink. 

Similarly, Mittleman split Boulenger's Rhodona into two genera: 
Rhodona with the frontoparietals and interparietal fused into a 
single scale (indicated as 1 in Table 2), and N odor ha in which the 
frontoparietals are distinct ( 2/ 1 ) or fused ( 1 / 1 ) but always distinct 
from the interparietal. 

Mittleman removed the four species of Ophioscincus from the 
Rhodona-Nodorha complex and was certainly right in doing so. 
However, in reverting to Smith's 1935 treatment of the genus 
Ophioscincus, he has re-emphasized the anomalous geographic 
distribution of that genus. The disjunct Australian, southeast Asian 
distribution, once again, makes the "naturalness" of the genus 
suspect. 

Although Mittleman's arrangement provides groupings which are 
of value in identifying and cataloging skinks, it fails to demonstrate 

1 In 1887 the only known species of Ophioscincus Peters, 1873 was 
australis, and this was put in the subgenus Lygosoma by Boulenger. 



1967 AUSTRALIAN SKINKS: LERISTA 3 

any greater awareness for the evolution of higher taxa than did 
earUer pre-BouIengerian classifications. This has been an a priori 
criticism of Mittleman's lygosomine classification since its publi- 
cation, and perhaps explains why his system has been passed over 
in favor of a Boulenger-Smith arrangement by most present day 
authors. Even these two earlier authors must have been aware of 
many of Mittleman's "generic characters" but chose not to use them. 

In my own research on the higher taxa of the skinks I have 
attempted to correlate characters of external morphology and 
skull morphology. It is my contention that when a suite of external 
and skull characters can be found in only a certain group of species, 
there is strong evidence that the group is "natural," i.e., mono- 
phyletic. The geographic distribution of the taxon is used as a 
further check of the validity of the arrangement. If the distribution 
of the taxon is consonant with the general geographic barriers in- 
fluencing the distribtuions of other taxa, then the argument for 
the validity of the new taxon is strengthened. 

A consideration of the external morphology has, of course, been 
the standard approach in skink taxonomy, although, like Boulenger 
(1887:131), I believe I have found new characters which are 
important. 

Little has been done with skull morphology in relation to skink 
classification. Boulenger examined the relationships of the bones 
of the palate as seen through the epidermis of the mouth and 
utilized what he saw at the generic level. This method led to some 
serious misinterpretations of the anatomical facts and these have 
been perpetuated (see Smith, 1935, and Mittleman, 1952, for 
example) until the present time. By removing the epidermis over- 
lying the palate, Waite (1929) and Mitchell (1950) made a tech- 
nical advance, but failed to take full advantage of it in their own 
taxonomic work. 

My own approach (outlined above) has provided evidence which 
convinces me of the previously only suspected polyphyletic nature 
of many groupings in all three earlier classifications (Boulenger, 
1887; Smith, 1937; and Mittleman, 1952). Many of the single 
characters which were utilized exclusively or heavily in diagnosing 
skink taxa simply do not correlate with any other characters (inter- 
nal or external), as one would expect them to do in a natural group. 

Other correlations utilizing skull and external characters are 
possible, however, and will form the basis of a new classification 
of skinks. This brief study delimiting a newly recognized taxon 
and giving it generic rank is an example of the new approach and 



4 BREVIORA No. 267 

is a first contribution toward the distant goal of a complete generic 
revision of the skinks. 

The taxon to be discussed here consists of all the species of 
Rhodona (sensu Boulenger) and those species of Australian Able- 
pliarus that conform to the descriptions of Rhodona except for 
the "ablepharine" eye (Table 2). The earliest named species of 
the group {Ablephanis liueata) was described by Bell in 1833 
(see below) as the type species of Lehsta. The new taxon should 
therefore be properly known as Lerista, with the following primary 
generic synonymy. 

Lerista Bell 

Lerista Bell, 1883, Proc. Zcol. Soc. London, p. 99 (Type species. Lerista 

lineata Bell, 1833, by monotypy) 
Rhodona Gray, 1839, Ann. Mag. Nat. Hist., vol. 2, p. 335 (Type species, 

Rhodona punctata Gray, 1839, by monotypy). 
Soridia Gray, 1839, Ann. Mag. Nat. Hist., vol. 2, p. 335 (Type species, 

Soridia lineata Gray, 1839, by monotypy). 
Brachystopus Dumeril and Bibron, 1839, Erpetologie generale, vol. 5, p. 778 

(Type species, Brachystopus lineopunctulotus Dumeril and Bibron, 

1839 = Rhodona punctata Gray, 1839, by monotypy). 
Praepeditus Dumeril and Bibron, 1839, Erpetologie generale, vol. 5, p. 787 

(Type species, Soridia lineata Gray, 1839, by monotypy). 
Ronia Gray, 1841, Ann. Mag. Nat. Hist., vol. 7, p. 87 (Type species, Ronia 

catenulata Gray, 1841 = Rhodona punctata Gray, 1839, by monotypy). 
Leptosoma Fitzinger, 1843, System reptilium, p. 23 (Type species, Riopa 

bougainviUii Dumeril and Bibron, 1839, by monotypy). 
Ophiopsis Fitzinger, 1843, Systema reptilium, p. 23 (Type species, Lerista 

lineata Bell, 1833, by monotypy). 
Miculia Gray, 1845, Catalogue of the specimens of lizards in the collection 

of the British Museum, p. 66 (Type species, Miculia elegans Gray, 1845, 

by monotypy). 
Pholeophilus A. Smith, 1849, Illustrations of the zoology of South Africa, 

p. 15 of Appendix (Type species, Pholeophilus capensis Smith, 1849 = 

Soridia lineata Gray. 1839, by monotypy). 
Phaneropus Fischer, 1881, Arch. Naturg. Jhrg. 47, p. 236 (Type species, 

Phaneropus inuelleri Fischer, 1881, by monotypy). 
Nodorha Mittleman, 1952, Smithsonian Misc. Coll., vol. 17, p. 14 (Type 

species, Riopa bougainviUii Dumeril and Bibron, 1839, by original 

designation). 

DIAGNOSIS OF THE GENUS LERISTA 

There is no single diagnostic character that will distinguish the 
species of Lerista from all other skink taxa. This is, however, not 



1967 AUSTRALIAN SKINKS: LERISTA 5 

to say that once one is familiar with the diversity of skinks that a 
species of Lerista cannot be identified as such at a glance. 

On the basis of external characters, the taxon may be diagnosed 
as follows: no supranasals; nasals enlarged so that they just fail to 
touch or do touch medially; prefrontals reduced and widely sepa- 
rated or absent; frontal generally broader than supraocular region; 
frontoparietal/interparietal configuration = 2/1, 1/1, or 1; 2-5 
pairs of nuchals; lower eyelid with a transparent disc and either 
movable or fused to form a permanent spectacle; no row of scales 
separating the small scales of the eyelid from the subocular supra- 
labial; ear opening small to minute, but always present; a single 
pair of enlarged preanals; body form attenuate; limbs diminutive 
or rudimentary, but never totally lacking (digital formula from 5-5 
to 0-1). 

DESCRIPTION AND RELATIONSHIPS OF THE GENUS lERiSJA 

The following description of the taxon Lerista is based on an 
examination of whole skulls, alcoholic specimens (see Materials 
section of paper), and also of type descriptions when species have 
not been available. 

The skull is considered first, as skull morphology has generally 
proved more useful in determining the broader relationships of 
skinks than has external morphology. 

Skull morphology: The skull characters of the species are re- 
markably uniform, although the species adapted to a more burrow- 
ing way of life (e.g., L. bipes and L. praepeditu) show certain 
modifications in the skull that are often seen in burrowing skinks. 

To facilitate comparisons with its near relatives, a brief descrip- 
tion of the skull characters of the genus is given. 

The whole skull tends to be somewhat conical in shape. The 
palatines are approximately 2-2 '/a times as long as broad and meet 
along their entire medial edge. The palatal rami of the pterygoids 
are "squared-off'" and in some species slightly emarginate posteri- 
orly, and they also meet along their medial edge (Fig. 1). The 
palatines and palatal rami of the pterygoids thus form an extensive 
"secondary palate." Among lizards the tendency toward the forma- 
tion of a bony secondary palate is unique to skinks, and the 
extensive palatine-pterygoid palate of Lerista is, in turn, a rather 
"advanced" development of the palate in skinks. 

An ectopterygoid process extends forward along the lateral edge 
of the palatal ramus of the pterygoid, past the palatine-pterygoid 
suture and onto the posterolateral edge of the palatine, thereby 



BREVIORA 



No. 267 







Figure 1. Ventral view of the palate of A, Lerista elegans (FMNH 
11319), Lake Toolburnup (= Tool Brumup), Western Australia; B, Spheno- 
morphus pardalis (MCZ 35413), Coen, Cape York, Queensland; C, Lerista 
boiigainvilli (MCZ 61403), Tallarock, Victoria; D, "Ahlepliarus" lineo- 
ocellotiis (MCZ 33144), Margaret River, Western Australia. Abbreviations 
for this figure: e, ectopterygoid; p, palatine: pt, pterygoid, A, C, and D 
drawn to scale. 



1967 AUSTRALIAN SKINKS: LERISTA 7 

excluding the pterygoid from a position on the infraorbital fenestra 
(Fig. 1). This anteriorly projecting ectopterygoid process is also 
unique to skinks, among lizards, although it is not found in all 
skinks. 

The premaxillae and nasals are paired; the vomer, frontal, and 
parietal are single. There is a parietal foramen in the parietal. 
The frontal and maxilla form a surface suture or the frontal, pre- 
fontal, maxilla, and nasal meet at a point. 

The temporal arch and the postorbital bar (formed by the 
jugal) are present. A small supratemporal fenestra may be pres- 
ent but is more frequently covered by the posterior extension of the 
postfrontal. A postorbital is lacking. 

The bones of the skull and mandible generally lack pigment. 

Teeth are present on the maxillae, premaxillae, and dentaries. 
There are usually 7 (5 in L. bipes and 6 in L. lineopimctulata) 
teeth on the premaxillae and 9-13 (usually 11-12) on each maxilla. 

Meckel's groove may be present or partially overlapped by the 
dentary. 

Relationships as inferred from skull morphology. On the basis 
of the relationships of the bones of the palate, Lerista is most 
similar to Hemiergis and one group of Sphenomorphus species 
(Fig. 1 ) . For convenience, this latter group may be referred 
to as the solomonis group; it encompasses so far as is known 
the following species: solomonis, emi grans, pardalis, australe, 
antimorum, crassicauda, tenuis, and tryoni. The species of all 
three groups, that is Lerista, Hemiergis, and the solomonis group, 
have the palatines and the pterygoids meeting along the midline, 
and, except for three species of Hemiergis and S. australe, the 
species of all three groups possess an ectopterygoid process which 
extends forward to the palatine to exclude the pterygoid from the 
infraorbital vacuity. 

In Hemiergis peroni, H. tridactylus, and H. initiale the anteriorly 
projecting ectopterygoid process has been secondarily reduced to 
a small nubbin which fails to exclude the palatal ramus of the 
pterygoid from a position on the infraorbital vacuity. The process 
is better developed in 5. australe where it extends anteriorly about 
half way along that outer edge of the palatal ramus that would, 
in the absence of the process, border the infraorbital fenestra. 

The species of the genus Hemiergis and the solomonis group, 
however, differ from the Lerista skinks in possessing more teeth on 
the premaxillae (8-9) and in having a long thin postorbital.^ 

1 The number of teeth on the premaxillae and the absence or presence and 
shape of the postorbital are of greater taxonomic significance in skinks than 
in most other lizard families. 



8 BREVIORA No. 267 

The relationship of the bones of the palate described above for 
Lerista and its near relatives is not found in any Ablepharus other 
than in the "ablepharine" Lerista. In all other Ablepharus the 
palatal rami of the pterygoids are separated (contrary to Boulen- 
ger's 1887 diagnosis of the genus), and there is no ectopterygoid 
process (Fig. 1). 

Furthermore, in all Ablepharus other than the "ablepharine" 
Lerista there are usually 9 (or more) teeth on the premaxillae, and 
the prefrontal often forms a broad suture with the nasal. 

External morphology: In gross aspect the species of Lerista are 
attenuate skinks in which the tail comprises fully half the total 
length. The appendages are diminutive and widely separated when 
adpressed to the body. The head tends to be somewhat conical 
to wedge shaped. 

Upon closer inspection it can be seen that the nasals are en- 
larged in a dorsoventral aspect so that they meet mid-dorsally 
behind the rostral or just fail to do so. 

There are no supranasals and the frontonasal is wider than long. 
The prefrontals are small and widely separated or, in some species, 
they are absent. The frontal tends to be wider than the supraocular 
region and is in contact with 1 or 2 (usually 2) of the 2 to 4 
(usually 3) supraoculars. 

The frontoparietals and interparietal display the three possible 
degrees of fusion. The degree of fusion is indicated in Table 2 
by a simple notation in which the number of frontoparietals is given 
on the left side of the slash mark (/) and the interparietal is indi- 
cated on the right side. The numeral one means that all three scales 
are fused into a single scale. 

It can be seen from the data in Table 2 that there is a very 
approximate correlation between the state of fusion of the fronto- 
parietals and interparietal and the reduction of the appendages and 
digits. Reduction of the appendages and digits is in turn correlated 
with burrowing habits. It would appear, therefore, that in species of 
more burrowing habits there is a greater tendency for the fronto- 
parietals and interparietal to fuse. This is just one of several evolu- 
tionary trends (see below) in the group and, to me, it seems 
pointless to break the concept of this trend by splitting the spec- 
trum into two parts as Mittleman (1952) did when he set up 
Nodorha as distinct from Rhodona. 

The parietals meet behind the interparietal, and there are 2-4 
pairs of nuchals. 

There is a well defined transparent area or "window" in the lower 



1967 AUSTRALIAN SKINKS: LERISTA 9 

eyelid. In most species the lower eyelid is movable, but in a few 
it has become fused to the upper resulting in an "ablepharine" eye 
(Table 2). In all species of Lerista a supralabial contacts the 
small scales of the lower eyelid below the transparent area and 
thereby acts as a subocular. There are 2-4 (usually 3) supralabials 
anterior to the subocular (= a supralabial). 

The external ear opening is small, that is, it is smaller than the 
transparent area in the lower eyelid, but although the opening is 
minute in several species, it is never completely covered over by 
the epidermis. 

The scales around the midbody are smooth and number 16-24. 
There is a pair of very large preanals. The three midventral series 
of subcaudals are subequal. 

The digital formula (fingers-toes) ranges from 5-5 to 0-1 (Table 
2). Only two species have 5-5 while all others show a reduced 
number of digits, although no species is limbless. The digits are 
covered above by a single row of scales (Group I of Brongersma, 
1942). 

Color patterns in the genus are variations on a striped motif 
(Table 2). A solid, dark, dorsolateral stripe is common and often 
occurs in conjunction with either a third mid-dorsal dark stripe or 
with a pair of mid-dorsal broken dashed stripes on the two dor- 
salmost scale rows. A few species, on the other hand, lack broad 
dorsolateral or mid-dorsal stripes and simply have broken dashed 
stripes on each dorsal scale row. There seems to be little intra- 
specific variation in regard to pattern type. 

In addition to the osteological differences mentioned above, two 
features of the external morphology serve to further separate the 
"ablepharine" Lerista from other Ablepharus and unite them with 
Rhodona in a common genus. These are ( 1 ) the enlarged nasals 
and (2) the reduced number of digits. Of all other Ablepharus only 
A. greyi and A. burnetti have a reduced number of digits (4-5). 
These two Australian species are, however, only distantly related 
to Lerista and will be discussed at a later date. 

It might also be mentioned here that the species of Lerista differ 
from those attenuate, burrowing skinks of the subgenus Lygosoma 
which occur in Australia in having transparent eyelids and external 
ear openings rather than having scaly eyelids and completely cov- 
ered ear openings. 

The differences in skull morphology between Lerista, Hemiergis 
and the solomonis species group have been mentioned above. 



10 BREVIORA No. 267 

Table 1 summarizes the external differences between the three 
groups. 

Geographic distribution of Lerista and its relatives: It is signifi- 
cant that the genus Lerista is found only in Australia and Tasmania. 
If we discount, for reasons given above, any close affinity with 
Ablepharus and Ophioscincus, then it is clear that there are no 
close relatives of Lerista in either Africa or Eurasia. 

The closest relatives of Lerista, namely Hemiergis and the 
solomonis assemblage, are quite obviously a product of the Aus- 
tralian Region. The species of Lerista represent an exclusively 
Australian adaptive radiation into the "surface litter and subsur- 
face burrowing niche." They are certainly not the only lizards to 
have entered this niche — Hemiergis and some of the solomonis 
group species mentioned above also occupy it — but Lerista is one 
of the most successful groups to exploit it, if diversity and abun- 
dance are criteria for such success. Furthermore, it is the most 
specialized species of this genus that have exploited the relatively 
new (geologically speaking) arid areas of Australia. 

EVOLUTIONARY TRENDS WITHIN THE GENUS lERlSlA 

The species of the genus as it is understood here can be arranged 
according to certain structural sequences that are undoubtedly 
indicative of the evolutionary trends that have occurred within 
the group. 

One of these trends has been a reduction in the size of the limbs 
and the number of digits on the fore and hind feet (Table 2). In 
this regard it should be noted that the number of digits on the 
forefront are equal to or less than, but never greater than, the 
number of digits on the hindfoot. This pattern of digit reduction 
is common among lizards, the most noteworthy exceptions being 
the amphisbaenid Bipes biporus (digital formula = 5-0) and the 
scincid genus Opiiiomorus. In the latter genus the number of 
fingers is equal to or greater than, but never less than the number 
of toes (Anderson and Leviton, 1966). 

The reduction of the digits in Lerista is not unexpectedly cor- 
related with other structural changes indicative of greater adapta- 
tion to burrowing habits (Mitchell, 1958); e.g., the size of the 
eye is reduced somewhat, the lower jaw becomes more counter- 
sunk into the upper, and the ear openings become more minute 
but never disappear. 

Although there is a tendency toward smaller external ear open- 
ings in the species of Lerista more closely adapted to a burrowing 



1967 AUSTRALIAN SKINKS: LERISTA 11 

existence (e.g., L. bipes and L. praepedita), the quadrate bone 
undergoes no striking changes in shape from that found in most 
Hzards with external ear openings. The bone is excavated posteri- 
orly, and the tympanum attaches to the lateral rim. By way of com- 
parison, the five species of Hemiergis are less closely adapted to a 
fossorial life than many of the Lerista skinks but have developed 
a simple rod-shaped quadrate concomitant with the loss of the 
external ear opening. 

Several characteristic changes in the skull are found in the ex- 
treme burrowers of the group (L. bipes and L. praepedita) . The 
skull posterior to the orbits is proportionately elongated, and the 
supra- and post-temporal fenestrae are often completely closed over, 
although no elements in the arches are lost. The prefrontal also 
tends to be reduced in its anterior extension, and posteriorly may 
touch the postfrontal to exclude the frontal from the orbit. In 
L. bipes and L. praepedita the premaxillae project forward well 
beyond the level of the premaxillary teeth, a modification which is 
undoubtedly an advantage to these species which employ the snout 
as a burrowing wedge. 

The very approximate correlation between the loss of digits and 
the fusion of the frontoparietals and the interparietal (Table 2) has 
already been mentioned. 

Most of the species of Lerista that have followed these trends to 
the extreme are among Australia's most closely adapted arid area 
dwellers. In this regard they are to arid Australia what Ophiomorus 
and Scincus are to arid southwest Asia and North Africa. The 
similarities are, of course, entirely convergent, for Ophiomorus and 
Scincus, which are rather closely related, are fairly distantly re- 
lated to Lerista. 

MODE OF REPRODUCTION IN lEMSlA 

Nothing has been published concerning the breeding habits of 
these skinks, although from the meager data presented below, it 
is evident that there are both live-bearing and egg-laying species 
in the senus. 

A single Lerista microtis (MCZ 24577) with a snout-vent length 
of 51 mm contains three young in which scales and color pattern 
are evident, indicating that the young would probably be born. 

L. timida, on the other hand, is apparently oviparous. A gravid 
female (snout-vent length = 40 mm) in the Field Museum of 
Natural History (97739) contains two oblong, whitish, leathery 
shelled eggs. An untagged female (snout-vent length = 46 mm) in 
the MCZ collection is likewise gravid with two shelled eggs. 



12 BREVIORA No. 267 

Single females of L. bougainvilli (MCZ 6751; snout-vent length 
■= 60 mm) and L. pimctatovittata (FMNH 911 Al; snout-vent 
length = 85 mm) each contain three spherical, yolky oviducal 
eggs. These eggs were not far enough along in their development 
to allow speculation as to the mode of reproduction in either species. 

THE SPECIES OF LERISTA 

Although a thorough revision of the species in the genus is sorely 
needed, the following table (Table 2) may be of some use to those 
faced with the tedious curatorial duties of identifying and organiz- 
ing collections. The table makes no pretense of being a summary of 
a generic revision and is based solely on the species as they are 
currently recognized in the literature. Two characters important in 
species diagnoses, namely the fusion or independence of the 
frontoparietals and the number of digits, are in the greatest need 
of re-evaluation on the intraspecific level. 

The number of digits is, in fact, already known to vary some- 
what within a single species. Boulenger (1887:335) recorded 
R. gerrardi as having either mono- or didactyl forelimbs, and King- 
horn (1924:180) reported on a specimen of R. pimctatovittata 
with the "forelimb distinctly didactyl" rather than monodactyl. 
Loveridge (1934:372) also found a "rudimentary stump of a sec- 
ond toe" in R. miopa, a species which had previously been consid- 
ered as having only a single toe on the hind foot. 

Mitchell (1955:402) studied the variation in the tarsal (carpal) 
and metatarsal (metacarpal) bones in nine species of Rhodona 
and came to the conclusion that "among specimens of the same 
species . . . from diflferent localities ... the loss of digits and 
digital bones may not be always indicative of specific variation. 
The genus may be found to contain polytypic species within which 
the number of digits varies." 

ACKNOWLEDGMENTS 

Broad revisionary studies within the Scincidae would be im- 
possible without the interest and cooperation of the curators of 
many institutions. The present study has been most generously 
supported through loans and encouragement from the following 
people and their respective institutions. 

Miss A. G. C. Grandison, British Museum (Natural History) 
(BM); Drs. R. G. Zweifel and C. M. Bogert, American Museum 
of Natural History (AMNH); Dr. R. F. Inger and Mr. H. Marx, 
Field Museum of Natural History (FMNH); Drs. A. E. Leviton 



1967 AUSTRALIAN SKINKS: LERISTA 13 

and Steve C. Anderson, California Academy of Sciences (CAS); 
Dr. C. M. Walker, University of Michigan Museum of Zoology; 
Dr. G. S. Myers, Division of Systematic Biology, Stanford Uni- 
versity. 

A special word of thanks goes to Dr. Ernest E. Williams of the 
Museum of Comparative Zoology for his enduring interest in this 
continuing research on the relationships of skink taxa. His 
generosity with the reptile collection under his charge and his 
indefatigable assistance in obtaining specimens from which skulls 
could be prepared have, in fact, made this research possible. 

Both Dr. Williams and my wife Carol have read and criticized 
the manuscript in many helpful ways. 

MATERIALS 

Complete skulls of the following specimens of Lerista have been examined: 
hipes (AMNH 86089); boiigainvilli (MCZ 61403); elegans (FMNH 
11319; fragilis (CAS 77190; MCZ 42988); gerrardi (MCZ 33255); lineo- 
pimctiikita (BM 1902.7.30.5); miopa (MCZ 33260); mueUcri (MCZ 
86699); planiventnile (BM 1954.1-2.21); praepcdita (MCZ 33265); punc- 
tatovitlaui (MCZ 79494); telrudaciyla (BM 1902.7.30.6); timida (MCZ 
X- 13246). 

In addition, the relationship of the palatine, palatal ramus of the ptery- 
goid, and the ectopterygoid were examined in the following specimens of 
Lerista: hipes (MCZ 3^3251, 35349, 35350. 79570); boiigainvilli (MCZ 
67149): elegans (MCZ 33126, 33136); tetradactyla (MCZ 51893); 
timida (MCZ 33152, 33153). 

Nineteen of the 28 species of Lerista were available for examination as 
alcoholic specimens: allanae (2); hipes (20); hougairirilli (20); distin- 
guenda (1); elegans (17); fragilis (many); gerrardi (12); lineata (2); 
lineopunctulata (1); microtis (2); miopa (3); muelleri (18); nichollsi (1); 
planiventrale (2); praepcdita (6); piinctatovittata (24); stylis (1); tetra- 
dactyla (6); timida (20). 

For comparative purposes the following skulls of Lerista's near relatives 
were examined: Hemiergis decresiensis (MCZ 49173); H. initiate (MCZ 
33210); H. peroni (MCZ 24648, 24652); H. quadrilineatum (MCZ 33210); 
H. tridactyliim (MCZ 24595). 

Sphenomorphus antimoriim (MCZ 25374); S. australe (MCZ 24568); 
S. crassicauda (AMNH 82606); S. emigrans (MCZ 27043); 5. pardalis 
(MCZ 35413); S. tenuis (MCZ 35398); S. tryoni (MCZ 35387; 35388); 
5. solomonis (MCZ 72618, 72626, 72664, 72665, 77373, 77374). 



14 BREVIORA No. 267 

LITERATURE CITED 

Anderson, S. C. and A. E. Leviton 

1966. A review of the genus Ophiomorns (Sauria: Scincidae), with 
descriptions of three new forms. Proc. Calif. Acad. Sci., 33(16) : 
499-534. 

BOULENGER, G. A. 

1887. Catalogue of the lizards in the British Museum (Natural His- 
tory). 2nd. ed., vol. 3. xii + 575 pp., British Museum, London. 

Brongersma, L. D. 

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

KiNGHORN, J. R. 

1924. Reptiles and batrachians from south and southwest Australia. 
Rec. Australian Mus., 14(3): 163-183. 

LOVERIDGE, A. 

1934. Australian reptiles in the Museum of Comparative Zoology, 
Cambridge, Massachusetts. Bull. Mus. Comp. Zool., 77(6): 
243-383. 

Mitchell, F. J. 

1950. The scincid genera Egernia and Tiliqiia (Lacertilia). Rec. South 
Australia Mus., 9: 275-308. 

1955. Preliminary account of the Reptilia and Amphibia collected by 
the National Geographic Society — Commonwealth Govern- 
ment — Smithsonian Institution Expedition to Arnhem Land. 
Rec. South Australia Mus., 11: 373-408. 

1958. Adaptive convergence in Australian reptiles. Australian Mus. 
Mag., 12: 314-317. 

MiTTLEMAN, M. B. 

1952. A generic synopsis of the lizards of the subfamily Lygosominae. 
Smithsonian Misc. Coll., 117(17): 1-35. 
Smith, M. A. 

1935. The fauna of British India. Reptilia and Amphibia. Vol. II — 
Sauria., xii + 440 pp., Taylor and Francis, London. 

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

1929. The reptiles and amphibians of South Australia. 270 pp., 
Harrison Weir, Govt. Printer, Adelaide. 



1967 AUSTRALIAN SKINKS: LERISTA 15 







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BREVIORA 

MMseum of Comparsitive Zoology 

Cambridge, Mass. 29 June 1967 Number 268 



THE ECOLOGY AND BEHAVIOR OF TWO 
SYMPATRIC LYGODACTYLUS GECKOS 

Allen E. Greer 

During a two-month field study (23 June-25 August 1965) of 
the behavior and ecology of the reptiles and amphibians of south 
Turkana, Kenya, data were collected on the two closely related 
and ecologically similar Lygodactylits geckos (L. picturatus kenien- 
sis and L. somalicus batter sbyi) which occur sympatrically in the 
area. 

Both species are small as geckos go. Adult L. pictunitus kcnien- 
sis Parker range in snout-vent length from about 26-35 mm and 
are dark grayish brown above with longitudinal light and dark 
markings on the head and shoulders. Adult L. somalicus battersbyi 
Pasteur are smaller, with snout-vent lengths of 21-27 mm. and are 
uniformly light gray to sandy beige on the dorsum. In the field 
it was virtually impossible to tell the sexes of either species apart 
solely on the basis of color pattern. 

Both species, L. picturatus keniensis Parker and L. somalicus 
battersbyi Pasteur, are diurnal as are all the other species in the 
genus, although the ophthalmological evidence indicates that the 
group has reacquired diurnal habits from a nocturnal ancestry 
(Underwood, 1954:470). 

As most geckos are primarily nocturnal, it is not surprising that 
the information which does exist on gecko behavior deals largely 
with the more easily observed diurnal species. Kastle's ( 1964) ter- 
rarium studies o/ Lygodactylus p. picturatus and three species of 
diurnal Phelsuma are among the most comprehensive available for 
geckos. His comparisons between these two groups of diurnal 
geckos should be a stimulus for obtaining more information by 
which the comparisons may be extended to other diurnal (e.g., 
Gouatodes and Sphaerodactylus) and nocturnal species. 



BREVIORA 



No. 268 



Distribution: Lygodactylus is an African-Malagasy genus in both 
origin and distribution. L. pivturatus^ is widely distributed through- 
out east and central Africa and is the most widespread of the 13 + 



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Figure 1. Map showing the distribution of Lxi^iuUictxhis pictiinitiis 
keniensis (area bounded by the dashed hne), and locahties outhning the 
distribution of L. sonutUcits (black dots). The westernmost locaHty for 
L. somaliciis corresponds to the study area where the data for the paper 
were collected. L. piitiiratiis (subspp.) occurs throughout the area shown 
in the map. except on the Arabian Peninsula. The distribution of L. p. 
keniensis is adapted from Pasteur (1964:77, fig. 20). The abbreviation F.S. 
stands for French Somaliland. 



1 I follow Loveridge's (1947) and Pasteur's (1960) taxonomic treat- 
ment of the genus. By simply calling L. pictiiratus a superspecies and 
raising subspecies to species rank, Pasteur (1964) has not substantially 
improved our understanding of the situation. 



1967 I.YGODACTYLUS GECKOS 3 

species in the genus found on continental Africa. L. p. keniensis 
is a subspecies occurring in tiie northern two-thirds of Kenya and 
the bordering edges of the neighboring countries. L. sonuiUcus is 
an endemic of the arid Somali region. L. s. battersbyi ranges from 
the vicinity of the Somaliland-Ethiopian border south into northern 
Kenya (Fig. 1 ). 

Turkana is a district in the extreme northwestern corner of 
Kenya, lying between Lake Rudolf on the east and the Uganda- 
Kenya border on the west. Both geographically and ecologically 
the area is a western outlier of the arid Somali region to the east 
(Schmidt. 1923; Parker. 1932. 1936, 1942). 

As various parts of Turkana have been described in some detail 
elsewhere (von Hohnel, 1894; Worthington. 1932; Fuchs, 1935. 
1939; Buxton, 1937), no more than a brief description of the area 
need be given here. 

South Turkana. where these observations were made (Figs. 
2-3), is extremely arid and consists of numerous rugged plateaus 
and broken hills. Between these extend vast lava and quartzite 
pebble sheets and barren windswept sand flats. Goats and drought 
have kept vegetation to a minimum except along the larger water 
courses, which never carry a permanent supply of water. Acacia 
is by far the most predominant plant group, and it is in the Acacia 
trees that one finds the Lxi^odactyliis geckos. 

The local distribution of the two species in south Turkana was 
a bit peculiar. Both species were quite common in the study area 
except along the Kerio River at Lokori (Fig. 3). This area was 
visited several times during the course of my two-month stay, but 
only three or four individuals of each species were ever seen. This 
scarcity was not too unusual in the case of L. s. battersbyi as it 
was always found less frequently in the large Acacia tortilis which 
is dominant along the larger water courses such as the Kerio, 
Kakurio. and Kalabata. L. p. keniensis, however, was usually very 
common in these acacias, except along the Kerio — an exception 
that merits special consideration below (see Competition section). 

Food: Movement of prey organisms seems to be a prerequisite 
for eliciting a gecko's attention. Food items of L. p. keniensis, as 
determined by field observations and an analysis of the contents of 
ten stomachs, included a wide variety of invertebrates ranging in 
size from Diptera less than three mm long to mantids as long as 
the snout-vent length of the gecko itself. Most food items, how- 
ever, are small (5 mm or less in length). 

Perhaps the most common arthropod in the geckos' microhabi- 
tat and yet one that is never considered as food is an ant species 



4 BREVIORA No. 268 

of the genus Crematogastcr (species near paolii) — large numbers 
of which occur on every Acacia. The ants of this genus are well 
known for their fetid secretions (W. L. Brown and E. O. Wilson, 
personal communication), which may account for their being gen- 
erally avoided by geckos. However, the geckos often "ambush" 
columns of ants returning from a raid on a tree termite nest and 
pluck the largest morsel from the clutches of a particularly suc- 
cessful raider. At other times, the ants are actively avoided, and if, 
on being wounded and disabled, the gecko adheres to the tree, he 
is soon covered with the swarming ants. 

The termites of the Acacia always cover their paths on the sur- 
face of the tree with thin, dirt arches. When one of these arches 
is accidentally broken open, the workers immediately begin to 
repair it. During this process, a worker must expose himself to 
some degree for a few seconds while at the breach, and there is 
rarely a breach without a gecko patiently partaking of the feast. 
There is usually intense competition for the single position at the 
breach and once the dominant animal is removed, his place is 
quickly taken. 

Both species possess great visual acuity, individuals often being 
attracted by a movement of a small (1-2 cm) object 2-2 '/2 meters 
distant. 

Both species of geckos spent a good deal of time licking the 
extremely viscous, amber colored exudate of the Acacia trees, al- 
though the reasons for such actions are unknown. Like Kastle's 
( 1964:489) Phelsuma, the Lygodactylus geckos under my obser- 
vation also licked a variety of jams and jellies, apricot being their 
favorite flavor. 

Water economy with these geckos, as with all reptiles of the 
area, must be of crucial importance. No dew was ever formed 
during the two months of field work, and we encountered rain only 
twice (1, 14 August); both times the amount was insufficient to 
cover the ground. One day a L. s. battersbyi was found licking 
the moisture from near the mouth of a canvas water bag that was 
customarily hung from the tree it inhabited. The gecko appeared 
practically oblivious to movement around him and continued his 
licking for approximately 15 minutes. 

Activity temperatures and periods: Cloacal temperatures of 
three active L. p. keniensis taken during the morning were 32.9, 
34.5 and 35. TC. Air temperatures at the positions from which 
these geckos were taken were 30.8, 32.9 and 35.5'C. respectively. 
Cloacal temperatures for six geckos on the initial loss of the right- 
ing reflex ranged from 43.3-44.3'C. (average 43.7°C.). 



1967 LYGODACTYLUS GECKOS 5 

During the preliminary stages of heat distress L. p. keniensis 
behaves in a manner characteristic of many lizards, i.e., the body is 
raised oflf the ground on straightened legs, the mouth is held open, 
and the respiration rate increases. 

Both L. /;. keniensis and L. s. battersbyi are most active during 
the morning and late afternoon. During the mid-day they usually 
remain stationary, sometimes remaining in the same spot for an 
hour or more. I rarely saw a Lygodactylus on the sunny side of 
a branch. They usually chose to move along the underside of 
branches although, if this side were exposed to the sunlight, they 
would move along the upper, shaded side. 

Approximately 15 to 30 minutes after the sun has set, foraging 
individuals of L. /;. keniensis suddenly begin to ascend the 
branches of their trees. Some seek out a cavity in which to pass 
the night, but the majority of them climb to the peripheral 
branches and, turning so as to face the quickest avenue of escape, 
settle down for the night. In the morning, about 15 to 30 minutes 
before sunrise, they suddenly "come to life" again and quickly 
scurry down the branches to the main part of the tree to begin the 
day's foraging. L. s. battersbyi was also seen ascending and de- 
scending the terminal branches at approximately the same time 
as L. p. keniensis, although no individual of this species was ever 
observed assuming his sleeping position. 

These were by far the longest activity periods of any of the 
diurnal lizards in the area. The lacertids and agamids rarely ap- 
peared until about an hour and a half after sunrise and retired 
shortly before sunset. It thus seems as if the diurnal Lxgochictyliis 
geckos have retained enough of their family's nocturnal heritage to 
enjoy the best of both worlds. 

External niorpliology: Before proceeding with a discussion of 
the behavior of the two species, it will be helpful to describe 
briefly some of the more important aspects of color pattern and 
external morphology (Table 1 ). 

Hatchling L. p. keniensis are 13-14 mm in snout-vent length. 
The dorsum is grayish brown with whitish spots which are bor- 
dered with black along their anterior edges. These whitish spots 
extend onto the dorsal surface of the tail where they become wider 
than long but are still bordered on the anterior edge by black. 
The result is a generally mottled and cryptic appearance. The tail 
is brownish orange above and orange-red below. This bright ventral 
color extends forward along the midventral line to the pectoral 
region. The dark chevron markings of the throat are faintly evident. 



6 BREVIORA No. 268 

The change to the adult pattern entails the loss of the bright 
color on the tail, the change from an orange stripe to a yellow one 
along the ventral midline, the development of roughly longitudinal 
light and dark markings on the head and shoulders, and an 
intensification of the black on the throat. 

Adult males are usually slightly larger and stockier than females. 
The largest male of our specimens has a snout-vent length of 35 mm; 
that of the largest female is 34 mm. The black on the throat of 
the males also tends to be slightly more extensive than in females. 
The light and dark chevrons and white. O-shaped postmental spot, 
said to be characteristic of this subspecies (Loveridge, 1947: 198, 
228), are lacking in some adults of both sexes that have almost 
solid black throats. In the majority of the specimens, however, the 
"typical"' pattern can be seen. 

Preanal pores and escutcheon scales appear in males at a snout- 
vent length of about 26-27 mm. There are usually 6-8 preanal 
pores arranged in an anteriorly projecting, obtuse chevron. The 
grayish escutcheon scales are distributed in four separate areas: the 
underside of the thighs, directly anterior to the preanal pores and 
between the preanal pores and vent. Females lack these secondary 
sexual characters. 

Both sexes have a light yellow midventral stripe on the body 
bordered laterally by a light gray area which extends to the dark 
brown dorsal color of the sides. When the animal is cold, how- 
ever, the pale midventral yellow becomes a very bright mustard 
yellow and the dark chocolate brown color of the dorsum extends 
ventrad to the edge of the bright yellow stripe. 

Hatchling L. .v. battershyi are about 1 1 .5 mm in snout-vent 
length. The dorsum is a cryptic light brown with scattered white 
and dark brown flecks. The tail is faintly tinged with reddish 
orange, and the entire venter is immaculate. 

Adults show little or no sexual dichromatism. Both sexes are 
uniformly brown above with dark brown spots on each shoulder 
and a few less well-defined dark brown spots on the sides. The 
white flecking and reddish tinge to the tail of juveniles are lack- 
ing. There is a thin dark brown horizontal stripe through the eye 
onto the side of the head. 

Adult females average slightly larger than adult males. The 
largest male and female in our sample each measures 27 mm. 

The smallest male with preanal pores and escutcheon scales 
measures 21 mm in snout-vent length. There are usually 6 pre- 
anal pores (two specimens have 5 pores) arranged in an anter- 
iorly projecting obtuse chevron. Only the undersides of the 



1967 LYGODACTYLUS GECKOS 7 

thighs bear escutcheon scales. Females lack these secondary 
sexual characters. 

Both L. p. keniensis and L. s. battersb\i have black pigment in 
the bones of the cranium and a jet black peritoneum. These 
characters have been tentatively associated with diurnal habits 
among geckos (Kiuge, 1967). although experimental work at- 
tempting to elucidate the adaptive significance of this pigment 
distribution has been inconclusive (Hunsaker and Johnson, 1939). 

Adults and hatchlings alike of both species have the tip of the 
tail very slightly spatulate and provided ventrally with a double 
row of lamellae. In general appearance this structure closely re- 
sembles the ventral surface of the expanded digital tip. The tip of 
a regenerative tail has the differentiated ventral "scansorial" scales 
of an original tail, but it lacks the ordered arrangement of a 
double row. 

Loveridge (1947:195), Kiistle (1964:487-488). Mertens 
(1964), and Pasteur (1964:12.76) discuss this structure on 
the underside of the tip of the tail in terms of its "adhesive" or 
"holding" advantage for these arboreal lizards. Such a function 
is suggested by its gross similarity to the digital pads, the fact that 
resting animals often have only the extreme terminal end of the 
tail in contact with the substrate (see Kastle. 1964. fig. 2a) but 
move about with the subterminal area dragging on the substrate 
and the terminal area curved slightly upward, and the fact that 
dead animals have been seen hanging by the tail. 

It is difficult to watch these two species of geckos actively forag- 
ing, especially during the late afternoon when they are most active, 
and not see another possible function of this terminal subcaudal 
expansion. During the times when an individual is very active, 
the tail is frequently raised in a deep arch and the tip of the tail 
is quickly touched to the substrate, withdrawn, and touched down 
again several times. In this manner the gecko quickly makes con- 
tact with parts of the surface behind him through almost 180". In 
this action it seems as if the tail functions less as a grasping organ 
than as a tactile sensory organ. Such a tactile sensory function 
would, of course, be an added advantage in an arboreal life. 

Display: From Kastle's ( 1964:494-495) description of the intra- 
specific display of L. p. picturalus it would appear that the dis- 
play of L. p. keniensis is very similar. In "full display" an adult 
gecko simultaneously raises the body slightly on straightened legs, 
arches the back (exposing the yellow midventral stripe?), bows 
the neck with the snout pointing slightly down and distends the 



8 BREVIORA No. 268 

black throat. The display may be given with the body in a head- 
on or broadside position relative to the individual eliciting the 
response. Often both body positions are assumed alternately or 
successively, with the displaying gecko moving closer to the antag- 
onist in the head-on position. It is also in the head-on position 
that the head is sometimes quickly jerked from side to side through 
a small arc. 

Among adults this display may be given by males to males in 
fighting, by males to females in courtship activities, by females to 
females in fighting, and by females to males in repulsing courtship 
overtures. Except in courtship, the displays never lasted more than 
a few seconds and in only two instances except for courtship 
did they lead to actual physical contact. 

One of these cases was brought about by introducing a foreign 
male into a small scrub Acacia tree occupied by two adult females 
and a single male. The resident male approached the quiescent 
foreign male while displaying head-on and broadside until, after 
a close broadside display, he suddenly attacked the foreign male 
with such force that both geckos fell out of the tree. The resident 
male immediately regained his tree, but the introduced male made 
his way back to the tree slowly and stayed around the base for 
some time. Several days later, however, he was still in the same 
tree, but he always elicited displays from the original male when- 
ever the two met. 

The only component of the display commonly encountered sep- 
arately was the distended black throat. This was often given in 
"mild distress" situations. Another, more common, means of 
aggression was a simple attack or rush at an individual. This be- 
havior was seen in juveniles as well as adults, especially when there 
was a large size difference — in favor of the attacker. 

During the 15 to 30 minutes after sunset that the geckos re- 
mained active, the intensity of the day's aggression between indi- 
viduals declined or diminished altogether. In one instance in 
which two females and one male were occupying a single, small 
Acacia tree, the larger female had been chasing the smaller female 
and male from a favorite food supply at the base of the tree all 
during the afternoon. Shortly after sunset, however, and until 
they retired, all three adults were feeding within centimeters of 
each other at the site. 

It is interesting to note that L. .v. battersbyi, which is so different 
in color pattern from L. p. keniensis, displays in the same way, i.e., 
the body is slightly raised on straightened legs, the back arched. 



1967 LYGODACTYLUS GECKOS 9 

[he neck bowed with his head pointing downward, and the immacu- 
late throat distended. The body may assume the head-on or 
broadside position alternately, and head jerking can be a part of 
the head-on approach. 

Although both species display in the same manner and some- 
times occur in the same tree, the display was never used inter- 
specifically. Confrontations did occur, however, and they always 
resulted in the L. s. battersbyi simply being chased from the 
scene by a larger or more bulky L. p. kcniensis. 

The hatchlings and juveniles of both species were always com- 
pletely ignored by the adults, although the adults often ate moving 
objects larger than the little geckos. The smaller individuals 
showed little fear of the adults, but they always moved aside when 
they found themselves in the path of an adult. 

Foreign objects never elicited a display response from cither 
species. 

Courtship and mating: Courtship and mating were observed 
only once in L. p. keniensis, but the behavior was sufficiently dif- 
ferent from Kasde's description (1964:497-498) to warrant a 
complete description based on my observations. 

The pair was first seen about 12:35 PM on the dark vertical 
trunk of an Acacia tort His along a large watercourse. The male 
was posturing in the typical display, slightly broadside to the fe- 
male. She moved toward his flanks and nipped at the basal third 
of his tail. The male immediately completed the circle by coming 
around to her tail which she would then twitch back and forth 
across his face while moving away with him following. The court- 
ship was interrupted several times at this point, but it was always 
initiated again by the female. After separating 70 to 100 cm. it 
appeared as if the female began searching for the male. She would 
approach his tail, and the male would stiffen slightly and give a 
few quick sideways jerks of the head. The female then nipped at 
his tail and he gave a full display. The female would again ap- 
proach his tail, and the male, completing the circle, would come up 
behind her and follow closely behind as the female moved slowly 
away wagging her tail across his face. 

Mating took place when the male continued creeping forward 
over the female after she had stopped, and grasped the female 
about the body approximately 2 mm posterior to the axilla with 
his forefeet. Holding the skin of the right side of the female's nape 
in his mouth, the male twisted his tail up and under hers from 
the right side, bringing their vents together with the male's tail on 
the female's left side. The left hind foot of the male was held on 



10 BREVIORA No. 268 

the dorsal side of the female's left thigh at its base, and his right 
hind foot was on the tree trunk. After a firm union was obtained, 
the male dropped his forefeet from around the body of the female 
onto the surface of the tree. Matins lasted for about 40 minutes, 
after which time the female flicked her tail sideways several times 
and the male disengaged himself, arching his tail upward in a deep 
bow. Each individual proceeded to lick the area around his vent. 

The cyclic turning of the female on the tail of the male and 
being turned upon by the male, the wagging of the tail in the face 
of the following male and the recurrent initiation of courtship 
activities by the female are elements in the courtship and mating 
of L. p. keniensis not observed by Kastle in his L. /;. picturatiis. 

Males were seen to initiate courtship activity on several occa- 
sions, but were quickly repulsed by a display from the female. 

Only a single mated pair of the more elusive L. s. battcrsbyi was 
observed. The pair was found about 10:30 AM in a shadow on 
the side of a scrub Acacia. 

The male had grasped the skin of the right side of the female's 
nape in his jaws and had brought his vent next to hers from the 
left side. The male's back was slightly hunched and his tail trailed 
behind on the left side. The male's forefeet were employed in 
grasping the female around the pectoral region behind the shoul- 
ders and were not placed on the trunk. His right hind foot rested on 
the dorsal surface of her right thigh at its base, and his left foot 
was in contact with the trunk. 

The female seemed to be made quite nervous by my presence. 
She wagged her tail from side to side, often touching the male's 
back. The female stayed at the same locus but frequently turned 
about 180\ always carrying the male with her and keeping the 
pair oriented along the longitudinal axis of the branch. 

After breaking off copulation, the female quickly disappeared, 
but the male began licking his vent. 

Reproduction: Both L. p. keniensis and L. .v. battersbyi lay two 
white, hard-shelled eggs in cavities of the tree which they inhabit. 
Usually the termites have excavated cavities in one or more 
branches, and, when these are abandoned, they become favorite 
sites for egg deposition (Fig. 4). Eggs will be laid in a particu- 
larly favored spot year after year, the newest pair of eggs being 
deposited on top of the pile. In this way as many as 40 egg shells 
can accumulate in a single cavity. 

L. s. battersbyi lays slightly smaller eggs than L. p. keniensis 
and the eggs of both species can be found in the same cavity. 



1967 LYGODACTYLUS GECKOS 11 

Whole egg shells were frequently found with a small circular 
hole through the shell and the yolky contents completely removed. 
The most likely explanation for this loss is predation by the ants 
which invariably inhabit the Acacia trees. 

Another possible predator of Lygodacrylus eggs or young is 
Homopholis fasciata, a gecko which inhabits, almost exclusively, 
these same cavities in which the Lygodactylus species lay their eggs. 

Developing eggs, hatchlings and gravid females of both species 
were encountered throughout July and August. The smallest fe- 
males of L. p. keniensis and L. s. battersbyi with large, shelled 
oviducal eggs nearly ready for deposition measured 28 and 24Vi 
mm, respectively, in snout to vent length (Table 1 ). In all females 
carrying two large oviducal eggs, the egg in the right oviduct was 
placed more anteriorly in the body cavity than the egg in the left 
oviduct. 

Popiilaiion structure: Almost any scrub Acacia over a meter in 
height was likely to have one or more Lygodactylus inhabiting it. 
In an attempt to get some idea of the population structure of the 
L. p. keniensis in these trees, several of them were held under 
observation and the adults collected as they came to light until 
it was felt that the entire adult complement had been collected. 
In one instance, a large Acacia tortilis was also "collected out." 
The results of such collecting (Table 2) showed that females out- 
number males 2 to 1 . 

Whether the 2 to 1 sex ratio is a reflection of a disproportion- 
ate birth ratio or the result of heavy selection pressure on the 
males cannot be decided at present. 

L. s. battersbyi is so secretive that it is impossible to satisfy one- 
self that the entire adult population of any one tree has been col- 
lected. The general impression, as indicated by the sex ratio of 
the specimens collected (18 males. 26 females), is that females 
may also outnumber males. 

On the open expanses, where the scrub Acacia predominates, 
the population inhabiting a single tree usually consisted of a 
single male, one to three females and a few juveniles. This uni- 
formity of structure is undoubtedly a reflection of the uniformity 
in size and form of the Acacia scrub. Very few scrub Acacia were 
large enough to support two or more males, although this was 
the rule in the much larger A. tortilis. 

In the few instances in which more than one male inhabited a 
tree small enough to allow continuous observation, it was evident 
that males were highly territorial. Females in trees with two or more 
males generally restricted their foraging areas to coincide approxi- 
mately with the territories of the males and were rarely seen far 



12 BREVIORA No. 268 

outside this area. No territorial display of females was ever ob- 
served, although a female will display to another female in certain 
instances, such as confrontation over a food source or a cavity for 
egg deposition. 

Confrontations over a favorite food source of limited size, such 
as a partially opened termite nest, indicated that dominance is 
largely a matter of size and not sex: a large female for instance will 
dominate the position at a breach in a termite archway (see above) 
to the exclusion of the male within whose territory the breach 
may be. 

Kastle (1964:491) reported that his L. pictitratus were never 
observed to come to the ground. Observations on L. p. kcniensis 
and L. .v. battersbyi showed that, although both species are highly 
arboreal, they will come to the ground when attracted by some 
small movement. If the tail of an individual were accidentally 
broken ofT during capture, another gecko in the tree would usually 
come to the ground for the thrashing tail. Geckos could also be 
enticed two or three feet away from the base of the tree by scatter- 
ing wood termites on the ground. During the late afternoon or 
early evening the geckos would often stay near the ground in the 
base of the tree and make forays onto the ground after a small, 
passing invertebrate. 

Individuals that were marked artificially (with paint spots) or 
naturally (lost tail, scars, etc.) were kept under observation for 
as long as three weeks, and in no case was there emigration from 
the "home tree." Strong arboreal tendencies and indications of 
territoriality make it seem likely that most individuals, once they 
have become established in a tree, rarely leave the tree for the rest 
of their lives. 

Competition: As these two species obviously occupy similar 
niches, the ways in which competition may be reduced are of in- 
terest. 

Although it was by no means unusual to find both species occu- 
pying the same tree and foraging within inches of each other, the 
general situation was one species to a tree. In this regard there 
seemed to be some correlation between the bark color of the 
Acacia inhabited and the occupant species, although no systematic 
analysis was undertaken to quantify this impression. 

In the relatively dense stands of the dark-barked A. tortilis along 
the larger watercourses, L. p. keniensis, the darker species, oc- 
curred almost to the exclusion of L. s. battersbyi. Even in the few 
A. tortilis found along the smaller watercourses, L. p. keniensis 
greatly outnumbered L. .v. battersbvi which might also be found 
in the same tree. 



1967 LYGODACTYLUS GECKOS 13 

On the more open flats, the scrub A. etbiaca is the dominant 
tree. The bark of this species is smooth and ranges in color from 
light gray to dark reddish brown. In pure stands of the gray- 
barked trees, L. s. battersbyi, the lighter species, was found to 
the exclusion of L. p. keniensis. 

Between these two extremes, however, the correlation became 
less exact and more difficult to establish. Whether this correlation, 
to the extent that it does exist, is due to habitat selection or random 
dispersal and natural selection for body color on the given substrate, 
or both, cannot be decided on the basis of the data at hand. 

Whenever the two species were found together in the same tree, 
it was more frequently the case that a L. s. battersbyi would be 
found in a "L. p. keniensis tree" than vice versa. In such situations, 
the smaller L. s. battersbyi tended to occupy the more peripheral 
branches, while the larger L. p. keniensis would be found on the 
larger central trunks and branches. 

Adult L. s. battersbyi also inhabited smaller trees and shrubs 
than adult L. p. keniensis. 

Both species were often encountered separately on single trees 
that were quite isolated by stretches (100-200 meters) of lava or 
quartz pebble sheets. In one instance a male and female L. p. 
keniensis and developing eggs were found on a windswept flat in 
a dark-barked Acacia that some time previously had been stripped 
of its foliage leaving only the trunks and main branches (Fig. 5). 
During the afternoon that I worked in this area the tree was hit 
by three large dust devils. 

Beyond these differences, however, there was little indication 
of any other ecological or behavioral differences between the two 
species. This is not too disturbing as many of these differences, 
which are expected on the basis of theory, are hinted at by dif- 
ferences in size and color patterns of the two species. For ex- 
ample, the size difference may be reflected by average differ- 
ences in the size and kind of prey organisms taken by the two 
species. 

It remains to explain the absence of L. p. keniensis from the 
Kerio River at Lokori (Fig. 3), while it is so common along all 
other large river courses investigated. It seems to me that this 
most peculiar local distribution may perhaps be due to the exclu- 
sion of L. /;. keniensis by the somewhat larger Hemidactyliis brooki 
angulatus. This widespread nocturnal gecko was very abundant 
in numerous situations along the Kerio (under the exfoliating bark 
of A. tortilis as well as in earth crevices) but was extremely rare 
beyond the confines of the Kerio except in the chimneys of the 
ubiquitous termite nests. 



14 BREVIORA No. 268 

The Kerio has its headwaters far to the south in the high central 
plateau of Kenya and is undoubtedly the most nearly permanent 
river in this region of south Turkana. It may be that some en- 
vironmental factor such as humidity is more nearly optimal for 
H. b. angulatus in south Turkana only along the Kerio and in the 
termite nests. Under such special conditions in an otherwise hostile 
environment the species may be able to occur to the exclusion of 
L. p. keniensis. 

ACKNOWLEDGEMENTS 

My thanks go first and foremost to Professor Bryan Patterson of 
the Museum of Comparative Zoology for permitting me to accom- 
pany his paleontological expedition to Turkana. Professor Patter- 
son and the other members of the expedition, Messrs. Arnold D. 
Lewis, Muteti Nthale, Daniel Ngumi, William D. Sill, Clyde T. 
Williams, and Roger C. Wood, aided in making the collections 
and observations. Without the encouragement and assistance of 
Dr. E. E. Williams of the Museum of Comparative Zoology, the 
idea of accompanying the Patterson expedition would never have 
come to fruition. Dr. Williams has also supplied many worthwhile 
suggestions during the course of his many readings of the manu- 
script. 

Dr. W. L. Brown of Cornell University very kindly identified 
the single species of ant which I collected, and both he and Dr. 
E. O. Wilson of the Biological Laboratories, Harvard University, 
supplied me with information on the biology of the genus Crema- 
togaster. 

Mr. Thomas Schoener of the Biological Laboratories at Har- 
vard made a thorough examination of the contents of 10 stomachs 
of L. p. keniensis. 

Mr. J. B. Gillet of the East African Herbarium identified the 
plants which I obtained in the Turkana country. 

Dr. A. S. Rand, Mr. George Gorman and my wife Carol have 
read the manuscript and offered several helpful suggestions. 

On behalf of the whole expedition I would like to express our 
appreciation and thanks for the many kindnesses extended to us 
by Dr. and Mrs. R. J. Drysdale-Anderson and their mission staff 
at Lokori. 

My field work in Turkana was supported by a grant from the 
Evolutionary Biology Fund, which is administered for the National 
Science Foundation by the Biological Laboratories at Harvard Uni- 
versity. Professor Patterson's expedition was supported by NSF 
Grant G. P. 1188. 



1967 LYGODACTYLUS GECKOS 15 

LITERATURE CITED 

Buxton, D. R. 

1937. A natural history of the Turkana fauna. J. East Africa and 
Uganda Nat. Hist. Soc, 13 (3-4) . -85-104. 
FucHS, V. E. 

1935. The Lake Rudolf Rift Valley Expedition. 1934. Geograph. J.. 
86 (2):114-142. 

1939. The geological history of the Lake Rudolf Basin, Kenya 
Colony. Phil. Trans. Roy. Soc. London, Ser. B. 229 (560): 
219-274. 

HOHNEL, L. VON 

1894. Discovery of Lakes Rudolf and Stefanie. Longmans, Green 
and Co., London, 2 vols. Translated by Nancy Bell. 
HuNSAKER, D. and C. Johnson 

1959. Internal pigmentation and ultraviolet transmission of the in- 
tegument in amphibians and reptiles. Copeia, 1959, No. 4: 
311-315. 

Kastle, W. 

1964. Verhaltensstudien an Taggeckonen der Gattungen Lyxodcuuylii.s 
und Plielstima. Zeitschrift flir Tierpsychologie. 21 (4):486-507. 
Kluge, a. G. 

1967. Higher taxonomic categories of gekkonid lizards and their 
evolution. Bull. Amer. Mus. Nat. Hist.. 13S (1 ):l-65. 

LOVERIDGE, A. 

1947. Revision of the African lizards of the family Gekkonidae. Bull. 
Mus. Comp. Zool.. 98 { 1) : 1-469. 
Mertens, R. 

1964. Der Eidechsenschwanz als Haftorgan. Senck. Biol., 45 (2): 
1 17-122. 
Parker, H. W. 

1932. Scientific results of the Cambridge Expedition to the East 
African Lakes, 1930-1931. 5. Reptiles and amphibians. J. Linn. 
Soc. London, Zool., 38 (258) :2 13-229. 

1936. Reptiles and amphibians collected by the Lake Rudolf Rift 
Valley Expedition, 1934. Ann. Mag. Nat. Hist., (10) 18:594- 
609. 

1942. The lizards of British Somaliland. Bull. Mus. Comp. Zool., 
91 (1 ):1-101. 
Pasteur, G. 

1960. Notes preliminaires sur les lygodactyles (Gekkonides). L — 
Remarques sur les sous-especes de Lyi^iutdcfyln.s pictiiratiis. 
Bull. LF.A.N., Ser. A, 22 (4) : 1441-1452. 

1964. Recherches sur revolution des lygodactyles, lezards .Afro- 
Malagaches actuels. Trav. Inst. Scient. Cherif., Ser. Zool., No. 
29:1-^32. 



16 



BREVIORA 



No. 268 



SCHMIDI, K. P. 

1923. Contributions to the herpetology of the Belgian Congo based 

on the collection of the American Museum Congo Expedition. 

1909-1915. Part II.— Snakes. Bull. Amer. Mus. Nat. Hist.. 

49:1-146. 
Underwood, G. 

1954. On the classification and evolution of geckos. Proc. Zool. .Soc. 

London. 124 (3):469-492. 

WORTHINGTON, E. B. 

1932. The lakes of Kenya and Uganda, investigated by the Cambridge 
Expedition, 1930-1. Geograph. J., 79 (4):275-297. 



TABLE 1 



Summary of the size and color pattern differences between 
Lygodactylus pictwatus keuicnsis and L. somalicus battersbyi 



pictiiniliis 



Snout-vent length (in mm.) of: 
Hatchling 

Largest adult male 

Largest adult female 

Smallest male with 
escutcheon scales 

Smallest female with 
oviducal eggs 



13-14 (N = 10) 
35 (N = 53) 
34 (N = 79) 

26-27 (N = 53) 

28 (N=12) 



sonuiliiiis 



111/2 (N=3) 
27 (N=17) 
27 (,N = 26) 

21 (N=18) 

241/2 (N=3) 



Color pattern of: 
Juvenile 



Adult 



Throat with faint 
dark chevrons; venter 
with orange-red mid- 
ventral stripe. 

Longitudinal light 
and dark markings on 
head and shoulders: 
dark black throat 
chevrons; venter 
with yellow mid- 
ventral stripe. 



Throat and venter 
immaculate. 



Head and shoulders 
uniform — color 
of dorsum; throat 
and midventer 
immaculate. 



1967 



LYGODACTYLUS GECKOS 



17 



TABLE 2 



Summary of data on sex ratios of total adult complement of 
Lygodactylus pictwatiis keniensis inhabiting single trees 



Number 
adult lua 


of 
les 




Number of 
adult females 


Number of 
juveniles 









6 
8 
3 
1 

3 
2 
3 

2 
2 
2 
3 




1 or more 

1 





2 

1 or more 

2 



2 

4 or more 


15 







21 ( 
2 

2 

1 




+ 47?) 


several 



1 



4 or more 




30 






1 

64 






20 + 


Ratio males/f 


emales = 


1/2 


1 







18 



BREVIORA 



No. 268 





Figure 2. Typical view of the open flats in the study area. 




Figure 3. The dry bed of the Kerio River near Lokori. 



1967 



LYGODACTVLUS GECKOS 



19 




Figure 4. Two pairs of Ly^odacyliis puiiiniiiis kciiiciisis eggs in the 
termite-excavated cavity of a scrub Aciuiu. Part of the dead branch has 
been broken away to expose the eggs. 




Figure 5. Defoliated Acacia, on a wind blown fiat, harboring a male 
and female Lxt^ocldcixli/.s picturniiis kenieasis and developing eggs. 



BREVIORA 

Mmsemm of Coimiparative Zoology 

Cambridge, Mass. 29 June 1967 Number 269 



TWO NEW SPECIES OF AMPHITARSUS (OPHIUROIDEA) 
FROM THE WESTERN NORTH ATLANTIC 

Amy Schoener 

INTRODUCTION 

The brittlestars discussed in this paper were recently obtained 
from the Western North Atlantic, at depths of between 200 and 
535 m. Two new species are described and both are referred 
to the previously monotypic genus Amphitarsus H. L. Clark 
(1941). The type species of Amphitarsus is A. mirabilis (by orig- 
inal designation), remarkable in possessing peculiar winglike 
structures on the oral surface of the disk; these structures are in 
reality lateral arm plates which have become greatly expanded in 
the genital region. The two new species have similar structures 
(Figs. 1, 2, 3, B). Both species are apparently rare, since only 
seven specimens in all have been obtained in collections which 
included several thousand brittlestars. 

In 1941 H. L. Clark described the monotypic ophiuroid genus. 
Amphitarsus, from the Caribbean region. The familial relationship 
of the genus was ambiguous, since the mouth papillae, which are 
extremely important diagnostic characters, varied considerably, 
and Clark observed that "no two of the jaws look exactly alike." 
Clark tentatively placed it in the Amphiuridae. No further contri- 
bution towards the solution of this problem has since been made. 
Recent collections obtained by Drs. Robert Hessler and Howard 
Sanders (Woods Hole Oceanographic Institution), as part of a 
detailed study of benthic assemblages of the Western North Atlan- 
tic (Sanders, Hessler and Hampson, 1965), have yielded two new 
species of this genus, one of which apparently sheds light on its 
familial relationship. 



2 BREVIORA No. 269 

Amphitarsus mirabilis Clark 
Figure 1 

Amphitarsus mirabilis H. L. Clark, 1941, Mem. Soc. Cubana Hist. Nat., 
15 (1): 83-85, pi. 8. Holotype, MCZ 6232 from off Cayo Coco, Camaguey 
Province, Cuba, from 230 fathoms (420m). Two paratypes, MCZ 6398, 
6399, from off Santa Clara Province, Cuba, from 175-235 fathoms (320- 
430m). 

Description: Disk: Diameter ca. 6 mm. Aboral disk surface (Fig. 
1, A) covered with fine granulation; very short radial areas 
alternate with much longer and concave interradial areas; radial 
shields small and narrow. Oral interradial surface (Fig. 1, B) 
densely covered with minute spinules or pointed granules. Oral 
shields rounded, pentagonal, slightly swollen, wider than long, and 
not granulated. Oral papillae in two groups: inner oral papillae 
typically two; short, swollen, and located on each side of the jaw 
angle, the innermost forming the pair characteristic of the Amphi- 
uridae, but on several of the jaws two of the papillae have fused 
laterally into a single wide papilla; outer oral papillae consist of a 
group guarding the large outer oral tentacle. Genital plate appears 
to be fused with four basal winglike arm plates. 

Arms: Reaching a length of 50-60 mm. Arm spines, 9. Ven- 
tral arm plates rather poorly defined, more or less pentagonal. 
Tentacle scales: 1 very large nearly round scale present through- 
out most of the arm; 1-3 smaller scales on basal arm joints (Fig. 
1,C). 

Remarks: Clark's original description was accompanied by 
photographs of the holotype but no detailed illustrations were 
included. 

Amphitarsus nike' new species 
Figure 2 

Holotype: MCZ 6797, about 140 miles N. of Surinam; Latitude 
1° 53.5' N; Longitude 54° 33.3' W. Depth: 535 m. Woods Hole 
Oceanographic Institution CHAIN cruise 35, Dredge Station No. 
33. April 25, 1963. 

Paratypes: From the same locality, MCZ 6799, 6800. 

Description: Disk: Diameter of 7.4 mm measured from outer 
radial shield edge to opposite edge of disk. Disk subpentagonal, 
flattened, and covered with fine granulation. Primary and other 

^ The specific name, a nominative in apposition, refers to the Winged 
Victory (Nike of Samothrace). 



1967 



NEW SPECIES OF AMPHITARSUS 



.. • •'. < — ^v.. 



'•r-:"t-.'-'":'Uly(^ .■'■.■;. / radial shield 
V'.-'.'" '" i^---- ''"'■'! 

dorsal arm plare 

lateral arm plate 




0,5nim 



.arm spme 



J.v';.-;,^ latere I arm plate 



u_arm spine 



tentacle scale 



ventral arm plate 



Figure 1. Ainphiiursiis mirabilis H. L. Clark, 1941. A, Aboral view of 
arm and part of disk. g.Oral view of two arms and part of disk. C, Lateral 
aspect of arm. 



4 



BREVIORA 



No. 269 



C ^ 



6 mm 



arm spme 











oral papilla 




oral shield 
genital area 



lateral arm plate 



tentacle scale 
ventral arm plate 



Imm 



Figure 2. Amphitarsus nike n.sp. A, Aboral view of arm and part ol 
disk. B, Oral view of arm and part of disk. C, Lateral aspect of arm. 



1967 NEW SPECIES OF AMPHITARSUS 5 

plates not visible, and only the tips of the radial shields exposed. 
Midline between pairs of radial shields barely distinguishable. Oral 
interradial surface of disk covered with fine granulation; oral 
shields small, subtriangular with rounded angles and a slightly de- 
pressed convex base; length about 0.9 mm. and width 0.8 mm. 
Adoral shields narrow proximally, becoming much wider at base 
between second oral tentacle pore and oral shields. A single papilla 
generally present at the jaw apex, but in other cases it is difficult 
to distinguish whether one, two, or none of the papillae are 
occupying this position. Inner oral papillae: four to five, conical, 
becoming broader and oval towards the outer edge of the jaw. A 
vertical series of teeth occurs at the jaw apex. Oral tentacle pores 
partially visible, opening into the mouth angle. Genital clefts 
elongate, flaring from the ventral arm plate and narrowing at the 
margin of the disk; they extend from the adoral shield to the margin 
of the disk. These clefts are superficially partitioned by five over- 
lapping winglike lateral arm plates; these plates begin at the second 
ventral arm plate and are attached to the lateral arm plate and 
not directly to the disk itself. 

Arms: Arms incomplete, at least 36 mm beyond the disk on 
7.5 mm specimen. At the arm base, the fifth (outermost) over- 
lapping winglike lateral plate is divided into two or three flat, 
finger-like projections which are visible from the aboral side of 
the disk. Dorsal arm plates angular, broadened ovals, i.e. with 
obtuse angles inward in the proximal segments, becoming widelv 
oval after the first six segments. Narrow lateral arm plates meet 
at the midline on both oral and aboral surfaces, thereby separating 
the ventral and dorsal arm plates, respectively. Arm spines: 
three occur on most of the lateral arm plates; four are present 
on those joints which carry the winglike lateral plates. Beyond 
the disk, upper arm spine ca. 0.8-1.0 arm joints in length; lower 
two spines increasing slightly in length, the most ventral spine 
being ca. 1.2 arm joints in length. Ventral arm plates indistinct, 
somewhat broadened, hexagonal, with narrower proximal portions 
and wider, distal, convex portions beyond the disk. Tentacle pore 
large, and conspicuous, round-oval; on the basal pores there are 
two to three tentacle scales of reduced size. Thereafter, there is a 
single, flat, tentacle scale covering the entire pore. 

Color in life: Unknown; color in alcohol and dry specimens: 
whitish. 

Specimens examined: Holotype and paratypes. 



6 BREVIORA No. 269 

Amphitarsus spinifer^ new species 
Figure 3 

Holotype: MCZ 6798, about 200 miles east of New York City; 
Latitude 40 01.8' N; Longitude 70 42' W. Depth: 200 m. Woods 
Hole Oceanographic Institution ATLANTIS cruise No. 283; 
WHOI Benthic Slope Station No. 2. August 28, 1962. 

Paratypes: From the same locality, MCZ 6801. 

Description: Disk: Disk diameter 3.8 mm from the outer edge 
of the radial shields to the opposite disk margin. Short radial areas 
alternate with longer interradial areas; disk subpentagonal, gen- 
erally flat, covered with multitudinous somewhat poorly-defined 
scales with isolated spines present on many of them; primary 
plates clearly visible, much larger than the surrounding scales, 
irregular in shape and separated from each other by the smaller 
disk scales. Radial shields clearly exposed, closely paired but not 
touching, and less than half the disk radius in length; length about 
0.6 mm and width 0.2 mm. Oral disk interradial surface covered 
with scales and isolated spines similar to those on the aboral sur- 
face; the spines are largest on the centro-dorsal disk surface {ca. 
0.4 mm long), becoming slightly smaller and narrower on the disk 
aboral margin {ca. 0.3 mm long), and decreasing to 0.25 mm 
on the oral surface. Genital clefts elongate, flaring from the first 
lateral arm plate, narrowing at the disk margin, extending from 
the oral shield to the margin of the disk. Clefts partitioned super- 
ficially by two overlapping plates which begin at the second ventral 
arm plate, to whose surface these winglike lateral plates are joined. 
Oral shields small, length ca. 0.3 mm and width ca. 0.5 mm, 
rounded triangular, with a slightly convex distal margin which 
buds off from the triangle base thereby creating two laterally pro- 
truding bulges just proximal to it. Adoral plates long, narrow, 
slightly widened where they meet on the midline. Two conical oral 
papillae present, on each side of the jaw, the terminal one form- 
ing one side of the pair characteristic of the Amphiuridae. The 
second papilla is separated by a diastema and is closest to the 
papillae encircling the second tentacle pore. Jaw character, nearly 
constant in specimens examined. 

Arms: In a smaller specimen (of 3.4 mm disk diameter), the 
arms, though broken, extend 20 mm beyond the disk edge; in the 
specimen described, the break is 6 mm from disk edge. At the 



1 The specific name is intended to call attention to the spiny character 
of the disk.  



1967 



NEW SPECIES OF AMPHITARSUS 



0.4inm 



B 



— — radial shield 





'i^^— arm spine 
dorsal arm plate 



arm spme 



oral papilla 

-s^oral shield 




disk spine 



5^ 

-^ — lafcral arm plate 



arm spme 



0.5mm 



Figure 3. Amphitarsus spinifer n.sp. A, Aboral view of arm and part of 
disk. B. Oral view of two arms and part of disk. C. Lateral aspect of arm. 



8 BREVIORA No. 269 

arm base, second (outermost) winglike lateral plate, visible from 
the aboral side, is broken into five, flattened, finger-like projections. 
Dorsal arm plates rounded hexagonals, length exceeding width. 
Arm spines: six at arm base, five to four further out on arms; 
two spines on joints carrying winglike lateral plates. Beyond disk, 
arm spines ca. 0.4 mm, equalling approximately the length of an 
arm joint; dorsalmost spine is shorter and flat. Tentacle scales: 
two on proximal joints; one externally placed, rounded, leaflike 
scale; one internally placed erect spine, attached to the median edge 
of the ventral arm plate (present only in first six-seven segments). 
Ventral arm plates basically hexagonal, with deeply emarginated 
lateral areas where one tentacle scale attaches. 

Color in life: Unknown; color when dry: whitish. 

Remarks: Three of the four paratypes were fragmented, but it 
was clear that they were not from the same individual, since each 
bore an almost complete set of primary plates on the aboral disk 
surface. The fourth was badly mangled. 

Specimens examined: Holotype and paratypes. 

DISCUSSION 

The three species of the genus Amphitarsus illustrated and dis- 
cussed above possess the singularly striking character of winglike 
lateral arm plates in the genital region. Although variable in num- 
ber, these plates distinguish the genus from all others. If the main 
characteristic relating these species is taken to be these unusual 
winglike plates, then the genus could be placed in the family Am- 
phiuridae, on the basis of the jaw structure of A. spinifer. The 
presence of two papillae at each jaw angle in A. spinifer is quite 
constant, even though the species is represented by smaller indi- 
viduals than either of the other two species; perhaps the younger 
stages of A. spinifer differ less from the adult condition, or it may 
not be as immature as is suspected on the basis of size alone. The 
variation in the oral papillae evidenced in A . mirabilis and A . nike 
may be due to the immaturity of these specimens. All individuals 
of this genus so far as known are fairly small, although one arm 
fragment possibly belonging to A. spinifer suggests that a larger 
size is attained. Thus it is possible that these are young individuals, 
and if this is the case, they might belong to the Ophiacanthidae or 
even to the Ophiocomidae (Fell, personal communication). 

At present the genus is known only from the Western North 
Atlantic. Whether or not it is truly confined to this region, only 
continued investigations of other areas of the ocean will tell. 



1967 NEW SPECIES OF AMPHITARSUS 9 

This work was carried out at the Museum of Comparative Zool- 
ogy during the tenure of fellowships from the International Wom- 
en's Fishing Association and from Harvard University; research 
facilities were provided under NSF grant GB-3532, and speci- 
mens were collected by the Woods Hole Oceanographic Institution 
under NSF grants GB-15838 and GB-563. I wish to thank Dr. 
H. B. Fell of this department for his guidance and advice, Dr. R. 
Turner for reading the manuscript, and Drs. R. Hessler and H. 
Sanders for allowing me to study their collections. 

BIBLIOGRAPHY 

Clark, H. L. 

1941. Report of scientific investigations of the Atlantis expedition to 
the West Indies. Mem. Soc. Cubana Hist. Nat. 15 (1):83-85. 
Fell, H. B. 

1960. Synoptic keys to the genera of Ophiiiroidea. Zool. Piibl. Vic- 
toria Univ., Wellington, No. 26:1-44. 
Sanders, H. L., R. R. Hessler, and G. R. Hampson 

1965. An introduction to the study of deep-sea benthic faunal 
assemblages along the Gay-Head — Bermuda transect. Deep 
Sea Res. 12: 845-867. 



BREVIORA 



MiLiseiiaiM of Comparsitive Zoology 

Cambridge. Mass. 29 June 1967 Number 270 

ADDITIONS TO THE UNIONID FAUNA OF THE GULF 

DRAINAGE OF ALABAMA, GEORGIA AND FLORIDA 

(MOLLUSCA: BIVALVIA) 

Richard I. Johnson 



More than a decade has passed since Clench and Turner ( 1956) 
published their pioneering work, "Freshwater MoUusks of Ala- 
bama, Georgia, and Florida from the Escambia to the Suwannee 
River." It is not surprising that in the time since the publication 
of that paper certain additions and corrections have been, and 
can be, made concerning the Unionidae. 

Athearn (1964) added two new species, Lcunpsilis haddleton'i 
and Villosa choctcnvcu.sis. both from the Choctawhatchee River, 
Alabama. Johnson (1965) described Auodonta peggyac which 
ranges from the Choctawhatchee River, Alabama, to the Hills- 
borough River, Florida. Athearn (1964) convincingly differenti- 
ated Lcunpsilis jonesi Vander Schalie from L. australis Simpson, 
both from the Choctawhatchee River. Alabama, and regarded as 
synonymous by Clench and Turner. 

In 1964 Athearn collected a live specimen of Lcunpsilis bimym- 
inatiis Simpson in the upper Flint River drainage of Georgia, which 
confirmed its distinctness from Lcunpsilis cxccivaius (Lea) and vali- 
dated the earlier records thought to be in error by Clench and 
Turner. An examination of the type, and of subsequently col- 
lected specimens, has clearly shown that Alasmidonta wrii^/uicuui 
(Walker) of the Ochlockonee River, Florida, is distinct from A. 
iricuigulata (Lea) of the Apalachicola River system of Georgia 
and Florida. Under Amulontoides radicitus (Conrad), referred 
to as A. elliotti (Lea) by Clench and Turner, are included records 
of Strophints subvexus (Conrad) from the Chipola River, Florida. 
Obovarici mtulota (Wright) from the Escambia River, Florida, 
was overlooked and has recently been rediscovered. 

This paper is intended to be supplemental to the work of Clench 
and Turner. All of the species mentioned here are figured except 



2 BREVIORA No. 270 

Anodonta peggyac Johnson which I described and figured in 
1965. Strop/lit us subvexus (Conrad), Anodontoides radiatus 
(Conrad), Lampsilis binominatus Simpson, and L. jonesi Van- 
der SchaHc are fully described. Remarks are given on Alasmidonta 
wrightiana (Walker), Obovaria rotulata (B. H. Wright), Lampsilis 
haddk'toni Athearn, and Villosa choctawcnsis Athearn. 

All of the extant types of the various named forms mentioned 
here have been examined and photographed. I wish to thank Mr. 
Herbert D. Athearn for allowing me to examine his specimen of 
Lampsilis binominatus Simpson. Thanks are also extended to Drs. 
K. J. Boss, W. J. Clench, and R. D. Turner for reading the manu- 
script and suggesting improvements. 

The following abbreviations have been used. 

ANSP — Academy of Natural Sciences of Philadelphia, Penn- 
sylvania. 
MCZ — Museum of Comparative Zoology, Harvard Uni- 
versity, Cambridge, Massachusetts. 
MZUM — Museum of Zoology, University of Michigan, Ann 

Arbor, Michigan. 
USNM — United States National Museum, Washington, Dis- 
trict of Columbia. 
All specimens mentioned in the text are in the Museum of Com- 
parative Zoology unless otherwise noted. 

Strophitus SUBVEXUS (Conrad) 
Plate 1, fioures 1-3 

o 

Anodonta siihvexa Conrad 1834, .Amer. Jour. Sci.. 25: 341, pi. !. fig. 12 
(Black Warrior River [Alabama] type, ANSP [lost] ). 

Mciri^arifdna connasangaensis Lea 1858. Proc. Acad. Nat. Sci. Phila., 10: 
138 (Connasaiiga River, one of the headwaters of the Alabama River, 
Gilmer Co.. Georgia). Lea, 1859, Jour. Acad. Nat. Sci. Phila.. (2)4: 
229, pi. 32. fig. 113; figured holotype USNM 86277. Lea 1859, Obs. 
Unio. 7: 47. 

Margcirilana spilUnunii Lea 1858, Proc. Acad. Nat. Sci. Phila., 10: 138 
([Magbys Creek, written on type, not located on modern county map] 
Tombigbee River, near Columbus [Lowndes Co.[ Mississippi). Lea, 
1862, J^our. Acad. Nat. Sci. Phila., (2) 5: 105, pi. 17, fig. 252; figured 
holotype USNM 86278. Lea, 1862, Obs. Unio, 8: 109. 

Muri^aritana tombecbeensis [sic\ Lea 1858, Proc. Acad. Nat. Sci. Phila., 
10: 138 (Tombecbee River, Columbus [Lowndes Co.] Mississippi). 

Margcirilana tombigbeensis Lea 1862, Jour. Acad. Nat. Sci. Phila. (2) 5: 
107, pi. 18, fig. 255; figured holotype USNM 86253. Lea, 1862, Obs. 
Unio, 8: 111. 



1967 GULF DRAINAGE UNIONIDS 3 

Margaritana gesnerii Lea 1858, Proc. Acad. Nat. Sci. Phila., 10: 138 ( Up- 

haupee Creek [Macon Co.] Alabama; Chattahoochee River, below 

Columbia [sic] [Columbus, Muscogee Co.] Georgia). Lea, 1862. 

Jour. Acad. Nat. Sci. Phila. (2 ) 5: 211. pi. 32, fig. 280; figured holo- 

type USNM 86212 from Uphaupee Creek. Lea. 1863. Obs. Unio. 

9: 33. 
Morgaritaiia aJahamensis Lea 1861, Proc. Acad. Nat. Sci. Phila.. 13: 41 

(Talladega Creek [Talladega Co.] Alabama). Lea. 1862. Jour. Acad. 

Nat. Sci. Phila.. (2)5: 104. pi. 16. fig. 249; figured holotype USNM 

86262. Lea, 1862. Obs. Unio, 8: 108. 
Margaritana coliimheiisis Lea 1867, Proc. Acad. Nat. Sci. Phila.. 19: 81 

(Tombigbee River, near Columbus [Lowndes Co.] Mississippi, type not 

in USNM [lost]). 
Strophitus conasaiigaensis [sic] (Lea). Ortmann. 1923. Nautilus, 36: 130. 
Strcjpliifiis subvexiis (Conrad). Frierson. 1927. Check List North American 

Naiades, p. 23. 
Strophitus spillnuinii (Lea). Vander Schalie, 1940. Lloydia. 3: 197. 
Anodontoides elliotti (Lea), partini. Clench and Turner. 1956. Bull. 

Florida State Mus.. 1: 182. 

Description. Shell medium in size, reaching about 100 mm 
in length. Outline subrhomboidal, becoming subelliptical or trape- 
zoidal with age. Valves inequilateral, subcompressed to rather 
inflated, thin. Anterior end regularly rounded, posterior end 
obliquely subtruncate above, rounded or subtruncate below. Ven- 
tral margin almost straight, often becoming slightly arcuate with 
maturity. Dorsal margin short, forming a broad angle with the 
obliquely descending posterior margin. Hinge ligament short but 
prominent. Posterior ridge generally quite broad and very faintly 
doubie. Posterior slope flat or slightly concave. Umbos moder- 
ately swollen, slightly raised above the hinge line, located from 
about the center to the anterior third of the valves, their sculpture 
consisting of a few very strong ridges running parallel to the growth 
lines, somewhat interrupted at the posterior ridge. Surface of the 
shell smooth, generally shiny, waxy yellow, greenish yellow, 
brownish, often rayless, though sometimes with green rays espe- 
cially on the posterior slope. Occasionally the surface has numerous 
broad green rays which give the whole shell a greenish appearance. 

Left valve with a stumpy, vestigial pseudocardinal tooth directly 
below the umbo, sometimes with traces of first and third teeth, 
before and behind it. No lateral teeth. Right valve with one rather 
well developed pseudocardinal, triangular and compressed, or 
tubercular, knob-like and stumpy. 

Umbonal cavities moderately deep with dorsal muscle scars. 



Length 


Height 


Widtl 


mm 


mm 


mm 


59 


36 


25 


62 


36 


24 


77 


52 


37 


92 


56 


39 


93 


51 


40 


116 


54 


38 



4 BREVIORA No. 270 

Anterior adductor muscle scars well impressed, posterior ones less 
so. Pallial line faint. Nacre bluish, dull salmon or purplish, often 
spotted with yellow. 

Measurements 



Topotype of S. suhrcxiis (Conrad) MCZ 

146655. 

Holotype of M. c:>niui.sau^aen.sis Lea 

Holotype of M. tombigheensis Lea 

Holotype of M. gesnerii Lea 

Holotype of M. spillnumii Lea 

Holotype of M. alabamensis Lea 

Remarks. Strophitus siihvexiis (Conrad) of the Gulf drainage, 
shows some of the same diversity of shape as does Strophitus undu- 
kitus (Say) of the Atlantic drainage and interior basin. It is read- 
ily distinguished from itndiilaius which has entirely rudimentary 
pseudocardinals, represented, if at all, by slight swellings. In S. 
siibve.xus there is at least one pseudocardinal in each valve, and 
while small, they are usually well developed, triangular and com- 
pressed, or tubercular, knob-like and stumpy. It has also been 
confused with Anodontoides radiatus (Conrad), but that shell is 
always regularly elliptical with rays over the entire surface, while 
subvexus is subelliptical, inclined to be biangulate behind, seldom 
with rays over the entire surface, and with pseudocardinals that 
are less rudimentary. 

Vander Schalie (1940) correctly reports this species from two 
localities on the Chipola River as Strophitus spillmanii. Clench 
and Turner ( 1956) assumed that Vander Schalie's specimens were 
Anodontoides elliottii (Lea) \= radiatus (Conrad)], but all of 
their records of radiatus from the Chipola River are S. subvexus. 

Ortmann (1923) reviewed some of the names applied to the 
forms of this species, but did not include those from the Tombigbee 
River drainage. S. subvexus (Conrad), like its synonyms con- 
nasaugaensis Lea and tombigbeensis Lea, was founded on rather 
trapezoidal inflated individuals, while other names were given 
to the more common subelliptical specimens. Conrad's figured 
type which was about 2 inches (51 mm) in length, was specifically 
stated to be in the Academy of Natural Sciences of Philadelphia, 
but it has been lost. He did not give a precise locality, but in all 



1967 GULF DRAINAGE UNIONIDS 5 

probability the type came from the Black Warrior River in the 
vicinity of Tuscaloosa, Tuscaloosa Co., Alabama, since Conrad 
is known to have been there in June of 1833 (Wheeler, 1935: 38). 
Range. Gulf drainage, from the Pascagoula River system of 
Mississippi to the Apalachicola River system of Georgia and 
Florida. 

SPECIMENS EXAMINED 

PASCAGOULA RIVER SYSTEM 

Mississippi: Edahoma Creek, 6 mi. NW Soso, Jones Co. 
ALABAMA COOSA RIVER SYSTEM 

Tonibigbee River Drainage. — Mississippi: Tombigbee River, 
Columbus, Lowndes Co. Alabama: Coalfire Creek, Coalfire. Pick- 
ens Co. Bodka Creek, 5 mi. NW Gainsville, Sumter Co. Okatuppa 
Creek, 4 mi. SE Toxey, Choctaw Co. Black Warrior River. Tusca- 
loosa Co. 

Cahawba River Drainage. — Alabama: Black Creek, St. Clair 
Co. Shoal Creek, Montevallo, Shelby Co. 

Coosa River Drainage. — Georgia: Teloga Creek; Chattooga 
River, Summersville; both Chattooga Co. Alabama: Mills Creek, 
Cherokee Co. Tennessee: Conasauga River, Conasauga, Polk Co. 
Georgia: Conasauga River, Gregorys Mill, 10 mi. N Eaton, Mur- 
ray Co. Conasauga River, Gilmer Co. (USNM). Conasauga River. 
1 .4 mi. N Resaca, Gordon Co. Black Creek. St. Clair Co. Choc- 
ciocca Creek, 3 mi. S Lincoln; Talladega Creek; both Talladega 
Co. 

Tallapoosa River Drainage. — Alabama: Uphaupee Creek, 
Macon Co. 

Alabama River Drainage.- — Alabama: Cub Creek, Pine Hill, 
Wilcox Co. 

APALACHICOLA RIVER SYSTEM 

Chipola River Drainage. — Florida: Big Creek, 8 mi. W 
Malone, Jackson Co. Cowarts Creek, nr. Florida state line, Hou- 
ston Co. (MZUM). Reedy Creek, 6 mi. W Malone; Chipola River. 
1 mi. N Marianna; both Jackson Co. Chipola River. Scotts Ferry, 
Ca'hounCo. (MZUM). 

Chattahoochee River Drainage. — Georgia: \ Chattahoochee 
River] below Columbus. Muscogee Co. (USNM). 

Flint River Drainage. — Georgia: Kinchafoonee Creek, 4 mi. N 
Bronwood, Terrell Co. Ichawaynochaway Creek. 3 mi. N Morgan. 
Calhoun Co. 



6 BREVIORA No. 270 

Anodontoides radiatus (Conrad) 
Plate 2, figures 1-4 

Alusinidonta ladiata Conrad 1834, Amer. Jour. Sci., 25: 341, pi. 1, fig. 10 
(small streams in South Alabama; measured holotype ANSP 41147 
labeled, Greene Co. ) 

Margahtana elliottii Lea 1858, Proc. Acad. Nat. Sci. Phila., 10: 138 (Chat- 
tahoochee River (below Uchee Bar] near Columbus [Muscogee Co.] 
Georgia). Lea, 1859, Jour. Acad. Nat. Sci. Phila.. (2)4: 226, pi. 31, 
fig. 108. Hgured holotype USNM 86257. Lea, 1859. Obs. Unio, 
7: 44. 

Margaiitana clUpticci Lea 1859, Proc. Acad. Nat. Sci. Phila., 11: 113 
(Tombigbee River, Columbus [Lowndes Co.] Mississippi). Lea, 1862. 
Jour. Acad. Nat. Sci. Phila., (2)5: 106, pi. 18, fig. 254; figured holo- 
type USNM 86258. Lea. 1862, Obs. Unio, 8: 110.^ 

Anodonta showcdterii Lea 1860, Proc. Acad. Nat. Sci. Phila.. 12: 
307 (Coosa River Wetumpka [Elmore Co.] Alabama). Lea, 1862, 
Jour. Acad. Nat. Sci. Phila., (2)5: 215, pi. 33, fig. 284: figured holo- 
type USNM 86487. Lea, 1863, Obs. Unio, 9: 37.^ 

Strophitiis radiatus (Conrad). Frierson, 1927, Check List North American 
Naiades, p. 23. 

Anodontoides elliotti (Lea), pariini. Clench and Turner, 1956, Bull. Flor- 
ida State Mus., 1: 182, pi. 5. fig. 3. 

Description. Shell generally small to medium in size, reaching 
72 mm in length. Outline elliptical. Valves inequilateral, inflated, 
thin. Anterior end regularly rounded, posterior end rather 
pointed. Ventral margin slightly curved. Dorsal margin slightly 
curved scarcely forming an angle with the obliquely descending 
posterior margin. Hinge ligament covering most of the dorsal mar- 
gin. Posterior ridge generally broad and rounded, occasionally 
subangulate. Posterior slope flat. Umbos moderately swollen, 
slightly raised above the hinge line, located toward the anterior 
third of the shell, their sculpture consisting of a few moderate 
ridges that curve up sharply behind. Surface of the shell smooth, 
brownish or brownish green, generally with bright or dark green 
rays of different widths over the entire surface of the shell. 

Left valve with a rudimentary, laminate, somewhat bifurcated, 
pseudocardinal tooth. Hinge plate very narrow. No lateral teeth. 
Right valve with one long, narrow, low, pseudocardinal tooth, 
anterior to the umbo. 

Umbonal cavities rather shallow with dorsal muscle scars. An- 
terior adductor muscle scars clearly outlined, posterior ones ob- 
scure. Pallial line faint. Nacre bluish white, often with yellow 
spots toward the umbonal cavities, iridescent posteriorly. 



!967 GULF DRAINAGE UNIONIDS 



Length 


Height 


Widtli 


mm 


mm 


mm 


50 


29 


19 


64 


36 


22.5 


71 


35.5 


25.5 



Holotype of M. clliptica Lea 
Holotype of A. radkita Conrad 
Mosquito Creek, Chattahoochee, Gadsden 
Co., Florida. 

72 43 31 Holotype of /I. i/n^nY/Z/i^r/V Lea 

Remarks. Anodontoides radiatus (Conrad) of the Gulf drain- 
age, shows the same consistency of shape as does Anodontoides 
jerussacianus (Lea) of the interior basin. It is readily distinguished 
from jerussacianus which is toothless, by the presence of a vestigial 
tooth in each valve. A. radiatus is, in general, more consistently 
elliptical, and has brighter, more clearly defined rays. In the Gulf 
drainage. A. radiatus has been confused with Strophitus siibvexus 
(Conrad). Clench and Turner's (1956:183) records of elliotti 
\= radiatus] from the Chipola are all suhvexus. 

A. radiatus differs from S. suhvexus by being a smaller species 
with a consistently elliptical shell, having bright green rays over 
the entire surface of the shell, whereas siibvexus is subelliptical. 
inclined to be biangulate behind, seldom with rays over the entire 
surface. The hinge teeth of radiatus are laminate and delicate, 
the left valve with a single bifurcated tooth, the highest portion 
often being posterior to the umbo, the right valve with one long 
narrow tooth, parallel to the anterior dorsal margin. The hinge 
teeth of siibvexus are less delicate, the left valve with a stumpy, 
vestigial, pseudocardinal directly below the umbo, sometimes with 
the trace of a first and third tooth before and behind it, the right 
valve with a triangular, or knob-like tooth which is not parallel to 
the anterior dorsal margin. 

Frierson ( 1927:23) gives the correct synonymy of this species. 
Conrad's holotype of Alasniidonta radiata is labeled as coming 
from Greene Co.. Alabama. In all probability the type is from a 
tributary of the Black Warrior River near Erie | a town, no longer 
extant, about 10 mi. W Greensboro, Hale Co.. Alabama] since 
Conrad is known to have been there around the first of June 1833 
(Wheeler. 1935:38). 

Range. Gulf drainage, from the Alabama-Coosa River system 
of Alabama to the Apalachicola River system of Georgia and 
Florida. Absent from the intervening Choctawhatchee River sys- 
tem and the Chipola River of the Apalachicola River system. 



8 BREVIORA No. 270 

SPECIMENS EXAMINED 
ALABAMA COOSA RIVER SYSTEM 

Tonibigbee River Drainage. — Mississippi: Tombigbee River, 
Columbus, Lowndes Co. (USNM). Akibanui: [Tributary of Black 
Warrior River] Greene Co. (ANSP). | Tombigbee River] Mar- 
geno Co. 

Coosa River Drainage. — Alabama: Coosa River, Wetumpka, 
Elmore Co. (USNM). 

Alabama River Drainage. — Alabama: Pine Barren Creek, 
Camden, Wilcox Co. 

ESCAMBIA RIVER SYSTEM 

Conecuh River Drainage. — Alabama: Oakywoods Creek, But- 
ler Co. Dry Creek, Brantly, Crenshaw Co. 

APALACHICOLA RIVER SYSTEM 

Chattahoochee River Drainage. — Georgia: Chattahoochee River, 
West Point, Troup Co. (USNM). Chattahoochee River, Uchee 
Bar, below Columbus, Muscogee Co. (USNM). Big Uchee Creek, 
Young's Bridge, 6 mi. NE Scale; Big Uchee Creek, below Fort 
Mitchell; both Russell Co. 

Apalachicola River Drainage. — Florida: Mosquito Creek, 1 mi. 
S. Chattahoochee, Gadsden Co. 

Alasmidonta wrightiana (Walker) 
Plate 2, figure 5 

Stropliitiis wriglitiaiiiis Walker 1901, Nautilus, 15: 65, pi. 3 (Tributaries of 
the Flint River, Baker Co., Georgia; holotype MZUM 74938. Type 
locality corrected to Ochlockonee River, Florida, on the authority 
of Walker, by Simpson, 1914, Cat. Naiades, 1: 357). 

AUisnudoiita triangiilala (Lea), partiin. Clench and Turner, 1956, Bull. 
Florida State Mus., 1: 180. 

Remarks. In the Gulf drainage, Alasmidonta wrightiana (Wal- 
ker) has been confused with A. triangulata (Lea) but it differs 
from that species by being elliptical, rather than triangular, by 
having a posterior ridge that is rounded rather than sharp, and 
especially by having a very characteristic sculpture on the posterior 
slope which consists of a number of ridges that extend from the 
posterior ridge to the dorsal and posterior margins. A. triangulata 
is not sculptured on the posterior slope. 

Long known only from the holotype, the Museum of Compara- 
tive Zoology is grateful to the Florida State Museum for a more 
recently collected specimen. 



1967 GULF DRAINAGE UNIONIDS 9 

Range. Gulf drainage, restricted to the Ochlockonee River, 
Florida. 

SPECIMENS EXAMINED 

OCHLOCKONEE RIVER SYSTEM 

Ochlockonee River Drainage. — Florida: Ochlockonee River, 
bridge, 8 mi. W Tallahassee, Leon Co. 

Obovarfa rotulata (B. H. Wright) 
Plate 3, figure 1 

Unio rotulatiis B. H. Wright 1899. Nautilus, 13: 22 (Escambia River, 
Escambia Co., Florida: holotype USNM 159969, figured by Simpson, 
1900. Proc. Acad. Nat. Sci. Phila., p. 78, pi. 4. fig. 2). 

OhoYurici rotiihita (Wright). Simpson, 1914, Cat. Naiades, 1: 297. 

Remarks. Obovaria rotulata (Wright) of the Escambia River, 
differs from Obovaria imicolor (Lea) of the Alabama-Coosa 
River system by being almost circular in outline, the latter being 
elliptical or ovate. Overlooked by Clench and Turner, rotulata 
is the easternmost representative of Obovaria in the Gulf drainage. 
Formerly known only from the holotype, the Museum of Zoology, 
University of Michigan, has recently acquired specimens of this 
species collected by William H. Heard. 

Range. Gulf drainage, restricted to the Escambia River, Florida. 

SPECIMENS EXAMINED 

ESCAMBIA RIVER SYSTEM 

Escambia River Drainage. — Florida: Escambia River, Escam- 
bia Co. (USNM). 

Lampsills BiNOMiNATUS Simpsou 
Plate 2. figure 6; Plate 3, figure 2 

Unio lineatiis Lea 1840, Proc. Amer. Philos. Soc, 1: 287 (Chattahoochee 

River, Columbus [Muscogee Co.] Georgia). Lea, 1842, Trans. Amer. 

Philos. Soc, 8: 206, pi. 12, fig. 20; figured holotype USNM 84884. 

Lea, 1842, Obs. Unio, 3: 44, non Unio lineata "Valenciennes' Bory de 

St. Vincent 1827. 
Lcunpsilis hinoininatus Simpson 1900, Proc. United States Natl. Mus., 22: 

528, new name for Unio lineatiis Lea 1840. 
Lanipsilis e.xcavatiis (Lea), partiiu. Clench and Turner, 1956, Bull. 

Florida State Mus., 1: 198. 



10 BREVIORA No. 270 

Description. Shell rather small in size, not reaching over 58 mm 
in length. Outline of male elliptical; of female obovate. Valves 
inequilateral, somewhat inflated, and rather thin. Anterior end 
regularly rounded, posterior end distinctly pointed in the male, 
somewhat truncated in the female. Ventral margin straight or 
slightly curved in males, often extending below the base line in 
females. Dorsal margin short and straight, forming a broad angle 
with the obliquely descending posterior margin. Hinge ligament 
short but prominent. Posterior ridge faintly double, rather rounded 
and poorly defined. Posterior slope slightly concave, usually with 
a few faint ridges and wrinkles. Umbos moderately swollen, slightly 
raised above the hinge line, located toward the anterior third of 
the shell, their sculpture not observed. Surface of the shell smooth 
and shiny, waxy yellow, greenish yellow, sometimes brownish, with 
very fine, slightly interrupted, dark green rays over the entire sur- 
face of the shell. 

Left valve with two serrated pseudocardinals, one in front of 
the other, the anterior one somewhat triangular, the hinder one 
much lower. Hinge line short but with a considerable interdentum, 
before two short straight lateral teeth. Right valve with two 
pseudocardinals, separated by a deep pit, the anterior tooth ves- 
tigial, the hinder one rather chunky and triangular; one lateral 
tooth. 

Umbonal cavities rather deep with dorsal muscle scars. An- 
terior adductor muscle scars well impressed, posterior ones less so. 
Pallial line faint. Nacre bluish white, or salmon colored 

Measurements 

Length Height Width 
mm mm mm 

Holotype of U. lineatus Lea, female 
Chattahoochee River, Columbus, Musco- 
gee Co., Georgia, female 

Ibid., male 

Line Creek, Vi mi. W Digbey, Spaulding 

Co., Georgia, male 

Remarks. Lanipsilis binominatns Simpson of the upper Flint 
River drainage, has been confused with Latnpsilis excavatus (Lea) 
which ranges from the Amite River in Louisiana to the Escambia 
River of Alabama and Florida. L. binominatns appears to be a 
much smaller species. When specimens of the same size of both 



33 


23 


16 


37 


28 


19 


38 


25 


18 


58 


41 


32 



1967 GULF DRAINAGE UNIONIDS 11 

species are compared, the males of hinominatus are more ellipti- 
cal, and specimens of both sexes have narrower, sharper, darker 
green rays which are consistently linear and do not broaden and 
become diffuse as do the rays of L. e.xcovatus. 

Clench and Turner did not find L. excavatus east of the Escambia 
River, and since they did not recognize L. bmominatus, they as- 
sumed its type-locality to have been in error. Subsequent collect- 
ing has substantiated the original locality. 

Range. Gulf drainage, limited to the Upper Apalachicola River 
system. 

SPECIMENS EXAMINED 

APALACHICOLA RIVER SYSTEM 

Chattahoochee River Drainage. — Georgia: Chattahoochee 
River, Westpoint, Troup Co. (H. D. Athearn, formerly Boston 
Soc. Nat. Hist. 3526). Chattahoochee River, Columbus, Muscogee 
Co. 

Flint River Drainage. — Georgia: Line Creek, Vi mi. W Dig- 
bey, Spaulding Co. (H. D. Athearn). 

Lampsilis jonesi Vander Schalie 
Plate 3, figure 5 

Lampsilis jonesi Vander Schalie 1934. Nautilus, 47: 125, pi. 15, figs, la, 
lb, 2, 3a, 3b (Pea River, Priston's Mill, Dale Co., Alabama; holotype, 
Alabama Mus. Nat. Hist.) 

Lampsilis aiistralis Simpson, partini. Clench and Turner, 1956, Bull. Florida 
State Mus., 1: 182. 

Lampsilis jcmesi Vander Schalie. Athearn, 1964. Nautilus, 77: 138. 

De.scription. Shell generally small to medium in size, reaching 
48 mm in length. Outline subovate or elliptical, moderately elon- 
gate. Valves inequilateral, not much inflated, and rather solid. 
Anterior end regularly rounded, posterior end generally distinctly 
biangulate. Ventral margin slightly curved. Dorsal margin slightly 
curved, forming a sharp angle with the obliquely descending, 
doubly scalloped, posterior margin. Female shells show general 
swelling of the disc below the posterior ridge, and the ventral mar- 
gin is more angular than that of the male shell. Hinge ligament 
short. Posterior ridge broad, but distinctly double. Posterior slope 
slightly concave. Umbos moderately swollen, slightly raised above 
the hinge line, located toward the anterior quarter of the shell, 
their sculpture not seen. Surface of the shell smooth, shining. 



12 BREVIORA No. 270 

olivaceous green or olivaceous yellow, usually with irregularly 
distributed green rays. 

Left valve with two solid, compressed, jagged pseudocardinals. 
a slight interdentum, and two thin slightly granular lateral teeth. 
Right valve with two pseudocardinals, the more posterior one rudi- 
mentary, the anterior one well developed, stumpy, and jagged; one 
lateral tooth. 

Umbonal cavities shallow, with traces of dorsal muscle scars. 
Anterior muscle scars well impressed, posterior ones less so. Pal- 
lial line distinct anteriorly. Nacre bluish-white, thickened anter- 
iorly; thinner and slightly iridescent posteriorly. 

Measurements 



Length 


Hci^lit 


Width 


mm 


mm 


mm 


46 


22.5 


16 


48.5 


23 


18 



Lectotype of L. jonesi Vander Schalie 
Allotype 

Remarks. Lampsilis jonesi Vander Schalie of the Choctawhat- 
chee River system has been confused with Lampsilis australis 
Simpson, a species which occurs in the Choctawhatchee and Es- 
cambia drainage systems. The posterior ridge of L. jonesi is 
double, often with a distinct tertiary ridge above. The extremities 
of the ridges form a scalloped edge posteriorly, a unique character. 
The posterior ridge of australis is poorly defined and ends in a blunt 
point. The periostracum of jonesi is somewhat coarse, while that 
of australis is often smooth and glossy. L. jonesi is somewhat cylin- 
drically shaped, australis is more elliptical and less inflated. Females 
of jonesi exhibit a swelling of the disc below the posterior ridge, 
rendering the ventral margin slightly angular, while those of 
australis only show a greater rounding of the ventral margin. 

Vander Schalie compared jonesi to Lampsilis subangulata (Lea) 
which he said occurred with it in the Choctawhatchee River system, 
but these were actually specimens of australis since subangulata 
is restricted to the Apalachicola and Ochlockonee River systems. 
The two records of Clench and Turner (1956:198) from the 
Choctawhatchee River system listed as subangulata are australis. 

Lampsilis subangulata and australis are allopatric. Lampsilis 
subangulata is generally more attentuate and pointed posteriorly, 
more inflated, with a sharper posterior ridge and a more excavated 
posterior slope. The surface is more highly polished and the 
green rays are sharper and brighter. 

Vander Schalie did not specifically designate a holotype for L. 
jonesi though he makes it clear that it is in the Alabama Museum 



1967 GULF DRAINAGE UNIONIDS 13 

of Natural History. It is assumed that he intended the male 
specimen shown on his plate 15, figures la, lb, to be the holotype. 
It is here selected as the lectotype. 

Range. Gulf drainage, restricted to the Choctawhatchee River 
system of Alabama and Florida. 

SPECIMENS EXAMINED 
CHOCTAWHATCHEE RIVER SYSTEM 

Pea River Drainage. — Alabama: Pea River, Elamville, Bar- 
bour Co. Pea River, Priston's Mill. Dale Co. Pea River, Flemings 
Mill, Coffee Co. 

Choctawhatchee River Drainage. — Alabama: East Fork Choc- 
tawhatchee River, Midland City, Dale Co. East Fork Choctawhat- 
chee River, 8 mi. W Abbeville, Henry Co. West Fork Chocta- 
whatchee River, 7 mi. SE Ozark; Choctawhatchee River. 1 mi. N 
Newton; both Dale Co. 

Lampsilis HADDLETONi Athcam 
Plate 3, figure 3 

Lampsilis haddletoni Athearn 1964, Nautilus. 77: 135. pi. 9, figs. g. h. 
(Choctawhatchee River, West Fork. 7 mi. SW Ozark, Dale Co.. Ala- 
bama; holotype. National Museum of Canada 20095 ). 

Remarks. Lampsilis haddletoni Athearn is known only from 
the two specimens collected at the type locality. It "appears some- 
what similar to L. ochracea (Say) |of the Atlantic drainage]. It 
is smaller than that species, less inflated, the shell thicker, and the 
cardinal teeth are much larger. The species differs from Villosa 
choctawensis Athearn by its orbicular outline, flesh colored nacre 
and in particular in the coloration of the epidermis [periostracum]. 
The I periostracum I of L. haddletoni is darker anteriorly on the 
disc while it is lighter near the ventral margin and on the posterior 
slope. Rays are prominent only on the posterior slope. The 
[periostracum] of adult V. choctawensis is dark throughout except 
in the vicinity of the umbones where it appears lighter and exhibits 
fine rays." (Athearn) 

Villosa choctawensis Athearn 
Plate 3, figure 4 

Villosa choctawensis Athearn 1964, Nautilus, 77: 137, pi. 9, figs, c, d. e, f. 

(Choctawhatchee River, 2 mi. SW Caryville; about 1 mi. downstream 

from U. S. Highway 90. Holmes Co., Florida; holotype National 

Museum of Canada 20096). 
Pleiirobema strodeanuin ( B. H. Wright), parfini. Clench and Turner, 

1956, Bull. Florida State Mus.. 1: 161. 



14 BREVIORA No. 270 

Remarks. "Villosa choctawensis has probably been m'staken 
for Pleurobcma .siroclccuium B. H. Wright by collectors in the past. 
The males of V . choctawensis are strikingly similar to that species. 
However, they lack the consistently well defined posterior ridge of 
P. strodeanuni. The male and female shells of P. strodeanum are 
essentially alike as is characteristic of shells of the genus Plcuro- 
bema. Sexual dimorphism is well defined in specimens of V. 
choctawensis. 

"The nacre of the posterior area of V. choctawensis lacks most 
of the bluish appearance found in that area of P. strodeanum. V . 
choctawensis is in some respects similar to V. villosa B. H. Wright 
but is much shorter than that species." (Athearn) The records 
beiow are supplemental to those given by Athearn. 

Range. Gu'f drainage, restricted to tlie Choctawhatchee River 
system. 

SPECIMENS EXAMINED 

CHOCTAWHATCHEE RIVER SYSTEM 

Pea River Drainage. — Alabama: Pea River, Vi mi. SW Geneva, 
Geneva Co. 

Choctawhatchee River Drainage. — Florida: Choctawhatchee 
River, 8 mi. W Miller's Cross Roads, Holmes Co. 

LITERATURE CITED 

Athearn. Herbert D. 

1964. Three new unionids from Alabama and Florida and a note on 
Lampsilis jonesi. Nautilus, 77: 134-139. pi. 9. 

Clench, William J. and Ruth D. Turner 

1956. Freshwater mollusks of Alabama. Georgia, and Florida from 
the Escambia to the Suwannee River. Bull. Florida State Mus., 
1: 97-239, pis. 1-9. 
Frierson, Lorraine S. 

1927. A classified and annotated check list of the North American 
Naiades. Bavlor Univ. Press, Waco, Texas, 111 pp. (errata et 
corrigenda. 1 p. ). 
Johnson, Richard I. 

1965. A hitherto overlooked AnoiloiiUi (Mollusca; Unionidae) from 
the Gulf drainage of Florida. Breviora. Mus. Comp. Zool., No. 
213: 1-7. pis. 1-2. 

Ortmann, Arnold E. 

1923. The anatomy and taxonomy of certain Unionidae and Anjdon- 
tinae from the Gulf drainage. Nautilus, 36: 130-132. 



1967 GULF DRAINAGE UNIONIDS 15 

Vander Schalie, Henry 

1940. The naiad fauna of the Chipola River, in northwestern Florida. 
Lloydia. 3: 191-208, pis. 1-3. 
Wheeler, Harry E. 

1935. Timothy Abbott Conrad with particular reference to his work 
in Alabama one hundred years ago. Bull. Amer. Paleont.. 
23(77): 1-157, 27 pis. 



16 BREVIORA No. 270 



PLATE 1 

Fig. I, Strophitiis stihve.xii.s (Conrad). Black Warrior River, Tuscaloosa 
Co., Alabama. Topotype MCZ 146655. Length 59, height 36, width 25 mm 
(nat. size). 

Fig. 2, Srrophitus siihrcxns (Conrad). |Magbys Creek of] Tombigbee 
River, near Columbus | Lowndes Co.] Mississippi. Holotype USNM 86278 
of Mari^aritaiui spillnuinii Lea. Length 93, height 51. width 40 mm 
(slightly reduced ). 

Fig. 3, Stropliiiiis suhvcxus (Conrad). Cowarts Creek, nr. Florida state 
line, Houston Co., Florida. MZUM 138467. Length 86, height 47. width 
37 mm (slightly reduced). 



1967 



GULF DRAINAGE UNIONIDS 



17 




18 BREVIORA No. 270 



PLATE 2 

Fig. 1, Anodontoides rcidiutiis (Conrad). [Tributary of Black Warrior 
River, Greene Co., Alabama]. Holotype ANSP 41147. Length 64, height 
36, width 22.5 mm (slightly reduced). 

Fig. 2, Anodontoides radiotus (Conrad). Chattahoochee River [below 
Uchee Bar] near Columbus [Muscogee Co.] Georgia. Holotype USNM 
86257 of Marf^ctrilaiKi clliottii Lea. Length 45, height 26, width 18 mm 
(slightly reduced). 

Fig. 3, Anodontoides radiatiis (Conrad). Tombigbee River, Columbus 
[Lowndes Co.] Mississippi. Holotype USNM 86258 of Margarilmui ellip- 
tica Lea. Length 50, height 29, width 19 mm (slightly reduced). 

Fig. 4, Anodontoides radiutiis (Conrad). Coosa River, Wetumpka ] El- 
more Co.] Alabama. Holotype USNM 86487 of Anodonta showalterii Lea. 
Length 72, height 43, width 31 mm (slightly reduced). 

Fig. 5, Alasmidonta wrightiaiui (Walker). Ochlockonee River, Florida. 
Holotype MZUM 74938. Length 54, height 38. width 31.5 mm (nat. size). 

Fig. 6, Lampsilis hinoni'natns Simpson. Chattahoochee River, Columbus 
(Muscogee Co.] Georgia. Holotype USNM 84884 of Unio lineatns Lea. 
Length 33. height 33, width 16 mm. female (slightly reduced). 



967 



GULF DRAINAGE UNIONIDS 



19 




20 BREVIORA No. 270 



PLATE 3 

Fig. 1, Ohovaria rotitlata (B. H. Wright). Escambia River, Escambia Co., 
Florida. Holotype USNM 159969. Length 48, height 45, width 32 mm 
(nat. size). 

Fig. 2, Lampsilis binominatus Simpson. Line Creek, Vi mi. W. Digbey, 
Spaulding Co., Georgia. Collection of H. D. Athearn. Length 58, height 
41, width 32 mm, male (slightly reduced). 

Fig. 3, Lampsilis haddletoni Athearn. Choctawhatchee River, West Fork, 
7 mi. SW Ozark, Dale Co., Alabama. Holotype National Museum of Can- 
ada 20095. Length 30, height 23, width 12.5 mm (slightly enlarged). 

Fig. 4. Villosa choctawensis Athearn. Choctawhatchee River, 2 mi. SW 
Caryville; about I mi. downstream from U. S. Highway 90, Holmes Co., 
Florida. Holotype National Museum of Canada 20096. Length 37, height 
26, width 16.5 mm, male (slightly enlarged). 

Fig. 5, Lampsilis jonesi Vander Schalie. Pea River, near Elamville, Bar- 
bour Co., Alabama. While not exactly from the type locality, this specimen 
was received from Vander Schalie labeled as a paratype, MCZ 98802. 
Length 48, height 24, width 16 mm, female (slightly enlarged). 



967 



GULF DRAINAGE UNIONIDS 



21 




BREVIORA 

MmseMinti of Comparative Zoology 

Cambridge, Mass. 17 November, 1967 Number 271 



CHELONIA DEPRESSA GARMAN RE-INVESTIGATED 

Ernest E. Williams, 
Alice G. C. Grandison,' 
and Archie F. Carr, h.^ 

The pantropical sea turtle genus Chelonia is characterized by 
the tendency to mass at restricted breeding places — islands or bits 
of shore — where copulation as well as nesting takes place. Desul- 
tory nesting by females going separately ashore occurs in inter- 
vening regions, but by far the greater part of the reproduction of 
the genus takes place in aggregations. Results of extensive recent 
tagging studies have revealed that members of a nesting as- 
semblage may converge from several different year-around feeding 
grounds, some of which may lie as far as a thousand miles away. 
This site tenacity in breeding is so strong that one wonders how 
new nesting colonies are ever established. The genus may there- 
fore be strongly fragmented genetically, and nascent species, or 
even quite genuine species, may currently be confused under the 
collective name Chelonia mydas. The surprising thing is that this 
isolation has produced so little well-marked morphologic di- 
vergence. One example of the degree of differentiation that occurs 
is shown by Carr and Hirth (1962). but material representing the 
numerous isolated breeding populations of the genus is still too 
meagre to support an extensive revision of the group. One local 
population, however, is morphologically so distinct that it may be 
tentatively regarded as a species, even though only about 40 speci- 
mens are at present known to have been preserved in museums. 
This sharply distinguished taxon is Chelonia depressa Carman, 
described in 1880. 

The type of C. depressa is a mounted adult male in the Museum 
of Comparative Zoology (MCZ 4473), with the locality "North 
Australia." It is a flat-shelled turtle with short flippers, and with 



^ British Museum (Natural History), London S.W. 7, England. 

- Department of Zoology, University of Florida, Gainesville, Florida. 



2 BREVIORA No. 271 

only three postoculars on either side. The squamation of both tore 
and hind limbs is notably weak. In superficial appearance this 
could be equally well a freakish specimen of Chclonia mydas or 
some sort of Lepidochelys. 

The type is still one of the few adults known. However, our own 
examination of the type and of hatchlings and young turtles from 
the vicinity of the type locality and data generously provided for 
specimens in Australian museums (see Table 1 ) leave little doubt 
that there exists a distinct north and northeast Australian popula- 
tion of Chelonia, clearly differentiated from other known forms of 
the genus, for which the name dcprcssa is available. 

Boulenger in 1889 placed C. dcprcssa in the synonymy of C. 
nixdas, although he had one juvenile specimen of the Carman 
species at hand, in the British Museum collections. A year later, 
believing that Boulenger was in error, Baur proposed elevating 
dcprcssa to generic rank on the basis of trivial features of the 
lower jaw seen in the type. Baur's opinion was generally ignored, 
and C. inxdas and C. dcprcssa have almost universally continued 
to be regarded as conspecific. Probably the principal reason for 
the rejection of Carman's species by the majority of herpetologists 
has been the absence from most institutions of any material of the 
form. Even the careful re-appraisal of the characters of dcprcssa 
by D. B. Fry in 1913. on the basis of seven specimens from north- 
ern Australia and adjacent New Cuinea waters, did not carry the 
day. The issue may have been somewhat clouded by McCulloch's 
(1908) attempt to erect a new genus and species {Natator tessel- 
latus) for a specimen which Fry was able to show was clearly 
identical with dcprcssa. However, it was also unfortunate that Fry, 
after providing a superlative case for the distinctness of dcprcssa 
on the basis of external characters, went on to place emphasis on 
certain skull characters (he had only one subadult skull of dc- 
prcssa) which are subject to considerable individual variation. 
Skepticism was only reinforced when Barbour (1914) revealed 
that Carman's own original series was composite and that the 
young specimen (MCZ 1413) was a typical mydas according to 
Fry's newly provided characters. 

The down-grading of Fry's study — much more than the neglect 
of Carman's inadequate description — was truly unfortunate. 
More than 50 years later we can add no new really useful charac- 
ters to those offered by Fry. Most of Fry's characters are indeed 
subject to some individual variation and others are difficult to 
state objectively, but in aggregate thev — at least those based on 
externals — build a clear diagnosis of dcprcssa and suggest that 



1967 CHELONIA DEPRESSA CARMAN 3 

it may be the most trenchantly ditTerentiated local population of 
any of the five sea turtle genera anywhere in the world. 

We have checked 1 5 characters in the material examined. We 
list these characters below, with the condition characteristic of 
depressa in parenthesis in each case. 

1. Shape of the carapace (oval in depressa). 

2. Contact of first vertebral and first marginal (rarely present 
in depressa). 

3. Postanal plastral scutes (frequent in depressa). 

4. Brachial plates (often absent in depressa). 

5. Shields between first inframarginals and humeral (one only 
in depressa). 

6. Relative length of pastral plates (femoral longest in de- 
pressa ) . 

7. Size of forelimb (short in depressa). 

8. Scales of forelimb (wrinkled skin overlying phalanges in 
depressa ) . 

9. Scales of hind limb (wrinkled skin overlying phalanges in 
depressa ) . 

10. Number of postoculars (3 in depressa). 

11. Prefrontal length relative to supraocular (equal or less in 
depressa ) . 

12. Contact of prefrontal and maxillary sheath (no or very lim- 
ited contact in depressa). 

13. Number of postparietals ( 1 or 3 in depressa. not symmet- 
rically divided). 

14. Upper eyelid scales (scales uniformly small in depressa). 

15. Number of scales posterior to postoculars (subtcmporals) 
(numerous in depressa). 

Each of these characters is of some use for the recognition of 
depressa but most are not key characters. It is clear from the 
statement of some of them that there is individual variability (e.g. 
marginal vertebral contact, postanals). Others, such as the shape 
of the carapace or the size of the forelimbs, are relatively diflficult 
to make objective since allometry will modify the condition at 
various sizes. Still others (e.g. upper eyelid scales) are obvious 
when comparative material is available but in the absence of 
readily repeatable counts are sometimes equivocal. 

Two characters in combination, however, do appear absolutely 
and unequivocally to define depressa as against other Green 
Turtles anywhere. These are the postocular scale count of 3. plus 
the areas of wrinkled skin distally on fore and hind limbs. In 



BREVIORA 



No. 271 



reality, the last character may be sufficient, since we know of no 
overlap or approach in this feature at all. 

There is, unhappily, rare overlap in the postocular count, al- 
though this also is usually diagnostic. All known depressa have 
3 postoculars. In all other populations sampled by us and in 
material available in the British Museum, the Museum of Com- 
parative Zoology, the United States National Museum, and the 
American Museum of Natural History a count of 3 postoculars on 
each side, without evidence of aberrant fusion with other scales, 
is very rare indeed, almost inconsequential (but see below). 
The extreme infrequency of 3 postoculars in non-depressa popu- 
lations is shown with great clarity in Table 2, in which counts 
from 3,000 specimens from one East Pacific and two Atlantic lo- 
calities are given; at least one other aberrant count in the table is 
a more frequent variation than the 3-3 count. Only one British 
Museum specimen, one from the Great Barrier Reef, is equivocal 
in these two crucial characters. It has postocular counts of 3-4 
and counts across the forelimb of 6-7. It seems, however, on most 
characters to be nndas, and the three other members of the small 
series collected at the same time are unquestionable mydas, though 
one other has a 3-3 postocular count. (One Museum of Compara- 
tive Zoology specimen | MCZ 947 1 | from Murray Id., Torres Strait, 
likewise has a 3-3 postocular count.) 




Figure 1. Forelimbs of Cheloiiia mydas (A) and Clwlonid (U'[)rc.yui (B) 
compared to show suggested places for counting scales across limb. (Modi- 
fied from Fry, 1913.)"^ 



1967 CHELONIA DEPRESSA GARMAN 5 

The disturbing element here is not the occasional overlap in 
characters but that the overlap occurs in the relatively small 
sample we have from the known range of depressa. The dearth 
of information on the distribution of breeding grounds of Clwlonici 
in Australia makes it difficult to judge whether, and to what de- 
gree, depressa and mydas may nest sympatrically. There is an 
important nesting colony of mydas-Wkt turtles on Capricorn Reef 
at the southern end of the Great Barrier Reef — especially on 
Heron Island and Northwest Island, at 23)°2)0'S (Moorhouse, 
1933). The islands of Torres Strait are. or used to be. heavily 
used for nesting by one or more forms of Chelonki, and Stokes 
(1846) reported abundant nesting both in the Gulf of Carpentaria 
and at Baron Island on the western Australian coast, at 20°45'S, 
11530' W. In none of these cases is it known whether the turtle 
involved is mydas or depressa} 

The presence of the areas of wrinkled skin on the flippers of 
depressa is an even better method of diagnosing depressa than the 
three postoculars. The condition can clearly be shown in a compar- 
ative figure (such as Fry, 1913. fig. 46 a and b. and our photo- 
graphs). However, a simple count will solve the problems of 
those who may feel, in the absence of actual comparative ma- 
terial, that they have an ambiguous or intermediate condition in 
front of them. A count of scales across the dorsal surface of the 
forelimb. two scales distal to the claw, will in C/wlonia nixdas 



1 Harold Cogger (Australian Museum) writes in a letter of March 9, 
1967, "I'm afraid that I can give you little information re breeding sites of 
this species or of C. ?nyclas along the north coast of Australia. C. depressa 
certainly breeds in many areas, and at the moment David Lindner, who 
collected our adult, currently has several depressa nests under observation 
at Port Essington. Despite frequent reports of mydas breeding in the area, 
we have no authentic records. As you know we have no specimens of 
mydas sympatric with our dozen or so specimens of depressa. I have writ- 
ten to David Lindner asking him to obtain all possible information on the 
occurrence of mxdas in the area." 

For the northeast coast of Australia — the other portion of the known 
range of C. depressa — J. T. Woods, Director of the Queensland Museum, 
reports sympatry between C mydas and C. depressa over a stretch between 
Townsville and Heron Island (the latter approximately opposite Keppel 
Bay) but no overlapping records further north. Data that Mr. Woods has 
generously provided indicate that the eastern Queensland specimens of C. 
mydas adjacent to C. depressa localities are quite typical. However, since 
breeding range and residence range are for these turtles usually quite dis- 
tinct, further information is much to be desired. 



6 BREVIORA No. 271 

almost always be 4 or 5, while in Chelonia clepre.s.sa the same 
count (including always the small weak scales between the larger 
ones) will be 7 to 10. 

Most of the characters of depressa involve what appears to be a 
general weakening or reduction of the scalation. Besides the 
fewer postoculars, the lateral temporals and brachials are reduced 
in number, and the scales of the eyelid and forelimb are reduced 
in size. Series of specimens of depressa of several size groups 
will be required for final evaluation of certain of the other dif- 
ferentiating characters; for example, the broader and shorter fore 
flipper, the greater width of the head and the special texture of 
the areolar areas in the laminae of the hatchling. 

A survey of the breeding and feeding range of depressa will 
reveal aspects of its ecology that will surely help clarify the status 
and relationships of the population. The short foreleg and broad 
skull, for example, if fully verified, may prove to be features of a 
carnivorous forager, like Caret ta and Lepidochelys which poke 
about rocky places preying on crabs and mollusks, and hence do 
not make regular journeys between underwater pastures of 
turtle grass or algae and sleeping places and nesting beaches as 
typical populations of Chelonia do. That depressa may be a 
carnivorous Chelonia is also suggested by Fry's citing (1913:165) 
of the opinion of Hugh Christie that its meat is disagreeable in 
taste. However, it should be noted that Christie insisted that, "C. 
depressa is purely a vegetarian as far as my observations go. ..." 
As Fry stated (1913, footnote, p. 165), the disagreeable taste may 
well have kept depressa out of turtle markets and thus out of the 
sight of zoologists. Green turtles with inferior eating qualities 
turn up in other places too, and these seem consistently to have a 
tendency to feed on invertebrates instead of plants. While the 
dietary divergence is, in other populations, not correlated with 
strong morphological differentiation, this may simply mean that 
depressa, with its putatively peculiar feeding regimen, has been 
isolated for a longer period of time than they. *Studies of the 
feeding habitats and habits of depressa as well as of its sexual cycle 
and nesting ethology are needed; this work when done will greatly 
augment our understanding of a long neglected species. 

ACKNOWLEDGMENTS 

For the opportunity to examine material or for critical infor- 
mation on specimens not borrowed, we are indebted to L. D. 



1967 CHELONIA DEPRESSA CARMAN 7 

Brongersma (Leiden Museum), R. Inger (Field Museum. Chi- 
cago), H. Cogger (Australian Museum, Sydney), J. L. Woods 
(Queensland Museum). W. D. L. Ride. G. M. Storr (Western 
Australian Museum). M. Tyler (South Australian Museum). 
Data on Atlantic and Caribbean Clielonici were obtained with the 
support of the National Science Foundation, the Office of Naval 
Research, and the Caribbean Conservation Corporation. 

TABLE 1 
The known specimens of Chelonia depressa^ 

'■'I. Type, MCZ 4473, "North Australia" — adult 

*2. BM 87-5.16.3, "N. W. Australia" — juvenile 

"3. FMNH 97086, Green Island near Cairns, Great Barrier Reef, 

Queensland, Australia — juvenile 

*4. MCZ 54713 = J 1111, Queensland Mus., no data — juvenile 

— not seen by Fry but the fourth Queensland Museum speci- 
men cited by him 

•=5. MCZ 54714 - J 3065, Queensland Mus., Cape Bowling Green, 

mid-East Queensland — juvenile 

*6. Type, Natator tessellatiis — Port Darwin, Australian Mus. — 

Northern Territory. Australia = Fry specimen No. 6 — juve- 
nile 

*7. Australian Mus. — Port Darwin, Northern Territory, Australia 

= Fry specimen No. 1 — juvenile 

"8. Australian Mus. — Torres Strait = Fry specimen No. 4 — 

juvenile 

9. .Australian Mus. — Murray Island = Fry specimen No. 7, 

Torres Strait (skull removed, figured by Fry) — half grown 
10-11-12, Queensland Mus., J 184, J 185 (2), no data — not "New 
Guinea?" as stated by Fry = Fry specimens Nos. 2, 3, 5 — 
destroyed since Frv"s time because of poor condition — juve- 
niles 

13-14. Queensland Mus., J 1109, J 1110, Keppel Bay. mid-East 

Queensland — seen by Ogilby — "cannot be located in the 
present collections" — juveniles 

15-16-17. Queensland Mus.. J 3066-68, Cape Bowling Green. mid-East 
Queensland — -juveniles 

18-19-20. Queensland Mus., J 8551 (cast only). J 8575, J 8577, Thursday 
Island, Torres Strait — adult 

'^'Specimens examined by E. E. Williams 

1 MCZ = Museum of Comparative Zoology; BM = British Museum 
(Natural History); FMNH = Field Museum of Natural History. 



8 BREVIORA No. 271 

TABLE 1 (Cont.) 

*21-30. Australian Mus., R.81 15-16, 8909-13, 9012, Sir Edward Pellew 

Group, Northern Territory — juveniles 
*31. Australian Mus.. R.I 1756, Bathurst Island, Northern Territory 

— juveniles 
-32. Australian Mus., R. 25691, Cape Don, Northern Territory — 

juvenile 
33. Western Australian Museum, R 773, Cape Don, Northern 

Territory — adult 
*34-37. Western Australian Museum. R 682-85, "N W Coast" — 

juveniles 
*38-39. Western Australian Museum, 10423-24, "from the Northwest" 

— ^ juveniles 
40. Australian Mus., R. 26347, Port Essington near Cape Don, 

Northern Territory — adult 



1967 



CHELONIA DEPRESSA GARMAN 



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10 BREVIORA No. 271 

PAPERS CITED 

Barbour, T. 

1914. On some Auslralasian reptiles. Pioc. Bidl. Soc. Washington. 
27: 201-206. 

Baur. G. 

1890. The genera of the Cheloniidae. Amer. Nat., 24: 486-487. 

BOULENGER, G. 

1889. Catalogue of the chelonians, rhynchocephalians and crocodiles 
in the British Museum (Natural History). London, x + 3 1 1 
pp. 

Caldwell. D. K. 

1962. Sea turtles in Baja Californian waters (with special reference to 
those of the Gulf of California) and the description of a new 
subspecies of northeastern Pacific Green Turtle. Los Angeles 
County Mus. Contrib. Sci.. No. 61 : 1-31. 

Carr, Archie and Harold Hirth 

1962. The ecology and migrations of sea turtles. ?. Comparative 
features of isolated Green Turtle colonies. Amer. Mus. Novit., 
No. 2091: 1-42. 

Fry. D. B. 

1913. On the status of Clwlonlci clcprcssci Garman. Rec. Austral. 
Mus.. 10: 159-185. 

Garman, S. 

1880. On certain species of Chelonioidae. Bull. Mus. Comp. Zool.. 
6: 123-126. 

McCulloch, a. R. 

1908. A new genus and species of turtle from North Australia. Rec. 
Austral. Mus.. 7: 126-128. 

MOORHOUSE, F. W. 

1933. Notes on the Green Turtle (Chclonia Diydas). Great Barrier 
Reef Committee Reports, Brisbane, vol. I, pt. 1. 23 pp. 

Stokes, J. L. 

1846. Discoveries in Australia; with an account of the coasts and 
rivers explored and surveyed during the voyage of H.M.S. 
Beagle. London, 2 vols. 

(Received 6 January 1967.) 



967 



CHELONIA DEPRESSA GARMAN 



11 




Figure 2. Distribution of Chelonia deprcssa in Australian waters. 



12 BREVIORA No. 271 



Plate 1. Clielonia dcpres.sa, the British Museum hatchhng. Top: dorsal 
view of head to show asymmetrical division of postparietal, and prefrontals 
equal in length to supraoculars. Bottom: dorsal view of body to show rela- 
tive shortness of forelimbs and their weak scalation. 



1967 



CHELONIA DEPRESSA CARMAN 



13 




14 BREVIORA No. 271 



Plate 2. Clielonia depressa, the British Museum hatchling. Top: lateral 
view of head to show three postoculars and small number of subtemporal 
scales. Bottom: ventral view of body to show single brachial scale and 
weakness of scalation on forelimbs. 



!967 



CHELONIA DEPRESSA CARMAN 



15 




BREVIORA 

Museiiijm of Comparative Zoology 

Cambridge, Mass. 17 November, 1967 Number 272 



THE ECOLOGICAL DISTRIBUTION OF THE ANOLINE 
LIZARDS AROUND KINGSTON, JAMAICA 

A. Stanley Rand ' 

In each of the Greater Antilles and the islands of the Great 
Bahama Bank it is usual to find several species of Anolis very 
common in a small area. Though taken in the same habitat or 
even from the same tree, the species differ in where they live 
within the habitat. Differences in microhabitat have been de- 
scribed for the four species on Bimini by Oliver (1948); on Cuba 
for five species in Camaguey and Oriente by Ruibal (1961), and 
for five species near Havana by Collette (1961); for seven of 
the ten species on Puerto Rico by myself (Rand, 1964); and for 
the three commonest species on Hispaniola (Rand, 1962). No 
comparable study has been published for Jamaica, though the sys- 
tematics of the anoline lizards have been described by Underwood 
and Williams (1959), and some notes on their ecology given by 
Grant (1940) and Underwood (1951). This paper attempts to 
fill this gap in our knowledge by presenting data on the dif- 
ferences in microhabitat among five Jamaican anoles. I restrict 
myself to the anoles near the city of Kingston because micro- 
habitats do differ geographically in Jamaican anoles — particularly 
in A nolis lineatopus. 

THE JAMAICAN ANOLES 

Seven species of the genus Anolis are known from Jamaica: 
Anolis valencienni, gannani, opalinus, grahami, lineatopus, recon- 
ditus, and sagrei. The last of these, sagrei, occurs widely on 
Cuba and the Bahamas and may have been introduced by man into 
Jamaica. It is restricted to the west end of the island. Anolis 
reconditus is known from only a few specimens from the Blue 

1 Smithsonian Tropical Research Institute, Box 2072 , Balboa, Canal Zone 



2 BREVIORA No. 272 

Mountains and is closely related to lineatopus. The remaining 
five species are widespread on the island. According to Etheridge 
(1960), all five are quite closely related, and four of them, gar- 
mani, opalinus, grahami, and lineatopus, are more closely related 
to one another than they are to species occurring elsewhere. 
Among these four, opalinus and grahami are particularly close 
(Underwood and Williams, 1959). 

Except for sagrei, only grahami occurs outside of Jamaica. It 
has been introduced into Bermuda, and a subspecies or related 
species occurs on Grand Cayman. 

I have followed the nomenclature of Underwood and Williams 
except in considering valencienni a member of the genus Anolis 
rather than of Xiphocercus; in this I follow Etheridge cited in 
Williams (1962). 

METHODS 

The observations, summarized, on the ecological distribution 
of the five species of Anolis occurring in the vicinity of Kingston 
were made during a ten month stay. 

Kingston, on the south coast of Jamaica, in the rain shadow 
of the Blue Mountains, has a climate which is tropical and semi- 
arid. Probably completely covered by forest or scrub originally 
(Asprey and Robbins, 1953), there are now no completely un- 
disturbed areas in the vicinity of the city. Present habitats range 
from moderately tall forest through dense, thorny second growth 
to open grassy pastures, and include a variety of edificarian situa- 
tions, including well-watered gardens and parks. 

The distributions of the Anolis are not uniform through these 
habitats. Each species occupies a different microhabitat though 
these overlap widely and in some places all five species are pres- 
ent. I have seen four diflferent species in one tree at the same 
time and individuals of two species within a few inches of one 
another. This overlap between species is striking and tends to 
obscure differences in their ecological distribution. 

The microhabitats of species around Kingston differ in at least 
two ways: in the sorts of perches which they use, and in the sorts 
of habitats where they are common. 

To show these differences, censuses were run in four diflferent 
areas selected to cover the most frequent habitats around King- 
ston. Two of these areas were much modified by man, the other 
two less so. 



1967 ECOLOGY OF JAMAICAN ANOLES 3 

The first area, "Barbican, open," was in a new residential 
suburb north of Kingston. This had recently been cleared of thorn 
scrub second growth and built up to new houses with moderate 
to extensive grounds. There were some small trees but few large 
ones, many fences, hedgerows, grassy lawns, and buildings. It was 
the most open area censused. 

The second area, "Mona, park," was part of the grounds of 
the University of the West Indies at Mona, east of Kingston. 
This was park-like with quite well-kept lawns and many larger 
trees, most of these scattered and isolated from one another but 
some growing in small groves. There were not many smaller 
trees and relatively few bushes. 

The third area, "Barbican, bush," was dense thorny scrub sec- 
ond growth joining the first area. There were scattered larger 
trees but most were only 15-30 feet tall with much undergrowth 
and many tangled vines. The ground was covered with dry leaf 
litter. 

The fourth area, "Mona, bush," was taller forest at the base of 
Long Mountain near the second area. This was also second 
growth, though older than that at Barbican, and may never have 
been clean felled. There were many tall trees, 40-60 feet, form- 
ing a nearly complete canopy, and the undergrowth, though quite 
dense, was more open than in the Barbican second growth and 
in one place had been cleared completely leaving only the larger 
trees. There were many vines and a litter of dead leaves on the 
ground. 

Each area was carefully searched repeatedly, and each lizard 
recorded where it was first seen. No area was censused more 
than once a day and a lizard was only counted once per census 
even if it changed its position radically. 

Even with binoculars, many of the lizards present in an area 
were not seen, and those seen are not a random sample of those 
present. In all species the males are larger and more con- 
spicuous than the females. Certain species are more easily seen 
than others. The large green garmani is more conspicuous than 
any other; the slow-moving grey valencienni is the least con- 
spicuous. A lizard on a bare tree trunk at eye level is not apt to 
be overlooked, but one on top of a branch high overhead fre- 
quently escapes notice. The further up a lizard is, and the further 
out among the foliage, the less likely it is to be seen. Because of 
this bias, the census results are not fully valid representations of 
the distribution of the lizards, but since this bias acts equally for 
each species, the differences between them should be real. This 



4 BREVIORA No. 272 

censusing technique and its weaknesses have been discussed more 
fully in my paper (Rand, 1964) on the Anolis of Puerto Rico. 

Of the various possible characteristics of the lizards' perches, 
two were selected for analysis: the height above the ground at 
which each lizard was seen and the diameter of its perch. 

The data on height of perch and perch diameter did not seem 
to vary significantly from habitat to habitat and these have been 
combined in Table 1. The statistical significance of the differ- 
ences in perch height is given in Table 2. The abundance of the 
various species in the difl'erent habitats is given in Table 3. 

In the following accounts of the species, the census data are 
supplemented by a summary of more casual observations on these 
species, both within and outside of the census areas. 

OBSERVATIONS 

Anolis lineatopus lineatopus 

This form is moderate in size (males reaching 70 mm snout- 
vent length) and a strongly patterned brown in color. This is 
probably the most abundant species in the Kingston area and 
certainly the most commonly seen. 

Anolis lineatopus occurs in the widest variety of habitats of all 
of the Kingston anoles. As Table 3 shows, it was common in all 
four census areas, but least common in the Mona bush. It was 
also common in old shady gardens, in fencerows through grazed 
pastures, and was the only species seen in the edge of the man- 
grove swamps on Palisadoes. It was quite commonly seen on 
porches and verandas and sometimes entered houses. 

Like the other Anolis, lineatopus spends most of its waking 
time sitting on a perch. Unlike the others, it is commonly seen 
on a perch where it could not climb higher than a couple of feet. 
Individuals were seen on trees, on fence posts, on stone walls, on 
rocks both large and small, on ruined houses and on the walls of 
occupied dwellings, as well as in hedges and bushes. Of all the 
species it was most frequently seen on the ground (6 during cen- 
sus), though individuals spent only short periods there. 

Most Anolis lineatopus recorded during censuses were close to 
the ground and on moderate to large diameter perches (Table 1). 
Seventy-six per cent were below 6 feet and about 83 per cent on 
perches larger than V2 inch. Large individuals (adult males) 
usually perched somewhat higher (only 26 7^ below 3 feet) than 
did smaller individuals (adult females and sub-adults, 60% below 
3 feet). This difference is significant at the 1 per cent level 



1967 ECOLOGY OF JAMAICAN ANOLES 5 

(Table 3). The maximum height at which a large individual was 
seen during census was 9 feet. Individuals were rarely seen 
higher than this at any time. Larger lineatopus were less fre- 
quently seen on small diameter perches (13% on perches less 
than Vi inch) than were small ones (207^ on perches less than 
V2 inch). 

Very young lineatopus seemed to avoid larger trees and were 
usually seen close to the ground in bushes, hedges, brush piles, 
or sometimes perching on large dead leaves on the ground. 

Anolis grahami grahami 

This species is of moderate size (males reaching 70 to 75 nam 
snout- vent length), predominantly light green in color but can 
change to dark brown. There is also a common finely spotted 
phase. 

Where it occurs, grahami may rival or even surpass lineatopus 
in abundance. However, as Table 3 shows, it is common only 
in the more open areas both at Mona and in Barbican, and quite 
rare in the bushy or forested areas. It occurs in grazed pastures. 
It was also common in the shady, well-watered gardens around 
Kingston as well as in the newer more open ones, and it was this 
species which most commonly entered houses, climbing on the 
walls and hiding behind pictures, though usually venturing outside 
the house during the day. 

This species is most frequent on trees and walls and, when 
seen on fence posts and stones and in brush heaps, there is usu- 
ally the opportunity to climb to a fair height. Individuals, par- 
ticularly smaller ones, were not infrequently seen in hedges. 
Individuals were seldom seen on the ground and none during 
census taking. 

Most Anolis grahami (Table 1) recorded during census were 
well above the ground and on moderate to large diameter 
perches. Sixty-six per cent were above 5 feet and 92 per cent 
on perches of a diameter greater than Vi inch. Large individuals 
(adult males) and smaller ones (adult females and sub-adults) 
show litde difi'erence (not significant at the 5% level. Table 
2) in the height at which they were seen, though there is a 
curious and unexplained small number of smaller lizards seen 
between 6 and 10 feet. There is some tendency for smaller in- 
dividuals to be seen on small diameter perches more frequently 
than are larger individuals. 

The maximum height at which large individuals were seen 
during census was 25 feet, for smaller individuals 30 feet. Such 
heights were not infrequently seen during casual observations. 



6 BREVIORA No. 272 

Very young grahami were seldom seen and usually were up 
well above the ground. 

Anolis opalinus 

This is a small species (adult males reaching between 45 and 
50 mm snout-vent length), grey or brown in color, usually reticu- 
lated, with a prominent light lateral stripe. 

Though quite common in the habitats which it occupies, it is 
much less conspicuous than either grahami or lineatopus, in part 
because of its small size and drab coloration and in part because 
it is less common around houses. 

Anolis opalinus is primarily a lizard of bushy or forested areas. 
Grant reports that it is called "coffee lizard" for this reason. As 
Table 2 shows, it was completely absent from the residential 
area in Barbican and rarely seen in the park-like area in Mona. 
These census results are substantiated by other observations, and 
this species was never seen around the houses in Mona and Bar- 
bican and only rarely on the University of the West Indies 
campus at Mona, and then only in one group of tall trees, with 
dense foliage, grown close together and close to the forested area. 
It was quite common in both bushy study areas, less in the dense 
second growth at Barbican than in the more open and taller 
forest at Mona. 

This species was never seen in open pasture situations around 
Kingston and curiously enough it was rare or absent in the old, 
well shaded gardens in the residential areas we examined. 

Around Kingston, this species was almost always seen on trees 
or bushes, though sometimes on hanging vines. It was seldom 
seen on rocks, logs, or stumps even in the areas where it was 
quite common. It was also seldom seen on the ground and never 
so during censusing. 

Most opalinus recorded during census were well above the 
ground and on moderate to large diameter perches (Table 1). 
Only 11 per cent were recorded below 3 feet and about 63 per 
cent were recorded above 5 feet. About 94 per cent were on 
perches of a diameter greater than Vi inch. Large individuals 
(adult males) and smaller ones (adult females and sub-adults) 
show somewhat similar distributions in height, though the dif- 
ference is significant at the 1 per cent level (Table 2). More 
of the smaller individuals were seen on small diameter perches 
(11% on perches less than Vi inch) and were less frequently 
seen more than 6 feet above the ground. 

No very young individuals were seen close to the ground. 



1967 ECOLOGY OF JAMAICAN ANOLES 7 

Anolis garmani 

This species is the largest of the Jamaican Anolis (males reach 
120 mm in snout- vent length). It is usually bright green in color 
but can change to dark brown. 

In the Kingston area this species is nowhere near as abundant 
as the three preceding species. An hour or two of searching was 
usually required to find even a single individual, and during most 
census periods none were seen at all. As Table 2 suggests, this 
species was most common in the bushy or forested areas, and it 
was seen during census in neither the open residential area at 
Barbican nor the park area at Mona. However, during casual 
observation, occasional individuals were seen in both these latter 
areas. Individuals of this species were never seen on isolated 
trees in open pastures, even on large trees. They were, however, 
seen occasionally in the older, shady, well-established gardens in 
Mona. 

Though occasionally seen in bushes or vine tangles, and once 
on a telephone pole, most individuals were seen on large trees. 
Though they do occur in gardens occasionally, none were seen 
to enter houses or on stone walls or ruins. None were seen on 
the ground. 

Data from individuals collected to determine temperatures have 
been added to those collected during censuses, in Table 1. The 
data are still meagre but suggest that garmani occurs primarily high 
in the trees, as 61 per cent of the records are over 6 feet and 44 
per cent over 10 feet. All individuals seen were on moderate to 
large perches. These data accord with general observations that 
this species is one which lives well above the ground. 

No very young individuals were seen. 

Anolis valencienni 

This species is second in size only to Anolis garmani (adult 
males reach about 80 mm snout- vent length). It is a mottled grey 
in color. 

Though seldom seen and certainly less common than any spe- 
cies except garmani, this species occurs in a wide variety of 
habitats. It is very cryptically colored and usually slow moving 
so that it is seldom seen and is certainly more common than the 
census figures indicate. As Table 2 shows, it was taken in 
Mona both in the park area and in the forest. None were seen 
during censusing at Barbican but during casual observations they 
were noted occasionally in both the open residential area and in 
the dense second growth. Almost everywhere we watched Anolis, 



8 BREVIORA No. 272 

this species eventually turned up — except in the mangroves and 
in the open pastures. It certainly occurred in the old established 
gardens in the residential areas. 

This species was almost always seen on trees, though one was 
seen on a fence post, and occasionally in taller bushes. This 
species is most unlike the others in its movements and posture for 
it is a slow-moving animal which spends much of its time pressed 
close to its perch. 

The few census records (to which have been added data col- 
lected during temperature studies) show that this species occurs 
over a wide range of heights and with a less clear height prefer- 
ence than the other species (Table 1). The census data suggest 
that it uses mostly moderate to large diameter perches. This is 
the only point where my casual impressions contradict the census 
results. My impression, based on casual sightings and on watch- 
ing individuals, is that this species spends much of its time out 
among the smaller branches and twigs far from the trunk, and 
that in this it differs from the other species. 

The few very young individuals seen were on trees and bushes 
several feet above the ground. 

DISCUSSION 

A comparison of the three common species of Anolis in the 
Kingston area shows that each occupies a different microhabitat, 
though overlap occurs between them. 

These differences are of two types, differences in the usual 
perch height and differences in the type of habitat occupied. 

Each species of Anolis spends most of its time on certain kinds 
of perches. The characteristics which these perches have in 
common describe the structural niche of the species. Important 
among these characteristics is perch height. The types of habitats 
frequented by a species seem related to the amount of sun avail- 
able and in turn to temperature. This aspect of the distribution 
can be called a climatic niche. These terms have been discussed 
at greater length with respect to the Anolis of Puerto Rico (Rand, 
1964). 

In perch height, (Table 1), Uneatopus differs from both grahami 
and opalimis in perching closer to the ground, while grahami and 
opalinus are almost identical in their perch height distribution. 
Table 2 shows the differences between male Uneatopus and male 
grahami, female grahami, and male opalinus significant at the 
0.1 per cent level. The differences between female Uneatopus 



1967 ECOLOGY OF JAMAICAN ANOLES 9 

and both sexes of opalinus and grahami are significant at the 0.1 
per cent level. The differences between grahami and opalinus are 
nowhere significant at the 1 per cent level, though that between 
female grahami and female opalinus is significant at the 5 per cent 
level. Though there are differences among these species in an- 
other aspect of structural niche, i.e. perch diameter, these are 
small and probably of much less importance. 

The difference in perch height is probably more important than 
these figures indicate. Much of the food which Uneatopus catch 
is on the ground, while the food of the other two species tends 
instead to be insects which are either on the lizard's perch or on 
nearby branches or leaves. 

Two of these three species show intraspecific differences in 
perch height. In both Uneatopus and opalinus small individuals 
perch closer to the ground than do large ones. The difference in 
Uneatopus is striking, in opalinus less so. Where these two species 
occur together the intraspecific differences assure that the greatest 
overlap between them involves the large individuals of the larger 
species {Uneatopus) and the small individuals of the smaller 
species (opalinus). Those individuals which are closer in size, 
large opalinus and smaller Uneatopus, overlap least. 

In grahami there is a slight tendency for the smaller individuals 
to range higher than do the large ones, so that where grahami 
and Uneatopus occur together the overlap is greater between the 
large individuals (adult males) of each species and less between 
the smaller individuals (females and sub-adults). A study of the 
social behavior of Uneatopus (Rand, 1967) has shown that adult 
males have much larger territories (defended home ranges) than 
do females and sub-adults. A. grahami behave similarly. It was 
also found that female Uneatopus and grahami defend their terri- 
tories against other lizards of their own size regardless of species 
while the males seem to be more tolerant of other species. 
Where Uneatopus and grahami occur together, the overlap is 
greatest between the individuals with the largest home ranges and 
least between those with the smaller home ranges. The spatial 
separation between the smaller grahami and Uneatopus may be re- 
inforced by their interspecific territorial defense. 

The other differences between these three species are in the 
habitats which they occupy. Anolis Uneatopus occurs throughout 
the habitats studied around Kingston and so overlaps com- 
pletely the other two species. However, opalinus and grahami oc- 
cupy quite different habitats and show relatively little overlap. 
In general, opalinus occurs in bushy or forested areas while 



10 BREVIORA No. 272 

grahami occurs in more open areas and in gardens. Differences 
of this sort between other species of Anolis on other islands 
have been correlated with differences in amount of shade and 
with the preferred temperatures of the lizards, by Ruibal (1961) 
for several Cuban species, and by Rand (1964) for the Puerto 
Rican species. 

Certainly the forested areas where opalinus is most common are 
more shaded than the open areas where grahami occurs. Some 
very scant data on temperature indicate that these areas are also 
cooler. Rand (1964) found that in Puerto Rico the species which 
lived in the shade at low elevations ranged much higher into the 
mountains than did the species which lived in the sun at low 
elevations. In Jamaica, opalinus reaches much higher elevations 
than does grahami (Underwood and Williams, 1959). 

Data on the body temperatures of these three species were col- 
lected at Kingston. Lizards were noosed and their cloacal tem- 
peratures taken with a Schulthies Museum Special Thermometer, 
within a few seconds after capture. The temperature of the air was 
then taken in the immediate vicinity. Temperatures were taken 
only during sunny periods when the lizards had the opportunity to 
thermoregulate. These data are shown in Figure 1. This figure 
shows that grahami has an appreciably higher body temperature 
than does opalinus. This suggests that the difference between 
them in habitat distribution may be associated with their dif- 
ferent temperature preferences, opalinus with a lower body tem- 
perature preference living in more shaded forest areas and 
grahami with a higher preferred body temperature living in more 
open, sunnier areas. The occurrence of grahami and not opalinus 
in some of the very shady gardens in residential areas does not fit 
this picture but may be due to the small extent of each garden. 

The body temperature of lineatopus, whose habitat range over- 
laps those of both the other tv.'o species, might be expected to be 
intermediate. It is clearly lower than that of grahami. Associated 
with this is the fact that it becomes active earlier in the morning 
than does grahami and even more noticeably moves out of the 
sun earlier in the day. The opalinus temperatures appear lower 
than those of lineatopus, though too few to be really conclusive. 

To summarize, the three common species of Anolis in the 
Kingston area differ from one another in their ecological distri- 
bution. One species, Anolis lineatopus, differs from the other two 
in one aspect of its structural niche, living much closer to the 
ground than do the other two. These two have very similar struc- 
tural niches but one of them, opalinus, lives in the forested areas 



1967 ECOLOGY OF JAMAICAN ANGLES 11 

and the other, grahami, in the more open areas. A. opalinus has 
a lower preferred body temperature than does grahami, which 
suggests that the difference between them is one of a climatic 
niche. 

The relationships of the two rarer species of Anolis, garmani 
and valencienni, between themselves and to the three common 
species, are less well documented than are the relationships among 
the three common species. 

The largest species, garmani, lives high in trees and thus sel- 
dom comes in contact with lineatopus. The difference in perch 
height between male lineatopus and garmani is significant at the 1 
per cent level; between smaller lineatopus and garmani at the 0.1 
per cent level (Table 2). Like opalinus, garmani is most com- 
mon in the forested or bushy areas and avoids the more open 
areas, though it occurs in shaded gardens with grahami. There 
is a suggestion that garmani lives even higher in the trees than do 
opalinus and grahami, and also that it is much more closely 
restricted to large trees than they are. It is probable that in 
Anolis the size of the prey caught is strongly correlated with 
the size of the lizard (Schoener, 1967). It, therefore, may be 
important that the two species which overlap most widely (garmani 
and opalinus) are the most different in size, garmani being the 
largest of the Jamaican species and opalinus the smallest. 

The census data on the final species, A. valencienni, suggest 
that this species overlaps both the climatic and structural niches 
of all of the other species, though overlapping lineatopus least. 
However, the census data are few and, as mentioned before, I 
have a strong impression that this species spends much more time 
out among the smaller branches than do any of the others. It is 
rather different from the other species in its slow motion and in 
its appearance, and Underwood (1951) reports: "I have kept 
alive a number of these lizards, and it seems to me that they prefer 
somewhat larger insects than do other anoles of comparable size." 

The situation described for Kingston seems to be repeated all 
along the dry south coast of Jamaica from about the Morant 
River in the east nearly to Mandeville in the west and a short way 
up into the foothills of the mountains. A similar situation may 
exist on the north coast from about Port Antonio west to near 
Lucea. However, over the rest of the island the situation is com- 
plicated by the replacement of ^. /. lineatopus by other forms of 
A. lineatopus with rather different habitat preferences. Unfortu- 
nately, there is not enough information to describe these situations 
adequately. 



12 BREVIORA No. 272 

The Jamaican anoles can be compared with those of Puerto 
Rico (Rand, 1964), an island of comparable size and with a simi- 
lar number of species, but ones that are only very distantly related 
to those on Jamaica (Etheridge, 1960), the Jamaican animals 
belonging in the beta section of the genus while the Puerto Rican 
species are members of the alpha section. 

With respect to the structural niche, two Puerto Rican species 
(A. cristatellus and gimdlachi) are very similar to A. lineatopus; 
two species {A. evermcmni and stratulus) are very similar to A. 
opalinus and grahanii; and one species {A. cuvieri) is apparently 
like A. gannani. There is on Puerto Rico no species comparable 
to A. valeucienni in structural niche, while on Jamaica there is no 
species comparable to the Puerto Rican grass and twig living spe- 
cies A . krugi, pulcheUiis and poncensis. 

As in Jamaica, the most closely related Puerto Rican species 
occupy similar structural niches but differ in microclimatic niches. 
In Puerto Rico, two pairs of species and a triplet do this, while in 
Jamaica only one (possibly two) such pair exists. The eccritic 
temperatures of the Puerto Rican species are similar to those of 
their Jamaican counterparts but not identical. 

Thus the anoline faunas of these two islands, though only dis- 
tantly related, show similarities in their microhabitats. Certain of 
the structural and climatic niches occupied on both Jamaica and 
Puerto Rico are very similar. Even more striking is the similarity 
in the kinds of differences in microhabitats between anoles. On 
each island very closely related species are paratopic or nearly so 
and differ in climatic niche and eccritic temperature, while syntopic 
species are less closely related and usually occupy different 
structural niches. There has been considerable parallelism in at 
least certain of the ecological aspects of the independent adaptive 
radiations of the Anolis on Puerto Rico and on Jamaica. 

ACKNOWLEDGMENTS 

I am indebted to Professor David Steven and the staff of the 
Zoology Department of the University of the West Indies for 
allowing me to use their facilities and for helping me in many 
other ways during my stay in Jamaica. I must also thank Dr. 
E. E. Williams for his advice and criticism throughout this proj- 
ect, and my wife Patricia for her help both in the field and in 
preparing the manuscript. This study was supported in part by 
National Science Foundation Grant No. G- 16066. 



1967 ECOLOGY OF JAMAICAN ANOLES 13 

SUMMARY 

During a ten month field study in the vicinity of Kingston, 
Jamaica, four different habitats were censused for comparative 
data on perches used (structural niches) and habitats occupied 
(climatic niches) by five species of Anolis. 

In the Kingston area there are three very common species: 
lineatopus, opalinus, and grahami. 

Anolis lineatopus lives near the ground on moderate to large 
diameter perches, female and young perching lower and on smaller 
perches than adult males. It ranges from the most densely forested 
through the most open habitats. The eccritic temperature range 
seems intermediate between those of the following species. 

The closely related species A. opalinus and grahami both use 
moderate to large perches and perch higher than does lineatopus. 
A. opalinus lives in forest and grahami in the more open habitats. 
They replace one another with relatively little overlap. A. 
grahami has a higher eccritic temperature range than does opalinus. 

Of the two less common species, A. garmani lives high in trees 
in the forest. This is the largest of the Jamaican species and it 
may be significant that the common species that it overlaps most 
widely is the smallest, opalinus. 

Anolis valencienni ranges through all habitats and occurs on a 
variety of perches, probably most frequently on the branches of the 
crown. This species differs from the others in shape and in its 
slow movements, and there is some indication that it also takes 
larger prey. 

Though the Jamaican anoles are only distantly related to those 
of Puerto Rico and represent the results of independent adaptive 
radiations, there has been considerable parallelism in the parti- 
tioning of the habitat with respect to both structural and climatic 
niches on the two islands. 



14 



BREVIORA 



No. 272 



TABLE 1 

Structural niche distribution in Kingston anoles based on data 
collected during the censuses described on page 2. The number 
of individuals seen is expressed as a percentage of the total num- 
ber of individuals of that species (and size group in the first 
three species). 



Anolis lineatopus 



Perch Large individuals 
diameter > 3" 1/2 -3" <i/2" 

'|i>10ft 5.27o 
6-10 ft 7.8 
20.8 
11.7 



% 3- 5 ft 
fe < 3ft 



1.3 
15.6 
11.7 
13.0 



1.3 
6.5 
3.9 



N=77 
leaf Total 

7.8 

29.9 

36.4 

1.3 26.0 



Smaller individuals 

>3" 1/2-3" <i/2" 



7.7% 
12.8 
24.4 



1.3 
16.7 
16.7 



1.3 



19.2 



Anolis grahami 



N=78 
leaf Total 

10.3 
29.5 
60.3 



Large individuals N=62 



Smaller individuals N=40 



>3" 1/2 
>10ft 19.4% 
6-10 ft 25.8 
3- 5 ft 14.5 
< 3 ft 6.5 



■3" <'/2" 

9.7 1.6 
9.7 1.6 
8.1 
3.2 



leaf Total 

40.7 

37.1 

22.6 

9.7 



>3" 

27.5% 

2.5 
20.0 

5.0 



1/2-3" 

10.0 

12.5 
5.0 

2.5 



< 1/2 "leaf 



10.0 

2.5 



2.5 



Total 

37.5 
25.0 
27.5 
10.0 



Anolis opalinus 



Large individuals N=46 
>3" 1/2-3" <i/2" leaf Total 
>10ft 13.07o 23.9 36.9 

6-10 ft 28.3 10.9 39.2 

3- 5 ft 6.5 6.5 13.0 

< 3 ft 10.9 10.9 



Smaller individuals N=64 

>3" 1/2-3" <i/2"leaf Total 

4.7% 9.4 14.1 

15.6 15.6 7.8 39.0 

21.9 12.5 1.6 36.0 

6.2 3.1 1.6 10.9 



A nolis garmani 

All individuals N=18 

>3" 1/2-3" <i/2" leaf Total 

>10ft 33.3% 11.1 44.4 

6-10 ft 11.1 5.6 16.7 

3- 5 ft 11.1 11.1 22.2 



< 3 ft 11.1 



5.6 



Anolis valencienni 
All individuals N=23 
>3" 1/2-3" <i/2"leaf Total 
17.4% 4.3 4.3 26.0 

17.4 8.7 26.1 

17.4 13.0 30.4 



16.7 13.0 



4.3 



17.3 



1967 ECOLOGY OF JAMAICAN ANOLES 15 

TABLE 2 

Statistical significance of differences in perch height in Kingston 

Anolis. Chi squares calculated by P. E. Vanzolini. 

Significance level: * =5%, ** = 1%, 

*** = 0.1%. 

valen- 
lineatopus grahami opalinus garmani cienni 

large smaller large smaller large smaller 

large 24.630 17.556 17.059 22.743 6.289 15.618 5.750 

lineatopus 

smaller 59.108 48.510 60.512 16.462 40.800 30.027 

large 1.683 1.674 5.884 3.184 1.981 

grahami 

smaller 3.595 7.862 1.125 1.247 

large 11.252 3.219 4.257 

opalinus 

smaller 9.496 2.923 

garmani 1.661 



16 BREVIORA No. 272 

TABLE 3 

Habitat distribution: the number of Kingston Anolis seen in four 
different habitats during a series of censuses. 

lineatopus grahami opaliniis garmani valencienni 



Barbican, open 
(Residential area) 

Mona, park 

fUWI Campus) 

Barbican, bush 

(Dense second growth) 

Mona, bush 

(Tall second growth) 



44 20 



65 69 



52 3 25 



19 5 77 



1967 ECOLOGY OF JAMAICAN ANOLES 17 

LITERATURE CITED 

AsPREY, G. F. and R. C. Robbins 

1953. The vegetation of Jamaica. Ecol. Monogr., 23: 359-412. 

COLLETTE, B. B. 

1961. Correlations between ecology and morphology in anoline liz- 
ards from Havana, Cuba and southern Florida. Bull. Mus. 
Comp. Zool., 125: 137-162. 

Etheridge, R. 

1960. The relationships of the anoles (ReptiUa: Sauria: Iguanidae). 
An interpretation based on skeletal morphology. University 
Microfilms, Inc., Ann Arbor, Michigan. 235 pp. 

Grant, C. 

1940. The herpetology of Jamaica. II. The reptiles. Bull. Inst. Ja- 
maica Sci. Ser., 1: 61-148. 
Oliver, J. A. 

1948. The anoline lizards of Bimini, Bahamas. Am. Mus. Novit., 
No. 1383: 1-36. 
Panton, E. S. 

1928. Eight months with the green lizard or: the history of Anolis 
garmani. Jamaica Times. 13 pp. 
Rand, A. S. 

1962. Notes on Hispaniolan herpetology. 5. The natural history of 
three sympatric species oi Anolis. Breviora. No. 154: 1-15. 

1964. Ecological distribution in anoline lizards of Puerto Rico. 

Ecology, 45: 745-752. 
1967. Ecology and social organization in the isuanid lizard Anolis 

lineatopiis. Proc. U. S. Nat. Mus., 122: 1-79. 
Ruibal, R. 

1961. Thermal relations of five species of tropical lizards. Evolution, 
15: 98-111. 

Schoener, T. W. 

1967. The ecological significance of sexual dimorphism in size in the 
lizard Anolis conspersus. Science, 155: 474-477. 
Underwood, G. 

1951. Introduction to the study of Jamaican reptiles. Part VIII. 
Nat. Hist. Notes, Nat. Hist. Soc. Jamaica, 51: 57-65. 
Underwood, G. and E. E. Williams 

1959. The anoline lizards of Jamaica. Bull. Inst. Jamaica Sci. Sen, 
9: 1-48. 

Williams, E. E. 

1962. Notes on the herpetology of Hispaniola. 7. New material of 
two poorly known anoles: Anolis monticola Shreve and Anolis 
cliristophei 'VJWWams. Breviora, No. 164: 1-11. 

(Received 8 March, 1967.) 



18 



BREVIORA 



No. 272 



35" 


C 




/ 


33 


- 




":./ 


• 






• •• X 


<. 








3 








|3; 






• '•'• • •"• • / " • 


• 

ex 


• 


•*. 


••• :•: x</ 


1^ 


•• 


• 


. :; ::: v^ 


K. 


'~ 




V^ 


127 


- ', 


« 


../ ' 


N 


— 




/ 


•^ 


• 


• 


/ • 


25 


• 


/ 


• 




" / 




Mnolii lintatopus 


23 


/\ 





1 1 1 1 1 1 1 1 1 1 



23 25 27 29 31 33 35 'C 
Air Temp»rature 



3S»C 




/ 


33 


- 


• • • 


. / 


ft 


— 




/ 


«. 


• 


• X * * 


• • • / 


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• 




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1- 


- 


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2S 














Anolis grahami 


23 


/i 1 


1 1 1 1 


1 1 i 1 1 I 



23 25 27 29 31 33 
Air T»mp»rature 



35 "C 



35»C 
33 



3 

fe 31 

ft 

I ^ 
%27 



25 
23 



J I I 1 I L 



Anolis opalinus 

J I I \ \ I 



23 25 27 29 31 33 35'C 
Air Temperature 



Figure I. Anole body temperature plotted against ambient air tem- 
peratures. 



BREVIORA 

Miaseiiiiii of Coinniparative Zoology 



Cambridge, Mass. 17 November, 1967 Number 273 



STUDIES ON NEOTROPICAL POMPILIDAE 

(HYMENOPTERA) 
ADDITIONAL NOTES ON EPiPOMPILUS KOHL 

Howard E. Evans 



Epipompilus is one of the more exciting genera of Pompilidae, 
not only because these are among the most brilliantly colored of 
spider wasps, but also because of the many unusual and apparently 
primitive structural features of the genus. A few years ago the 
group was known from about six specimens, all females. In 1961 
(Psyche. 68: 25-37), I described the male and reviewed the 
American species, basing my review on 18 specimens representing 
nine species. I have now accumulated more than 100 specimens 
and feel in a position to make some important additions to knowl- 
edge of the genus. Three species are described as new, all from 
South America, and the males of three species are described for 
the first time (nigribasis, aztecus, and pulcherrimus) . The last- 
named species is the only one known from the United States, and 
I am indebted to R. W. Dawson of Washington State University 
for sending me an excellent series collected at the Archbold Bio- 
logical Station in Florida. For much new South American material 
I am indebted to Fritz Plaumann, of Nova Teutonia, Brazil, and 
to Charles C. Porter of Harvard University. 

Thirteen American species of Epipompilus are now known. 
Seven of these are known from both sexes, five from females only, 
and one from males only. All species are included in the following 
key, but only those species are discussed in the text which are 
described as new or for which new information is available. I 
have placed the species in two groups on the basis of an apparent 
concordance of the shape of the male subgenital plate with the 
nature of the convergence of the eyes of the female. I am hopeful 
that this may assist in making future sex associations. 



BREVIORA No. 273 

KEY TO SPECIES 
Females 

Eyes converging at the top, the upper interocular line distinctly less 
than the lower interocular line; front fairly wide, middle interocular 
line at least about 0.57 x width of head (aztecus group) 2 

Eyes parallel or diverging above, the upper interocular line equal to or 
exceeding the lower; front barely wider than the two eyes taken 
together, middle interocular line at most about 0.55 x width of head 
{delicatus group ) 6 

Wings wholly fuliginous; abdomen wholly rufous; middle and hind 

tibiae with numerous short spines above 

pulcherrimus (Evans) 

Wings hyaline, bifasciate; abdomen black with whitish spots; middle 
and hind tibiae not or exceedingly weakly spinose 3 

Abdomen with whitish spots only on tergites two and five, none on 
sternites aztecus (Cresson) 

Abdomen with whitish spots on tergites two through five or six, also 
on some of the sternites 4 

Greater part of thorax and propodeum rufous; apical tergite black 
bijasciatus (Ashmead) 

Greater part of thorax and propodeum black; apical tergite with whit- 
ish markings 5 

Pronotum entirely rufous; hind femora black; propodeum, in profile, 
nearly flat in front, obliquely declivous behind inca n. sp. 

Pronotum mostly black, with a whitish band along the posterior mar- 
gin and with whitish and rufous markings anteriorly; hind femora 

ferruginous; propodeum, in profile, forming a smooth arc 

jocosus n. sp. 

Body and legs wholly black except for a pair of small whitish spots on 

second tergite; pronotum short, subangulate behind 

williamsi (Banks) 

Body and legs variously marked with rufous and whitish; pronotum 
arcuate or very weakly subangulate behind 7 

Thorax wholly rufous (or with limited black markings on sides of 
pronotum or on venter) (Central and South American species) 
8 

Thorax with extensive black markings (South American species) .... 
10 

Front angle of ocellar triangle less than a right angle, postocellar and 
ocello-ocular lines subequal; second abdominal tergite yellowish 
brown insolitus Evans 

Front angle of ocellar triangle greater than a right angle, postocellar 
line much exceeding ocello-ocular line; second tergite with a 
whitish band or spots 9 



1967 NOTES ON EPIPOMPILUS 3 

9. Postnotum a very narrow band; face and clypeus testaceous, antennae 

wholly testaceous; length of fore wing under 4 mm .nigribasis (Banks) 

Postnotum medially about half as long as metanotum; face and clypeus 

black, antennae infuscated on apical third; length of fore wing 

5-6 mm delicatus Turner 

10. Tergites one and two mostly black, the latter with white markings; 

pronotum with extensive pale markings; postocellar line only slightly 

exceeding ocello-ocular line tuciimaniis n. sp. 

Basal 1.5-2.5 tergites rufous, second sometimes with white markings; 
pronotum wholly black; postocellar line much exceeding ocello-ocular 
line 11 

1 1 . Propodeum black except for limited whitish markings; antennae black; 

radial vein angulate at second intercubital vein haiipti (Arle) 

Propodeum ferruginous; antennae ferruginous except darker apically; 
radial vein arcuate excelsus (Bradley) 

Males 

1. Subgenital plate with a broad, truncate base from which arises a 
slender, parallel-sided spine-like process (as in Figs. 1, 3); known 
species either with the thorax extensively ferruginous or the femora 

and tibiae almost wholly ferruginous {azteciis group) 2 

Subgenital plate tapering gradually to a point from a relatively narrow 
base (as in Fig. 5); known species without ferruginous markings on 
the thorax and with the hind legs, at least, mostly fuscous {delicatus 
group) 5 

2. Length of fore wing 4.5 mm or less; transverse median vein of hind 

wing erect, nearly perpendicular to anal and median veins; body 
black except clypeus and pro- and mesonota contrastingly ferruginous 

pidcherrimns (Evans) 

Length of fore wing 5.5 mm or more; transverse median vein of hind 
wing oblique; body color not as above 3 

3. Thorax black, with limited whitish markings on pronotum and propo- 

deum; hind femora ferruginous; aedoeagus very long and slender 

(Fig. 2) jocosus n. sp. 

Thorax in considerable part rufous; hind femora black; aedoeagus not 
as above 4 

4. Wings strongly twice-banded; middle legs mostly fuscous; basal third of 

antennae light ferruginous aztecus (Cresson) 

Wings weakly once-banded; middle femora and beyond light reddish 
brown; antennae brownish basally innubus Evans 

5. Eyes strongly hairy; length of fore wing not over 3 mm; aedoeagus 

exceedingly short (Fig. 6) nigribasis (Banks) 

Eyes with very short, barely noticeable hairs; length of fore wing over 
3 mm; aedoeagus longer than above 6 



4 BREVIORA No. 273 

6. Front coxae with at least the basal 0.4 black; mesopleura rather dull, 

the punctures coarse, separated for the most part by less than their 
own diameters; parameres slender, somewhat tapering (Brazil) .... 

excelsiis ( Bradley) 

Front coxae with at most the basal 0.3 black; mesopleura shining, 
finely punctate; parameres broader and more coarsely setose 7 

7. Third submarginal cell barely wider than high; parameres blunt apic- 

ally, volsellae moderately slender (Central America) delicatus Turner 

Third submarginal cell much wider than high; parameres acute apic- 

ally, volsellae unusually slender (Argentina) tucumaniis n. sp. 

AZTECUS SPECIES-GROUP 
Epipompilus pulcherrimus (Evans) 

This species has been known from two females, one from 
southern Florida and the other from Andros Island in the Bahamas. 
R. W. Dawson collected one female and five males in a Malaise 
trap at the Archbold Biological Station, Lake Placid, Florida, in 
the winter of 1965-1966 (29 Oct.-Jan. 6). The female is small 
(fore wing 4.5 mm) and differs from the two females previously 
known in having the pronotum wholly black except the collar, the 
remainder of the thorax and propodeum wholly black. 

Description of male. — Length 4.5-5.0 mm; fore wing 3.5-4.5 
mm. Head black except clypeus and mandibles in large part 
testaceous; antennae black; palpi brown; thorax black except 
entire pronotum, mesoscutum, and scutellum bright ferruginous 
(these areas with some black blotching in one specimen); legs 
black except all spurs whitish, front femora and tibiae suffused 
with testaceous, hind tibiae with a basal white spot; wings clear 
hyaline, with dark veins and stigma. Body clothed with short, 
whitish hairs, these hairs rather long on posterior slope of 
propodeum. 

Clypeus truncate, twice as wide as high. Head 1.15 X as wide 
as high; front broad for the genus, middle interocular line 0.62- 
0.65 X width of head; eyes rather weakly convergent below, 
lower interocular line 0.95 X upper interocular line; ocelli in a 
broad triangle, postocellar line about 1.5 X ocello-ocular line. 
Front strongly alutaceous, with shallow but rather large, setigerous 
punctures. Antennae compact, segments five and beyond slightly 
produced below, so that the flagellum is crenulate in profile; first 
four antennal segments in a ratio of about 16:6:11:10, segment 
three about 1.5 X as long as thick. Setae of eyes exceedingly small. 
Pronotum short, its posterior margin arcuate; mesonotum closely 



1967 



NOTES ON EPIPOMPILUS 







Fig. 1. Subgenital plate of male Epipompilus jocosiis n. sp., paratype. 
Fig. 2. Genitalia of same specimen, ventral aspect. Fig. 3. Subgenital plate 
of male E. piilcherrimus (Evans). Fig. 4. Genitalia of same specimen, 
ventral aspect. Fig. 5. Subgenital plate of male E. nigribasis (Banks). Fig. 
6. Genitalia of same specimen, ventral aspect. 



6 BREVIORA No. 273 

punctate; postnotum nearly as long as metanotum, polished, de- 
pressed and slightly contracted medially. Propodeum shining, the 
setigerous punctures small; slope of profile low and even. Middle 
and hind tibiae with numerous short spines above; claws with a 
weak, erect tooth, except front tarsal claws more strongly toothed, 
the tooth subparallel to the outer ray, the outer claws of the front 
tarsus strongly curved. Fore wing with basal vein arising well 
beyond transverse median vein; radial vein arcuate, with only a 
faint angulation at the second intercubital vein; hind wing with 
transverse median vein erect, nearly perpendicular to anal and 
median veins, anal lobe large for the genus, half the length of the 
submedian cell. Abdomen rather small, depressed; subgenital 
plate in the form of a slender, setigerous process arising from a 
broad plate (Fig. 3); genitalia as shown in Figure 4. 

Epipompilus aztecus (Cresson) 

This species has been known only from southern Mexico. I 
have recently studied a male from Guatemala which represents the 
first known male of this species, as well as two females from 
Panama. The two females were collected on Barro Colorado 
Island, Canal Zone, one in April 1926 by C. T. Greene and one 
in May 1939 by J. Zetek [both U. S. National Museum]. They 
are relatively small for this species (fore wing 7.5-8.0 mm) and 
somewhat darkly colored: the middle femora are fuscous, and the 
propodeum and metapleura are wholly fuscous in one specimen, 
fuscous except at the anterior margin in the other. The male is 
from Santa Adelaida, Guatemala, collected at 1000 meters eleva- 
tion in March and April, 1931, by J. Bequaert [coll. of Mus. 
Comp. Zool.]. All three of these specimens were borrowed many 
years ago by the late R. R. Dreisbach and have only recently 
become available for study. Dreisbach made a slide of the male 
terminalia, but it was poorly prepared, and in my opinion the 
subgenital plate on the slide (and perhaps the genitalia) do not 
properly belong with this specimen. I have therefore omitted men- 
tion of the terminalia in the short description that follows. 

Description of male. — Length 7.0 mm; fore wing 5.7 mm. 
Head black, except lower inner orbits with a whitish streak, 
antennal sockets, face below sockets, and clypeus wholly light 
ferruginous; mouthparts largely light ferruginous, basal segments 
of maxillary palpi slightly infuscated; antennae ferruginous, apical 
two-thirds somewhat infuscated on upper surface; pronotum, 
mesoscutum, scutellum, upper half and posterior margin of meso- 
pleura, and a small blotch on the metapleura, light ferruginous, 



1967 NOTES ON EPIPOMPILUS 7 

remainder of thorax and propodeum fuscous (posterior corners of 
propodeal rim white; disc of scutellum and lateral posterior mar- 
gins of scutum slightly infuscated); front legs wholly light fer- 
ruginous, middle and hind legs fuscous except spurs white, tibiae 
with a white spot, and posterior coxae with a large white apical 
spot; abdomen fuscous, shining; fore wings strongly twice-banded 
much as in female. Body clothed with short, whitish hairs, these 
hairs rather coarse and conspicuous on the front and on the 
posterior part of the propodeum. 

Clypeus large, measuring 1.85 X as wide as high. Head 1.12 X 
as wide as high, the vertex between the eyes forming a rather 
strong arc; front broad, middle interocular line 0.60 X width of 
head; inner orbits strongly convergent below, lower interocular 
0.78 X middle interocular line, 0.93 X upper; ocelli in a broad 
triangle, postocellar line twice the ocello-ocular line. Front dull, 
with subcontiguous punctures on an alutaceous background. An- 
tennae moderately long for the genus, weakly crenulate in profile; 
first four segments in a ratio of about 17:6:10:10, segment three 
1.6 X as long as thick. Eyes not hairy. Pronotum short, with an 
abrupt anterior declivity, its posterior margin arcuate; mesonotum 
dull, closely punctate; postnotum considerably shorter than metano- 
tum; propodeum with a strong median impression, in profile the 
slope very low. Hind tibiae with a few weak spines which barely 
extend above the pubescence; tarsal claws as described for pul- 
cherrimus. Fore wing with the basal vein arising very slightly 
beyond the transverse median vein; radial vein slightly angulate 
at second transverse cubital vein; maximum width of third sub- 
marginal cell 1.6 X its maximum height; hind wing with transverse 
median vein oblique; length of anal lobe about 0.4 that of sub- 
median cell. 

Epipompilus bifasciatus (Ashmead) 

This species has been known only from the type female from 
Bahia, Brazil. I have recently studied a second specimen from a 
locality not far distant: Corema, Paraiba, Brazil, June 1957 (no 
collector given) [coll. Dept. Zool., Sec. Agricultura, Sao Paulo]. 
It is slightly smaller than the type (fore wing 7.4 mm), but there 
are no important differences in color or structure. 

Epipompilus jocosus new species 

Holotype. — $ , URUGUAY: Florida, Feb. 1952 (no collector 
stated) [MCZ, No. 31321]. 



8 BREVIORA No. 273 

Description of type female. — Length 9.5 mm; fore wing 7.5 
mm. Head black, except mandibles, clypeus, and lower half of 
front light ferruginous, the ferruginous extending up the inner 
orbits to their point of greatest emargination; lower outer orbits, 
malar space, and labium also ferruginous, the palpi partially 
infuscated; pronotum black, except collar whitish and posterior 
margin with a whitish band (including the posterior lobes), the 
anterior margin, laterad of the collar, ferruginous; remainder of 
thorax and propodeum black, except for paired whitish spots 
just above the middle and hind coxae; abdomen black, with paired 
ivory-white spots on tergites two through six and sternites two 
through five, all of these spots partially or completely enclosing 
a black spot within them; antennae and front legs wholly ferru- 
ginous except somewhat dusky apically; middle and hind coxae 
black with whitish apices; middle and hind femora and middle 
tibiae ferruginous; hind tibiae fuscous, middle and hind tarsi 
dusky ferruginous; middle and hind tibiae with whitish spots near 
the base, tibial spurs whitish. Wings hyaline, the fore wing with 
two prominent brownish bands. Body uniformly clothed with 
pale pubescence. 

Clypeus shallowly emarginate, measuring about 2.5 X as wide 
as high. Head 1.23 X as wide as high; eyes covered with short 
hairs; front moderately broad, middle interocular line 0.60 X 
width of head; eyes convergent above, lower interocular line 1.1 
X upper; ocelli in a broad, flat triangle, postocellar line 2.2 X 
ocello-ocular line. First four antennal segments in a ratio of 
about 20:8:18:21, segment three 0.47 X upper interocular line. 
Front shining, closely punctate. Pronotum fairly long, broadly 
subangulate behind; mesoscutum closely punctate; postnotum 
wholly concealed dorsally; propodeum, in profile, smoothly con- 
vex. Front femora weakly incrassate; middle and hind tibiae with 
some very short spines intermingled with the pubescence; claws 
with the tooth long, erect. Fore wings with the radial vein quite 
strongly angled at the second intercubital vein, hind wing with the 
transverse median vein oblique, the submedian cell very sparsely 
and weakly setose, the anal lobe about 0.3 as long as the sub- 
median cell. 

Allotype. — $ , BRAZIL: Nova Teutonia, Santa Catarina, 23 
Feb. 1961 (F. Plaumann) [Mus. Comp. Zool.]. 

Description of allotype male. — Length 8.5 mm; fore wing 
7.5 mm. Body black, with a bluish cast, marked with ivory-white 
as follows: apical three-fourths of mandibles, apical half of cly- 
peus, spots on inner orbits opposite bases of antennae, pronotal 



1967 NOTES ON EPIPOMPILUS 9 

collar, band along posterior margin of pronotum, interrupted 
medially, sides of posterior rim of propodeum, apical margin of 
last abdominal tergite; antennae ferruginous, first two segments 
black above, flagellum moderately infuscated above, especially 
apically; coxae black, with apical white markings; trochanters 
black; femora ferruginous, tibiae also of this color except middle 
and hind tibiae infuscated at extreme base and toward apex and 
with a whitish spot on the outer side near the base; spurs whitish; 
tarsi fuscous. Wings lightly tinged with brownish, with a faint 
darker cloud in the marginal and second and third submarginal 
cells, also at the extreme wingtip. Body pubescence light brown 
to silvery, especially long and semierect on posterior slope of 
propodeum. 

Clypeus weakly emarginate, 2.1 X as wide as long. Head 1.16 
X as wide as high, the vertex arched very weakly above the eye 
tops; front narrow, middle interocular line 0.58 X width of head; 
inner orbits weakly convergent below, lower interocular line 0.95 
X upper; ocelli in a broad triangle, post-ocellar line 1.7 X ocello- 
ocular line. Eyes with only extremely short, inconspicuous setae. 
Front alutaceous and uniformly punctate. Antennae rather long, 
the flagellum crenulate in profile; first four segments in a ratio 
of about 17:7:14:13, segment three 1.8 X as long as thick. 
Pronotum of moderate length, broadly subangulate behind; meso- 
scutum closely punctate; postnotum shining, depressed medially, 
about two-thirds as long as metanotum; slope of propodeum low 
and even, the median line weakly impressed. Middle and hind 
tibiae with a large number of short spines above; tooth of claws 
strong, sloping outward somewhat, outer claws of front tarsus 
strongly curved, bifid. Features of wings as in female, except anal 
lobe of hind wing larger, about 0.4 as long as submedian cell. 
Subgenital plate and genitalia as shown in Figures 1 and 2; the 
very long setae at the apex of the parameres protrude from the 
apex of the abdomen even when the genitalia are in resting position. 

Paratypes. — BRAZIL: 3 S S, same data as allotype except 
dated 10-15 Feb. 1964, Jan. 1965 (F. Plaumann) [Mus. Comp. 
ZooL, U. S. Nat. Mus.]. 

Variation. — The paratypes resemble the allotype closely in 
color except that one of them has the whitish band on the prono- 
tum obsolescent. The fore wing varies from 6.5 to 7.7 mm in 
length; the middle interocular line varies from 0.56 to 0.59 X 
the head width, the lower interocular line from 0.85 to 0.95 X 
the upper interocular line. 



10 BREVIORA No. 273 

Epipompilus inca new species 

Holotype. — 9 , PERU: Machu Picchu, 1900 meters, 4-19 
Sept. 1964 (C. C. Porter) [MCZ, No. 31322]. 

Description of type female. — Length 13.5 mm; fore wing 11 
mm. Head black except lower inner orbits with a whitish streak, 
apical third of clypeus testaceous, mandibles testaceous except 
darker apically, palpi fuscous; pronotum wholly ferruginous; re- 
mainder of thorax and propodeum black except for paired small 
whitish spots on lower posterior angles of mesopleura and on 
upper metapleura, and much larger spots at the posterior angles 
of the propodeum; abdomen black, with large, paired whitish spots 
on tergites two through six and on sternites two through four, with 
much smaller spots on the last two sternites; antennae ferruginous, 
somewhat dusky apically; front legs ferruginous, the tarsi dusky; 
middle and hind coxae black, with white apices; middle femora 
ferruginous, the middle tibiae ferruginous except for a black 
streak on the outer surface; hind femora and tibiae black, the 
middle and hind tarsi also nearly black; hind tibiae with white 
spots near the base; tibial spurs dusky ferruginous. Wings hyaline, 
fore wing with two prominent brownish bands. Entire body 
clothed with short, pale pubescence. 

Clypeus shallowly emarginate, measuring about 2.5 X as wide 
as high. Head 1.2 X as wide as high; eyes densely short-haired; 
front moderately broad, middle interocular line 0.60 X width of 
head; eyes convergent above, lower interocular distance about 
1.1 X upper; ocelli in a broad triangle, postocellar line 1.6 X 
ocello-ocular line. First four antennal segments in a ratio of 
about 29:10:23:27, segment three 0.51 X upper interocular dis- 
tance. Front shining, very closely punctate. Pronotum rather long, 
its posterior margin subangulate; mesoscutum closely punctate; 
postnotum complete, only about a third as long as metanotum. 
Propodeum, in profile, sloping very weakly on the anterior two- 
thirds, then abruptly declivous behind; surface of propodeum 
covered with fine, closely parallel transverse striae. Front legs 
weakly incrassate; middle and hind tibiae not at all spinose; claws 
with the tooth strong, erect, the outer ray curved so as to be nearly 
parallel to it. Basal and transverse median veins of fore wing 
interstitial; radial vein strongly angulate at second intercubital 
vein; transverse median vein of hind wing strongly oblique, the 
submedian cell almost devoid of setulae; anal lobe small, about 
0.3 as long as submedian cell. 



1967 NOTES ON EPIPOMPILUS 11 

Remarks. — This large species appears closely allied to bifascia- 
tus and to jocosus, but there are prominent color differences as 
well as differences in the postnotum, shape of the propodeum, 
spines of the hind tibiae, and so forth. Although innubus belongs 
to this group and is known from the male only, from a locality 
only a few hundred miles from the type locality of inca, it seems 
much too small and differently colored to represent the male of 
this species. 

DELICATUS SPECIES-GROUP 
EPIPOMPILUS DELICATUS Turner 

The male of this species was described by Evans, 1966, Mem. 
Amer. Ent. Soc, 20: 33. The genitalia are most similar to those 
of piilcheirimus, although the subgenital plate is very different 
and much like that of excelsus Bradley. As a result of other new 
sex associations in the genus, I feel more confident that this male 
is correctly associated with the female delicatus than I did when 
describing it. 

I have seen one additional female of this species, collected at 
Turrialba, Costa Rica, by Kenneth Cooper [collection of U. S. 
National Museum]. This specimen is of the same size as the type 
and is very similar structurally, but there are some minor color 
differences: the greater part of the second tergite is rufous, upon 
which two whitish spots are imposed; tergite five also has large, 
paired whitish markings against a rufous background (these may 
be present but concealed in the type); the legs are mostly rufous 
but are irregularly tinged with brownish. 

EPIPOMPILUS NiGRiBASis (Banks) 

This species was transferred to Epipompilus by Evans, 1966, 
Mem. Amer. Ent. Soc, 20: 31. It has been known only from 
the type female, from Panama, but I have recently discovered two 
males which unquestionably belong here. One is from Colombia: 
Bonda, August, Ace. no. 1999 [Acad. Nat. Sci. Philadelphia], 
the other from Rio de Janeiro, Brazil, 7 March 1966 (H. and M. 
Townes) [coll. H. K. Townes]. The small size and close coin- 
cidence of the wing venation with that of the female lead me to 
feel confident of this association. The eyes of the female are 
unusually strongly hairy, and these are the only known males 
having strongly hairy eyes. Although the two males are from 
widely separated localities, the genitalia and all other features are 
closely similar. 



12 BREVIORA No. 273 

Description of male. — Length 2.5-3.5 mm; fore wing 2.2-3.0 
mm. Head black; thorax and abdomen dark castaneous to fuscous, 
without white or rufous markings; antennae Hght brown, the fiagel- 
lum darker on the upper surface; coxae and hind femora and 
tibiae dark brown, legs otherwise light to medium brown, the spurs 
brownish; wings subhyaline, the fore wings with a brownish cloud 
at the marginal and submarginal cells. Body extensively clothed 
with short, light brown setae; eyes densely covered with short setae. 

Clypeus truncate, slightly wider than lower front. Head about 
1.10 X as wide as high, the vertex forming an even arc above 
the eye tops; eyes strongly convergent below, lower interocular 
distance only 0.8 X upper interocular; middle interocular line 
0.57-0.61 X width of head; ocelli in an obtuse triangle, post-ocellar 
line 1.2-1.6 X ocello-ocular line. Front alutaceous and with shal- 
low punctures. Antennae rather long, the flagellum crenulate in 
profile; first four segments in a ratio of about 6:2:5:6, segment 
three 1.6 X as long as thick; flagellar pubescence coarse. Prono- 
tum very short, arcuate behind; pro- and mesonota somewhat 
shining, with shallow, irregular punctures; postnotum very short, 
concealed medially by the overhanging metanotum; slope of pro- 
podeum low and even. Hind tibiae with some unusually long 
spines for the genus; claws with the tooth short, erect, except outer 
claws of front tarsus strongly curved, bifid. Basal vein of fore 
wing arising well beyond transverse median vein; marginal cell 
large, the radial vein angulate at second intercubital vein; hind 
wing with transverse median vein weakly oblique, almost perpen- 
dicular to median vein; anal lobe very small, hardly more than a 
minute scale with a fringe of setae (as it is in the female). Sub- 
genital plate shaped much as in excelsiis, but with fewer setae 
(Fig. 5). Genitalia with the aedoeagus exceedingly small; para- 
peniai lobes abruptly expanded on apical half; parameres short, 
setose (Fig. 6). 

Epipompilus tucumanus new species 

Holotype. — 9 , ARGENTINA: Horco Molle, Tucuman, 25 
March-30 April, 1966 (C. C. Porter) [MCZ, No. 31320]. 

Description of type female. — Length 8.5 mm; fore wing 7.0 
mm. Head black, except antennal lobes with ivory-white spots, 
apical half of mandibles testaceous; pronotum black, its posterior 
margin with a whitish band which does not include the posterior 
lobes but extends broadly across the shoulders to the anterior 
margin, the bands irregularly bordered with rufous; mesoscutum 



1967 NOTES ON EPIPOMPILUS 13 

black except ferruginous on the midline posteriorly; remainder of 
thorax and propodeum entirely ferruginous; abdomen black except 
basal third of first tergite as well as basal 1.3 sternites ferru- 
ginous, and with the following ivory-white: large, paired spots on 
tergite two, large, connected spots on tergite five, and a median 
spot toward the base of tergite six; antennae fuscous except each 
of the first three segments with an ivory-white spot above; front 
coxae black, with a whitish spot apically, middle and hind coxae 
ferruginous, with black and white markings apically, middle and 
hind femora ferruginous except black apically, legs otherwise 
fuscous (including spurs) except middle and hind tibiae with 
whitish markings on outer surface. Fore wings strongly twice- 
banded, faintly luteous between bands, but apices hyaline; hind 
wings hyaline, the tips clouded. Body wholly clothed with short, 
pale pubescence. 

Clypeus arcuately emarginate, measuring 2.3 X as wide as its 
median length. Head 1.2 X as wide as high; eyes wholly covered 
with short hairs; front narrow, middle interocular line 0.56 X 
width of head; eyes convergent below, lower interocular line 0.9 X 
upper; front angle of ocellar triangle slightly exceeding a right 
angle, postocellar line 1.2 X ocello-ocular line. First four antennal 
segments in a ratio of about 19:7:18:24, the third segment 0.53 X 
upper interocular line. Front somewhat shining, the punctures 
rather shallow. Pronotum arcuate behind; mesoscutum minutely, 
shallowly punctate; postnotum a very narrow transverse band; 
slope of propodeum low and even, the median line not impressed. 
Front femora not incrassate; hind tibiae with numerous short, 
dark spines; claws with the tooth strong, erect. Fore wing with 
the basal and transverse median veins interstitial; radial vein not 
at all angled at second intercubital vein; hind wing with the trans- 
verse median vein oblique, the anal lobe very small, about 0.3 the 
length of the submedian cell. 

Allotype. — 6 , ARGENTINA: Horco Molle, Tucuman, 15-21 
May 1966 (L. Stange) [Coll. H. K. Townes]. 

Description of allotype male. — Length 5.0 mm; fore wing 4.7 
nmi. Head black, except marked with whitish around antennal 
insertions, on extreme lower inner orbits, and on basal half of 
clypeus; thorax and propodeum black, with some bluish reflec- 
tions, except posterior margin of pronotum white (not reaching 
lateral lobes); abdomen black, with bluish reflections; antennae 
dark brown, blackish on upper surface; palpi light brown to testa- 
ceous; front coxae black on basal third, middle section light fer- 
ruginous, apex white, middle coxae black at base but mostly light 



14 BREVIORA No. 273 

ferruginous, hind coxae black with white tips; all trochanters 
fuscous; front legs beyond trochanters light ferruginous, also mid- 
dle femora; middle legs beyond femora lightly infuscated, also 
middle spurs; hind femora, tibiae, tarsi, and spurs black, except 
tibiae with a basal white spot; wings hyaline, fore wings weakly 
infuscated on outer third. Body pubescence pale, silvery. 

Clypeus arcuate emarginate, measuring twice as wide as high. 
Head 1.1 X as wide as high, the vertex elevated in an even arc 
above the eye tops; inner orbits converging below, lower interoc- 
ular line only 0.7 X upper; middle interocular line 0.60 X width 
of head; ocelli in an obtuse triangle, postocellar line only 1.1 X 
ocello-ocular line. Eyes not hairy. Front alutaceous, punctate. 
Antennae moderately elongate, weakly crenulate in profile; first 
four segments in a ratio of about 13:5:9:8, segment three nearly 
twice as long as thick. Pronotum short, its posterior margin 
broadly arcuate; mesoscutum strongly punctate; median portion 
of postnotum nearly as long as metanotum, smooth and polished; 
slope of propodeum very low. Claws dentate except outer claws 
of front tarsi strongly curved, bifid; hind tibiae with a number of 
spines extending above the coarse pubescence. Wing venation not 
differing appreciably from that of female. Subgenital plate essen- 
tially the same as in excelsus (Bradley) (see Evans, 1961, Psyche, 
p. 34, fig. 3). Genitalia differing from those of excelsus in only 
a few details: the volsellae are considerably more slender and 
elongate, and the parameres are slightly broader, tapering to a 
subacute point on the apical third, and slightly more strongly setose. 

Par city pes. — 4 9 9, same data as type [Mus. Comp. Zool., 
U. S. Nat. Mus., and Inst. Miguel Lillo, Tucuman]. 1 9 , Jujuy, 
Argentina, 15 Jan. 1966 (H. and M. Townes) [Coll. H. K. 
Townes]. 

Additional specimen (not designated a paratype). — 1 9 , San 
Esteban, near Puerto Cabello, Venezuela, 14 Jan. 1940 (P. J. 
Anduze) [Coll. H. K. Townes] . 

Variation. — The paratypes vary in length from 7.0 to 8.5 mm, 
fore wing from 6.0 to 7.0 mm. The third antennal segment varies 
from 0.45 to 0.55 X the upper interocular line, the lower interoc- 
ular fine from 0.85 to 0.95 X the upper. There are no differences 
in color or structure worthy of note. 

The Venezuela specimen is similar to the type series in many 
respects, and it seems to me very probable that it belongs with 
this species. In size and all standard measurements it falls within 
the ranges expressed above, and the wing venation is similar, 
although the wings have more of a yellow tinge toward the base. 



1967 NOTES ON EPIPOMPILUS 15 

There are several color differences: the mandibles are mostly 
testaceous; the antennae are medium brown except the first three 
segments yellowish below, white above; the pronotum has a very 
broad ivory-white band, though occupying the same position as 
in the types; the front femora are brownish, the middle and hind 
femora fuscous, the coxae rather extensively mottled with black; 
the first abdominal segment is wholly black, the spots on tergite 
two confluent medially. 

EPIPOMPILUS EXCELSUS (Bradley) 

This species is characteristic of southeastern Brazil, and appears 
to be relatively common for the genus. Fritz Plaumann, of Nova 
Teutonia, Santa Catarina, has now sent me 13 females and 33 
males from that locality, collected August through April. I have 
also seen one female and 16 males from Rio de Janeiro, collected 
in March, 1966, by Henry and Marjorie Townes, as weU as 10 
males from Alto da Serra, Morretes, Parana, collected by the 
Townes in February 1966 [Coll. H. K. Townes]. I now feel 
convinced that the male I described in 1961 does in fact go with 
this species. 

The series from Rio de Janeiro differs somewhat from specimens 
from Parana and Santa Catarina, but the male genitalia are 
identical and other structural differences minor. The female has 
a pair of white spots superimposed on the rufous base of tergite 
two, and the ocellar triangle is a little less broad, the postocellar 
line being only about 1.5 X the ocello-ocular hne. The males 
from this locality are rather small (fore wing 3.3-4.4 mm) and 
also tend to have a less broad ocellar triangle; they also have 
somewhat more white on the legs, the front coxae having the 
apical 0.3-0.5 white. The genitalia of the specimens examined 
proved to resemble very closely the figure in my 1961 paper (p. 
34, fig. 4). 

(Received 8 March 1967) 



BREVIORA 

Mmseiiimi of ComparatiYe Zoology 

Cambridge, Mass. 17 November, 1967 Number 274 



THE GENUS HOPS (ARANEAE, CAPONIIDAE) IN PANAMA 

AND THE WEST INDIES 

Arthur M. Chickering 

The Caponiidae include a comparatively small group of genera 
now known to be distributed over much of Africa, South America, 
Central America, southwestern part of the United States, and the 
West Indies. For many years I have had a continuous interest in 
this family and during this period I have accumulated a rather large 
collection of certain genera in the family. I have had a special inter- 
est in the genus Nops and this seems to be a convenient time to 
record my understanding of this genus as it is now believed to 
occur in Panama and the West Indies. 

Again, I wish to express my appreciation and gratitude for the 
aid and encouragement in the pursuit of my studies extended by the 
staff of the Museum of Comparative Zoology for many years. My 
thanks are also extended to Dr. W. J. Gertsch, American Museum 
of Natural History, for the loan of numerous Caponiidae from Cen- 
tral America and the West Indies. Grants GB-1801 and GB-5013 
from the National Science Foundation have made it possible for me 
to continue my studies in the Museum of Comparative Zoology and 
to spend a total of nearly eleven months collecting in Panama and 
the West Indies. 

With the exception of the type of Nops gertsch i sp. nov., all types 
named in this paper together with all other specimens belonging to 
the genus Nops in my personal collection will be deposited in the 
Museum of Comparative Zoology. 

Genus Nops MacLeay, 1839 

Type species is Nops guanabacoae by monotypy. 

The genus Nops was established on the basis of a female from 
Cuba. Since that time a dozen additional species have been recog- 
nized from South and Central America and the West Indies. The 



2 BREVIORA No. 274 

most important features of the genus observed during my study of 
the group may be stated as follows: Carapace low; widest opposite 
or just behind the second coxae; considerably narrowed at anterior 
end; median thoracic groove or pit obscure. Eyes: two, on a slightly 
raised tubercle; separated by less than their diameter. Chelicerae 
of moderate size; with no teeth along fang groove but with a rela- 
tively large, soft lobe on the promargin (Fig. 11); fang evenly 
curved; with conspicuous striations on lateral surfaces. Maxillae: 
robust; lobate at base; strongly convergent; obtusely terminated dis- 
tally and not narrowed as in Caponina; somewhat concave near 
middle. Lip: usually firmly united to sternum; about as broad as 
long. Sternum: with a well marked margin; raised somewhat from 
margin and then flat throughout; nearly oval in outline; widest 
opposite interval between second and third coxae. Legs: 4123 in 
order of length; coxae 1243 in order of length; all tarsi with three 
claws each but third c'aw often hidden; first and second tarsi with 
a proximal, ventral, membranous lobe; all tarsi bisegmentate; first 
and second metatarsi with a median, ventral, membranous fold 
nearly or quite throughout the segment; first and second coxae 
lobate at base. Palp: femur in both sexes with a small, prolateral 
cusp at its base in opposition to the cheliceral striations; with a 
more or less conspicuous, tibial brush on the prolateral side near 
distal end in both sexes; in both sexes the tarsus bears a dorsal, 
distal, oval pad of fine hairs; in females the tarsus lacks a terminal 
claw; in males the tarsus is simple but usually more or less distinc- 
tive. Abdomen: elongate; ovoid; with four spiracles clearly shown; 
six spinnerets with anterior and median pairs nearly in a transverse 
row; females usually with a moderately well developed scutum from 
genital groove to pedicel but this is usually poorly developed in 
males; females have no definite epigynum. 

I have been obliged to transfer Caponina blanda Bryant to the 
genus Nops on the basis of what I regard as very clear and definite 
evidence. After a period of uncertainty I have also come to the 
conclusion that Nops coccineus Bryant belongs to Nops blandus. 
Caponina darUngtoni Brvant, known only from a female taken in 
the Dominican Republic, has been rather carefully examined with 
the conclusion that it probably deserves to be placed in a new 
genus. Its status can be determined only when males are available 
for study. The total list of species belonging to the genus Nops, 
as presented in this paper, may be given as follows: Nops blandus 
(Bryant); Nops coccineus Simon; Nops craneae sp. nov.; Nops 
flutillus sp. nov.; Nops gertsclii sp. nov.; Nops glaucus Hasselt; 



1967 NOPS IN PANAMA AND THE WEST INDIES 3 

Nops guanabacoae MacLeay; Nops largus sp. nov.; Nops simla 
sp. nov.; Nops toballus sp. nov.; Nops ursumus sp. nov. With the 
exception of Nops gertschi sp. nov., all new species listed above 
are known from both sexes. 

Key to male Nops from Panama and the West Indies 

la. Species with embolus nearly as long as or longer than the cymbium 
(craneae, guanabacoae, largus, simla) 2 

lb. Species with embolus distinctly shorter than the cymbium (blandus, 
coccineus, flutillus, gertschi, toballus, ursumus) 5 

2a. Species with distinct dorsal abdominal spots on a light back- 
ground {craneae, simla) 3 

2b. Species without distinct dorsal abdominal spots (guanabacoae, 
largus) 4 

3a. Abdomen with a central, longitudinal, irregular, grayish stripe and 

and a series of four irregular, grayish spots on each side . . craneae, p. 6 
3b. Abdomen with a series of paired purplish spots along the dorsal 

surface; without a central stripe simla, p. 13 

4a. Embolus somewhat sinuous largus, p. 1 1 

4b. Embolus not sinuous; somewhat swollen near middle 

guanabacoae, p. 1 1 

5a. Species with at least somewhat definite dorsal abdominal spots 

(coccineus, gertschi, ursumus) 6 

5b. Species with no definite dorsal abdominal spots (blandus, flutillus, 
toballus) 8 

6a. Abdomen with a central, dorsal, indented, brownish stripe consisting 
of four interconnected, irregular angular spots; each lateral side with 
a brownish stripe coccineus, p. 5 

6b. Abdomen lacking such a color pattern as given above (gertschi, 
ursumus) 7 

7a. Abdomen with a series of five pairs of dark grayish, dorsal spots 
ursumus, p. 16 

7b. Abdomen grayish in general, with two light greenish bars in an- 
terior two-thirds; without any such distinct series of spots as given 
above gertschi, p. 9 

8a. Embolus short; about half as long as tarsal bulb blandus, p. 4 

8b. Embolus longer; nearly or quite as long as tarsal bulb (flutillus, 
toballus) 9 

9a. Abdomen purplish dorsal ly flutillus, p. 7 

9b. Abdomen gray dorsally toballus, p. 14 

Except for the females with a distinct abdominal color pattern this sex 
is conspicuously lacking in characters with can be used in constructing a 
key for their separation into species. For this reason no satisfactory key 
for their separation has yet been devised. 



4 BREVIORA No. 274 

Nops BLANDUS (Bryant) 
Figures 1-2 

Caponina blonda Bryant, 1942: 328, pi. 2, fig. 17. The male holotype from 
St. Croix, U. S. Virgin Islands, is in the Museum of Comparative 
Zoology. 

Nops coccineus, — Bryant, 1948: 344. (Not Nops coccineus Simon.) 

Caponina blanda Bryant must be transferred to Nops because 
it very clearly exhibits the characters of the latter genus. Until 
recently I have considered the specimens called Nops coccineus by 
Miss Bryant, from Haiti, as a new species and intended to describe 
it as new. Recently, however, following careful comparison with 
N. blandus, I have been forced to place them together. There are 
minor differences but, in my judgment, no greater than are com- 
monly found among individuals of the same species (Figs. 1-2). 




Figure 1. Nops blandus (Bryant). Left male palp; prolateral view. Fig. 
2. Nops coccineus Bryant (= Nops blandus (Bryant)). Left male palp; 
prolateral view. 



I had hoped to collect specimens of Nops blandus during my two 
recent visits to St. Croix but I failed to find any members of the 
genus. I now have two males from St. John together with several 
immature specimens collected in July, 1966; I also have a female 
together with two immature specimens collected in February, 1964. 
These are all tentatively placed in Nops blandus pending more data 
on the Nops population of these islands. During my recent visit to 
Virgin Gorda, British Virgin Islands. I collected two mature males, 
one female, and two immature specimens. These were all taken on 
August 18, 1966. and are also tentatively placed in this species. 



1967 



NOPS IN PANAMA AND THE WEST INDIES 



A female Nops and several immature specimens collected on St. 
Thomas in February, 1964, and in July, 1966, are not yet definitely 
assigned to species. 

Nops coccineus Simon 
Figure 3 

Nops coccineus Simon, 1891, fig. 18, pi. 42. Male and female syntypes from 
St. Vincent, B. W. I., are in the British Museum (Natural History). 
Simon, 1893: 324; Petrunkevitch, 1911: 133; Lutz, 1915: 82; Berland, 
1932: 339; Roewer, 1942: 316; Bonnet, 1958: 3114. 

While working in the British Museum in the summer of 1958, 
I had an opportunity to examine both sexes of Nops coccineus 
Simon. The following notes were written during my examination of 
these specimens. Female: length from anterior border of clypeus 
to posterior end of anal tubercle 8.78 mm; carapace 2.99 mm long; 
2.26 mm wide opposite interval between second and third coxae 
where it is widest; regularly rounded posteriorly but much nar- 
rowed in front; about 1.3 mm tall. Two eyes circular and separated 
by a little more than the radius of one. Height of porrect clypeus 
3.25 times the diameter of an eye. Chelicerae with a fleshly lobe 
just promarginal to fang; no teeth observed; markedly striate along 




Figure 3. Nops coccineus Simon. Left male palp; prolateral view. Fig. 
4. Nops craneae sp. nov. Left male palp; prolateral view. 



outer surfaces; opposing palpal, femoral cusps moderately devel- 
oped. Tarsi 1 and 2 with a small third claw; tarsi 3 and 4 with a 
somewhat more prominent third claw; all tarsi bisegmentate; tarsi 
1 and 2 with the usual ventral, proximal lobe. Metatarsi 1 and 2 



6 BREVIORA No. 274 

with the usual ventral, median, longitudinal, membranous fold. 
Sternum only extended to bases of fourth coxae. Epigynal area 
lightly cornified. Color in alcohol: cephalothorax very reddish; legs 
somewhat lighter; abdomen yellowish dorsally with a central, in- 
dented, brownish stripe consisting of four connected, irregularly 
angular spots. Male: essential features of the male palp shown in 
Figure 3; total length from anterior border of porrect clypeus to 
posterior end of posterior spinnerets 5.6 mm; color in general like 
that of female but clearer; each lateral side of abdomen with a 
brownish stripe, very irregular; venter yellowish. 

NOPS CRANEAE Sp. nOV. 

Figure 4 

Holotype. The male is from Simla, Arima Valley, Trinidad, 
W. I., in the close vicinity of the Wm. Beebe Tropical Research 
Station, April 17, 1964. This species is named after Miss Jocelyn 
Crane, director of the station. 

Description. Total length from anterior border of clypeus to 
posterior end of extended spinnerets 7.8 mm; length from anterior 
border of clypeus to posterior end of abdomen 7.21 mm. Carapace 
3.25 mm long; 2.34 mm wide opposite interval between second and 
third legs where it is widest; only about 0.55 mm tall; only a 
slight indication of a median, thoracic fovea; surface very finely 
granulate. Eyes: two as usual on a slightly raised tubercle; sepa- 
rated from one another by slightly less than the radius of one; 
height of clypeus equal to nearly 2.75 times the diameter of an eye. 
Chelicerae, maxillae, lip and sternum essentially as given in state- 
ment of characters of the genus. Legs: tibial index of first leg 13, 
of fourth leg 9; no spines observed on legs; typical of the genus in 
regard to tarsal bisegmentation, tarsal claws, tarsal basal, ventral 
lobes and metatarsal ventral, longitudinal folds. Palp: essential 
features shown in Figure 4; cusp at base of femur on prolateral 
surface; oval, dorsal, tarsal pad of short, fine hairs as usual; all 
segments simple except the tarsus. Abdomen: typical of the genus. 
Color in alcohol: carapace and sternum a dark reddish; consider- 
able black pigment in ocular area; legs and mouth parts reddish 
brown in general but with variations; metatarsi 1 and 2 and the 
proximal divisions of tarsi 1 and 2 very light yellowish. Abdomen: 
light greenish with dark gray spots; an irregular, median, dark gray 
stripe extends throughout the dorsum; on each dorsolateral side 
there is a series of 4 irregularly elongated, dark gray spots the last 
of which spreads and unites with the median stripe; the venter has 



1967 NOPS IN PANAMA AND THE WEST INDIES 7 

two small and very irregular, dark gray spots and a pair of larger 
gray spots in a ventrolateral position; the greenish coloration is 
somewhat darker in the region of the tracheal spiracles. 

Female paratype. The described female paratype measures 
10.34 mm from anterior border of clypeus to posterior end of 
abdomen; from anterior border of clypeus to posterior end of 
spinnerets 11.25 mm. Carapace 4.03 mm long; 3.06 mm wide op- 
posite second coxae where it is widest; otherwise essentially as in 
male holotype. Eyes: two, separated by about Vs of their diameter; 
height of clypeus equal to about 3.5 times the diameter of an eye. 
Chelicerae, maxillae, lip and sternum essentially as in male. Legs: 
tibial index of first leg 14, of fourth leg 10; tarsal claws, tarsal, 
basal, ventral lobes, metatarsal ventral, membranous folds and 
other features as stated for the genus all typical. Abdomen: es- 
sentially as in male; scutum only moderately developed anterior to 
genital groove. Color in alcohol: nearly identical to that of male; 
abdominal color pattern clear and definite; venter with three very 
irregular, dark spots on each ventrolateral side; the first of these is 
lateral to the scutum, the second is a little behind the middle and 
somewhat nearer the middle line, while the third is anterior to the 
spinnerets and still nearer the middle line. 

Records. The described female paratype is also from Trinidad, 
W. I., with no date of collection (N. A. Weber). Two females are 
in the collection with the described female paratype; two immature 
specimens from Balandra Bay, Trinidad, April, 1922 (Reynolds 
collection). I took eight specimens in April, 1964, in the vicinity of 
Simla, Arima Valley, Trinidad, W. I. One immature specimen from 
Piarco (A. M. Nadler), January, 1955, and a female from Gas- 
paree, November, 1944 (T. S. Jones) are tentatively placed here. 

NOPS FLUTILLUS Sp. UOV. 

Figures 5-6 

Holotype. The male holotype and described female paratype 
are from the Dutch West Indies, Curagao, Sint-Nicolaas; St. 
Marthaa, December 29, 1962 (B. de Jong and H. W. Levi). 

The specimens referred to this species were tentatively placed 
under the name Nops gloiicus Hasselt — a species based upon a 
female from the West Indian island of Bonaire. After a period of 
some uncertainty the decision has been made to regard the species 
as new. The name of the species is an arbitrary combination of 
letters. 



8 



BREVIORA 



No. 274 



Description. Total length of male from anterior border of cly- 
peus to posterior end of abdomen 5.79 mm. Carapace 2.5 mm 
long; 1.87 mm wide opposite second coxae where it is widest; about 
0.55 mm tall; otherwise essentially typical of males of the genus. 
Eyes: two as usual; separated by about %4 of their diameter. 
Chelicerae, maxillae and lip essentially typical of males of the 
genus. Sternum: longer than wide in ratio of about 6:5; somewhat 
more oval in outline than usual; fourth coxae separated by about 
%o of their width; otherwise quite typical of the genus. Legs: 
tibial index of first leg 14, of fourth leg 10; tarsi typical with respect 
to bisegmentation, claws, and ventral, proximal, membranous lobes; 
metatarsi also typical with respect to the ventral, medial, membra- 
nous fold; no spines but with many hairs and bristles; trichobothria 
observed on tarsi, metatarsi, tibiae and, possibly, also on palpal 
segments. Palp: essential features shown in Figures 5-6. Abdomen 
apparently typical of the genus in all essential features. Color in 
alcohol: all parts except abdomen essentially as described for N. 
largus sp. nov.; abdomen purplish over most of the dorsum, be- 
coming darker and broader posteriorly; lateral sides yellowish, be- 
coming purplish posteriorly; venter with a fairly well defined 
scutum anterior to genital groove and with light green color around 
a broad margin; remainder of venter yellowish with purplish dots, 
becoming more definitely purplish toward posterior end. 




Figures 5-6. Nops fliitilliis sp. nov. Fig. 5. Left male palp; prolateral view. 
Fig. 6. Tip of embolus, more enlarged; retrolateral view. Figs. 7-8. Nops 
gertschi sp. nov. Fig. 7. Right male palp; prolateral view, Fig. 8. Tarsal 
bulb, more enlarged; retrolateral view. 



1967 NOPS IN PANAMA AND THE WEST INDIES 9 

Female paratype. The described female paratype has a total 
length of 8.45 mm from anterior border of porrect clypeus to pos- 
terior end of abdomen. Carapace 4.03 mm long; 3.04 mm wide op- 
posite posterior border of second coxae where it is widest; 0.96 mm 
tall; otherwise essentially typical of the genus. Eyes: two as usual 
on a low tubercle; separated by nearly % of the diameter of an eye; 
height of porrect clypeus equal to about 4.5 times the diameter of 
an eye. Chelicerae, maxillae, lip and sternum all essentially typi- 
cal of females of the genus. Legs: tibial index of first leg 18, of 
fourth leg 13; tarsi typical with respect to claws, bisegmentation, 
and basal, ventral lobes; metatarsi typical with respect to ventral, 
medial, longitudinal folds; claw tufts moderately developed; with no 
palpal tarsal claws; with the usual, distal, prolateral, tibial, palpal 
brush. Abdomen: essentially as in male; a well defined scutum oc- 
cupies the whole area of tracheal spiracles continued to the pedicel; 
what has frequently been termed the epigynum is a slightly re- 
curved slit between anterior spiracles. Color in alcohol: essentially 
as in male holotype with minor variations; with considerable light 
green color in parts of the ventral scutum. 

Records. One male paratype, two female paratypes and two im- 
mature specimens taken with the holotype. One male paratype and 
eleven immature specimens from several localities on the island of 
Curagao collcted by Dr. and Mrs. H. W. Levi and Mr. B. de Jong 
in December, 1962. 

NOPS GERTSCHI sp. nOV. 

Figures 7-8 

Holotype. The male holotype is from Loma Cibao, La Vega, 
Dominican Republic. August 9, 1956, 600-1600 m elevation (Dr. 
Allan F. Archer, E. B. M.). The holotype will be deposited in the 
American Museum of Natural History, New York, N. Y. 

Until recently the specimen described here as a new species was 
regarded as another specimen of Nops blandus (Bryant) but study 
has convinced me that it must be regarded as new. The species is 
named after Dr. W. J. Gertsch. 

Description. Total length from anterior border of clypeus to 
posterior end of abdomen 5.98 mm; from tip of somewhat por- 
rect chelicerae to posterior end of abdomen 6.18 mm. Carapace 
2.86 mm long; 2.08 mm wide opposite second coxae where it is 
widest; narrowed to about 0.78 mm at anterior end. Eyes: two as 
usual on a low tubercle; separated by their diameter; surrounded 



10 BREVIORA No. 274 

by black pigment, lighter between the eyes; height of porrect 
clypeus equal to nearly 3.75 times the diameter of an eye. 
Chelicerae, maxillae, lip and sternum all essentially typical of males 
of the genus; fourth coxae separated by their width. Legs: tibial 
index of first leg 14, of fourth leg 9; tarsi typical as stated in de- 
scription of the genus; metatarsi 1 and 2 also typical with respect 
to the ventral, longitudinal, membranous fold; typical with respect 
to other observed features. Palp: essential features shown in Fig- 
ures 7-8; palpal, tibial, prolateral brush apparently lacking on left 
palp. Abdomen: essentially typical of males of the genus. Color in 
alcohol: carapace and sternum an orange yellow; with consider- 
able black pigment in ocular area; legs nearly the same as carapace 
dorsally but lighter ventrally; lip and chelicerae like sternum; 
maxillae lighter; palp like legs with variations. Abdomen: dorsum 
grayish in general but with two irregular, light greenish, dorsolateral 
bars in anterior two-thirds; lateral sides dark grayish with ventral 
extensions of the lighter dorsal, irregular bars; venter yellowish 
with a faint greenish tint anterior to genital groove; posterior to 
genital groove light greenish with irregular dark spots for nearly 
three-quarters of the distance to the spinnerets; the area im- 
mediately in front of the spinnerets is dark like most of the dorsum. 

Nops GLAUCUS Hasselt 

Nops glaiiciis Hasselt, 1887: 77, figs. 1-4, 10. Female holotype from Bonaire, 
Dutch West Indies, in the Natural History Museum, Leiden. Simon, 
1892: 449; Petrunkevitch, 1911: 134; Roewer, 1942: 316; Bonnet, 
1958: 3114. 
Through the courtesy of Dr. van der Hammen of the Natural 
History Museum in Leiden I have been able to examine the holo- 
type of this species from which I have derived the following: total 
length 9.17 mm from anterior border of clypeus to posterior end 
of abdomen; carapace 3.9 mm long and 2.92 mm wide opposite 
interval between second and third legs. Eyes: two as usual on a 
low tubercle; separated by nearly three-fourths of the diameter of 
an eye; height of clypeus nearly equal to four times the diameter 
of an eye. Legs: typical of the genus in all respects as given in the 
description of the genus. The usual palpal, tibial brush is present. 
The color appears to have faded somewhat during long preserva- 
tion. Legs and palps light yellowish; carapace, sternum, and other 
mouth parts all reddish brown; abdomen light grayish with a pur- 
plish tint dorsally and yellowish ventrally. I consider the original 
drawings very good with one exception; in figure 1 the second femur 



1967 NOPS IN PANAMA AND THE WEST INDIES 11 

is over-inflated; in the holotype under observation the second 
femur is only normally robust. Simon reported this species from 
Venezuela in 1892. Apparently it has not been definitely recorded 
since that time. 

NoPS GUANABACOAE MacLcay 
Figures 9-10 

Nops guanabacoae MacLeay, 1839: 1-14, pis. 1-2. The female holotype 
from Cuba is probably in the British Museum (Natural History), 
London. Walckenaer, 1842: 442; Hasselt, 1887: 71; Simon, 1893: 
328; Petrunkevitch, 1911: 133; Bryant. 1940: 271; Roewer, 1942: 
316; Bonnet, 1958: 3114. 

The male was unknown until 1940 when Miss Bryant described 
it and had both sexes for study. As a result of my examination of 
these specimens I am obliged to disagree with several of the pub- 
lished statements concerning the male. These may be briefly 
stated as follows: eyes separated from one another by nearly % of 
the diameter of one of them; chelicerae with the usual fleshy lobe 
on the promargin of the fang groove; lip firmly united to sternum 
and only a little longer than wide; legs 4123 in order of length; all 
tarsi with three claws; tarsi 1 and 2 with the typical ventral, proxi- 
mal lobe; metatarsi 1 and 2 with the ventral, median, membranous 
fold; the usual palpal, tibial brush is present in both sexes. Es- 
sential features of the male and female palps are shown in Figures 
9-10. There are now two mature males together with several fe- 
males and immature specimens from several localities in Cuba, and 
the species is, apparently, known only from that island. 

Nops largus sp. nov. 
Figures 11-15 

Holotype. The male is from the Panama Canal Zone Forest Pre- 
serve, January 21, 1958. The name of the species is a Latin ad- 
jective referring to its abundance in its natural habitat. 

Description. Total length 6.7 mm from anterior border of cly- 
peus to posterior end of abdomen; from anterior border of slightly 
extended chelicerae to posterior end of spinnerets 6.89 mm. Cara- 
pace 3.15 mm long; 2.34 mm wide opposite second coxeae where 
it is widest; about 0.7 mm tall. Eyes: two as usual on a slightly 
raised tubercle; separated by slightly less than their radius; 
height of porrect clypeus nearly equal to 2.66 times the 
diameter of an eye. Chelicerae, maxiUae and lip essentially 



12 



BREVIORA 



No. 274 



typical of the genus; cheliceral fleshy lobe shown in Figure 
11. Sternum: surface rather coarsely granulate; coxae 1 and 2 
basally lobed as usual; terminated just posterior to bases of fourth 
coxae which are separated by about their width (Fig. 12). Legs: 
tibial index of first leg 14. of fourth leg 11; spines lacking; tarsal 
claws, ventral, proximal lobes on tarsi 1 and 2 and ventral, medial. 




Figures 9-10. Nops giianabacoae Macleay. Fig. 9. Left male palp; pro- 
lateral view. Fig. 10. Left female palp; prolateral view. Figs. 11-15. Nops 
largus sp. nov. Fig. 11. Left chelicera of male. Fig. 12. Sternum of male. 
Fig. 13. Left male palp; prolateral view. Fig. 14. Tip of embolus; en- 
larged. Fig. 15. Left female palp; prolateral view. 



1967 NOPS IN PANAMA AND THE WEST INDIES 13 

longitudinal folds on metatarsi 1 and 2 all typical of the genus. 
Palp: essential features shown in Figures 13-14. Abdomen: quite 
typical of the genus; ventral scutum rather poorly developed; with 
a copious supply of short hair. Color in alcohol: carapace and 
sternum a bright orange red; legs generally somewhat lighter 
dorsally and still lighter ventrally; ail coxae and mouth parts yel- 
lowish with variations. Abdomen: greenish gray with many nar- 
row, light colored lines running in a variety of directions; lateral 
sides grayish with the gray areas extending to ventral surface and 
covering the posterior fourth of the venter with the remainder nearly 
white with greenish spots around spiracles. Considerable variation 
in coloration has been noted among paratypes with a tendency for 
a purplish color to replace the greenish gray. 

Female paratype. Total length of described female paratype 
8.58 mm from anterior border of clypeus to posterior end of abdo- 
men; from anterior border of slightly porrect chelicerae to pos- 
terior end of spinnerets 8.78 mm. Carapace 3.84 mm long; 2.86 
mm wide opposite second coxae where it is widest; 0.58 mm tall; 
posterior end transversely rugulose; with a series of light colored, 
fine, intricately branched lines covering entire surface; otherwise as 
in male. Eyes: two as usual; separated by about % of their di- 
ameter; height of porrect clypeus equal to about 3.3 times the di- 
ameter of an eye. Chelicerae, maxillae, lip and sternum essentially 
as in male and typical of females of the genus. Legs: tibial index 
of first leg 15, of fourth leg 9; essentially as in male with respect 
to the ventral lobe on tarsi 1 and 2, the median, ventral fold on 
metatarsi 1 and 2, and tarsal claws. The palp lacks a tarsal claw 
but has the usual tibial brush and the femoral, prolateral, basal 
cusp. Abdomen: essentially as in male. Color in alcohol: es- 
sentially as in male with minor variations. 

Records. The described female paratype is from Barro Colorado 
Island, Panama Canal Zone, January, 1958. About forty mature 
males together with many females and immature specimens as- 
signed to this species have been taken in several localities in the 
Canal Zone and in Panama proper in El Valle, El Volcan, and 
Arraijan. I have found it especially abundant on Barro Colorado 
Island, Canal Zone. 

NoPS SIMLA sp. nov. 
Figures 16-17 

Holotype. The male is from Simla, Arima Valley, Trinidad, W. 
I., in the close vicinity of the Wm. Beebe Tropical Research Sta- 
tion, March 31, 1964. The name of the species is a noun used in 
apposition after the locality where the holotype was collected. 



14 BREVIORA No. 274 

Description. Total length 4.94 mm. Carapace 1.95 mm long; 
1.45 mm wide opposite second coxae where it is widest; 0.44 mm 
tall; median thoracic fovea barely indicated. Eyes: two as usual; 
separated by about %i of their diameter; with a moderate amount 
of black pigment around eyes; height of porrect clypeus nearly 
equal to 2% times the diameter of an eye. Chelicerae, maxillae 
and lip with only minor variations from the typical. Sternum: 
very granulate; obtusely terminated between bases of fourth coxae 
which are separated by about ^;-, of their width. Legs: tibial index 
of first leg 14, of fourth leg 10; tarsi typical of the genus with re- 
spect to bisegmentation and claws; tarsi 1 and 2 typical with re- 
spect to ventral, basal lobe; metatarsi 1 and 2 essentially typical 
with respect to ventral, medial, longitudinal fold except that this 
seems to be somewhat reduced in prominence. Palp: essential 
features shown in Figure 16; embolus very long. Abdomen es- 
sentially typical of the genus. Color in alcohol: carapace, lip and 
sternum a medium reddish brown; palp and other mouth parts 
yellowish with variations; all coxae light yellowish; other segments 
of legs reddish brown with variations; legs 1 and 2 somewhat the 
darkest. Abdomen: background color light yellowish; dorsum with 
a series of five pairs of light, irregular, purplish spots (Fig. 17); at 
posterior end there is a pair of elongated, lateral, purplish spots 
extending forward from the spinnerets; venter generally yellowish. 

Female paratype. Total length 5.92 mm. Carapace 2.15 mm 
long; 1.69 mm wide opposite posterior border of second coxae 
where it is widest; about 0.44 mm tall. Eyes essentially as in male. 
Chelicerae, maxillae, lip and sternum essentially as in male. Legs: 
tibial index of first leg 16, of fourth leg 10; other features of legs 
essentially as in male and typical of the genus. Abdominal features 
and color essentially as in male. 

Records. The female paratype was taken in the same locality as 
the holotype on April 25, 1964, together with an immature speci- 
men. The following have been on loan from Dr. W. J. Gertsch: 
two males from Arima, Trinidad, May, 1953 (N. L. H. Kraus) and 
an immature specimen from Piarco, Trinidad, January, 1955 (A. 
M. Nadler). 

Nops TOBALLUS sp. nov. 
Figures 18-19 

Holotype. The male holotype is from St. Catherine Parish, near 
May Pen, Jamaica, W. I., November 22, 1957. The name of the 
species is an arbitrary combination of letters. 



1967 



NOPS IN PANAMA AND THE WEST INDIES 



15 




Figures 16-17. Nops simla sp. nov. Fig. 16. Left male palp; prolateral 
view. Fig. 17. Abdomen: dorsal view. Figs. 18-19. Nops tobaUus sp. nov. 
Fig. 18. Left male palp; prolateral view. Fig. 19. Left female palp; pro- 
lateral view. Figs. 20-21. Nops iirsitmus sp. nov. Fig. 20. Male carapace; 
dorsal view. Fig. 21. Left male palp; prolateral view. 



Description. Total length 3.94 mm. Carapace 1.87 mm long; 
1.47 mm wide opposite second coxae where it is widest; 0.51 mm 
tall; surface very finely granulate; with no definite thoracic fovea. 
Eyes: two as usual, separated by about their radius; height of por- 
rect clypeus equal to a little less than three times the diameter of 
an eye. Chelicerae, maxillae, lip and sternum essentially typical of 



16 BREVIORA No. 274 

the genus with minor differences; fourth coxae separated by shghtly 
more than their width. Legs: tibial index of first leg 15, of fourth 
leg 10; with very slender spines, hardly more than robust bristles; 
tarsal claws, tarsal ventral lobes and metatarsal, ventral folds on 
legs 1 and 2 typical of the genus. Palp: essential features shown 
in Figure 18. Abdomen: ovoid; with many short hairs; surface 
very granular; otherwise quite typical of the genus. Color in alco- 
hol: carapace a bright orange with eyes on a black spot divided in 
front; legs nearly the same color above but lighter below; sternum 
somewhat lighter than carapace; abdomen with dorsum and lateral 
sides gray, venter nearly white anterior to genital groove, greenish 
from genital groove to near spinnerets where it becomes grayish. 

Female paratype. Total length 6.18 mm, exclusive of somewhat 
extended posterior spinnerets; 6.5 mm long to tip of posterior spin- 
nerets. Carapace: 2.73 mm long; 2.02 mm wide opposite second 
coxae where it is widest; about 0.65 mm tall; otherwise essentially 
as in male. Eyes, chelicerae, maxillae, lip and sternum essentially as 
in male with minor differences. Legs: tibial index of first leg 16. of 
fourth leg 10; essentially typical of the genus in respect to other 
features. Abdomen also essentially typical of the genus. 

Records. The described female paratype is from Clarendon 
Parish, 3 miles north of May Pen, Jamaica, W. L, November 10, 
1963. Additional specimens are in the collection from Jamaica as 
follows: a female from Blue Mts., Main Range, August, 1934 (P. 
J. Darlington, Jr.); a male from St. Thomas Parish, Morant Bay, 
May, 1956 (C. C. Hoff); several immature specimens from St. 
Andrew, St. Catherine, and St. Thomas parishes taken in 1956, 
1957 and 1963. 

Nops URSUMUS sp. nov. 
Figures 20-21 

Nops maculata, — Banks, 1929: 53-96, 4 pis. (Not A'^. maciilata Simon). 

Holotype. The male holotype was taken in a Berlese funnel on 
Barro Colorado Island, Panama Canal Zone, between April and 
June, 1950, by Dr. James Zetek. The name of the species is an 
arbitrary combination of letters. 

Description. Total length from anterior border of clypeus to 
posterior end of abdomen 4.94 mm; from anterior border of 
clypeus to posterior end of spinnerets 5.14 mm. Carapace 2.07 
mm long; 1.52 mm wide opposite posterior border of second 
coxae where it is widest; 0.59 mm tall; with no discernible thoracic 



1967 NOPS IN PANAMA AND THE WEST INDIES 17 

fovea; surface finely granulate; with a sparse covering of hair (Fig. 
20). Eyes: two as usual; separated by about %3 of their diameter; 
height of porrect ciypeus about equal to 2.5 times the diameter of 
an eye. Chelicerae, maxillae, lip and sternum apparently quite 
typical of the genus as observed in this study. Legs: tibial index 
of first leg 13, of fourth leg 8; legs 1-3 only slightly different in 
length; typical of the genus with respect to tarsal claws, ventral, 
proximal lobes on tarsi 1 and 2 and ventral, median fold on 
metatarsi 1 and 2. Palp: essential features shown in Figure 21. 
Abdomen: all features essentially typical of the genus. Color in 
alcohol: carapace and sternum a rich, dark orange with a moderate 
amount of black pigment in ocular region. Mouth parts and legs 
reddish brown with variations. Abdomen: background color of 
dorsum a light greenish with five pairs of dark grayish spots; the 
first pair of spots are broad and extend posteriorly for a consider- 
able distance and are separated by a narrow, greenish stripe in the 
middle; the remaining four pairs of spots are diagonal bars reach- 
ing forward and then downward for a very short distance ventro- 
laterally; the last two pairs are somewhat united and the fourth pair 
is considerably extended posteriorly; shortly anterior to the anal 
tubercle there is a single, small, irregular spot somewhat united to 
the most posterior pair of bars; the venter is whitish with a green- 
ish tint; a short distance anterior to the spinnerets there is a short, 
median line of irregular gray dots and lateral to these is an irregu- 
lar dotted area on each side. 

Female paratype. Total length from anterior border of ciypeus 
to posterior end of abdomen 9.42 mm; length from anterior border 
of ciypeus to posterior end of spinnerets 9.94 mm. Carapace 3.45 
mm long; 2.67 mm wide; 0.85 mm tall; otherwise essentially typi- 
cal of females of the genus. Eyes: two as usual; separated by about 
their radius; height of ciypeus nearly equal to 3.5 times the dia- 
meter of an eye. Chelicerae, maxillae, lip and sternum essentially 
typical of females of the genus; sternal suture line unusually 
clear but probably does not indicate freedom of movement for the 
lip. Legs: tibial index of first leg 12, of fourth leg 9; tarsal claws, 
ventral, proximal lobe on tarsi 1 and 2 and ventral, median fold on 
metatarsi 1 and 2 all typical of the genus. Abdomen: essentially 
typical of females of the genus; region from genital groove to 
pedicel rather poorly supplied with a scutum. Color in alcohol: 
essentially as in male with minor variations; here the two posterior, 
dorsal, abdominal bars are united to one irregularly shaped spot; 
the venter has scattered dark spots, the largest of which occur as a 



18 BREVIORA No. 274 

pair of very irregular spots shortly anterior to the spinnerets. 

Records. The described female paratype was taken in the same 
locality as the holotype, January, 1958. One mature male para- 
type was taken with the holotype; another male was taken by 
Dr. Zetek in the same locality and by the same method, Oct. -Dec, 
1941. A very few females which may be mature and nearly three 
dozen others in various stages of immaturity are in the collection 
from Barro Colorado Id., C. Z. Forest Preserve, Summit Gardens, 
and Pedro Miguel, all in the Canal Zone. The immature specimen 
reported by Banks (1929) as N. macidata Simon quite clearly be- 
longs in N. ursumus sp. nov. The following have been on loan 
from Dr. VV. J. Gertsch: a male and a female from Barro Colorado 
Id., Dec. 1937-Jan. 1938 (F. J. Ryan); three immature specimens 
from the same locality taken July, 1930, and July-August, 1938; 
an immature specimen taken in Santa Rosa, Colon Prov., Panama, 
Sept., 1945 (C. D. Michener). 

BIBLIOGRAPHY 

Banks, Nathan 

1929. Spiders from Panama. Bull. Mus. Comp. Zool., 69: 53-96, 4 pis. 
Berland, Lucien 

1932. Encyclopedic Entomologique. 16. Les Arachnides. Paris. 
Bonnet, Pierre 

1958. Bibliographia Araneorum. Toulouse. Vol. 2(4). 
Bryant, Elizabeth B. 

1940. Cuban spiders in the Museum of Comparative Zoology. Bull. 

Mus. Comp. Zool.. 86(7); 249-532. 22 pis. 
1942. Notes on the spiders of the Virgin Islands. Ibid., 89(7): 317- 

363, pis. 1-3. 
1948. The spiders of Hispaniola. Ibid.. 100(4): 331-447, 12 pis. 
Cambridge, F. O. P.- 

1897- Arachnida-Araneida. Vol. II. //;: Biologia Centrali-Americana. 
1905 Dulau & Co., London. 
Hasselt, a. W. M. van 

1887. Etudes sur le genre Nops. Tijds. Entomol., 30: 77-86, figs. 
1-4, 10. 
LuTZ, Frank E. 

1915. List of Greater Antillean spiders with notes on their distribu- 
tion. Ann. New York Acad. Sci., 26: 71-148. 
Macleay, W. S. 

1839. On some new forms of Arachnida. Ann. Natur. Hist. 2: 1-14, 
pis. 1-2. 
Petrunkevitch, Alexander 

1911. A synonymic index-catalogue of spiders of North, Central, 
South America, etc. Bull. Amer. Mus. Natur. Hist., 29: 1-809. 



1967 NOPS IN PANAMA AND THE WEST INDIES 19 

ROEWER, C. Fr, 

1942. Katalog der Araneae. Vol.1: 1-1040. Bremen. 
Simon, Eugene 

1891. On the spiders of the Island of St. Vincent. Pt. 1. Proc. 
Zool. Soc. London, Nov. 17, 1891: 549-575. 

1892- Histoire naturelle des Araignees. Deuxieme edition. Vol. 1. 
1895. Librairie Encyclopedique de Roret, Paris. 

1892. Voyage au Venezuela. Ann. Soc. Entomol. France, 1892: 449. 
Walckenaer, C. a. Baron de 

1842. Historie naturelle des insectes: Apteres. Vol.2. Paris. 

(Received 20 March 1967.) 



BREVIORA 



Mmiseiiiiim of Connparative Zoology 

Cambridge, Mass. 17 November, 1967 Number 275 

A NEW SCiNCID LIZARD FROM THE NORTHERN 
SOLOMON ISLANDS 

Allen E. Greer and Fred Parker' 



During a term of duty by Parker (1962-1963) and a subse- 
quent ( 1966) collecting trip by him to Bougainville and neighbor- 
ing islands in the extreme northern Solomon Islands, a previously 
undescribed species of skink was discovered on Bougainville and 
Shortland Islands. A single specimen of the new species from 
Choiseul is also known but has previously been reported in the 
hterature as Sphenomorphus solomonis (Burt and Burt, 1932:544). 

On the basis of current, but not necessarily phylogenetically 
correct, generic concepts, the new species is assigned to the genus 
Sphenomorphus and may be known as 

Sphenomorphus tanneri- new species 

Holotype: Museum of Comparative Zoology 76551; collected 
by Fred Parker at Kunua, Bougainville, on 8 June 1963. 

Pamtxpes (423 specimens) :"nORTHERN BOUGAINVILLE: 
KUNUA (100 feet above sea level): MCZ 76483, 76484, 22.vi, 
ll.vii.l962; MCZ 76486-76487, 25-30.xii.1962; MCZ 76488- 
76550. 76552-76623. 77308-77361, 78091, 78299, 84140, + 2 
untagged specimens, 12.v-29.vii.l963; MELILUP {ca. 3000 feet 
a.s.l.)^ MCZ 89593. 92295, 92326-92339, 92366-92368, 12- 
20.V.1966; MUTAHI (2700 ± 500 feet a.s.l.): MCZ 87570- 
87577, 88799, 89102-89126, 89594, 91430-91437. 92203-92228, 
92269-92294, 92296-92302, 92304-92325, 92340-92365, 92372- 
92377, 9-20.V.1966; RAMAZON RIVER (1600-2400 feet a.s.l.): 



^ Department of District Administration, Kimdiawa, Territory of New 
Guinea. 

"The new species is named for Charles Tanner, Honorary Curator of 
Reptiles at the National Museum, Melbourne, who introduced one of us 
(Parker) to reptiles several years ago and has since helped in many ways. 



2 BREVIORA No. 275 

MCZ 92229-92236, 19.V.1966; TOPANAS (500 feet a.s.l.): MCZ 
88443-88453, 8.V.1966. 

SOUTHERN BOUGAINVILLE: MATSIOGU (2100 feet 
a.s.l.): MCZ 92379-92382, 92522, 2.iv. 24.iii.1966; PAMAUITA 
(1200 feet a.s.l.): MCZ 92521, 92523, 23. ii, 2.iv.l966; TURI- 
BOiRU (260 feet a.s.l.); MCZ 87578-87580, 87727-87732, 
92496-92501, 92520, 92990-92992, 9-21. iii. 1966; MCZ 92369- 
92371, 26-27.iv.I966. 

SHORTLAND: KOLIAI (10 feet a.s.l.): MCZ 89736, 
1 l.iv.l966. 

CHOISEUL: American Museum of Natural History 44004. 

Dioij^nosis: S. kinneri differs from all other species with 28-32 
midbody scales in its species group (Table 4) in the following 
combination of characters: small size (snout-vent length 52 mm 
or less); prefrontals forming a median suture; adpressed limbs 
widely separated; no symmetrical pairs of nuchal scales; m!ddorsal 
scales subequal rather than with the scales of the two vertebral 
rows transversely enlarged; dorsum rich dark brown to brownish 
black with small paler spots on body which often coalesce on the 
anterior dorsolateral line to form a distinct light line. 

S. tanneri is most similar in squamation to S. nigriventre de Rooij 
(1915: 214-215) from southern New Guinea, and S. antoniorum 
Smith (1927: 216-217) from Timor. It differs from S. nigriventre 
in being much smaller in size (snout-vent length 90 mm for 
nigriventre) and lacking the dorsal transverse series of light, dark- 
edged spots of this species. It differs from S. antoniorum in having 
the prefrontals meeting medially (prefrontals separated or just 
touching one another in antoniorinn), and in lacking the light 
brown dorsum with a heavy clustering of darker brownish spots 
along the dorsolateral line. 

Description (Fig. 1): A cryptic skink ranging in snout-vent 
length from 23-52 mm; tail slightly longer than snout-vent length; 
head obtusely conical; limbs pentadactyl, failing to meet when 
adpressed to body. 

Rostral slightly wider than deep, projecting well onto dorsal 
surface of snout; external naris placed well forward and ventral in 
a single large nasal; supranasals lacking; frontonasal wider than 
long, forming a short suture with rostral but separated from frontal 
by prefrontals, i.e., prefrontals paired and meeting medially (ex- 
cept in one individual); single anterior and posterior loreals; fron- 
tal about as long as frontoparietals and interparietal measured 
along midline, obtusely rounded posteriorly and in contact with 



1967 



NEW SOLOMON ISLANDS SKINK 




B 




Figure 1. Splienoinoiplius taiincii (MCZ 92304, paratype): (A) Dorsal 
view of the head and nape, showing two successive "nuchal" scales on the 
right side of the nape; (B) nasal area on the left side of the head, showing 
the external naris situated anteroventrallv in the large nasal scale. 



2 anteriormost supraoculars; 4 supraoculars; lower eyelid scaly, 
scales of eyelid separated from supralabials by a complete row of 
subocular scales; frontoparietals and interparietal distinct, approx- 
imately subequal in size; parietals meet behind interparietal; no 
distinct, symmetrical series of nuchais; 6-7 supralabials, 4th or 
5th supralabial below center of eye (Table 1 ). 

Ear opening oval, tympanum slightly sunk below surface; auric- 
ular lobes lacking; midbody scales smooth, in 28-32 (usually 30- 
32, see Table 1 ) longitudinal rows; midbody scales of dorsum 
subequal in size; a pair of enlarged preanals; 3 median rows of 
subcaudal scales subequal in size; 13-20 (usually 15-19, see Table 
1 ) obtusely keeled lamellae beneath 4th (longest) toe; dorsal 
surface of 4th toe covered distally by a few single scales, medially 
by 3 longitudinal rows of scales and proximally by 4 or 5 rows 
(Group IV of Brongersma, 1942). 

Color (Fig. 3): In life the color pattern is relatively constant. 
The dorsum is a rich dark brown to brownish black with small 
paler spots on the body and tail. Larger light spots on the anterior 



4 BREVIORA No. 275 

dorsolateral line often coalesce to form a noticeably distinct line. 
The flanks are marked as the dorsum but usually lighter. The 
chin and throat are heavily marked with black to brown; the an- 
terior venter is whitish to yellowish; the posterior venter and ventral 
surface of the tail arc occasionally translucent yellow, but more 
often, translucent dull reddish. Some specimens have dark spots 
under the tail and others show a bar of fine spots on the posterior 
part of the throat. Preserved specimens lose the yellowish and 
reddish colors. 

Some individuals from the southern Bougainville (Turiboiru. 
Pamauita and Matsiogu) populations lack the light dorsal spotting 
and tend to have the dark pigment of the dorsum and sides con- 
centrated through the center of the scales. This gives the appear- 
ance of dark longitudinal stripes on an only slightly lighter back- 
ground. 

Distribution (Fig. 2): S. tanncri has been collected at both the 
north and south ends of Bougainville (422 specimens), as well as 
on Shortland (1 specimen) and Choiseul (1 specimen). Altitud- 
inally, the species is known to range from the coastal lowlands up 
to about 3000 feet. 

Variation: The variation of several meristic characters is sum- 
marized in Table 1 . 

The degree of contact between the prefrontals ranges from a 
broad median suture to a bare "point contact" at their inner 
angles. In only a single specimen of the whole series, however, do 
the prefrontals just fail to meet at their inner angles. In the ma- 
jority of specimens, the line of contact is at least one-third to one- 
half the length of the prefrontal. 

Large, symmetrical nuchal scales are definitely lacking, although 
the sporadic occurrence of one or more (serial) large, transverse 
scales in the 4-5 scale rows posterior to the parietals ( Fig. 1 ) sug- 
gests that perhaps the genetic basis for the formation of nuchal 
scales is still being variably expressed. Such a high degree of 
variability might well obtain in a previously well-ordered structure 
that was being secondarily lost. 

Either the 4th or 5th supralabial may be below the center of the 
eye. When it is the 4th supralabial, either the 2nd or 3rd supra- 
labials have fused to form a scale as long as the first supralabial 
or a small wedge-shaped scale may project between the 2nd and 
3rd supralabial, thus separating them except along the edge of the 
lip where they remain in contact. In most individuals, however, 
the 5th supralabial is below the center of the eye. 



1967 



NEW SOLOMON ISLANDS SKINK 



The 4th supralabial occurs below the center of the eye with 
greater frequency in the populations from southern Bougainville 
(Turiboiru. Pamauita and Matsiogu = 37% ) than in the'popula- 
tions from northern Bougainvihe (Kunua, Mutahi, Topanas, 



155° E 



156° E 



30 miles 



#Topanas 
.Ramazon River 



■6°S 



7°S 




6°S 



SHORTLAND I 



FAURO 



7°S- 



I55°E 



MONO 1 (^ 



156° E 



Figure 2. Bougainville and surrounding islands, showing the localities 
from which Splwiioniorpluis tcinncri is presently known. There is also a 
single specimen from an unknown iocalitv on Choiseul. 



6 BREVIORA No. 275 

Melilup and Ramazon River = \VA%). The frequency of this 
trait is probably subject to clinal variation from one end of Bou- 
gainville to the other, but the rate of change per unit distance will 
not be known until populations from the central two-thirds of 
Bougainville are sampled. 

This particular difference in squamation between northern and 
southern populations on Bougainville is made even more interest- 
ing by noting its correlation with the difference in color patterns 
between the two populations (see above) — the striped pattern 
occurring in some individuals in the south but never in the north. 
These differences may be taxonomically important, but their sig- 
nificance can not be properly evaluated until populations from 
intervening localities are studied. 

Type specimen: The holotype has 30 scales around midbody, 
16-16 lamellae beneath the 4th toe, and the 5th supralabial is 
below the center of the eye. 

Shortland Island specimen: The single specimen from Shortland 
Island (Fig. 2), the only other island besides Bougainville and 
Choiseul from which the species is known, does not differ substan- 
tially from the Bougainville specimens. There are 30 scales around 
midbody and 17-17 lamellae beneath the 4th toe. 

Comparisons between the Shortland population and the northern 
and southern populations on Bougainville would be interesting, 
but can hardly be carried very far at present with only one speci- 
men available to represent the Shortland population. It is interest- 
ing to note, however, that while the Shortland specimen lacks the 
striped pattern characteristic of some individuals from southern 
Bougainville, it does have the 4th supralabial below the center of 
the eye on both sides of the head. This condition, as discussed 
above, tends to occur more frequently in lizards from southern 
Bougainville (37^r) than those from northern Bougainville 
{WVi'c). 

Choiseul Island specimen: The single specimen from an un- 
known locality on Choiseul (AMNH 44004) is poorly preserved 
but appears to differ in no significant way from Bougainville and 
Shortland specimens. 

Habitat: On Bougainville the species is found in its greatest 
numbers in the swampy lowlands near the coast where small 
creeks spread out and intermix and where the swampy species of 
the Pandanus plant grows. The land is subject to occasional inun- 
dation during heavy rains, but there are always dry knolls onto 
which the reptiles and amphibians can retreat. The usual micro- 
habitat in the swampy lowlands is under and in rotten logs and 



1967 NEW SOLOMON ISLANDS SKINK 7 

thicker layers of decaying vegetable matter, while on higher ground 
and in the mountain ranges the species is usually found only living 
under and in rotten logs. 

The Shortland Island specimen was found under a pile of decay- 
ing fronds in a coconut plantation. 

Habits: S. tcmneri is a burrower and shows a preference for very 
damp country. Other than this, little is known about the 
species' habits. No individual was ever seen out in the open of its 
own accord. It escapes by burrowing when cover is removed and 
it is disturbed. 

Reproduction: S. tcmneri is oviparous and remarkably consistent 
in laying only 2 eggs at a time. Of the 49 females found to be 
gravid with oviducal eggs, 47 had snout-vent lengths ranging from 
41-50 mm (Table 1). Given the large total sample size (424 
specimens) from which these females were taken, the low end of 
the size distribution may represent the size at which females become 
capable of reproducing. 

Of the 49 gravid females, 47 contained 2 eggs; 46 of these held 
1 egg in each oviduct and a single specimen contained 2 eggs in 
the right oviduct alone. Of the 2 remaining females, one con- 
tained 3 eggs, 2 in the right oviduct and 1 in the left, and a second 
female contained only a single well-developed egg in the right ovi- 
duct. In all cases in which there were eggs in both oviducts (47), 
those eggs in the right oviduct were invariably situated more an- 
teriorly; this is apparently an accommodation for the stomach, 
which lies on the left side. 

The 49 gravid females were collected in March (2), May (20), 
June (5 ), July (21 ), and December ( I ). 

Eggs collected in the field on 17 May 1966 measured 14.8 x 8.0 
(with an embryo), 13.0 x 7.5, 13.4 x 7.0 mm (unopened). 

Morphological comparisons n-ith Bougainville relatives: The 
only other sympatric members of tanneri's species group (Table 
4) on Bougainville are Sphenomorphus solomonis and S. cranei. 
Both species can be easily distinguished on the basis of the char- 
acters listed in Table 2. 

Tanneri and solomonis are similar in their dorsal color patterns 
which consist of white flecking on a dark ground color. In tanneri, 
however, the ground color is a rich dark brown to brownish black 
while in solomonis the color of the dorsum ranges from brownish 
black to gray. The white spots along the dorsolateral line never 
coalesce to form a light line in solomonis as they do in tanneri. 

The ventral color patterns of tanneri and solomonis are quite 
distinctive. In tanneri the venter of the body is immaculate while 



8 BREVIORA No. 275 

the chin and throat are heavily pigmented. Solomonis, on the 
other hand, usually has the venter of the head and body uniformly 
spotted throughout, each ventral scale of the body containing a 
central dark spot. 

The dorsal ground color in cranci varies from very dark dull 
brown to blackish brown and is patterned with thin, whitish to 
yellow transverse bars from the nape of the neck onto the tail. 

In life the belly and underside of the tail in tanneri and cranei 
are often suffused with a translucent reddish color. In soloinonis 
the ventral surfaces lack any reddish coloration. 

Ecological comparisons with Bougainville relatives: The species 
most similar in ecology to tanneri on Bougainville is Sphenomor- 
phus solomonis, one of tanneri's closest relatives on the island. 
Both species, along with two species of Tribolonotus (blanchardi 
and an as yet unnamed species), are the most extreme burrowers 
in the Bougainville lizard fauna. Solomonis is much more common 
than tanneri and has a slightly greater altitudinal range — from the 
coast to at least 3000 feet (and perhaps up to 4000 feet) in the 
mountains. 

Solomonis occurs in a wider variety of habitats than tanneri, 
being most commonly found in the top few inches of soil and 
humus and under logs in open gardens, secondary scrub, tall pri- 
mary forest, dry coastal country and swampy areas. In the moun- 
tains, solomonis is much reduced in numbers as the altitude in- 
creases and is found mainly on the ridges and in secondary scrub. 
In the lowlands very large numbers of solomonis can be taken when 
logs and vegetation are being cleared off the ground for native 
gardens and when the topsoil is first worked. 

As fewer cranei have been collected than solomonis and tanneri, 
field observations for this species are less complete than for its two 
closest relatives on Bougainville. Cranei, however, also seems to 
be a burrower, although much less so than solomonis and tanneri. 
Unlike solomonis and tanneri, cranei has not been collected in the 
lowlands, but is known from the mountains and steep-sided valleys 
above 500 feet where there is tall, cool primary forest. Cranei 
is much more highly dependent on water and moisture than tanneri 
and solomonis and is most usually found under stones, wood, or 
debris in small creek beds with running water or soaked beds. In 
this habitat it is found in company with Tribolonotus blanchardi 
and Sphenomorphns (Parotosannis) concinnatns, but cranei is by 
far the least common of the three species. 



1967 NEW SOLOMON ISLANDS SKINK 9 

Tanneri and solonionis are both oviparous, but the mode of 
reproduction in cranei is unknown. As noted above, tanucri is re- 
markably consistent in producing only two eggs at a time. Soloni- 
onis, on the other hand, produces clutches with 3-6 eggs. 

Discussion: Current research (by Greer) on the delimitation and 
relationships of skink taxa indicates that there are several species 
groups now included in the genus Sphenomorphus. The two larg- 
est species groups of the genus are discussed below. 

One of these, which may be called the variegatus species group 
(Table 3), also includes skinks of the genera Otosaurus, Paroto- 
sauriis, and Insulasauriis and is characterized by the following suite 
of external characters: well developed digits and limbs, which 
overlap in most species when adpressed to the body; frontal gen- 
erally in contact with 3 or more of the anteriormost supraoculars; 
generally 5 or more supraoculars; supranasal scale (large in Ofo- 
sourus, small in Parotosauriis) or double anterior loreal present 
in most species, but some species with no supranasals and only a 
single anterior loreaP; nuchals lacking; high number of scales 
around midbody (usually 32 or more); dorsal scales subequal in 
size. 

The other large species group within Sphenomorphus, the one to 
which tanneri belongs and which may be called the fasciatus species 
group (Table 4), is characterized by the following external char- 
acters: digits and limbs usually less well developed, the limbs gen- 
erally not overlapping, or just slightly overlapping when adpressed 
to the body; frontal in contact with the anterior 2 supraoculars; 
4 (much less frequently 5) supraoculars; a single anterior loreal; 
no supranasals; usually a series of two or more symmetrical nuchal 
scales; fewer scales around midbody (generally 36 or fewer); 
scales of the 2 vertebral rows usually broader than the other dorsal 
scales, i.e., they are transversely enlarged. 

Whole skulls of several species of both species groups have also 
been examined. Both species groups display several important 
similarities in their skull morphology, but a discussion of these 



^ As the small supranasal of some Paiotosaunts and the double anterior 
loreal of some Sphenomorphus may be variable in shape and form, or 
even present on one side of the head and absent on the other in a single 
individual {Parotosauriis concinnaliis. personal observation and Burt and 
Burt, 1932:542), it does not seem justifiable to exclude from the variegatus 
species group those species which agree with the diagnosis except for lacking 
a supranasal or a double anterior loreal. 



10 



BREVIORA 



No. 275 



similarities is best deferred to a later date. The three important 
differences in skull morphology between the variei^afus and solo- 
monis groups are as follows: 



variegatus group 

1.) Postorbital bone lacking or 
very small. 

2. ) Supratemporal fenestra usu- 
ally lacking or very small. 



3.) 



No anteriorly projecting ec- 
topterygoid process to the 
palatine which would ex- 
clude the palatal ramus of 
the pterygoid from a posi- 
tion on the edge of the in- 
fraorbital vacuity. 



solomonis group 

1.) Postorbital bone usually 
long and thin. 

2.) Supratemporal fenestra 
generally well developed. 

3.) Two subgroups within the 
fascia t us group: solomonis 
subgroup (Greer, 1967) 
with ectopterygoid process 
to palatine which excludes 
all or most of pterygoid 
from a position on infra- 
orbital vacuity; fasciatus 
subgroup without anteriorly 
projecting ectopterygoid 
process to palatine.^ 



The two species groups may also be distinguished ecologically 
and geographically. The skinks of the variegatus group appear to 
be primarily surface dwellers, while those of the fasciatus group 
are, by and large, secretive burrowers as is tanneri. 

Geographically, the variegatus group ranges from southeast Asia 
and the Greater Sunda Islands northeast to the Philippines and 
east through Celebes and the Lesser Sunda Islands to New Guinea 
and the Solomons. The group is not found, however, in Australia. 
The center of abundance for the group appears to be the western 
part of the Indo-Australian archipelago and. perhaps to a lesser 
extent. New Guinea. 

The fasciatus group has its center of abundance on New Guinea, 
but extends northwest to the Philippines (fasciatus), west into the 
Lesser Sunda Islands (emigrans and antoniorum) , south into north- 
ern and eastern Australia (e.g., crassicauda, elegantuluin, punctu- 
latum and mjobergi), and east into the Solomon Islands (cranei, 
solomonis and tanneri). 



1 S. tanneri lacks the ectopterygoid process to the palatine and is therefore 
a member of the fasciatus subgroup of the fasciatus species group of 
SpiH'in'>inorphus. 



1967 NEW SOLOMON ISLANDS SKINK 11 

The two species groups considered here appear to be mono- 
phyletic. although convergent tendencies in both groups (e.g. the 
loss of nuchal scales, of transversely enlarged vertebral scales, and 
of the postorbital bone in the jasciaius group and the presence of 
only 4 supraoculars in the variegatiis group) make formal taxo- 
nomic treatment difficult. For this reason the ranking and further 
delimitation and subdivision of the species groups will be consid- 
ered at a later date. 

ACKNOWLEDGEMENTS 

Our thanks go first and foremost to Miss A. G. C. Grandison 
of the British Museum (Natural History) who made extensive and 
detailed comparisons between S. tanneri and several types in her 
charge. 

Dr. Ernest E. Williams of the Museum of Comparative Zoology 
has read the manuscript in several drafts and has offered many 
helpful criticisms. 

Mrs. Patricia H. Kerfoot made the line drawing for Figure 1 
and Mr. Rick Stafford took the photograph for Figure 3. 

Travel expenses incurred by Greer in the course of his reseaich 
for this paper have been defrayed by support from the Evolutionary 
Biology Fund which is administered for the National Science Foun- 
dation by the Evolutionary Biology Committee of the Biological 
Laboratories. Harvard University. 

LITERATURE CITED 

Brongersma, L. D. 

1942. On the arrangement of the scales on the dorsal surface of the 
digits in Lygosonui and allied genera. Zool. Meded. Leiden, 
24 (1-2): 153-158. 
Burt, C. E. and M. D. Burt 

1932. Herpetological results of the Whitney South Sea Expedition. VL 
Bull. Amer. Mus. Nat. Hist.. 63 (5):461-597. 
Greer, A. E. 

1967. A new generic arrangement for some Australian scincid lizards. 
Breviora, No. 267:1-19. 
RooiJ. N. DE 

1915. The reptiles of the Indo-Australian archipelago. L Lacertilia, 
Chelonia, Emydosauria. Leiden, E.J. Brill Ltd., 384 pp. 
Smith, M. A. 

1927. Contributions to the herpetology of the Indo-Australian region. 
Proc. Zool. Soc. London, 1927:199-225. 



(Received 23 March 1967.) 



12 BREVIORA No. 275 

TABLE 1 

Frequency distribution for several nieristic characters in Spheno- 
morphus tarmeri. N = number of specimens examined. 

Midbody Scales (N — 156) 

X 28 29 30 31 32 

f(x) 5 5 115 11 20 

Supralabial below Center of Eye (N = 415) 

X 5-5 4-5 4-4 

f(x) 337 43 35 

Number of Lamellae beneath 4th Toe (N = 228) 



X 


13 


14 


15 


16 


17 


18 


19 


20 


f(x) 


2 


3 


20 


62 


90 


32 


17 


2 



Snout-Vent Length of Gravid Females (N = 47) 

x 41 42 43 44 45 46 47 48 49 50 

f(x) I 3 5 8 7 6 7 5 3 2 



TABLE 2 



Morphological differences between S. tanneri and its two closest 
relatives on Bougainville. 



tcinnevi 


solomonis 




cranei 


Snout-vent length 23-52 mm 


27-72 mm 




29-1 \ mm 


Scales around 28-32 


24-30 




32-41 


mid-body (usually 30) 


(usually 2( 


6-30) 


(usually 34, 36) 



Prefrontals meet Yes, except in 1 No Yes, except in 2 

of 424 specimens of 23 specimens 

Supralabial below 4th (137f) or 4th (< 1/2%) or 5th {W/c ) or 

center of eye 5th{879r) 5th (> 991/2%) 6th (897r ) 

Nuchal scales Absent Present, 2-4 pairs Present, 3-5 pairs 

Subdigital lamellae 13-20, 13-17 20-29, most 

(4th toe) usually 15-19 usually 22-28 



967 



NEW SOLOMON ISLANDS SKINK 



13 




Figure 3. Holotype of Sphcnomorpluis tanncri (MCZ 76551). showing 
the light spotting on the dark brown dorsum. 



14 



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No. 275 



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No. 275 



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^•~-5-SS-"^ ~5=:5 5T^ ~$S 



BREVIORA 



Miasemim of Coimparsitive Zoology 

Cambridge, Mass. 24 November, 1967 Number 276 

THE GENERIC RELATIONSHIPS OF THE AFRICAN 
SCINCID GENUS EUMECIA 

Allen E. Greer 



In 1870 Bocage described a new skink (anchietae) from Africa 
that was peculiar in having an elongate body form, small ap- 
pendages with a reduced number of digits (2 fingers and 3 toes), 
a pair of supranasals meeting behind the rostral and a spectacle 
in the movable lower eyelid. Bocage placed the new species in a 
distinct genus which he named Eumecia, apparently to emphasize 
a similarity which he believed to exist with Eumeces. 

The three subsequent revisionary studies on scincid genera 
rightly recognized the closer afiinity of Eumecia with what have 
come to be called the lygosomine skinks rather than with Eumeces 
or its subfamily.^ 

Boulenger (1887) placed anchietae in the Riopa section of the 
genus Lygosoma, and in 1897 described a species (johnstoni) 
from Nyasaland that was distinguishable from anchietae primarily 
on the basis of a further reduction in the number of digits (1 
finger and 2 toes in johnstoni; 2 fingers and 3 toes in anchietae) . 
Like anchietae, johnstoni was placed in the section Riopa of the 
genus Lygosoma. 

Smith (1937) revived Bocage's name Eumecia for a subgenus 
of the genus Riopa and included in it only anchietae and johnstoni. 
Mittleman (1952) agreed with Smith's conception of the taxon 
Eumecia but gave it full generic rank in his classification. 

Loveridge (1953 and 1957) placed both anchietae and johnstoni 
in the genus Riopa without recognizing subgenera, thus adopting 



^ The subfamily Lygosominae is characterized by a single frontal bone, and 
palatine bones which meet along the midline of the palate to form a 
secondary palate. Eumeces is considered to be a scincine, a subfamily 
characterized, in part, by a divided frontal bone, and palatines which do 
not meet along the ventral midline of the palate (Greer, MS). 



2 BREVIORA No. 276 

a basically Boulengerian conception of the species' taxonomic po- 
sition. The same author (1953) pointed out the very real pos- 
sibility that jolinstoni was simply a race of ancluetae and further 
noted that some anchietae had three fingers instead of two. 

That anchietae and jolinstoni should be accorded separate supra- 
specific rank as recognized by Bocage (1870), Smith (1937). and 
Mittleman (1952) is suggested by the fact that they are the only 
African lygosomine skinks with supranasals to have fewer than 5 
fingers and 5 toes. Indeed it was on this criterion that Eumecia 
has been regarded as distinct from Riopa and its supposed relatives. 

In this paper, the two closely related species anchietae and 
johnstoni^ are considered to constitute a distinct genus Eumecia, 
for reasons that will become apparent in the following discussion, 
and the genus is shown to be most closely related to Mabuya and 
not Riopa. For the purposes of this discussion Riopa is understood 
in the sense of Smith's (1937) subgenus Riopa and Mittleman's 
(1952) genera Riopa, Squamicilia and Mochliis collectively. It 
is not clear to me why Eumecia was always thought by Boulenger 
and later authors to be more closely related to Riopa and its sup- 
posed relatives than to Mabuya. On the basis of externals there 
is no one character that will serve to distinguish all Riopa from all 
Mabuya. There is however one external character that will dis- 
tinguish some Mabuya from all Riopa, namely the relative posi- 
tion of the prefrontal scales, and in this regard Eumecia is like 
Mabuya rather than Riopa. The prefrontals are never in contact 
in Riopa, but they do meet medially in Eumecia and in about 
one-third of the species of Mabuya (surveyed from Boulenger, 
1887). 

Comparisons of the skulls of 30 species of Mabuya and 13 spe- 
cies of Riopa indicate that there are important differences in skull 
morphology between Mabuya and Riopa, and on the basis of 
skull characters Eumecia is clearly more closely related to Mabuya 
than to Riopa (Table 1). 

The single greatest difference between Mabuya and Riopa is in 
the relationships of the bones of the palate (Fig. 1). In both 



^ I have seen neither a skull nor an alcoholic specimen of jolinstoni, which 
is still known only from the type. I am assuming throughout this paper 
that the two species are so similar (conspecific?) that, unless stated other- 
wise, observations made on the two skulls and alcoholic specimens of 
anchietae are also valid for jolinstoni, and therefore for the whole genus 
Eumecia. 



1967 



GENERIC RELATIONSHIPS OF EUMECIA 




Figure 1. Ventral view of the secondary palate of: A) Eumecia anchie- 
tae (MCZ 41562), Kaimosi, Kakamega, Kenya: B) Mahuya polytropis 
(MCZ 8103), Krilii Cameroon; C) Riopa punctata (MCZ 3238). 70 miles 
SW of Amballa, India: D) Leptosiaphos hlochmanni (untagged MCZ 
specimen). Upper Mulinga, Idjwi, Id., Congo. Abbreviations: E, 
ectopterygoid; P, palatine; PT, pterygoid. A and B drawn to one scale and 
C and D drawn to another scale. 



4 BREVIORA No. 276 

Mabuya and Riopa the palatine bones meet along the ventral 
midline of the palate to form a secondary palate separating the 
air (above) and food (below) passages, a structure that is char- 
acteristic of the subfamily Lygosominae (Greer, MS). In Riopa 
the development of a secondary palate is more extensive than in 
Mabuya in that the palatal rami of the pterygoids approach but 
do not touch along the midline of the palate. The pterygoids are 
separated from each other by a pair of medial, posteriorly project- 
ing processes from the palatines which are closely applied to the 
inner edges of the palatal rami and which themselves touch along 
the midline to close the gap between the pterygoids. 

In Mabuya, on the other hand, and in Eumecia, the palatal rami 
of the pterygoids are widely separated and divergent, and there are 
no posteriorly projecting processes from the palatines (Fig. 1). 

In Eumecia there is a broad suture between the prefrontal and 
nasal bones which thus separates the frontal from the maxilla. In 
12 of the 23 species of Mabuya examined for this feature, the pre- 
frontal articulates with the nasal, but in Riopa only 2 of the 12 
species checked show a similar relationship of the prefrontal and 
nasal. 

Similarly. Eumecia shows a short longitudinal series of pterygoid 
teeth, a feature shared with 17 of the 30 species of Mabuya ex- 
amined for this character. However, Riopa bowringi was the only 
one of the 1 3 species of Riopa examined which was found to have 
pterygoid teeth. 

Another somewhat statistical difference that aligns Eumecia with 
Mabuya is the number of teeth on the maxilla. The two specimens 
of Eumecia anchietae examined had 24-26 teeth on the maxilla. 
Only one species of Mabuya (occidenfalis) of the 29 examined 
had fewer than 20 maxillary teeth; all others had 20 or more. Of 
the 13 species of Riopa examined for this character, however, 
only 4 species had 20 or more teeth on the maxilla. 

The one significant similarity between Riopa and Eumecia in the 
gross morphology of the skull is in the common absence of the 
postorbital bone. Eumecia anchietae and the 13 species oi Riopa 
examined lacked this bone while in all 29 species of Mabuya 
studied, a small, but discrete, postorbital bone was present. On 
the weight of the total evidence presented here, however, it would 
seem as if the postorbital of Eumecia had been lost independently 
of its loss in Riopa. 

Although it seems established, on the basis of the evidence dis- 
cussed above, that the relationships of Eumecia lie closer to Ma- 
buya than to Riopa, it is still reasonable to ask if there might not 



1967 GENERIC RELATIONSHIPS OF EUMECIA 5 

be a group even more closely related to Eumecia than is Mabuya. 

In short, this does not seem to be the case. In Africa the only 
lygosomine skink which has the palatal rami of the pterygoids 
widely divergent and which lacks medial palatine processes pro- 
jecting posteriorly as in Mabuya and Eumecia is Ablepharus bou- 
toni, a relatively recent immigrant from the Australian Region. 
This species, however, lacks pterygoid teeth and supranasals which 
make it an unlikely representative of a stock ancestral to Eumecia. 

In all African lygosomines other than Mabuxa, Eumecia and 
Ablepharus, the palatal rami of the pterygoids are more closely 
apposed along the ventral midline of the palate, and medial pala- 
tine processes project posteriorly between the pterygoids. Indeed, 
in some African lygosomine groups with this basically Riopa-VikQ 
palate (African Ablepharus, with the exception of A. boutoni; 
African Leiolopisma; Panapsis; Leptosiaphos). a further special- 
ization has been the deep posterior emargination of the palatal 
rami of the pterygoids. 

The only other groups of skinks with both supranasals and a 
palatal pattern generally similar to that of Mabuya and Eumecia 
are the genera Emoia and Eugongylus. Although these two genera 
are closely related^ and are very close relatives of Mabuya, it seems 
less likely that Eumecia has arisen from an Emoia-Eugongylus 
stock than from a Mabuya ancestry. 

Eumecia has pterygoid teeth and a broad surface suture be- 
tween the prefrontal and nasal bones while none of the 20 species 
of Emoia or 2 species of Eugongylus examined had pterygoid 
teeth or a broad prefrontal-nasal suture. Many Mabuya, on the 
other hand, have pterygoid teeth { 17 of 30 species examined) and 
a broad suture between the nasal and prefrontal (12 of 23 species 
examined). 

Eumecia also has only 9 teeth on the premaxillae whereas all 
the Emoia and Eugongylus studied have 11 or more (12) pre- 
maxillary teeth. However, only a few Mabuya (6 of 30 species 

^ The species of Eugongylus have also been treated as relatives of Riopa 
in the three most recent classifications dealing with lygosomine skinks. 
BoLilenger (1887) referred the species of Eugongylus to the section Riopa 
of the genus Lygosonia. Smith (1937) recognized the taxon as a subgenus 
of Riopa, and Mittleman (1952) gave the taxon full generic rank. Again, 
the reasons for supposing that the relationships of Eugongylus lay in the 
direction of Riopa are as unclear to me as those for aligning Eumecia with 
Riopa. On the basis of skull morphology, partly discussed here, Eugongylus 
is strikingly similar to Emoia. This similarity and its taxonomic sig- 
nificance will be discussed elsewhere. 



6 BREVIORA No. 276 

available) have 10 or more (11-12) teeth on the premaxillae, all 
others have 9 (or less frequently, 8). 

Furthermore, Eumecia is live-bearing in its mode of reproduc- 
tion, as are about half the species of Mabuya (10 of 21 species 
for which information is available). In contrast, the 17 species of 
Emoia and 2 species of Eugongylus for which the mode of repro- 
duction is known are all egg laying (Greer, personal observation). 

Emoia and Eugongylus do lack the postorbital bone, as does 
Eumecia, and although Mabuya invariably has a small postorbital, 
it seems most likely, on the weight of other evidence presented 
above, that Eumecia has lost the bone independently of its loss in 
Emoia and Eugongylus. 

The present distribution of Mabuya and Eumecia certainly sup- 
ports the derivation of Eumecia from a Mabuya ancestry. Mabuya 
has obviously been in Africa a long time — long enough to have 
evolved numerous and diverse species. Part of this diversity is 
manifest in the evolution of Eumecia — a "Mabuya" with reduced 
appendages. In that it is the only "Mabuya" to have lost the post- 
orbital bone, and to have reduced the number of fingers and toes 
from the primitive number of 5-5, the taxon clearly deserves 
generic rank. 

ACKNOWLEDGEMENTS 

My thanks go first and foremost to Dr. Ernest E. Williams of 
the Museum of Comparative Zoology (MCZ) who has been most 
generous in his policies regarding the preparation of skulls from 
alcoholic specimens in the herpetological collections of the 
Museum. 

The following people have also contributed greatly to the present 
study by making available to me skulls of species not represented, 
or only poorly represented in the Museum of Comparative Zoology 
collections: Dr. Charles M. Bogert and Dr. Richard G. Zweifel, 
American Museum of Natural History (AMNH); Dr. Alan E. 
Leviton and Dr. Steve C. Anderson, California Academy of 
Sciences (CAS); Dr. Robert F. Inger and Mr. Hymen Marx, Field 
Museum of Natural History (FMNH); Dr. Charles F. Walker, 
University of Michigan Museum of Zoology (UMMZ). 

This study was supported in part by a grant from the Evolu- 
tionary Biology Fund which is administered for the National Sci- 
ence Foundation by the Evolutionary Biology Committee of the 
Biological Laboratories, Harvard University. 



1967 GENERIC RELATIONSHIPS OF EUMECIA 7 

MATERIALS 

Complete skulls of the following species have been examined. 

RIOPA: ajer (MCZ 41517, 41519, 71881); bowringi (1 from 
the series MCZ 26501, 26512); femaudi (MCZ 49696); laeviceps 
(MCZ 71889); lineata (AMNH 46379); mabiiiifonnis (MCZ 
40267); pembana (MCZ 46106); popae (MCZ 44706); punctata 
(MCZ 3238); sunder alii (MCZ 41537, 41543); tanae (MCZ 
40256); vinciguerrae (MCZ 17892); albopunctata (MCZ 8360). 

EMOIA: adspersa (AMNH 29227); atrocostata (MCZ 15074, 
15080, 26476, 26419); boettgeh (MCZ 22014); callisticta (MCZ 
67203, 612>0^);cyanogaster (CAS 100684, MCZ 15121, 15135, 
72278, 72287); cyanura (MCZ 14582, 14584, 14586, 75954, 
75956); flavigularis (MCZ 65869); kordoana (MCZ 48603); 
kuekenthali (FMNH 134594); loveridgei (MCZ 49321); macu- 
lata (MCZ 49501, 49505 lot); mivarti (MCZ 73807, 75984); 
nigra (MCZ 15153, 15157,67770, 72510, 72514, 72515. 72517, 
72523, 75522); paUidiceps (MCZ 79856); physicae (AMNH 
95772), ruficauda (MCZ 26482, 2 specimens, 26492); sanjordi 
(AMNH 40169); submetallica (AMNH 59015); sorex (MCZ 
1105); samoensis (MCZ 16931). 

EUGONGYLUS: albojasciolatus (MCZ 4097, 72703); rufes- 
cens (MCZ 49341). 

EUMECIA: anchietae (MCZ 41557, 41562). 

MABVYA: aurata (MCZ 56550); bayoni (MCZ 39731); ben- 
soni (MCZ 22583); binotata (MCZ 22421); blandingi (MCZ 
55171); brachypoda (MCZ 71410); brevicollis (MCZ 41306); 
capensis (MCZ 21433); comorensis (MCZ 24151, 2 specimens, 
24155); dorsovittata (MCZ untagged specimen); elegans (MCZ 
67954) ; ^'/^g/t'/ (MCZ untagged specimen) ; fasciata (MCZ 37835+ 
2 untagged specimens); gravenhorsti (MCZ 11609); hildebrandtl 
(MCZ 70254, 70248); lacertijormis (MCZ untagged specimen); 
longicaudata (MCZ 25191); m«/>i/vfl (CAS 71456^ UMMZ 1047, 
MCZ 32040, 36617, 38935, 54201. 81182, 81184); macrorhyn- 
cha (MCZ 49551, 49552); maculaha (MCZ 3926); maculilabris 
(MCZ 24820. 24821); megalura (MCZ 47611); multifasciata 
(CAS 60692 + 2 untagged specimens, UMMZ S 1830, 1831, 
MCZ 25198, 25199, 37843); occidentalis (MCZ 43180); per- 
rcteti (MCZ 19711); planijrons (MCZ 85545); /^o/y^ro/^/^ (MCZ 
8103, 54559); qidnquetaeniata (MCZ 52424, 2 specimens, 55179, 
67838, 67840); sulcata (MCZ 21645); vw/fl (MCZ 18658, 
18668, 50823, 50824, 85543). 



8 BREVIORA No. 276 

LITERATURE CITED 

BOCAGE, J.V.B. DU 

1870. Description d'un "Saurien" nouveau de TAfrique occidentale. 
J. Sci. Lisboa, 3:66-68. 

BOULENGER, G. A. 

1887. Catalogue of the lizards in the British Museum (Natural 
History). London, Vol. Ill, xxi + 575 pp. 

1897. List of the reptiles and batrachians collected in northern Nyasa- 
land by Mr. Alex Whyte, F.Z.S., and presented to the British 
Museum by Sir Harry H. Johnston, K.C.B.; with descriptions 
of new species. Proc. Zool. Soc. London 1897: 800-803. 

Greer, A. E. 

ms A subfamilial classification of scincid lizards. 

LOVERIDGE, A. 

1953. Zoological results of a fifth expedition to East Africa. 111. 

Reptiles from Nyasaland and Tete. Bull. Mus. Comp. Zool., 

110(3): 143-322. 
1957. Check list of the reptiles and amphibians of East Africa 

(Uganda; Kenya; Tanganyika; Zanzibar). Bull. Mus. Comp. 

Zool., 117(2): 153-362 + xxxvi. 

MiTTLEMAN, M. B. 

1952. A generic synopsis of the lizards of the subfamily Lygosominae. 
Smithsonian Misc. Coll.. 117(17): 1-35. 

Smith, M. A. 

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

(Received 14 April 1967.) 



1967 



GENERIC RELATIONSHIPS OF EUMECIA 



TABLE 1 



Comparison of certain external scale (1) and skull (2-6) characters 
in the genera Mabuya, Eumecia and Riopa (scnsu Smith's (1937) sub- 
genus Riopa and Mittleman's (1952) genera Riopa, SquamiciUa and 
Mochhis collectively). 



Mabuya 

1. Prefrontal scales meet 
medially in about Vs of 
the species. 

2. Palatal rami of ptery- 
goids separated along 
midline of palate; no 
posteriorly projecting 
medial processes from 
the palatines ( Fig. 1). 

3. 12 of 23 species examined 
with a surface suture be- 
tween prefrontal and 
nasal bones to separate 
frontal and maxilla. 

4. 17 of 30 species examined 
have pterygoid teeth. 

5. 28 of 29 species examined 
have 20 or more teeth 
on the maxilla. 

6. Small to minute postor- 
bital bone present. 



Eumecia 

Prefrontals meet 
medially. 

As in Mabuya 
(Fig. 1). 



Surface suture be- 
tween prefrontal and 
nasal bones to sepa- 
rate frontal and 
maxilla. 

Pterygoid teeth 
present. 

24-26 teeth on maxilla. 



Postorbital bone lack- 
ing. 



Riopa 

Prefrontals always sepa- 
rated. 



Palatal rami of pterygoids 
separated along midline 
of palate by posteriorly 
projecting medial pro- 
cesses from the palatines 
(Fig. 1). 

Only 2 of 13 species ex- 
amined with a surface 
suture between prefrontal 
and nasal bones to sepa- 
rate frontal and maxilla. 

Only 1 {R. bowringi) 
species of 13 examined 
has pterygoid teeth. 

Only 4 of 13 species ex- 
amined have 20 or more 
teeth on maxilla. 



Postorbital bone lacking. 



BREVIORA 

MiLHseuiitM of Comparative Zoology 

Cambridge, Mass. 24 November, 1967 Number 277 



LESTIDIUM BIGELOWh A NEW SPECIES OF 
PARALEPIDID FISH WITH PHOTOPHORES 

Michael J. F. Graae^ 



ABSTRACT 

Two specimens of a new luminous paralepidid fish were taken 
in the Indian Ocean during Cruise VI of the R/V Anton Bruiin. 
Their main distinguishing feature is the presence of three separate 
and distinct, circular photophores along the ventral midline that 
are unlike the light organs found in other paralepidids. The two 
specimens are described here as a new species, and the structure 
of one photophore is examined in detail. 

INTRODUCTION 

Two evolutionary lineages, the myctophoids and the alepisau- 
roids, are generally recognized within the pelagic members of the 
Myctophiformes (Iniomi). The myctophoid families Myctophidae 
and Neoscopelidae have discrete photophores lying in rows on 
each side of the ventral midline, and it has been shown recently 
(Haneda, 1958; Rofen, 1965) that a few members of the alepi- 
sauroid family Paralepididae have elongated luminous glands im- 
mediately on, or adjacent to, the ventral midline. One genus also 
has a small finger-like projection, before the eye, that may be 
luminous, but this organ does not appear to have the structure 
found in the myctophoids. This paper is concerned with a unique 
new species of paralepidid which, though quite similar to other 
paralepidids in most characters, is distinguished from them by the 
discrete photophores on the ventral midline. 



1 1702 San Andres St., Santa Barbara. California. 



2 BREVIORA No. 277 

I am indebted to the National Science Foundation for its or- 
ganization of the American Program in Biology, International 
Indian Ocean Expedition, and to Dr. John H. Ryther of the 
Woods Hole Oceanographic Institution, the Scientific Director of 
the program. I also gratefully acknowledge the support of NSF 
Grant (jF-147 to Harvard University, which aided this study. I 
thank Dr. Giles W. Mead, Chief Scientist during Cruise VI of the 
Bruun, for his counsel during the course of this study and for his 
review of the manuscript. I am especially indebted to William 
O'Day, who so kindly and patiently prepared, sectioned, and 
photographed the photophore described herein. I also wish to 
extend my appreciation to those who have helped me with advice 
and encouragement, especially Dr. Basil Nafpaktitis and Mrs. 
Myvanwy M. Dick. This work is part of a Harvard senior honors 
dissertation presented in April of 1966. 

Lestidium bigelowi, new species 

Holotype: MCZ 44881, 37.2 mm in standard length; collected 
during the International Indian Ocean Expedition, R/V Anton 
Bruun, Cruise VI, trawl 339A; APB label 7227; hydrographic 
station 339; collected in 10' Isaacs-Kidd Midwater Trawl, depth 
0-61 5m; 30-V-64. 0140-0645 hrs; 4"01'-4° 14'S, 65°00'-65°02'E. 

Paratype: MC/ 44880, 28.4mm in standard length; R/V 
Anton Bruun, Cruise VI; trawl 340B; APB label 7241; hydro- 
graphic station 340; 10' IKMT, 0-746m; 31-V-64, 1945-0155 hrs; 
5°55'-6'^08'S, 64°48'-64"^58'E. 

Diagnosis: This species is the only paralepidid known to have 
light organs that are small and discrete, as opposed to long and 
tubular. There are three of these; all are directed ventrally. One 
is at the isthmus, one is between the bases of the pectoral fins, and 
one is just posterior to the bases of the ventral fins. In most 
characters, Lestidium bigelowi is almost intermediate between the 
genera Lestidium and Lestidiops (see Table 1). 

Description: The following description is based on both types. 
Body long and slender, very compressed. Keel present for short 
distance ventrally between anus and anal fin. Anus between tips 
of appressed ventral fins. Skin smooth; scales absent, with lateral 
line ossicles insufficiently ossified to be observed. Lateral line ex- 
tending along side of body from edge of operculum lo a vertical 
with first of procurrent caudal rays, and slightly above middle of 
body. 



1967 



NEW PARALEPIDID FISH 





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4 BREVIORA No. 277 

Three luminous organs on the midventral hne — one at isthmus, 
another between bases of the pectoral fins, and the third between 
bases of ventral fins and anus. Isthmal and pectoral organs im- 
bedded in flesh; anal organ close to skin, possibly open to surface 
or connected to rectum. In alcohol, these organs are black 
with a translucent, white layer at center (see discussion of luminous 
organs, below). 

Head pointed and moderately long, widest just posterior to eyes. 
Snout length slightly more than two in head, about three-quarters 
as wide as head. Eye and pupil large, about five in head length; 
eye and pupil round; adipose eyelid present. Interorbital slightly 
concave, with two lateral, compressed, longitudinal ridges on each 
side. Occiput rounded, convex. Nasal apertures approximately 
halfway between tip of snout and eye. Opercle with posterior edge 
moderately pointed and notched about pectoral fin base. Mouth 
large, lower jaw with a slight non-ossified projection. Premaxil- 
lary extending to a point beneath anterior edge of orbit. Premaxil- 
lary with two or three depressible canines, posteriorly followed by 
fixed teeth. Palatine with a few depressible teeth. Gill rakers 
present but poorly developed on first two arches. 

Dorsal fin origin slightly anterior to origins of ventral fins, a 
little more than halfway back on body. Posterior edge of adipose 
fin anterior to caudal peduncle, slightly before last anal fin ray. 
Inner rays of ventral fins distinctly longer than the outer. Ossifica- 
tion is too incomplete to permit radiographic study of vertebral 
column or study of lateral line ossicles through staining. 

In ethyl alcohol the body is largely colorless. Melanophores, 
most of them stellate, cover many parts of the fish: the occiput, 
the edges, ridges and sutures of the jaws, the angular bone, the 
dorsum, the posterior part of the caudal peduncle, the ventral 
midline between the anus and the anal fin, the first pectoral and 
anal fin rays, and the anal and caudal fin ray bases. The angular 
bone is also covered by heavy black pigment, and the dorsum is 
covered by light brown pigment in a band becoming denser and 
darker posteriorly. Three anterior and one posterior peritoneal 
sections are relatively clearly outlined; the others are indistinct. 

All measurements and counts are given in Table 1, and recon- 
structed lateral and ventral views of the holotype are shown in 
Figure 1. 

This new species is named in honor of Dr. Henry Bryant Bige- 
low, to whom I wish to express my sincere thanks for his kind- 
ness and generosity. 

Distribution: Taken only in the Indian Ocean, approximately 
650 miles northeast of Madagascar at a depth above 750m. 



1967 



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6 BREVIORA No. 277 

Relationships: In relative proportions and numbers of fin rays, 
Lestidium bigclowi closely resembles Lestidiuin atkmticiim Boro- 
din, Lestidiops jayakari (Boulenger), and Lestidiops indopacifi- 
cum (Ege). The ranges of meristic characters shown by the species 
in these two genera provide no grounds for separation. Analysis 
is further impaired by discrepancies evident in the monograph re- 
cently published by Rofen (1965), for the proportionality data 
provided in certain of his tables (e.g., table XXIV, pp. 302-303) 
are at variance with those reported in his description (e.g. that of 
Lestidium atlonticiun on pp. 308-309). The single valid and con- 
servative character which does separate Lestidium from Lesti- 
diops is the presence or absence of an internal light organ; hence 
the species described here is relegated to Lestidium. The meristic 
data taken from this species are compared in Table 1 with those for 
Lestidium atlanticum and Lestidiops jayakari taken from Rofen's 
description (1965: 308. 348). and for Lestidiops indopacificum 
taken from Ege ( 1953 ). 

Anatomy of the luminous organs: The three organs differ some- 
what from one another in gross morphology. The anterior one 
(the isthmal) is buried in the muscle of the isthmus (Fig. lA, B). 
It has a black cup-shaped structure with the opening directed 



A 

O 




B 




liM^i: 



Figure 2. Photophores of holotype: A and B, lateral and ventral views 
of isthmal photophore: C and D, lateral and ventral views of anal photo- 
phore; E and F, lateral and ventral views of pectoral photophore. 



1967 NEW PARALEPIDID FISH 7 

posteroventrally and is covered by a mass of translucent tissue 
(Fig. 2 A, B). The middle one (the pectoral) is nearer the surface 
than the isthmal (Fig. JA). It too has a black, ventrally directed 
cup, covered with a translucent cap; its black pigment seems to be 
associated with the peritoneal lining (Figs. lA, B; 2E. F). The 
posterior one (the anal) is close to, or partially embedded in, the 
skin (Fig. lA, B). As seen from the ventral side, it has an arrow- 
head-shaped mass of black tissue surrounding a central translucent 
cap (Figs. IB; 2C. D). There may be an opening directed pos- 
teriorly toward the anus or the surface from the central part of 
this photophore (Fig. 2D). 

The isthmal photophore of the paratype was removed, and 
sectioned by William O'Day, using the following procedure: the 
fish was fixed whole in formalin aboard ship, and then transferred 
to 12^/( ethanol for preservation. The photophore was excised 
and dehydrated for twenty-four hour periods in 100';^ ethanol, 
100% n-propanol, and 100% n-butanol. It was then infiltrated in 
one cc of pure glycol methacrylate with a polymerization catalyst, 
and transferred to fresh monomer and polymerized in a gelatin 
capsule at bO'C. After polymerization, the sections were cut 
vertically through the photophore at one micron thickness by a 
Porter-Blum microtome. The sections were stained first with 4% 
acid fuchsin in water for four minutes, and then with a concen- 
trated solution of toluidine blue in water for five minutes. The 
sections were mounted on a slide and covered with a Permount 
cover slip (see Ashley and Feder, 1966). 

The isthmal light organ (Fig. 3) is a classical example of a 
photophore (e.g. see Brauer, 1908, fig. XXX). It consists of a 
cup of heavy black pigment surrounding an internal body of 
granular tissue. On examination under high magnification the 
latter material does not seem to be associated with bacteria, im- 
plying that the luminescence of at least this organ is glandular in 
nature. The cup is closed ventrally by a lens-like layer of cells 
which separates the luminous tissue from a gelatinous cap. This 
cap completely covers the ventral aspect of the photophore and 
may serve to filter, focus, or disperse the light emitted. 

The structures of the light organ resemble those found in other 
fishes, especially the myctophoid inioms. They do not appear to 
be similar to those described by Haneda (1958) for Lestidiiim 
prolixum Harry or Lestrolepis japonicum (Tanaka). They do, 
however, look like those found in the family Myctophidae except 
for the presence of an internal lens. However, since the Para- 
lepididae are so dissimilar to the Myctophidae in other re- 
spects, the similarity of the photophore of L. bigelowi to those of 



BREVIORA 



No. 277 




Figure 3. A. a vertical section through isthmal photophore. B, the 
photophore above, enlarged. 



1967 NEW PARALEPIDID FISH 9 

the myctophids is assumed to be due to convergence rather than 
to close relationship. 

The anal photophore of Lcstidium bigeknvi is similar in shape 
to that discussed by Bertelsen, Theisen. and Munk ( 1965) for the 
argentinoid fish, Rhynchohyalus natalensis (Gilchrist and von 
Bonde). Like that of R. natalensis, the anal photophore of L. 
bigekm'i may be connected to the anus (though posteriorly in- 
stead of anteriorly), and its luminescence may thereby also be 
bacterial in nature. As the anal photophore on the holotype was 
the only one found, the paratype being mutilated in that region, 
no sections were made of it; thus the question of whether this 
luminescence is bacterial or glandular can not now be answered. 

SUMMARY 

A new species of Lestidiiim was collected in the Indian Ocean 
in 1964. It most nearly resembles Lestidium atlanticum Borodin 
in proportions and counts, and in some other respects is similar 
to Lestidiops jayakari (Boulenger) and L. indopacificuni (Ege). 
It differs from all other paralepidids in possessing discrete photo- 
phores. These photophores have a darkly pigmented cap, with 
opening directed ventrally, and are covered by a gelatinous, 
translucent mass of tissue. Photomicrographs of the isthmal photo- 
phore of the paratype show that it is very similar to those of the 
Myctophidae, differing significantly only in the possession of an 
internal lens. This similarity is attributed to convergence rather 
than to close relationship. 

LITERATURE CITED 

Ashley, C. A. and N. Feder 

1966. Glycol methacrylate in histopathology. Arch. Path. 81:391-397. 
Bertelsen, E.. B. Theisen, and O. Munk 

1965. On a postlarval specimen, anal light organ, and tubular eyes of 
the argentinoid fish Rliyiichohyciliis natalensis (Gilchrist and 
von Bonde). Vidensk. Medd. fra Dansk Naturh. Foren.. 
128: 357-371. 
Brauer, a. 

1908. Die Tiefsee-Fische II. Anatomischer Tell. Wiss. Ergebn. 
Deutsch. Tiefsee Exped. auf dem Dampfer Valdivia 1898-99, 
15(2); 1-153, pis. 11-30. 
Ege. V. 

1953. Paralepididae 1 (Paralepis and Lestidium) : taxonomy, onto- 
geny, phylogeny, and distribution. Dana Rep., 40: 1-184. 



10 BREVIORA No. 277 

Haneda, Y 

1958. Preliminary report on a luminous fish of the family Para- 
lepididae. Sci. Rep. Yokosuka City Mus. 3: 31-35. 
HUBBS, C. L. AND K. F. Lagler 

1947. Fishes of the Great Lakes region. Cranbrook Inst. Sci., Bull. 
No. 26: 1-186. 

ROFEN, R. R. 

1965. Paralepididae. Pp. 205-443, in Fishes of the Western North 
Atlantic (G. W. Mead, ed.), Mem. Sears. Found. Mar. Res., 
Yale Univ., No. 1, Pt. 5, 647 pp. 

(Received 3 May 1967.) 



BREVIORA 

MiiseiLim of Compsirsitive Zoology 



Cambridge, Mass. 24 November, 1967 Number 278 

NOTES ON THE EAABERiZINE SPARROW 
RHYNCHOSPIZA STOLZMANNI 

Raymond A. Paynter, Jr. 

Rhynchospiza stolzmanni Taczanowski has one of the most re- 
stricted ranges of any of the continental Emberizinae. It occurs 
on the arid western slopes of the Andes in Loja. southwestern 
Ecuador, and from Tumbes to Cajamarca, northern Peru, or from 
approximately Lat. 3° 30'S to Lat. 7° 15'S. Its altitudinal range is 
roughly from 100 to 1,100 meters. Judging from the comparatively 
few specimens in collections, the species appears to be uncommon, 
but this may be a deceptive indicator. It seems more probable that 
it has a restricted habitat and is secretive, two characteristics which 
would make it difficult to collect. 

FIELD NOTES 

In mid-October 1965 I saw the species briefly at Yamana (alt. 
ca. 1,100 m), near the head of the arid Casanga Valley in south- 
western Loja. Ecuador. Although these observations are only 
an introduction to the species, they supplement the short notes 
made by Jeiski and Stolzmann (Taczanowski, 1886) nearly a cen- 
tury ago, which are the only published field observations of this 
interesting bird. 

On several days while walking along a path on a steep hillside, 
my attention was occasionally attracted by the short loud calls of 
two or three birds concealed in sparsely distributed patches of thick 
brush on the lower side of the hill. A few times I was able to 
glimpse a dark, heavy-set bunting on the ground or in the lowest 
branches of the shrubs. The birds were extremely shy and silently 
moved downhill almost as soon as I sighted them. Late one morn- 
ing a flock of about six birds was seen at a distance on the path 
and adjacent to a fence made of piled thorny brush. The flock 
quickly darted into the fence in the manner of Zonotrichia capensis, 



2 BREVIORA No. 278 

which is surprisingly wary in rural areas, unlike its tame and trust- 
ing behavior in Andean cities and towns. Z. capensis had been 
collected in the fence previously and, believing this to be another 
flock of that species, a bird was shot as it emerged from the top 
of the brush pile. On recovering the specimen it was found to be 
Rhynchospiza stolzmanni, the elusive bird I had glimpsed on 
earlier occasions. The presence of both Z. capensis and R. stolz- 
manni in the same habitat is contrary to the observations of Jelski 
and Stolzmann (Taczanowski, 1886), who stressed the exclusion 
of one species in the presence of the other. 

That afternoon the flock was in the same area but was even 
more elusive than in the morning. On the following day a loose 
flock of about a dozen R. stolzmanni was discovered a short 
distance away in a weedy area interspersed with leafless, lightly 
branched shrubs about 10 feet tall. Some of the shrubs bore pea- 
sized dark purple fruit. The birds were feeding on the fruit high 
above the ground and were completely exposed, in striking con- 
trast to their earlier behavior. They were watched for about an 
hour and could be approached to within 20 or 30 feet before they 
were disturbed; then they merely moved a short distance and re- 
sumed feeding. Several specimens were collected here, and later 
a single bird was taken from a small, fully leafed tree standing 
alone in an area of dense underbrush. These observations are 
again at variance with those of Jelski and Stolzmann (Taczanowski, 
1886) who saw the species only on the ground or in the lowest 
branches of bushes; doubtless the presence of fruiting shrubs al- 
tered the usual retiring behavior of the bird. 

The birds were moderately vocal. The calls were a single loud, 
metallic cheep and, less frequently, a series of three or four cheeps 
given in rapid succession and descending tone. 

Six specimens were collected at Yamana. All were in very 
fresh plumage and their gonads were small. Five of the series had 
consumed the purple fruit which stained their intestines nearly 
black; the sixth bird was collected a day earlier and its stomach 
contained only a few small seeds. 

TAXONOMIC NOTES 

When describing stolzmanni, Taczanowski (1877) placed the 
species in the genus Haemophila (= Aimophila) but without giv- 
ing his reasons for doing so. Presumably he thought its relation- 
ship too obvious to require amplification. However, in 1898. 



1967 NOTES ON RHYNCHOSPIZA STOLZMANNI 3 

Ridgway created the monotypic genus Rhxnchospiza for stolz- 
manni, noting that while the species resembles the shorter tailed, 
stouter billed species of Aimophila in several respects, it differs 
from them in having a tail which is much shorter than the wing, 
in having a "nearly even" tail, and in having very small, circular 
nares nearly concealed by feathers. 

The generic allocation of the species has been generally accepted. 
Chapman (1926) apparently had unexpressed doubts about the 
validity of the genus when he remarked on the close resemblance 
between stolzmcmni and Aimophila sumichrasti of Mexico, and 
suggested that stolzmanni may have had a Middle American origin. 
Hellmayr (1938) also mentioned the similarity between these two 
species but felt the short, nearly square, tail and (adding two new 
characters) "broader and less rounded" rectrices were "good tax- 
onomic characters" for maintaining Rhxnchospiza apart from 
Aimophila. In his survey of Aimophila, Storer (1955) briefly 
noted Chapman's suggested origin of stolzmanni but did not fur- 
ther consider Rhynchospiza. 

The supposed generic characters of Rhynchospiza seem to me 
to be either inconsistent or of dubious value. The nares of Rhyn- 
chospiza are small, rounded, and rather concealed, but similar 
nostrils are found in several of the Aimophila, such as strigiceps 
and rujescens. I fail to appreciate why Ridgway and Hellmayr 
believed Rhynchospiza has a squarer tail than that of Aimophila. 
While it is true that most species of Aimophila have a compara- 
tively rounded tail, the outer rectrices are long in A. sumichrasti 
and A. ruficeps and their tails are as square as that of R. stolz- 
manni. Heilmayr's belief that Rhynchospiza could be distinguished 
from Aimophila by its broad, blunt rectrices seems to have re- 
sulted from an observational error. Several species of Aimophila 
have rectrices which are rather narrow and pointed, in contrast to 
the condition in most emberizines, but other members of the genus 
have "normal" rectrices. Among the species whose tail feathers 
are broad and blunt, like those of R. stolzmanni, are A. mystacalis, 
c/uinquestriata, sumichrasti, ruficeps, and rujescens. 

The only character clearly difl'erentiating R. stolzmanni from all 
species of Aimophila is the tail, which averages about 12 mm 
shorter than the wing, while in the aimophilas the tail is at least 
equal to the wing, and usually considerably longer. While a short 
tail may be biologically advantageous, or at least not deleterious, to 
R. stolzmanni, it does not seem that a single character of this nature 
is of value in delimiting genera. If one were to accept a short tail 



4 BREVIORA No. 278 

as a generic character it would follow that the notably long-tailed 
A. strigiceps, isolated in Argentina, would have to be set apart in 
a genus of its own, a suggestion no one has yet made. I propose, 
therefore, that stolzmanni be returned to the genus Aimophila. 

AFFINITIES OF AIMOPHILA STOLZMANNI 

Storer (1955) groups the species of Aimophila into three as- 
semblages. First, that of mystacalis, huineralis, ruficauda, sumi- 
chrasti, and strigiceps, all large forms inhabiting arid tropical scrub, 
which have dark transocular stripes, rufous shoulder patches, 
breast bands (either pronounced or muted), and inconspicuously 
colored feathers on the underside of the bend of the wing. Second, 
a group consisting of aestivalis, botterii, petenica (which I consider 
conspecific with botterii), and cassinii. These are all smaller birds, 
morphologically fairly similar, which inhabit temperate grasslands 
and which have no dark eye stripe, no breast bands, and no pro- 
nounced shoulder patches, but do have bright yellow feathers at 
the wrist joint. The third division consists of the dissimilar species 
qiiinquestriata, carpalis, ruficeps, notosticta, and rujescens, a 
group of uncertain affinities, which lack eye stripes, shoulder 
patches, and conspicuously colored wrist feathers. 

A. stolzmanni fits well with the first group, except for distinctive 
yellow feathering at the angle of the wing. It is doubtful, however, 
if the presence or absence of such markings may be used to assess 
phylogenetic relationships. Conspicuously colored (or patterned) 
underwing patches, which apparently function as signals in threat 
display, and perhaps in courtship display, are found in many fami- 
lies. They occur rather frequently in the Emberizinae and often 
are the only bright feathers on an otherwise sombre-colored bird. 
A cursory survey of the emberizines seems to indicate that this 
character is most pronounced in secretive species which inhabit 
grasslands, dense undergrowth, or dark habitats — areas where 
an inconspicuous species might effectively use a bright patch of 
color as a signal. If this is the case, such markings probably evolve 
rather readily in certain habitats and are useless to the taxonomist 
searching for evolutionarily conservative characters which may be 
clues to phylogenetic relationships. For this reason, the possession 
of yellow feathering at the bend of the wing is no deterrent for 
considering A. stolzmanni a member of the group containing 
mystacalis, humeralis, ruficauda, siunichrasti, and strigiceps. 

Although stolzmanni bears a close resemblance to sumichrasti, 
and occupies a similar habitat, the former may not necessarily have 



1967 NOTES ON RHYNCHOSPIZA STOLZMANNI 5 

arisen from the latter. A. strigiceps, the only other South Ameri- 
can Aimoplula, also bears a strong resemblance to sumichrasti. 
Thus we find three species {sumichrasti, stolzmanni, and strigi- 
ceps), all with restricted ranges, which are markedly similar to 
one another. The restricted ranges and morphological resem- 
blances suggest that these are relict forms; they may be older than 
the three boldly marked species with which they seem allied, viz. 
mystacalis, humeralis, and ruficauda. Their origin may be postu- 
lated as follows: In pre-Pleistocene times a simply patterned an- 
cestor, with an incipient breast band and rufous wing patches, may 
have ranged from Mexico south to southern South America. Dur- 
ing a Pleistocene interglacial period the Central American popula- 
tion became isolated, owing to the formation of water gaps, and 
differentiated into ruficauda. In South America the population 
retreated to arid refugia in Argentina and in southern Ecuador 
and northern Peru, forming strigiceps and stolzmanni, while the 
Mexican population {sumichrasti) was isolated to the north of the 
Isthmus of Tehuantepec. During a glacial period, when the sea 
receded, ruficauda spread northward into Mexico, occupying the 
range of sumichrasti and beyond, becoming isolated during later 
interglacials and further differentiating into the even more boldly 
patterned species mystacalis and lutmeralis on the Mexican Plateau. 
Movement of ruficauda to the south beyond northern Costa Rica 
was prevented because of the absence of dry scrub, thus leaving 
stolzmanni and strigiceps isolated in South America. 

ACKNOV^LEDGMENTS 

I thank Ernst Mayr and Francois Vuilleumier for their com- 
ments and suggestions. The field work for this study was sup- 
ported by Grant GB-4210 from the National Science Foundation. 

LITERATURE CITED 

Chapman. F. M. 

1926. The distribution of bird-life in Ecuador. Bull. Amer. Mus. 
Nat. Hist., 55:1-784. 
Hellmayr, C. C. 

1938. Catalogue of birds of the Americas. Field Mus. Nat. Hist., Zool. 
Ser.. 13(11 ): 1-662. 
Ridgway, R. 

1898. Descriptions of supposed new genera, species, and subspecies of 
American birds. I. Fringillidae. Auk, 15:223-230. 



6 BREVIORA No. 278 

Storer, R. W. 

1955. A preliminary survey of the sparrows of the genus Aimophila. 
Condor, 57:193-201. 

Taczanowski, L. 

1877. Liste des oiseaux recueillis en 1876 au nord du Perou occi- 
dental par MM. Jelski et Stolzmann. Proc, Zool. Soc. London, 
1877: 319-333. 

1886. Ornithologie du Perou, Vol. 3, 552 pp. Typography Oberthur, 
Rennes. 



(Received 23 May 1967.) 



BREVIORA 

Mmseuiinii of Coiniparative Zoology 

Cambridge, Mass. 24 November, 1967 Number 279 



A SECOND SKINK WITH FRAGMENTED HEAD SCALES 
FROM BOUGAINVILLE, SOLOMON ISLANDS 

Allen E. Greer and Fred Parker ' 



One ot the most unusual skinks to come out of the Solomon 
Islands is Sphenomorphus taylori from the island of Bougainville. 
At the time of its description (Burt, 1930; based on 2 individuals 
from "Bougainville") the species was unique among skinks in 
having many of the normally large and symmetrical head scales 
broken up into a less ordered array of smaller scales (Fig. 3; fig. 
21 in Burt and Burt, 1932). Eight more specimens of this species 
recently collected by Parker on Bougainville confirm that the 
fragmentation of the head scales is the normal condition for the 
species. 

During the same collecting efforts that yielded S. taylori, seven 
individuals of an as yet undescribed species of skink were taken 
by Parker on Bougainville. Among the distinctive features of this 
new species is the rather symmetrical fragmentation of the head 
scales on the dorsal and lateral surfaces of the snout. 

Although the presence of a supranasal scale would place the 
new species in either the genus Otosaurus or Parotosaurus in any 
of the most recent classifications of lygosomine skinks (Boulenger, 
1887; M. A. Smith, 1937; Mittleman, 1952), recent work (by 
Greer; see Greer and Parker, 1967) on skink systematics indi- 
cates that these genera (along with the monotypic Insulasaurus) 
are not readily separable from a large group of Sphenomorphus 
(also see Burt and Burt, 1932: 542). Until the systematics of this 
group is better understood, we choose to reject the names Oto- 
saurus, Parotosaurus and Insulasaurus in favor of Sphenomorphus. 



^ Department of District Administration, Kundiawa, Territory of New 
Guinea. 



2 BREVIORA No. 279 

The new species may therefore be known as: 

Sphenomorphus fragosus new species^ 

Holotype (Fig. 1): Museum of Comparative Zoology 92265, 
collected by Fred Parker on 31 March 1966 at Lake Loloru (elev. 
4300 feet), Bougainville, Solomon Islands. 

Paratypes: Mutahi area (4000 feet), Bougainville: MCZ 92268, 
coll. 13 May 1966. Lake Loloru: MCZ 92262-92264, 92266- 
92267, same data as holotype. 

Diagnosis: S. fragosus is a member of the variegatus species 
group (Greer and Parker, 1967), which is characterized by the 
presence of a large or small supranasal, and/or a double anterior 
loreal (although some species lack supranasals and have only a 
single anterior loreal ) ; usually 5 or more supraoculars of which 
usually 3 or more are in contact with the frontal; digits and limbs 
well developed and generally overlapping broadly when adpressed 
to the body; and the absence of transversely enlarged vertebral 
scales, i.e., the middorsal scales are subequal in size. 

The variegatus species group includes all the species previously 
referred to the genera Otosaurus, Parotosaurus and Insulasaurus 
as well as part of the genus Sphenomorphus. Those species of the 
variegatus species group with a supranasal and/or double anterior 
loreal are as follows: amblx placodes, annectens, anomalopus, cele- 
bense, concinnatus, cumingi, curtirostris, cyanolaemus, darlingtoni , 
granulatus, haasi, jobiense, kinabahtensis, maculicoUus, mimi- 
kanus, multisquamulatus , murudensis, nigrolabris, sabaniis, sara- 
sinorous, simus, stickeli, taylori, totocarinatus, tropidonotus, 
variegatus, wrighti. Those members of the variegatus species group 
lacking both a supranasal and double anterior loreal are: aignanus, 
arborens, boulengeri, dussumieri, florense, formosensis, indicus, 
kiihnei, lineopunctulatus, maculatus, melanochlorus , milnense, mis- 
olense, sanctus, striolatus.- 

S. fragosus can be easily distinguished from other species of 
its species group by means of the small symmetrical scales on the 

^ The species name calls attention to the rather symmetrical fragmentation 
of the head scales on the dorsal and lateral regions of the snout. 

" The variegatus species group is distributed from southeast Asia and the 
Greater Sunda Islands northeast to the Philippines and east through Celebes 
and the Lesser Sunda Islands to New Guinea and the Solomon Islands. 
The group is not found in Australia, however. The center of abundance for 
the group appears to be the western part of the Indo-Australian archipelago 
and, perhaps to a lesser extent, New Guinea. 



1967 



NEW BOUGAINVILLE SKINK 



dorsal and lateral surfaces of the snout (Fig. 1 ). In this more or 
less symmetrical fragmentation of the scales of the snout, S. jrago- 
sus is unique among lygosomine skinks with the exception of the 
much larger S. taylori, also from Bougainville (for other differ- 
ences between the two species, see Table 1 ) . 

Description: Body brownish above and in form tending toward 
stockiness; snout obtusely rounded and deep; limbs pentadactyl, 
well developed and overlapping when adpressed to the body (tip 
of 4th toe to forearm); snout-vent length of seven known speci- 
mens from 31-73 mm. 

Rostral as deep as wide, projecting only slightly onto dorsal 
surface of snout; nostril in a single nasal; nasal bordered ventrally 
by the first supralabial, anteriorly by the rostral, dorsally by a 
thin supranasal, which touches the rostral, and posteriorly by 2 
superposed anterior loreals, the upper of which contacts the supra- 
nasal; a small frontonasal is bordered anteriorly by the rostral and 
laterally by the supranasals; posteriorly the frontonasal is followed 
by a pair of small scales, and either a single small median scale and 
another pair of small scales, or simply by a single median scale; 
these small scales on the dorsal area of the snout separate the pre- 
frontals (Fig. 1 ). 





Figure 



1. Holotype (Museum of Comparative Zoology 92265) of 
Sphenomorphiis fragosus. A) Dorsal view of head. B) Lateral view of the 
left side of the snout anterior to the eve. 



4 BREVIORA No. 279 

Frontal scale contacts the last small scale or pair of scales of 
the snout; the 2 superposed anterior loreals are followed by a 
superposed pair of posterior loreals which are directly below the 
prefrontal; 5 supraoculars, the 3 anteriormost supraoculars in 
contact with the frontal; frontoparietals and interparietal distinct; 
parietals meet behind interparietal; no nuchals; lower eyelid scaly, 
separated from supralabials by a complete row of subocular scales; 
5th and 6th supralabials, or, less frequently, just 6th supralabial 
below eye; ear opening vertically elliptic, without lobules, and ap- 
proximately equal in size to eye opening. 

Scales in 42-46 longitudinal rows at midbody; dorsal body scales 
subequal, i.e., vertebral scales not transversely enlarged; a pair of 
enlarged preanal scales; 3 median series of subcaudal scales sub- 
equal in size; 4th (longest) toe with 18-21 laterally keeled sub- 
digital lamellae; upper surface of 4th toe covered by 1 or 2 scale 
rows distally, 3 scale rows along the mid-section and 4-5 scale 
rows basally (most similar to Group V of Brongersma, 1942). 

Color: In preserved specimens the upper surface of the head 
and body is brown with a series of more or less transversely con- 
fluent dark brown to black spots on the body which may be con- 
fined to the midline or, less frequently, extend the width of the 
brown dorsal area. The brown dorsum is bordered laterally by a 
black dorsolateral band, which is distinct but not sharply de- 
marcated, and extends from the posterior corner of the eye to the 
base of the tail. The upper part of this black dorsolateral band is 
bordered by thin longitudinal streaks of white which tend to be 
more confluent and well defined in the area just above and posterior 
to the shoulder. 

Below the black dorsolateral band the sides of the neck and body 
are grayish brown and. in some specimens, sporadically spotted 
with whitish flecks. The gray-brown of the sides fades to gray- 
white on the venter. The venter from the chin onto the tail is 
grayish white (yellow in life) and may be spotted with black. 

The upper surfaces of the limbs are brown but heavily spotted 
with black, and the lower surfaces are white and lightly spotted 
with black. 

The color pattern of the tail vaguely reflects the body pattern, 
i.e., brown middorsally with a series of dark brown to black spots 
dorsolaterally below which a gray ground color gives way to a 
gray-white venter. The regenerated tail is light reddish brown and 
generally unpatterned. 

In juveniles the dorsum is a lighter golden brown which ac- 
centuates the dark dorsal spots and the dark sides. 



1967 



NEW BOUGAINVILLE SKINK 



Distribution (Fig. 2): The seven known specimens of S. frago- 
sus are from between 4000-4300 feet, in the central highlands of 
Bougainville. Six specimens were taken in southcentral Bougain- 
ville (Lake Loloru, 4300 feet) and the seventh was collected in 
northeastern Bougainville (near Mutahi, 4000 feet). The 2 locali- 
ties are approximately 70 miles apart. 



156° E 



30 miles 



6°S 



7°S 



SHORTLAND 




6°S 



FAURO 



7°S- 



I55°E 



MONO I C^ 



156° E 



Figure 2. Map of Bougainville, Solomon Islands, showing the known 
collecting localities for Sphenomorphus fragosus (squares) and 5. taylori 
(triangles). 



6 BREVIORA No. 279 

Variation: The six specimens from southcentral Bougainville 
have 42(3), 43(1), 44(1), or 46(1) longitudinal scale rows 
around midbody. The single specimen from northeast Bougain- 
ville has 46 scales around midbody. 

The color pattern of the individual from northeast Bougainville 
differs from the patterns of the specimens from the southcentral 
part of the island in that the black dorsolateral band becomes less 
distinct on the posterior half of the body and the light throat and 
chest are reticulated with black. In addition there is a dark mid- 
ventral line. 

MORPHOLOGICAL COMPARISONS WITH RELATIVES 
ON BOUGAINVILLE 

The only species of S. fragosus' species group (see Diagnosis) 
on Bougainville, or in the Solomon Islands, for that matter, are 
5. concinnatus and S. taylori. The three species are readily dis- 
tinguished by the characters given in Table 1 . 

S. concinnatus is widely distributed throughout the Solomon 
Islands from Buka Island in the northwest to Guadalcanal in the 
southeast. Specimens from Bougainville and neighboring islands 
(e.g., Buka and some of the islands in Bougainville Straits) show 
a good deal of inter- and intra-population variation in squamation 
and color pattern, indicating that the systematics of the species 
in the Solomons is probably not as straightforward as the simple 
binomial would indicate. A discussion of the variation of the 
species on Bougainville and in other parts of the Solomons will 
be the object of a separate study. The data given in Table 1 for 
5. concinnatus were obtained from 60 specimens collected at Mu- 
tahi (2200-3200 feet elev.), Bougainville. 

To date, S. taylori (Fig. 3) has been known in the literature 
from only the two type specimens discussed in the original descrip- 
tion (Burt, 1930). Parker, however, has recently obtained 8 more 
specimens of this species from the following localities on Bougain- 
ville (Fig. 2): Aresi area (1 specimen, 2500 feet); Kieta area (1 
specimen, no exact locality data); Meliiup (2 specimens, 3000 
feet); Mutahi (3 specimens, 2200-3200 feet); Ramazon River 
(1 specimen, 1600-2400 feet). Data from these specimens are 
combined in Table 1 with the data from the original description 
of the species. 

The color patterns of the three species are distinctive and imme- 
diately diagnostic. 5. concinnatus shows some geographic and 
altitudinal variation in pigmentation and color pattern with speci- 
mens from the highlands of Bougainville most closely resembling 



1967 



NEW BOUGAINVILLE SKINK 







Figure 3. Dorsal view of the head of 4 specimens of Sphenoinorphiis 
taylori. A) Holotype, American Museum of Natural History 42018 from 
"Bougainville." B) Museum of Comparative Zoology 65857 from Kieta 
area. C) Paratype, American Museum of Natural History 42016 from 
"Bougainville." D) Museum of Comparative Zoology 78090 from Aresi 
area (2500 feet). 



5. jragosus. S. concinuutus from Mutahi (2200-3200 feet), for 
example, are as dark dorsally as S. fragosus, whereas S. concin- 
natus from many lowland areas of Bougainville, as well as the 
small off-shore islands, are generally a much lighter golden brown 
dorsally. 

The highland S. concinnatus also tend to show a middorsal se- 
ries of dark brown to black blotches somewhat similar to those 
of S. fragosus. In addition there is a diffuse series of small dark 
blotches along the dorsolateral line of highland S. concinnatus 
which is generally lacking or is much less well defined in the 
populations from the lowlands of Bougainville and the off-shore 
islands. Each of these dark blotches along the dorsolateral line 
of highland S. concinnatus is often associated with a small whitish 
spot situated anterior to and just above the dark blotch. 



8 BREVIORA No. 279 

This dorsolateral series of dark and light blotches in 5. concin- 
natus is rather similar to the continuous, dark dorsolateral line 
with its associated light line in 5. fragosus. One might think of 
the S. fragosus pattern as originating from the fusion (into a con- 
tinuous line) of the distinct blotches in S. concinnatiis, or, vice 
versa, the formation of the distinct blotches in S. concinnutus from 
the fragmentation of the continuous line in S. fragosus. 

S. concinnatus differs rather strikingly from S. fragosus in dis- 
playing a large black spot between the external ear opening and 
the point of insertion of the forelimb. This black spot is totally 
lacking in S. fragosus. Furthermore, in life, S. concinnatus lacks 
the yellow ventral coloring of S. fragosus. 

Juvenile and young adult S. taylori are quite unlike either of 
the species' two Bougainville relatives in displaying a color pat- 
tern of light transverse bars on the dark brown mid-dorsum. The 
flanks are a lighter grayish brown to beige with faint white spots 
which may coalesce to form distinct vertical light lines. In large 
adults all indications of the transverse and vertical bars may have 
vanished. 

05TE0L0GICAL COMPARISONS WITH 
BOUGAINViLLE RELATIVES 

Single skulls have been prepared of S. fragosus, S. concinnatus 
and S. taylori. 

Those features of the skull osteology that are characteristic of 
the variegatus species group to which the three species belong (see 
Diagnosis) are as follows: skull fairly deep throughout much of 
its length; 9 teeth on the premaxillae; palatine bones and palatal 
rami of the pterygoid bones meet along the midline to form an 
extensive secondary palate; no ectopterygoid process to the pala- 
tine which would exclude the palatal ramus of the pterygoid from a 
position on the infraorbital vacuity; postorbital bone lacking; 
supratemporal fenestra small or absent, in the latter case being 
obliterated by the backward extension of the postfrontal between 
the squamosal and parietal bones. 

Differences between the skulls of S. fragosus, S. concinnatus and 
S. taylori are minor except for the shape of the teeth on the anterior 
part of the maxillae and dentaries in S. taylori. In this species the 
anterior maxillary and dentary teeth are slightly pointed and curved 
posteriorly, whereas the corresponding teeth in S. fragosus and S. 
concinnatus, as well as in other members of the variegatus species 
group that have been examined, are straighter and more peg-like. 



1967 NEW BOUGAINVILLE SKINK 9 

The more pointed anterior maxillary and dentary teeth of S. taylori 
may be indicative of a shift to slightly more predaceous habits than 
either of its two Bougainville relatives. 

5. taylori also differs from both S. concinnatus and S. fragosus in 
that the frontal bone forms a surface suture with the maxilla to 
separate the prefrontal and nasal bones, whereas in S. concinnatus 
and S. fragosus the prefrontal makes contact with the nasal 
through a finger-like anterior projection. 

ECOLOGICAL COMPARISONS WITH 
RELATIVES ON BOUGAINVILLE 

Of the three species of the variegatus species group on Bougain- 
ville, very little is known about the ecology and habits of S. taylori 
or S. fragosus. The ecology and habits of S. concinnatus are, how- 
ever, somewhat better known. 

S. concinnatus is a common species on Bougainville from the 
coast to about 4000 feet, although it is occasionally found to an 
altitude of at least 5000 feet. The species inhabits dense primary 
forest and shows a marked preference for moisture. Although 
normally crepuscular, it is found in the open by day during rain 
and on damp overcast days. Only very rarely are individuals seen 
basking in small patches of weak sun. The species is not a bur- 
rower but lives in leaf litter on the forest floor. 

S. taylori is almost certainly a montane species, having been col- 
lected between 1600-3000 feet. Native collectors have obtained 
all the MCZ specimens, and where collectors could recall, the speci- 
mens were found under and beside decayed logs on the forest floor. 
The species does not seem to be a true burrower, however. 

S. fragosus has been collected between 4000-4300 feet in the 
central highlands of Bougainville. The specimens were under moss 
and fallen Pandanus leaves on the ground in fairly open areas. 
None were found out in the open in spite of the fact that both 
days on which the species was collected were dull and overcast. 

S. concinnatus is by far the most widespread and numerous of 
the three species of the variegatus species group on Bougainville. 
It is the only species of the three known to occur in the lowlands, 
where it is very common, and it extends through the known alti- 
tudinal range of its two relatives (1600-4300 feet) to an elevation 
of at least 5000 feet. Furthermore, S. concinnatus is the only one 
of the three species known to occur sympatrically with the other 
two. S. fragosus and S. taylori have never been found in the same 
area, although, admittedly, this may be a result of sampling error 
in populations of low density. 



10 BREVIORA No. 279 

Although only the low and middle (0-5000 feet) elevations of 
Bougainville have been adequately collected, it would appear that 
S. fragosus and S. taylori do, in fact, occur in less dense popula- 
tions than other Bougainville skinks (e.g., S. concinnatus, S. 
solomonis and S. tanneri) at these elevations. Thousands of 
reptiles and amphibians have been obtained by Parker and his col- 
lectors in all kinds of accessible habitats on Bougainville, and it is 
unlikely that animals as large or larger than these abundant skinks 
should not come to light in numbers approximately proportional 
to their relative frequency in the whole fauna. 

The possibility can not be excluded, however, that S. fragosus 
and S. taylori may occur in very peculiar habitats that were not 
widely sampled during the general collecting, or that both species 
occur in much higher population densities at elevations above those 
collected. 

Nothing is known of the mode of reproduction of S. fragosus or 
S. taylori, although S. concinnatus is live-bearing. Thirty gravid 
S. concinnatus from Kunua, Bougainville, contained 1-3 oviducal 
eggs or embryos in various stages of development. The snout-vent 
lengths of these 30 females ranged from 54-64 mm, and all were 
collected in the period May-June. 

EVOLUTIONARY HISTORY OF THE SKINKS OF THE VARIEGATUS 
SPECIES GROUP ON BOUGAINVILLE 

Within the variegatus species group, S. concinnatus, S. fragosus 
and S. taylori are most likely each other's closest living relatives. 

S. concinnatus and S. fragosus are more similar to each other in 
overall external morphology, skull osteology, and color pattern 
than is any other skink of the variegatus species group to either 
one of them. Furthermore, the fact that two such similar species 
occur together on Bougainville and not one on Bougainville and 
the other on Celebes, for example, lends support to the hypothesis 
of their close relationship. 

If it can be admitted that within the variegatus species group the 
fragmentation of the head scales as in S. fragosus is a specialization 
and the low supraocular number (4 as opposed to the more usual 
5 or more) of S. concinnatus is primitive, then an animal similar to 
S. concinnatus can easily be conceived of as being ancestral to S. 
fragosus. 

The evidence for the close relationship between S. fragosus and 
S. taylori rests primarily on the peculiar fragmentation of many 
corresponding head scales. The fact that among skinks this frag- 
mentation is unique to two species of the same species group on the 



1967 NEW BOUGAINVILLE SKINK 11 

same small island minimizes the possibility of convergence as an 
explanation. 

The large body size, the high number of midbody scales, and the 
slightly pointed anterior maxillary and dentary teeth of S. taylori 
must again be viewed as specializations for the variegatus species 
group, derivable, perhaps, from a stock similar to present-day S. 
fragosus. 

The structural changes in the evolution of the three species of 
the variegatus species group presently known from the Solomons 
may have proceeded along the general lines suggested by the 
sequence of living species: S. concinnatus -^ S. fragosus -^ S. 
taylori. 

The ancestor of these three living species would have un- 
doubtedly been recognized as an Otosaurus or Parotosaurus by 
Boulenger (1887), Smith (1937) or Mittleman (1952) and simply 
as a member of the variegatus species group by us. This ancestor 
most likely arrived in the Solomons from New Guinea or the 
Bismarck Archipelago, as the variegatus species group is unknown 
from Australia or the islands to the north, east or south of the 
Solomons. 

ACKNOWLEDGEMENTS 

Mrs. Patricia H. Kerfoot and Mr. Joshua B. Clark executed the 
line drawings of Sphenomorphus fragosus (Fig. 1) and S. taylori 
(Fig. 3), respectively. 

Dr. Ernest E. Williams of the Museum of Comparative Zoology 
has read the paper in many prenatal drafts and has, as usual, of- 
fered numerous helpful criticisms. 

Certain travel expenses incurred (by Greer) in the course of 
research pertinent to this paper have been defrayed by financial 
assistance from the Evolutionary Biology Fund. This fund is ad- 
ministered for the National Science Foundation by the Evolutionary 
Biology Committee of the Biological Laboratories, Harvard Uni- 
versity. 

LITERATURE CITED 

Boulenger, G. A. 

1887. Catalogue of the lizards in the British Museum (Natural His- 
tory). Vol. Ill, xii + 575 pp. Taylor and Francis, London. 
Brongersma, L. D. 

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



12 



BREVIORA 



No. 279 



Burt, C. E. 

1930. Herpetological results of the Whitney South Sea Expedition. IV. 
Description of new species of lizards from the Pacific islands 
(Scincidae). Amer. Mus. Novit., No. 427:1-3. 

Burt, C. E. and M. D. Burt 

1932. Herpetological results of the Whitney South Sea Expedition. VI. 
Pacific island amphibians and reptiles in the collections of the 
American Museum of Natural History. Bull. Amer. Mus. Nat. 
Hist., 63(5) :461-597. 

Greer, A. E. and F. Parker 

1967. A new scincid lizard from the northern Solomon Islands. 
Breviora, No. 275 : 1 -2 1 . 

Mittleman, M. B. 

1952. A generic synopsis of the lizards of the subfamily Lygosominae. 
Sniithsonian Misc. Coll., 117 (17): 1-35. 

Smith, M. A. 

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

(Received 23 May 1967.) 



TABLE 1 



Morphological comparisons between Sphenomorphus fragosus 
and the 2 members of its species group on Bougainville. 





fragosus 


concinnatus 


taylori 


Snout-vent 
length 


31-73 mm 


26-70 mm 


59-160 mm 


Scales around 
midbody 


42-46 


38-46 


53-60 


Number of supra- 
oculars (touching 
frontal) 


5 (3) 


4 (2) 


5-7 (3) 


Supralabials 
below orbit 


5th and 6th, less 
often just 6th 


5th, rarely 6th 


6th - 9th, although 
variable 


Subdigital lam- 
ellae (4th toe) 


18-21 


18-27, usually 
19-23 


27-35 


Nuchal scales 


None 


None 


1-2 pairs, ill- 
defined 



BREVIORA 

Mmsemim of Compsirative Zoology 

Cambridge. Mass. 19 March, 1968 Number 280 

THE NOTATUS GROUP OF SPHAERODACTYLUS 
(SAURIA, GEKKONIDAE) IN HISPANIOLA 

Benjamin Shreve 



INTRODUCTION 

Some years ago while identifying Sphaerodactylus from His- 
paniola. it came to my attention that several different forms had 
been allocated to "Sphaerodactylus difficilis," or "Sphaerodactylus 
notatus difficilis" and that this nominal species was a veritable waste 
basket for any species (or subspecies) showing any resemblance 
to it at all. I therefore tried to see what could be done to clear up 
this puzzle. It was soon seen that related forms occurred out- 
side of the region. However, in order that a beginning could be 
made, and so that the whole undertaking would not get out of 
hand, I decided to confine my study to the notatus group on the 
island of Hispaniola (and its nearby smaller islands). Extralimital 
forms were indeed studied to sain an idea of the status of the forms 
revised, but no extralimital forms were themselves reviewed, 
though references to them will be found in the text. 

Thirteen taxa are discussed herein, including not only forms 
obviously close to notatus but also those less closely related but 
formerly confused with notatus relatives. 

METHODS 

A number of scale counts are employed here which have not 
previously been commonly used in this genus. The dorsal scales 
are counted on a line from the rear of hind limb to the front of fore 
limb. This count is made just to one side of the vertebral line. 
Another scale count is made around the body at midbody. Mid- 
body is taken as the point midway between the rear limit of the 
insertion of the fore limbs and the forward limit of the insertion 
of the hind limbs. The standard distance, as used here, is about 



2 BREVIORA No. 280 

the same as previous usage for this term; i.e., the distance be- 
tween the end of the snout and the middle of the eye is projected 
on the dorsum, equally distant from the rearward limit of the fore 
limbs and the forward limit of the insertion of the hind limbs, 
respectively. Again, a position just alongside the vertebral line is 
taken for this longitudinal count. 

The lamellae under the fourth toe are counted from the first 
lamella at the separation of the digit, up to, but not including, the 
terminal "disk." Either right or left foot is used for this count, 
depending on the suitability and position of the fourth toe for 
the count. Sometimes, when doubt has existed, both sides have 
been counted. 

The phrase "scapular spot," as used here, applies to the dark 
marking often present on the back in the shoulder region, with 
frequently one or two white spots within it or else white border 
markings, developed in varying degree. 

ACKNOWLEDGMENTS 

I am grateful to the following institutions which have loaned 
material for study and to the persons in charge of their collections. 
The abbreviation of each institution as used in the text follows 
the name: American Museum of Natural History (AMNH), 
Charles M. Bogert; United States National Museum (USNM), Doris 
M. Cochran and James A. Peters; Stanford University (SU), 
George S. Myers; British Museum of Natural History (BM), Miss 
Alice G. C. Grandison; Field Museum of Natural History 
(FMNH), Robert F. Inger; Carnegie Museum (CM), Neil D. 
Richmond; University of Florida (UF), Coleman C. Goin. For 
Museum of Comparative Zoology, the abbreviation MCZ is used. 
I also wish to extend my thanks to the National Science Founda- 
tion for the grants (NSF 16066 and GB 2444) that made this 
study possible. 

Sphaerodactylus notatus difficilis Barbour 

Sphaerodactylus difficilis Barbour, 1914: 265, Santiago de la Vega, San 
Domingo = La Vega, La Vega, Dominican Republic; Barbour, 1921: 
241 (part); Mertens, 1939: 42 (part); Cochran, 1941: 115 (part). 

Description. The supranasal scales are usually separated, the 
granular dorsal neck scales do not extend to the insertion of the 
fore limb. The ventral scales are not usually keeled at all, although 
the dorsals are, as in all members of the group. The scale counts 



1968 SPHAERODACTYLUS NOTATUS GROUP 3 

for 137 individuals are: (5) 6-9(10) scales in standard distance, 
going to 11 in subadult specimens, 28-43 dorsal scales between 
rear of hind limb and front of shoulder, 39-54 scales around the 
body at midbody, (9)10-13(14) lamellae under the fourth toe. 
In coloration the female has three longitudinal dorsal dark stripes 
on the head and neck, sometimes extending further, with varying 
amounts of spotting elsewhere dorsally. The male color shows 
dorsal spotting, frequently confined to the neck and anterior part 
of the back but also quite often extending onto the head and the 
rest of the back. Sometimes dorsal marking is practically obso- 
lete. Below, spotting or darker washing is mostly confined to 
underside of head and throat but sometimes more extensive. The 
scapular spot present in females is frequently absent in the adult 
male coloration. The largest head and body measurement is 
33 mm (MCZ 63184 and 63191, both females from Bombardop- 
olis, Haiti). 

Originally described as a full species, difficilis appears barely 
separable from S. uotatus races (the type locality of typical 
uotatus is Key West). In general, Cuban specimens seem to re- 
semble those from Key West rather than those from the islands 
of the Great and Little Bahaman banks, although those from 
various localities in Cuba also show some geographical variation, 
the evaluation of which is beyond the scope of this work. 

My own comparison of Key West examples and Cuban speci- 
mens' from various localities with what I consider difficilis shows 
a usually less stocky build in the latter. The middle head stripe 
of the female and immature coloration of difficilis usually narrows 
or stays about the same width as it proceeds posteriorly and has 
decidedly uneven edges, whereas in Key West and Cuban speci- 
mens this stripe is likely to have more even edges and widen as it 
extends posteriorly. 

The scale counts of Cuban and Key West animals (6 of each) 
follow: 6-7 scales in standard distance, 26-30 dorsal scales be- 
tween rear of hind limb and front of shoulder, 37-47 scales 
around the body at midbody, (8)10-13 lamellae under the 4th 
toe, the figure in parentheses representing one specimen. The 



1 In a recent work, Schwartz, 1966, recognized a number of races of 
Cuban and Bahaman notatus. Actually, the notatiis specimens used are a 
composite of the races separated by Schwartz (1966). He confines typical 
notatus to Key West and part of the Florida mainland. 



4 BREVIORA No. 280 

corresponding counts for 137 difficilis follow: (5)6-9(10) (reach- 
ing 11 in subadult examples), 28-43, 39-54, (9)10-13(14). As 
can be seen, the dorsal count between the rear of hind limb and 
front of shoulder shows the least overlap. 

As just mentioned, there is considerable variation in the His- 
paniolan population. It was at first thought that the population 
from Haiti and the northwestern part of the Dominican Republic 
was separable from the rest, on lower scale counts, but there is 
considerable overlap. In Puerto Plata, Sosua, and to a less extent 
in Sabana de la Mar specimens, there is a tendency for head 
streaks of the female to break up into spots. In those from the 
first two localities, there is a tendency for the scapular spot not to 
persist in the adult. As in most examples of this form, there are 
two white spots within the scapular spot. Los Bracitos females, 
the two specimens from near Pefia (the latter subadult), and two 
of three females from Licey Almedio near TamboriP (the third 
with obsolete scapular spot), show only one white spot there. In 
those from Los Bracitos the spot appears to represent the fusion 
of two spots. One of the two males from Pascal Island shows 
black spotting on a whitish head. The specimen from the unlikely 
locality of Paradis will be discussed below under 5. n. randi. 

At all events, the variation mentioned above does not seem to 
be sufficient or consistent enough, geographically, to warrant fur- 
ther division into races. 

Material examined: HAITI. Departement du Nord Quest. Bom- 
bardopolis: MCZ 63174-63217; Port de Paix: MCZ 63219-21. 
Departement du Nord. Cap Haitien: AMNH 501 17, MCZ 63233; 
Grande Riviere: MCZ 9365-6; Ti Guinin near Cap Haitien: MCZ 
66283. Departement de L'Artibonite. Hinche: BM 1948.1.4.27- 
30. DOMINICAN REPUBLIC. Monte Cristi. El Ahogado, near 
Monte Cristi: AMNH 42142-3; Monte Cristi (from houses): 
AMNH 41995-6; 1 Vz miles west of Monte Cristi: AMNH 42148-9. 
and five untagged specimens; small island in Monte Cristi Bay: 
AMNH 42140; Muertos Island, Siete Hermanos: USNM 76718- 
23. Puerto Plata. Puerto Plata: MCZ 5444 (paratype), AMNH 
41340, 41975-7; Sosua: MCZ 13663. 43664-6, 44400, USNM 
121521-2. Samana. Pascal Island, Bay of Samana: AMNH 
41979-85; Sanchez: MCZ 44395-6. Santiago. Licey Almedio, 
near Tamboril: MCZ 78732-5; 3 km south of Peiia: MCZ 57822-3. 



1 Tamboril is apparently a different name for the locality referred to 
above as Pefia. 



1968 SPHAERODACTYLUS NOTATUS GROUP 5 

La Vega. La Vega: MCZ 7834-5 (type and paratype). Diiarte. 
Los Bracitos: AMNH 45201-2, 45204. 45206-7, 45209-12, 
45214-5, 45217-20. Seibo. Sabana de la Mar: AMNH 41992-3, 
50047, 50122-4, 50232, MCZ 57824-6, 57831-3. Barahona. 
Paradis: AMNH 51466. Distrito Nacional. Santo Domingo City: 
SU 14694. San Pedro de Macoris. 31 mi. east of Santo Domingo 
City: AMNH 49990-2. 

Field notes. Through the courtesy of Miss Alice G. C. Grandi- 
son, the following color note has been supplied for the series from 
Hinche by the collector, Ivan Sanderson: "Upper surfaces fawn 
brown, prominently spotted with jet black. Spots tend to run into 
three lines on head, while on shoulders are a pair of well defined 
jet black spots bordered by an intense white point on either side. 
Underside of head and forebody pale gray and semi-transparent. 
Of limbs, lower belly and tail, same, but minutely speckled with 
black." 

Identification aids. The three dorsal streaks in the female and 
immature coloration will serve to distinguish this form from all 
other Hispaniolan members of this group. Outside of its own 
races, probably most closely related to S. b. brevirostratus, so see 
identification aids for this form (p. 00). 

The following form is named for A. Stanley Rand, co-collector 
of the type series and student of West Indian herpetology. 

SPHAERODACTYLUS NOTATUS RANDI Subsp. UOV. 

Holotype: MCZ 57839, female, from Oviedo, Pedernales, Do- 
minican Republic, collected by Clayton E. Ray and A. Stanley 
Rand, August 20, 1958. 

Paratypes: MCZ 57840-5, with the same data as the type; 
MCZ 57838, also with the same data as the type, except that the 
collecting date is August 19, 1958. 

Diagnosis. The new form differs from S. notatus difftcilis in 
apparently slightly higher scale counts as follows (the count of 
the type followed by those of the paratypes in parentheses); stand- 
ard distance 8 (7-10); dorsal scales from rear of hind limb to 
front of shoulder 44 (36-40); around body at midbody 54 (48- 
52); lamellae under 4th toe 12 (12-13); also, randi appears to not 
grow quite as large as difficilis. In the female coloration there is 
no sign of the three head and nuchal stripes of typical difftcilis; 
instead, above, there is much spotting, marking, and washing of a 
light brown ground color with dark brown; back of head showing 
ground color more than is shown elsewhere, a poorly developed 



6 BREVIORA No. 280 

scapular spot, a dark brown marking preceded by a curved whitish 
transverse streak, convexly directed anteriorly, followed by a 
dark spot, marked posteriorly by whitish; below, ground color about 
the same as above, stippled and washed with dark brown. 

Most of the paratypes, above, show the same kind of markings 
as the type. However, the scapular spot is practically obsolete in 
some, and in only one, a male, takes the form of a black spot en- 
tirely borderd with whitish, which may represent the maximum 
in its development. There is also variation in the spotting, par- 
ticularly on the head and neck. Whitish spots and markings are 
present in some. Two show a faint marking on the back of the 
head and neck, somewhat reminiscent of that of S. b. brevirostratus, 
described below. Below, the markings are like those of the type, 
except that they are stronger in some. 

Of the four male paratypes, two apparently have not lost the 
female or inmiature coloration. The two showing what is ap- 
parently the adult male coloration, above, have lost much of the 
dorsal markings, the head being whitish with dark brown or black 
spots above and below, including throat; otherwise, below, marked 
similarly to the rest of the type series. Even the males with the 
female or immature pattern are decidedly spotted under the head 
and on the throat. The females show little of this. 

The male coloration of difficilis somewhat resembles the female 
coloration of randi, although it usually shows less dense markings. 
However, some males of difficilis resemble those of randi. 

Measurements (in mm). Type, MCZ 57839: length, head and 
body, 29; length of tail, 26. Largest paratype, MCZ 57840: head 
and body, 27; tail regenerated. 

Remarks. It might be supposed that this form is the representa- 
tive of difficilis in the Barahona Peninsula, but a specimen from 
Paradis (Paraiso) appears to be a male difficilis. It has the 
reduced spotting characteristic of a male of that form, especially 
reduced on the head, with the scale counts being a bit low for 
randi although barely within the limits of that form, except for 
the rear of hind limb to front of shoulder count which is one below 
the limit for randi. Also, the head and body length is one milli- 
meter more than the maximum for randi. The counts for this 
specimen, AMNH 51466, are: standard distance 8, dorsal scales 
between rear of hind limb to front of shoulder 35, around midbody 
48, lamellae under 4th toe 12. It is possible that the locality 
data for this animal may be incorrect. 



1968 SPHAERODACTYLUS NOTATUS GROUP 7 

Identification aids. This has essentially the scale characters of 
S. n. difficilis but differs in counts and color, as stated in the 
diagnosis. 

Another race of notatus, described below, is named for Jay M. 
Savage because of his interest in this group. 

SPHAERODACTYLUS NOTATUS SAVAGEI Subsp. UOV. 

Sphaerodacty Ills difficilis; Gv^ixxu 1952: 187; 1956: 86; Savage, 1954: 328. 

Holotype: SU 14695, a female, from La Romana, Altagracia, 
Dominican Republic, collected by Chapman Grant in December 
1946. 

Paratypcs: SU 14696-708, with the same data as the type; 
MCZ 73900 from the same place, collected by Clayton E. Ray 
and Robert R. Allen, March 27, 1963. 

Diagnosis. Apparently most closely related to S. notatus difficilis 
from which it differs in having larger scales, or at least lower 
scale counts, although there is considerable overlap. The scale 
counts follow, with those of the paratypes in parentheses: scales 
in standard distance 6 (6-7 and up to 8 in the immature); dorsal 
scales between rear of hind limb and front of shoulder 28 (27-34) ; 
scales around body at midbody 42 (36-44); lamellae under 4th 
toe 13 (11-14); and the limited extent of the male escutcheon, 
this not extending onto the leg. There is also a difference in color. 
The female coloration shows but little evidence of the head stripes 
of difficilis, only a rough tendency for the spots to form into lines, 
most notably the ones behind the eye. Males would appear to 
average more heavily spotted than those of difficilis. Subadult 
specimens, presumably of both sexes, instead of being spotted 
show a series of cross bars across the back, with two stripes ex- 
tending back from each eye and joining on the back of the head 
in somewhat the same manner as in typical brevirostratus (to be 
described), the central stripe being mostly obsolete. Subadults 
also have an ill-defined scapular spot, the foremost dark band 
on the neck being followed by two whitish spots, this band being 
more conspicuous than those following. The scapular spot ap- 
pears to disappear as the animal gets older, especially so in the 
male, but is entirely absent in the female type. 

Measurements (in mm). Type, SU 14695: length, head and 
body, 32; length, tail, 26 (apparently regenerated). Largest para- 
type, SU 14696: head and body. 30; tail missing. 



8 BREVIORA No. 280 

Identification aids. This is one of the two forms where the male 
escutcheon does not extend onto the leg. The other is 5. n. juanil- 
loensis. This, as far as known, is a character unique within the 
Haitian notatus group. Beyond this the racial diagnosis is covered 
under each form. 

Sphaerodactylus notatus juanilloensis subsp. nov. 

Holotype: MCZ 73901, a female, from Juanillo, Altagracia, 
Dominican Republic, collected by Clayton E. Ray and Robert R. 
Allen, March 29, 1963. 

Paratypes: MCZ 73893-98, UF 21970-75, with the same data 
as the type. MCZ 73899, Dominican Republic, Altagracia, 1 km 
north of Boca de Yuma, collected by Clayton E. Ray and Robert 
R. Allen, March 27, 1963. 

Diagnosis. Apparently most closely related to S. notatus savagei 
from which it differs in coloration. The coloration, notably the 
female and immature, resembles that of the immature savagei. It 
differs particularly in that the dorsal cross bands are arranged in 
pairs, the two bars of the pair separated by a narrow white area; 
the scapular spot in the female well developed, squarish, with two 
white spots in the middle, located as usual betwen the shoulders. 
In the male coloration there is also a tendency for the dark spots 
to form indistinct bars or at least to coalesce. In adults of both 
sexes of savagei there appears to be no tendency toward bar for- 
mation. The scale counts for juanilloensis follow, with those of 
the paratypes in parentheses: scales in standard distance 6 (5-7, 
one with 8, scale abnormality present, evidently scar tissue); dorsal 
scales between rear of hind limb and front of shoulder 29 (25-32) ; 
scales around the body at midbody 42 (40-48) ; lamellae under 4th 
toe 12 (12-14). 

Measurements (in mm). Type, MCZ 73901: length, head 
and body, 26; part of tail missing. Largest paratype, UF 21972: 
head and body, 31, a female; tail missing. 

Identification aids. See 5. n. savagei. 

The following form is named for James D. Lazell, Jr., the 
co-collector of the type specimen and other valuable herpetologi- 
cal material. 

Sphaerodactylus lazelli sp. nov. 

Holotype: MCZ 63218, a male, from Cap Haitien (under bark 
of tree in shady gully), Departement du Nord, Haiti, collected by 
A. Stanley Rand and James D. Lazell, Jr., July 16, 1960. 



1968 SPHAERODACTYLUS NOTATUS GROUP 9 

Diagnosis. Possibly most closely allied to S. notatus difficilis 
with which it is sympatric, differing in lower scale count from 
rear of hind limb to front of shoulder, in having distinctly keeled 
scales on the throat, and in coloration, with a very prominent 
scapular spot and two black spots preceding it (female coloration 
unknown). 

Description. Snout long, rather pointed, distance of eye from 
tip of snout greater than distance of eye to ear; rostral with 
median groove; nostril between rostral, first supralabial, a larger 
supranasal and three smaller scales, the third or lowest particu- 
larly small; supranasals separated by one scale; three large and 
one small supralabial to below center of eye, or a bit beyond; head 
above and on sides covered with small juxtaposed keeled scales, 
those on snout slightly enlarged; scales on back enlarged, some- 
what elongate, keeled and subimbricate; the more granular scales 
of the neck appear well anterior to the forward level of the 
shoulder; 5 scales in standard distance; 25 dorsal scales between 
rear of hind limb and front of shoulder; 42 scales around the body 
at midbody; 11 lamellae under the 4th toe; mental somewhat 
larger than rostral; two large infralabials, followed by a small one, 
to center of eye; two enlarged squarish chin shields behind mental; 
behind scutes just mentioned, scales decreasing in size posteriorly 
to throat, increasing in size on throat and chest, and again on 
belly, smooth, except keeled on throat and forward up to chin 
shields; tail above with scales like dorsum but smaller; below, 
with scales like belly at base, but beyond, those on midline en- 
larged, especially nearer terminus (tail apparently regenerated, pos- 
sibly twice). 

Coloration in alcohol. Above, light brown, a large diamond- 
shaped scapular spot about on a level with the anterior border of 
the shoulders, with a white spot near each lateral corner. Pre- 
ceding this are two prominent black or dark brown spots, both 
on the midline, one on the neck and the other on the occipital 
region of the head; an indistinct, dark brown streak from the rear 
of the supraocular region, on each side, to the anterior edge of 
the neck spot, another from each eye along the side of the head 
on to the neck. The body, limbs and tail are lightly spotted with 
black or dark brown, three spots just to the rear of the scapular 
spot being particularly prominent. There is a whitish chevron- 
shaped bar, with the apex directed rearward, a bit posterior to 
the base of the tail; a narrow crossbar of the same color on the 
tibia. Below, washed with dark brown, especially nearer the sides; 



10 BREVIORA No. 280 

also lightly spotted with dark brown nearer side of throat and 
chest. 

Measurements (in mm). Type, MCZ 63218: length, head and 
body, 31; length, tail, 29 (apparently regenerated). 

Identification aids. The keeled throat scales will serve to sep- 
arate this from all forms which have only smooth scales ventrally. 
From those that have keeled throat scales, the very large scapular 
spot and the two spots preceding it are a type of coloration not 
seen elsewhere in this group. However, the female of this form 
is not known. The dorsal scale count from the rear of hind limb 
to the front of the shoulder is lower than that of any other form, 
although with more material this might be found to overlap with 
S. b. enriquilloensis, a form with slightly keeled throat scales, in 
this character. 

Sphaerodactylus brevirostratus brevirostratus 

sp. and subsp. nov. 

Sphaerodactylus difficilis: Barbour, 1921: 241 (part); Cochran, 1941: 115 
(part). 

Holotype: MCZ 63234, a female, 5 km south of Dufort, south 
of Leogane. Departement de I'Ouest, Haiti, collected by A. Stanley 
Rand and James D. Lazell, Jr., July 31, 1960. 

Paratypes: All from HAITI. Departement de L'Artibonite. 
Saint Marc: AMNH 49680, William G. Hassler, April 1935. De- 
partement de r Quest. With the same data as the type: MCZ 
63235-8; Boutillier Road near quarry: MCZ 64514, Ernest E. 
Williams and A. Stanley Rand, August 15, 1959; west of Carre- 
four: USNM 117307-8, Anthony Curtiss, May 11 and 15, 1943; 
Carrefour Feuille, Port-au-Prince: MCZ 64812-3. Ernest E. Wil- 
liams and A. Stanley Rand, August 15, 1959; Damiens: AMNH 
50120-1, William G. Hassler, May 2, 1935; Diquini: MCZ 64815- 
21, Luc Whiteman, March 10-11, 1961; Eaux Gaillees: MCZ 
51315-6, Anthony Curtiss, April 22, 1950; "Ferronai^': MCZ 
13458, Glover M. Allen, August 1919; Furcy: USNM 124158, 
Anthony Curtiss, June 1944; Glore: USNM 118908-14, Anthony 
Curtiss, May 18, 1943; Hatte Latham: AMNH 90168-9, USNM 
117151, Anthony Curtiss, December 7, 1946, and February 13, 
1943; La Fond near Jacmel: MCZ 64822, Luc Whiteman, April 
20, 1961; Manneville: MCZ 64804-11, Ernest E. Williams and 
A. Stanley Rand, August 13-14, 1959, CM 38882, George 
Whiteman, 1963, BM 1948.1.4.31-33, Ivan Sanderson, January 



1968 SPHAERODACTYLUS NOTATUS GROUP 11 

23, 1937; Mon Repos: USNM 69124, Gerrit S. Miller, Jr., March 
5, 1925; Petionville: USNM 59185, Paul Bartsch and J. B. Hen- 
derson, April 1, 1917; Pont Beudet: BM 1948.1.4.34, July 12, 
1937; 8 miles north of Port-au-Prince (fig tree in cotton field): 
AMNH 49588, 49590-1, William G. HassleV April 18, 1935; Port- 
au-Prince: USNM 118867-70, Anthony Curtiss, USNM 59179 
Paul Bartsch and J. B. Henderson, April 24, 1917, SU 14688- 
14693, Major Chapman Grant. December 18, 1941; Source 
Puantes: USNM 117159-64, Anthony Curtiss, November 18, 
1943; Trou Caiman: USNM 117153-6, Anthony Curtiss, January 
16, 1943; Trou Forban: USNM 117157-8, 118894, Anthony 
Curtiss, October 1942. 

The locality "Ferronai" has not been located on the map, but 
has been listed under Departement de TOuest, as this seemed a 
likely place of origin, the majority of the localities being from 
there. Through the courtesy of Mrs. W. E. Schevill, I was able to 
examine the field journal of the collector, the late Glover M. Allen. 
The locality was not mentioned. 

Diagnosis. Possibly most closely allied to S. niclwlsi Grant from 
which the new form differs in having smaller scales (average larger 
scale counts), larger size, less short and blunt snout, and different 
coloration. Also related to, and in the past confused with, 5. notatus 
difficilis, from which the new form differs in its smaller size, shorter 
and more blunt snout, more stocky build, in having larger scales 
(smaller average scale counts) and in coloration. 

Description. (Paratype variation in parentheses.) Snout short, 
blunt; distance of eye from tip of snout slightly greater than dis- 
tance of eye to ear; rostral with median groove; nostril between 
rostral, first supralabial, a large supranasal and two smaller 
scales; supranasals separated by one scale (in some, separated by 
two or more scales) ; three large and one small supralabial to below 
center of eye, head above and on sides covered with small jux- 
taposed keeled scales, those on snout enlarged; scales on back 
enlarged, elongate, keeled and subimbricate; the more granular 
scales of the neck appear anterior to the forward level of the shoul- 
der (also in paratypes); 6 scales in standard distance (6-7 in 
adults, subadults going to 8); 31 dorsal scales between rear of 
hind limb and front of shoulder (28-35, and one example 38, 
with scale abnormality in the sacral region); 41 scales around the 
body at midbody (37-45); 10 lamellae under the fourth toe 
(9-12, only two instances of the last figure); mental somewhat 
larger than rostral; two large infralabials followed by a small one 



12 BREVIORA No. 280 

to center of eye; three small squarish chin shields behind mental; 
behind these the scales decreasing in size, posteriorly, to throat, 
increasing in size on throat and chest and again on belly, smooth 
(a very few throat scales very slightly keeled in a few of the para- 
types); tail, above, with scales like dorsum but smaller and more 
imbricate and not keeled near terminus; tail, below, with scales 
like those of belly. 

Coloration in alcohol. Above, ground color light grayish brown; 
on head a dark brown streak extending from end of rostral, where 
it is joined with its fellow on the other side of the head, the color 
on rostral being rather dark gray, extending through eye to back 
of head where it again joins its fellow, this time in a figure which 
resembles the top of a heart with the direction of the possible apex 
toward the snout. Within the area enclosed by these markings and 
to some extent just outside, the ground color becomes very pale; 
also several dark brown spots inside of this area, one behind eye 
and joining onto eyelid on each side, two others more or less 
between eyes and one on snout. Behind the top of heart-like 
figure on the back of the head is a second one, very similar to, 
but less well defined, and more or less joining the first and just 
behind it, both with a more or less distinct whitish border on the 
inner posterior margin. A distinct scapular spot consisting of an 
almost transversely oval blotch of black with a small white spot 
on each side near its lateral edge; just preceding the scapular spot 
is a thin, rearwardly curved, whitish, transverse line; elsewhere, 
back spotted with dark brown; the limbs spotted with the same, 
including white markings on the hind limbs; an obscure whitish 
streak at posterior side of body; a white streak bordered with 
dark brown along side of tail going on to rear of thigh; tail above 
spotted and marked with dark brown and white spots, no special 
marking at root of tail. 

Below, whitish; longitudinal dark brown lines on venter, be- 
coming more broken under head and neck. 

In the female paratypes the head markings may be less heart- 
shaped, less well defined or broken, the hinder one sometimes 
being obscure or absent. Sometimes the head spots inside are 
reduced or absent or united with each other and the enclosing 
streaks in various ways. 

In the males the head markings may be even more reduced or 
faint and in some are entirely absent, being replaced by small 
spots. In the males the scapular spot, although sometimes nearly 
obsolete, is very persistent; yet, USNM 117157, a female, is the 



1968 SPHAERODACTYLUS NOTATUS GROUP 13 

only one which shows no trace of it. The paratypes from Trou 
Forban and Source Puantes, however, often show less development 
of the scapular spot than those from elsewhere. 

Other variations are the reduction to almost total absence of 
the ventral markings in some specimens, and the variation in 
shape of the scapular spot which is sometimes more square 
or longitudinally elongate than in the type. 

Measurements (in mm). Type, MCZ 63264: length, head and 
body, 22; length, tail, 20. Largest paratype (one) MCZ 64805: 
head and body, 26; tail, 27. 

Remarks. In order that a comparison may be made with the 
scale counts of the new form, counts for nicholsi are supplied in 
the same order as they occur in the description: standard distance 
6-7 (two adults and one subadult 7); rear of hind limb to front 
of shoulder 25-32; around body at midbody 35-40; lamellae under 
fourth toe 9-12 (two examples at 12). These counts were based 
on 18 nicholsi. 

Some of the type series were found in leaf litter on the ground, 
while specimens AMNH 49588, 49590-1, from 8 miles north of 
Port-au-Prince, were collected on a fig tree in a cotton field. 

S. n. difficilis is not known to occur with this new form any- 
where. 

Field notes. Through the kindness of Miss Alice G. C. Grandi- 
son, the following field notes have been supplied for the British 
Museum specimens collected by Ivan Sanderson: The three from 
Manneville were collected among leaves under thorns. The color 
was described as ". . . brown lighter to almost white below. Tail 
small, yellowish. Throat spotted with black, spots run in lines 
on belly. Flanks and dorsum likewise spotted, tending to run in 
lines on flanks." The Pont Beudet specimen was collected in a 
small mud house. The color was described as fawn, tail unspotted, 
head with black reticulum. 

Identification aids. The usually prominent, dark, longitudinal, 
ventral streaks will serve to distinguish this form from all others in 
the present group, except the race to be described next, enriquillo- 
ensis. However, in some specimens these are not entirely evident. 
The latter form, however, differs in having a decided suppression of 
the dorsal pattern including absence of the scapular spot, which is 
almost always present in both sexes of the typical form. Also, 
enriquilloensis has keeled throat scales, atlhough less strongly 
keeled than other forms covered in this work. S. b. brevirostratus 
belongs to the stocky, small, blunt-snouted group which includes, 



14 BREVIORA No. 280 

aside from the race mentioned above, noblei and darlingtoni about 
to be described as new also. 5. armstrongi Noble and Hassler may 
possibly also belong to this group. The separation from S. n. diffi- 
cilis, with which it was formerly confused, is covered in the diag- 
nosis. 

Sphaerodactylus brevirostratus enriquilloensis subsp. nov. 

Holotype: MCZ 57846, a male, from 4 km east of La Descubi- 
erta, near Lake Enriquillo, Independencia, Dominican Republic, 
collected by Clayton E. Ray and A. Stanley Rand, August 22, 
1958. 

Paratypes: All from the DOMINICAN REPUBLIC. Inde- 
pendencia. With the same data as the type: MCZ 57847-8; La 
Descubierta: AMNH 45125, G. K. Noble, September 25, 1922; 
Las Baitoas, Lake Enriquillo: AMNH 50153-63, William G. 
Hassler, August 5, 1935, AMNH 50205-09, William G. Hassler, 
August 5-8, 1935. Barahona. Barahona: AMNH 50118, William 
G. Hassler, June 23, 1935. 

Diagnosis. This new form differs from the nominate subspecies 
in less blunt snout (overlap in this character), average larger scale 
counts as follows, the count of the type followed by those of the 
paratypes in parentheses: standard distance 7 (6-8), rear of hind 
limb to front of shoulder 31 (32-37), around body at midbody 46 
(40-47), lamellae under fourth toe 10 (9-10, with one specimen 
at 11), and general reduction of the color pattern (in alcohol). 
There is also more keeling of throat scales than in the typical 
form. In the males, female head pattern virtually non-existent, 
sometimes heavily spotted, especially above, or virtually unmarked. 
The type is much spotted above, and below, on the head and 
throat, also showing evidence of the ventral lines of the typical 
form. In general these seem less developed in enriquilloensis than 
in brevirostratus. The female often lacks the head markings and, 
when present, there is only a rather indistinct replica of those of 
the typical form. Females may be virtually unmarked. Both sexes 
lack any scapular spot. 

Measurements (in mm). Type, MCZ 57846: length, head and 
body, 23; tail missing. Largest paratypes, AMNH 50209, female, 
and AMNH 50118, male: head and body, 26; 50209 tail regen- 
erated, 50118 tail missing. 

Remarks. A specimen of Sphaerodactylus notatus difficilis, al- 
ready referred to, from Paradis (Paraiso), seems to be the nearest 
approach to sympatry between that taxon and brevirostatus. The 



1968 SPHAERODACTYLUS NOTATUS GROUP 15 

locality is, however, south of that of the nearest b. emiquilloensis 
(Barahona) on the Barahona peninsula, well south of the Cul de 
Sac Plain. No other difficilis is known from the "south island," 
and the record may be an error of locality labelling. 

Identification aids. As stated under the nominate form, this is 
one of those with keeled throat scales. This character, and the 
presence of longitudinal ventral streaks and suppression of dorsal 
color pattern, will serve to distinguish it from the other forms. 

The following species is named for Dr. P. J. Darlington, Jr., the 
collector and the major student of West Indian zoogeography. 

SPHAERODACTYLUS DARLINGTONI sp. nov. 

Sphaerodactyliis difficilis: Cochran, 1941: 115 (part). 

Holotype: MCZ 44380, a male, from Mt. Diego de Ocampo, 
summit dome, ca. 4000 feet, between Puerto Plata and Santiago, 
Dominican Republic, collected by P. J. Darlington, Jr., July 1938. 

Paratype: MCZ 44379, with the same data as the type. 

Diagnosis. Apparently most closely related to S. noblei, described 
below, differing in coloration and possibly slightly larger size, and 
also in the absence of distinctly keeled scales on the throat, in more 
lamellae under 4th toe, more dorsal scales from rear of hind limb to 
front of shoulder, and in having one small scale instead of two in 
contact with rear of nostril scale. 

Description. (Paratype variation in parentheses.) Snout short, 
blunt; distance from eye to tip of snout about equal to that of eye 
to ear; rostral with median groove; nostril between first supra- 
labial, a large supranasal and two smaller scales; supranasals 
separated by one scale; three large and one small supralabial to 
below center of eye (four and one on left side; small scale behind 
center of eye on right side, although the three and one formula 
mentioned for the type exists here); head above and on sides 
covered with small, juxtaposed, keeled scales, larger and mostly 
keeled on snout; scales on dorsum enlarged, elongate, keeled, and 
subimbricate; the more granular scales of the neck appear anterior 
to the forward level of the shoulder, the dorsals, however, decidedly 
smaller than those located more posteriorly, at the point just men- 
tioned; scales in standard distance 7 (9); 45 dorsal scales between 
rear of hind limb and front of shoulder (43); 54 scales around the 
body at midbody (48) (in both type and paratype there is an in- 
cision at the side of the belly where it is suspected some scales may 
be missing, hence the two counts may be lower than they should 



16 BREVIORA No. 280 

be); 12 lamellae under 4th toe; mental somewhat larger than 
rostral; two large infralabials followed by a small one to center of 
eye; two small squarish scales behind the mental (more irregularly 
shaped); somewhere behind the two just mentioned, scales de- 
creasing in size posteriorly to throat, increasing in size on throat 
and chest and again on belly, smooth, unless it can be said some of 
the throat scales are very weakly keeled; base of tail, above, with 
scales like those of dorsum but more imbricate and slightly larger; 
posteriorly, scales abruptly smaller where regenerated; tail, below, 
with scales like those of belly but abruptly smaller under regen- 
erated portion. It may be stated for the sake of clarity that both 
the type and the paratype have much of the tail regenerated. 

Coloration in alcohol. Above, light brown, a very few dark 
brown spots on back and on head, particularly in loreal region; a 
blurred dark brown scapular spot; below, brownish white, a few 
obscure dark brown markings on throat and underside of head; the 
paratype, also a male, shows a persistence of what is probably the 
female and immature pattern; above, ground color slightly darker 
than that of type; a brownish black scapular spot, rather indistinct, 
with an obscure whitish line just preceding it and following a dark 
brown line from nostril to eye, emerging on the posterior side of 
eye and curving to meet its fellow from the opposite eye, forming 
a marking on the back of the head like that of b. brevirostratus and 
noblei but not shaped like the top of a heart; posterior lines just 
mentioned likely to be bordered with whitish and broken; inside the 
figure produced by the lines, dark and light mottlings and markings; 
elsewhere on back, a few, more than in type, small dark brown 
spots; below, brownish white, more or less powdered and marked 
with darker brown, a reticulate pattern of whitish and dark brown 
on underside of head and throat. 

Measurements (in mm). Type, MCZ 44380: length, head and 
body, 26; tail regenerated. Paratype, MCZ 44379: head and body, 
22; tail regenerated. 

Identification aids. The very blunt snout, the absence of any 
ventral keeling, small size, and small scales are a combination of 
characters that will separate this species from all forms covered 
here. 

The next species is named for the late G. K. Noble, the collector, 
whose work in Hispaniolan herpetology is especially important. 



1968 SPHAERODACTYLUS NOTATUS GROUP 17 

Sphaerodactylus noblei sp. nov. 

Holotype: AMNH 45216, a male, from Los Bracitos, Duarte, 
Dominican Republic, collected by G. K. Noble, August 1929. 

Paratypes: AMNH 45203, 45205, 45208, 45213, with the same 
data as the type. 

Diagnosis. Allied to S. notatus difficilis Barbour from which the 
new form differs in more stocky build, smaller size, and colora- 
tion. Also allied to S. b. brevirostratus from which the new form 
differs in smaller scales or larger scale counts (slight overlap in 
longitudinal and standard distance counts), color, and seemingly 
smaller size. 

The new form also differs from both in possessing distinctly 
keeled throat scales. 

Description. (Paratype variation in parentheses.) Snout short, 
blunt; distance of eye from tip of snout slightly greater than dis- 
tance of eye to ear; rostral with median groove; nostril between 
rostral, first supralabial, a large supranasal, and two smaller scales; 
supranasals separated by two scales (separated by one scale, at 
least one specimen giving the impression that the one scale has been 
made by the fusion of two); three large and one small supralabial 
to below center of eye; head above and on sides covered with small, 
juxtaposed, keeled scales, larger and mostly keeled on snout, scales 
on dorsum enlarged, elongate, keeled and subimbricate; the more 
granular scales of the neck appear anterior to the forward level of 
the shoulder, the dorsals, however, decidedly smaller than those 
located more posteriorly, at the point just mentioned; scales in 
standard distance 8(8-9); 35 dorsal scales between rear of hind 
limb and front of shoulder (35-38); 51 scales around the body at 
midbody (50-55); 11 lamellae under the 4th toe (9-11); mental 
somewhat larger than rostral; two large infralabials followed by a 
small one to center of eye; two small, rather irregularly shaped 
scales behind the mental; behind the two just mentioned, the scales 
decreasing in size posteriorly to throat, increasing in size on throat 
and chest and again on belly, smooth, except that the throat scales 
and to some extent those further forward, nearly to chin shields, are 
distinctly keeled; tail, above, with scales like those of dorsum, more 
imbricate, but about the same size or possibly slightly larger at the 
base, not keeled near terminus; below, with scales like those of 
belly. 

Coloration in alcohol. Above, ground color brown, markings 
very similar to those of 5. b. brevirostratus but less distinct, a dark 
brown line extending from nostril through eye and joining its fellow 



18 BREVIORA No. 280 

in a marking which resembles the top of a heart with the direction 
of the apex toward the snout; within the area enclosed by the 
streaks just mentioned the ground color becomes pale, especially 
near the lines themselves; within this area a broken line proceeding 
backward from each eyelid on to the top of the head; other similar 
but obscure markings between the eyes and nearby and on the 
snout; a very indistinct suggestion of a second heart-shaped figure 
behind the first, with also a suggestion of lightening inside; on the 
neck an indistinct scapular spot consisting of a rather blurred dark 
brown spot preceded by a rather thin, rearwardly-curved, whitish, 
transverse line; elsewhere on back obscure dark brown spots with 
two very obscure whitish ones to the rear of two of the darker 
spots a bit behind the insertion of the forelimbs; an obscure whitish- 
bordered triangular spot at the base of the tail, the apex directed 
rearward; also obscure dark brown spots on the limbs and the rest 
of the tail, the latter rather densely spotted; a whitish dark brown 
bordered streak at the side of the base of the tail, extending on to 
rear of thigh; no whitish streak on side of body. Below, heavily 
dotted and dusted with brown, a line of small spots on the tail; 
throat and underside of head with a prominent black or dark brown 
and white reticulate pattern. Paratypic variation seems not ex- 
tensive and sexual dichromatism seems to be very slight, if it exists 
at all, the single female paratype showing a fainter development of 
the pattern on throat and underside of head; otherwise all para- 
types seem to be more obscurely marked above than the type, 
although one animal (not the female) shows two irregular white 
spots near the posterior edge of the indistinct scapular spot. Below, 
the only important difference, except that mentioned above, would 
be no line of spots under the tail (only two with tails, one de- 
tached). 

Measurements (in mm). Type, AMNH 45216: length, head and 
body, 22; length, tail, 23. Largest paratype, AMNH 45213, a 
male: head and body, 24; tail missing. 

Remarks. The following specimens, also from the Dominican 
Republic, tentatively referred to noblei, are not listed as paratypes. 
Boca del Infierno: USNM 74974-77, Gerrit S. Miller, Jr., Feb- 
ruary 28, 1928; Rio San Juan, Samana Peninsula: USNM 74943, 
Gerrit S. Miller, Jr., March 1928, but identified by Miss D. M. 
Cochran as S. difficilis ( 1941 : 1 15). These animals may belong to 
a distinct race. Thus the dorsal count from the rear of hind limb to 
the front of shoulder is 38-41 in the USNM examples above, 
whereas the count is 35-38 in the type series. The complete 



1968 SPHAERODACTYLUS NOTATUS GROUP 19 

counts, given in the same order, are: 7-9 (10 for one immature), 
38-41, 50-56, 9-11. Three of the former have this count of 38, 
although only one of the latter has that figure. The type series 
has all but one paratype with only one scale separating the supra- 
nasals rather than two, although one other animal has a single 
scale in this position which has the appearance of two fused. All 
of the non-paratypic series have the supranasals separated by two 
scales. In view of the overlap in characters in the two series, 
taxonomic separation seems undesirable with such a small number 
of specimens. On the other hand, the possibility that with more 
material further separation would be desirable prevented the in- 
clusion of these specimens as paratypes. It is believed, in any case, 
that the two series indeed are specifically identical, even if not 
subspecifically so. 

The USNM non-paratypic specimens show weak or obsolete 
pattern on throat and underside of head, but they are regarded as 
females except for one doubtful male and a female equally 
doubtful. 

Identification aids. The dark brown or black and white reticu- 
lation of the underside of the throat and head will separate this 
form from the others of the group, although in some individuals, 
as stated before, this pattern is weak or obsolete. 

SPHAERODACTYLUS ARMSTRONGi Noblc and Hassler 

Sphaerodactylus annstrongi Noble and Hassler, 1933: 5, mountain top on 
property of G. Herrmann, near Paradis, Barahona, Dominican Re- 
public, 2400 feet. Cochran, 1941: 119. 

MCZ 57820-21 are referred here with some doubt, as they do 
not entirely fit the characters of armstrongi. Available for com- 
parison is a paratype of armstrongi from the type locality. The two 
referred specimens differ from armstrongi in showing no keeled 
throat or chest scales, larger scales (smaller scale counts), and 
slightly different coloration: difference in the size and shape of the 
white scapular spots and the absence of the white streak near the 
back of the head in the male individual. 

These two individuals may possibly be hybrids between arm- 
strongi and S. brevirostratus enriquilloensis where the former meets 
the latter on descending to the lowlands. It is also possible that 
some other lowland form, such as 5. notatus randi, could be in- 
volved, instead, but this is considered more unlikely. Another 
possibility is that these represent a lowland race or species. 



20 BREVIORA No. 280 

Until more material can be examined, no conclusions about 
these specimens can be arrived at. The paratype (AMNH 51469) 
has the following counts: standard distance 8, dorsal scales from 
rear of hind limb to front of fore limb 46; scales around body at 
midbody 61; lamellae under fourth toe 10. The corresponding 
counts for MCZ 57820-1 are: 10, 42-43, 52, 10, the figure for 
57821, on the last count, being doubtfully 10. 

The type and paratype had head and body lengths of 27 and 28 
mm, respectively (both male), while the two referred specimens, 
in numerical order, had 21 and 22 for the same measurement (male 
and female). The first two measurements are those of Noble and 
Hassler. 

Material examined: All from the DOMINICAN REPUBLIC. 
Barahona. 5 kilometers south of Cabral: MCZ 57820-1; on 
mountain top on property of G. Herrmann, 2400 feet, near Paradis: 
AMNH 51469 (paratype). 

Identification aids. The white scapular spots will separate 
armstrongi from all others in the Hispaniolan no tat us group (female 
color unknown) except the two MCZ specimens discussed here. 

Sphaerodactylus altavelensis Noble and Hassler 

Sphaerodactylus altavelensis Noble and Hassler, 1933: 7, Alta Vela Island, 
Dominican Republic. 

This form appears related to S. notatus randi. There is great 
similarity in color pattern, the main difference being a decidedly 
more definite arrangement of head markings in the female and 
immature coloration of altavelensis. Scale counts of the two forms 
are close, but those around the body appear to average higher in 
altavelensis. However, altavelensis has distinctly keeled throat 
scales, unkown in any race of notatus. This difference seems to me 
sufficient to maintain altavelensis as a full species. S. notatus 
randi appears to differ from S. notatus difficilis chiefly in coloration, 
the coloration in fact being more different between randi and 
difficilis than between the former and altavelensis. 

The head and body length of MCZ 45947, a female paratype, is 
29 mm, while Noble and Hassler give the length of the type of 
altavelensis as 26.5 mm, whereas the largest randi (the type) was 
29 mm for this measurement. The scale counts follow: scales in 
standard distance 8-9; dorsal scales from rear of hind limb to 
front of fore limb 39-43; scales around body at midbody 54-56; 
lamellae under 4th toe 11-13. 



1968 SPHAERODACTYLUS NOTATUS GROUP 21 

Material examined: DOMINICAN REPUBLIC. Pedemales. 
Alta Vela Island: MCZ 45946-7 (paratypes). 

Identification aids. The only form with keeled throat scales with 
which there is overlap in the scale counts (except standard dis- 
tance) is noblei. The latter is distinguished by its blunt snout and 
distinctive ventral coloring and smaller size (see noblei). 

The next form is named for William J. Clench, the collector, 
who has always had a special interest in the West Indies. 

SPHAERODACTYLUS CLENCHI sp. nOV. 

Sphaerodactyliis difficilis: Mertens, 1939: 43 (part); Cochran, 1941: 115 
(part), including plate on page 116. 

Holotype: MCZ 43706, a male, from Samana (= Santa Barbara 
de Samana), Samana. Dominican Republic, collected by William 
J. Clench, August 1937. 

Paratypes: DOMINICAN REPUBLIC. All from Samana. With 
the same data as the type: MCZ 43707-9, 57151-65, USNM 
107621-30; the same locality, Samana (Santa Barbara de Samana) : 
AMNH 40630 and 19 untagged examples, William G. Hassler, 
October-December 1929, AMNH 40110, 90167, J. King, 1929; 
larger of three islands near Las Flechas: AMNH 41987, William 
G. Hassler, November 9, 1929; Rio San Juan: USNM 74942, Ger- 
rit S. Miller, Jr., March 1927; "Samana and Lasuna": USNM 
65781, W. L. Abbott, March 1923; Sanchez: CM''8169, FMNH 
5963 (4 specimens), Emil Kaempfer, March 1924; just Samana 
Province (no more exact locality): USNM 75001-2, Gerrit S. 
Miller, Jr., February 22, 1925, AMNH 49882, William G. Hass- 
ler, July 28, 1935; Samana, whether city or province unknown: 
MCZ 5451, M. A. Frazar, June 1882. 

Diagnosis. Apparently most closely related to S. caicosensis 
Cochran from which it differs in having a longer snout, a larger 
number of lamellae under the 4th toe (overlap in one specimen of 
caicosensis, i.e. 12-14 [15] in 63 specimens of clenchi as compared 
with 9-11 [12] in caicosensis); it also differs in coloration: both 
sexes of clenchi are variously spotted and marked with darker and 
sometimes lighter, while female caicosensis is barred, and markings 
are wholly or mostly confined to the head in the male. The new 
form is less closely related to S. notatiis difficilis but has been con- 
fused with that form, from which it differs in much smaller scales 
and in coloration. 



22 BREVIORA No. 280 

Description. (Paratype variation in parentheses.) Snout of 
moderate length and acuteness; distance of eye from tip of snout 
greater than distance of eye to ear; rostral with median groove; 
nostril between supranasals separated by one scale (one specimen 
with them in contact, three with them separated by two scales); 
three large and one small supralabial to below center of eye; head 
above and on sides covered with small juxtaposed keeled scales, 
larger and mostly keeled on snout; scales on dorsum enlarged, 
elongate, keeled, and subimbricate; the more granular scales of the 
neck appear anterior to the forward level of the shoulder, the 
dorsals, however, decidedly smaller than those located more pos- 
teriorly at the point just mentioned (also in paratypes); scales in 
standard distance 14 (11-14, subadults sometimes going to 16); 
52 dorsal scales between rear of hind limb and front of shoulder 
(45-60); 62 scales around body at midbody (53-68); 13 lamellae 
under the 4th toe ( 1 2- 1 4 [ 1 5 in one specimen ] ) ; mental somewhat 
larger than rostral; two large infralabials followed by a small one 
to center of eye; two small, rather irregularly shaped scales behind 
the mental; behind these the scales decrease in size posteriorly to 
throat, increase in size on throat and chest and again on belly; 
smooth, except some very slight keeling on throat (also in some of 
the paratypes); tail in type regenerated, that of paratype MCZ 
43707 apparently not, showing scales like those of dorsum, more 
imbricate and somewhat larger near base, not keeled near terminus; 
below, with scales like those of belly, but larger. 

Coloration in alcohol. Above, light brown, heavily dotted, 
spotted, and vermiculated with dark brown including head, limbs 
and tail. Below, brownish white, underside of head and throat 
dotted with dark brown, elsewhere slightly washed or marked with 
the same color, especially at the sides of the belly. 

In the paratypes, both above and below, often less heavily 
marked than the type, some of both sexes being virtually unmarked. 
Above, female and immature specimens tend to have the spots or 
dots on the neck and posterior part of the head arranged in longi- 
tudinal rows. Both sexes sometimes have whitish spotting on the 
body and one or more short longitudinal streaks of the same color 
starting at the posterior part of the eye and extending on to the 
neck. 

Measurements (in mm). Type, MCZ 43706: length, head and 
body, 33; tail regenerated. Largest paratypes, duplicates with 
AMNH 40630 (2) : head and body, 34; tail broken off and missing. 



1968 SPHAERODACTYLUS NOTATUS GROUP 23 

Remarks. At Sanchez, both the new form and S. notatus 
difficilis occur. 

It was first thought that the two specimens (MCZ 44395-6) 
from Sanchez might be intermediate between clenchi and difficilis; 
the scale counts of both seemed rather high. The higher dorsal 
count from rear of hind limb to front of forelimb, at 43, was one 
higher than the highest for other difficilis, while the higher count 
around the body, at 54, was just equaled by other difficilis. The 10 
for the standard distance of both these examples is one higher than 
the highest for the other difficilis. However, these two show typical 
difficilis coloration (male and female), and the examples of clenchi 
from Sanchez appear to be typical. There is, therefore, no real 
evidence of hybridization or intergradation. S. caicosensis appears 
to be the nearest relative of clenchi but presents enough differences 
to be regarded also as specifically distinct. 

Since what is known as the Samana Peninsula is at best in- 
completely joined to the main island of Hispaniola even now, and 
formerly, even in historic times, was completely separated from it, 
there is an evident bar to the dispersal of clenchi, especially in 
more modern times. See Chapman Grant (1956:89). 

Mertens (1939:43), under the heading of Sphaerodactylus 
difficilis, notes the smaller scales of Samana specimens as compared 
with the other specimens discussed. However, he did not believe 
he had sufficient material to name any of the populations discussed. 

Identification aids. This form is characterized by its small scales, 
showing but little evidence of overlap in scale counts with other 
forms of the group. Most overlap is shown with some of the group 
with keeled throat scales. 

Sphaerodactylus samanensis Cochran 

Sphaerodactylus samanensis Cochran, 1932: 183, Boca del Infierno, Samana 
Bay, Dominican Republic. 

Jay M. Savage (1954:328) regarded samanensis as a synonym 
of S. n. difficilis. However, what Savage regarded as difficilis has 
been here described as yet another race of notatus (savagei). No 
real scale differences between difficilis and savagei have been found, 
the differences being chiefly in coloration and in the shape of the 
male escutcheon. S. samanensis, on the other hand, differs from 
savagei in having the supranasals in contact, an extension rear- 
ward of the dorsal neck granules to the level of the insertion of the 
fore limb, and paired dorsal bands in the female (the only sex 
known). The scale counts for the three paratypes viewed are as 



24 BREVIORA No. 280 

follows: in standard distance 8-9; dorsal scales from rear of hind 
limb to front of shoulder 36-38; scales around body at midbody 
44-47; lamellae under 4th toe 12-13. The largest paratype had a 
head and body length of 28 mm, one more than Miss Cochran's 
measurement for the type. 

Material examined: DOMINICAN REPUBLIC. Seibo. Boca 
dellnfierno: USNM 74971-3 (paratypes). 

Identification aids. This form can be distinguished from all 
others treated here by having its supranasals in contact (a rare 
variation in S. n. difficilis), an extension rearward of the dorsal 
neck granules to the level of the shoulder, and the regular paired 
bands in the female (the only sex known). The bands of S. n. 
juanilloensis suggest these but are not evenly formed. 

Key to the Sphaerodactylus notatus group in Hispaniola 

1. Ventral scales smooth 6 

Ventral scales smooth except for keeled scales on throat or on throat 

and chest 2 

2. Scales around middle of body 40-47 3 

Scales around middle of body 50-61 4 

3. Throat scales slightly keeled over a limited area; lamellae under 4th 

toe usually 9-10; no scapular spot; maximum head and body length 

26 mm brevirostratus enriqnilloensis 

Throat scales well keeled over a considerable area; lamellae under 4th 
toe II; large, black square scapular spot; maximum head and body 
length 31 mm (only known from one male) laze Hi 

4. Scales around middle of body 50-56, only throat scales keeled 5 

Scales around middle of body 61; chest scales as well as throat scales 

keeled; a curved, white streak on back of head; two white spots on 
dorsum at level of shoulders (female color unknown) . .armstrongi^ 

5. Dorsal scale count from rear of hind limb to front of shoulder 35-41; 

lamellae under 4th toe 9-11; throat black, spotted or marked with 
white, intensity variable; maximum head and body length 24 mm 

noblei 

Dorsal scale count from rear of hind limb to front of shoulder 39-43; 
lamellae under 4th toe 11-13; throat whitish, speckled with dark 
brown; maximum head and body length 29 mm altavelensis 

6. Dorsal scale count from rear of hind limb to front of shoulder 25-45; 

scales around middle of body 37-54 7 

Dorsal scale count from rear of hind limb to front of shoulder 45-60; 
scales around middle of body 53-68 clenchi 



1 Not included in key are two specimens referred with doubt to anr^- 
strongi. See under species heading the account of their variation. 



1968 SPHAERODACTYLUS NOTATUS GROUP 25 

7. Dorsal coloration without paired bands (unpaired bands in immature 

of savagei), or if paired, separated by pale color; supranasals usually 
separated, dorsal neck granules not extending to level of front of 

fore limb 8 

Dorsal coloration with paired bands (male coloration unknown), 
supranasals in contact, dorsal neck granules extending to level of 
front of fore limb samanensis 

8. Usually well-defined, dark, longitudinal streaks on venter, curved heart- 

shaped streak on back of head (like top of heart with the possible 
apex toward the snout), pale color inside this marking, sometimes 
a second, less distinct streak, following first. In males these head 
markings may become entirely obsolete, reduced to a few specks; 

scapular spot quite persistent even in males b. brevirostratiis 

No longitudinal ventral streaks; if a curved streak on back of head, 
this not heart-shaped or enclosing a lighter area; scapular spot present 
or absent 9 

9. Above virtually unmarked, vestige of scapular spot (male coloration 

only known); snout very short; maximum length of head and body 

26 mm darlingtoni 

Above variously spotted, marked and banded with darker; snout not 
very short; maximum lengths of head and body 29-33 mm 10 

10. Above heavily spotted, banded or washed with darker, male sometimes 

unmarked, with light head spotted with dark; female also sometimes 
with curved band on back of head, sometimes melanistic without 

markings 11 

Three longitudinal dark streaks on dorsal aspect of head and neck 
in female. Body of male usually rather obscurely speckled with 
darker, and head unmarked. Sometimes head lighter, marked with 
darker or speckles both on head and body notatits difficilis 

11. Scales, rear of hind limb to front of fore limb 36-44; scales around 

body at midbody 48-54 notatiis randi 

Scales, rear of hind limb to front of fore limb 25-34; scales around 
body at midbody 36-48 12 

12. Scales around body at midbody 36-44; heavily spotted above includ- 

ing head. Females with some development of obscure head stripes. 

Immatures with bars on back notatits savagei 

Scales around body at midbody 40-48; females often showing paired 
crossbars separated by lighter on back, two head stripes joining 
on rear of head. Males spotted above, showing tendency for spots 
to form into crossbars notatus jnanilloensis 



26 BREVIORA No. 280 

REFERENCES CITED 

Barbour, Thomas 

1914. A contribution to the zoogeography of the West Indies, with 

especial reference to amphibians and reptiles. Mem. Mus. 

Comp. Zool., 44(2): 209-359, 1 pi. 
1921. Sphaerodactylus. Mem. Mus. Comp. Zool., 47(3): 217-278, 

pis. 1-26. 

Cochran, Doris M. 

1932. Two new lizards from Hispaniola. Proc. Biol. Soc. Washing- 
ton, 45: 183-188. 

1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus., 177: 
1-398. 

Grant, Chapman 

1952. Sexual and growth changes in the pattern of the gecko Sphaero- 
dactylus difficilis Barbour. Copeia, 1952: 187-188. 

1956. Report on a collection of Hispaniolan reptiles. Herpetologica, 
12: 85-90. 

Mertens, Robert 

1939. Herpetologische Ergebnisse einer Reise nach der Insel Hispani- 
ola. Westindien. Abh. Senck. Naturf. Ges., 449: 1-84, pis. 1-10. 

Noble, Gladwyn K. and William G. Hassler 

1933. Two new species of frogs, five new species and a new race of 
lizards from the Dominican Republic. Amer. Mus. Novit., No. 
652: 1-17. 

Savage, Jay M. 

1954. Notulae Herpetologicae 1-7. Trans. Kansas Acad. Sci., 57 (3): 
326-334. 

Schwartz, Albert 

1966. Geographic variation in Sphaerodactylus nolatus Baird. Rev. 
Biol. Trop.. 13: 161-185. 

(Received 6 January 1967.) 



1968 



SPHAERODACTYLUS NOTATUS GROUP 



27 







Fig. 1. A, Sphaerodactylus notatiis difficilis, MCZ 78733, Licey Almedio, 
Prov. Santiago, Dominican Republic. B, S. notatns notcitus, MCZ 31638, 
Key West, Florida. C, S. notatns jnanilloensis holotype, MCZ 73901, 
Juanillo, Prov. Altagracia, Dominican Republic. D, S. brcvirostratns hrcvi- 
rostratns holotype. MCZ 63234. 5 km S of Dufort, S of Leogane, Dept. de 
rOuest, Haiti. 



28 



BREVIORA 



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BREVIORA 



MiaseiuaM of Coiraipsirative Zoology 



Cambridge, Mass. 



5 April, 1968 



Number 281 



ADDITIONAL NOTES ON BATOID FISHES FROM 
THE WESTERN ATLANTIC 

Henry B. Bigelow and William C. Schroeder 



Introduction 

Order Batoidei . 

Suborder Torpedinoidea . 
Family Torpedinidae . 

Torpedo nobiliana . 

Diplobatis pictus . 
Suborder Rajoidea 
Family Rajidae . 

Raja bullisi . 

Raja cervigoni . 

Raja clarkii . 

Raja floridana . 

Raja fuliginea . 

Raja lentiginosa . 

Raja olseni . 

Raja teevani . 

Breviraja plutonia . 

Breviraja sinusmexicanus 

Breviraja spinosa . 

Cruriraja riigosa . 
Family Pseudorajidae . 

Pseudoraja atlantica . 
Family Anacanthobatidae 

A nacan thobatis american us 

Anacanthobatis longirostris 

Springeria jolirostris . 

References cited .... 



Page 



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18 

18 

18 

19 

20 

22 



'Contribution No. 1875 from the Woods Hole Oceanographic Institution. 



BREVIORA No. 281 



INTRODUCTION 



During the last fifteen years we have received for study a large 
number of interesting batoids, together with some cyclostomes, 
sharks, and chimaerids, trawled by U.S. Fish and Wildlife vessels, 
chiefly the "Oregon" and the "Silver Bay" but also the "Combat," 
"Pelican," and "George M. Bowers." Most of these were taken in 
the Gulf of Mexico, the Caribbean, and along the coast of South 
America as far south as 01°49'N off the mouth of the Amazon, 
with a few from along our eastern coast to the southward of Cape 
Hatteras. Bottom trawling depths ranged from 40 to 750 fathoms 
(65-1370 m), chiefly between 100-500 fathoms (183-915 m). 
Many new species are included in our various reports from 1951 
to 1965, listed in the references cited. 

The present account includes eighteen species of batoids, none of 
them new, but they yield additional information as to range, depths 
of capture, size, etc. They were taken at 95 "Oregon" and 10 
"Silver Bay" stations which fall within the following depth ranges: 
10 at 40-100; 11 at 101-200; 60 at 201-300; 13 at 301-400; 8 at 
401-500; and 3 stations at 501-600 fathoms, ranging from northern 
Florida to Venezuela, including the Gulf of Mexico. Also included 
is a summary of previously published capture records for each 
species. 

We thank the various investigators who collected and preserved 
these specimens, and Harvey R. Bullis, Jr., of the U. S. Fish and 
Wildlife Service for placing them at our disposal. 

Order BATOIDEI 

Suborder TORPEDINOIDEA 

Family TORPEDINIDAE 

Torpedo NOBiLiANA Bonaparte 1835 

Our recent accounts (1962, 1965) of this species include the 
capture of 12 specimens, ranging in length from 211 to 700 mm, 
trawled in various localities ranging from North Carolina to the 
Caribbean coast of Panama (9°00'N, 81°23'W), including the 
Gulf of Mexico, in depths of 10 to 290 fathoms ( 18 to 530 m). It 
is known in the western Atlantic from as far north as the offing 
of southern Nova Scotia and in the eastern Atlantic from northern 
Scotland to the Mediterranean, Azores, Madeira, and tropical 
West Africa. 



1968 ADDITIONAL BATOID FISHES 3 

The present collection includes 12 specimens, 205 to 393 mm 
long, 3 from the east coast of Florida, 1 from the east coast of 
Hispaniola, and 8 from off the coast of Venezuela, in 40 to 240 
fathoms (73 to 440 m), "Oregon" stations 4393, 4394, 4401, 
4402, 4418, 5037, 5101, 5109 and "Silver Bay" stations 5188, 
5530. 

An examination of a number of recently obtained specimens, 
necessitates a modification of two descriptive characters given in 
Fishes of the Western North Atlantic, Part 2 (Bigelow and 
Schroeder, 1953, p. 94), stated as "disc appreciably shorter (less 
than 80 per cent) than its breadth" and "height of caudal fin not 
greater than distance from its own upper origin to origin of first 
dorsal." 

Four males 215-900 mm long and seven females of 211-613 mm 
have a disc length ranging from 77 to 90 per cent of the disc width, 
only two individuals falling below 80 per cent. The height of the 
caudal is greater, 1.10 to 1.38 times, than the distance from the 
upper origin of the caudal to the origin of the first dorsal on 10 
specimens and less, 0.95 times, on only one. 

In color, this ray has been described as dark chocolate to purplish 
brown above, either uniform or with a few obscure darker spots, 
and white below, but with the edges of the disc and pelvics of the 
same hue as the upper surface, the tail with irregular dark markings. 
Our specimens likewise are so colored above but the markings 
below on several are grayish. It is reputed to reach a weight of 
about 200 pounds. 

DiPLOBATis PiCTUS Palmer 1950 

The type of this species was taken off Georgetown, British 
Guiana, a female 137 mm in total length. Our recent account 
(1962) includes 24 specimens, 77-164 mm long, from the mouth of 
the Amazon 02°29'N to British Guiana 09°3rN in 9-50 fathoms 
(16-92 m). 

The present collection includes 3 males of 85-159 mm and 3 
females of 140-196 mm from 12°19'-12°37'N, 70°34'-71°10'W, 
off the mouth of the Gulf of Venezuela, in 40-65 fathoms (73-119 
m), "Oregon" stations 4393, 4394, 4401, 4402. 

In our 1962 account we mentioned certain characters that 
supposedly differentiated D. pictus from D. guamachensis Martin 
1957. Thus we stated that in 21 specimens of pictus, 11 to 164 mm 
long, the width of disc ranges from 38.8 to 50.0 per cent, the length 



4 BREVIORA No. 281 

of disc from 38.8 to 48.0 per cent, and the breadth of pelvics from 
28.7 to 36.6 per cent of total length; also that the lateral folds 
originated between the rear end of the first dorsal base and the 
origin of the second dorsal. In a personal communication received 
from Dr. Martin in 1959, based on 4 specimens of guamachensis 
56.5-70.9 mm long (including the type), from the Gulf of Cariaco, 
Venezuela, the above proportions are given as 53.0 to 58.1, 50.3 
to 54.5, and (on two specimens) 23.8-26.0, respectively, the 
lateral folds originating "immediately behind the origin of first 
dorsal." 

The color and markings on pictus vary considerably and our 
earlier collection does not include any with the design of guama- 
chensis with its irregular dark bars and spots. However, in our 
present collection of 6 specimens, two males 85 and 196 mm long, 
respectively, are marked much like Martin's illustration, though 
somewhat more elaborately. This variation in the markings on 
pictus recalls that of Narcine brasiliensis. Furthermore, they fall in 
line with the proportional dimensions of all our other pictus and 
also confirm that the lateral folds originate between the dorsal fins 
and not immediately behind the origin of the first dorsal. We 
therefore consider D. guamachensis a synonym of D. pictus. 

The known range of this species now extends from the offing of 
the Gulf of Venezuela to that of the Amazon River and in depth 
from 9 to 65 fathoms (16-119 m). 

Suborder RAJOIDEA 

Family RAJIDAE 

Raja bullisi Bigelow and Schroeder 1962 

Specimens previously recorded (1962, 1965), 166 to 478 mm 
long, range from Dry Tortugas, Florida, 24°36'N, to the coast of 
Surinam, 07°15'N, in 110-300 fathoms (201-549 m). 

Three specimens are included in the present collection: a 205 
mm female from off Santa Marta, Colombia, 11°08'N, 74°23.8'W, 
in 100 fathoms ( 183 m), "Oregon" station 4856; a 280 mm female 
11°31'N, 60°5rW, in 240-250 fathoms (440-457 m), station 
5029; and a 370 mm male 11°36.5'N, 62°46.5'W, in 200-240 
fathoms (366-440 m), station 5037, from the offing of the Gulf 
of Paria, Venezuela. 

Bullisi closely resembles Raja teevani but its persistent nuchal 
thorn, somewhat broader tip of snout, and somewhat greater space 
between the dorsals (0.23 to 0.45 times the first dorsal base) 



1968 ADDITIONAL BATOID FISHES 5 

distinguish it from the latter which lacks a nuchal thorn, has a very 
sharp tip of snout, and dorsals that are confluent or with an inter- 
space seldom exceeding 0.15 times the first dorsal base. 

Raja cervigoni Bigelow and Schroeder 1964 

Previous records of this species include: an immature male of 
357 mm, the holotype, from 10 miles (16 km) northeast of 
Carupano, in 20-30 fathoms (37-55 m), Museo Historia Natural La 
Salle, Venezuela, No. 873; a male of 206 mm, and a female of 229 
mm from Punta Araya, Estado Sucre, in about 20 fathoms (37 m), 
off the eastern part of Venezuela; and a male of 235 mm from off 
the Guianas, 07°25'N, 54°35'W, in 75-80 fathoms (137-146 m), 
"Oregon" station 2289. 

The present collection includes 2 males, 144 and 506 mm long, 
respectively, the latter mature, and a female of 418 mm, from off 
the Peninsula de Guajira, Venezuela, 12°29'N, 71°54'W, in 95 
fathoms (174 m), "Oregon" station 5685. 

Description of mature male. Proportional dimensions in per 
cent of total length. 

Disc. Extreme breadth 69.5; length 50.2. 

Snout length. In front of orbits 10.7; in front of mouth 13.5. 

Orbits. Horizontal diameter 4.4; distance between 4.2. 

Spiracles. Length 2.8; distance between 6.1. 

Mouth. Breadth 8.1. 

Exposed nostrils. Distance between inner ends 7.9. 

Gill openings. Length 1st 1.7; 3rd 1.7; 5th 1.5; distance between 
inner ends, 1st 13.8; 5th 7.3. 

First dorsal fin. Height 2.8; length of base 4.9. 

Second dorsal fin. Height 2.8 length of base 4.9. 

Pelvics. Anterior margin 12.1. 

Distance. From tip of snout to center of cloaca 46.5; from 
center of cloaca to 1st dorsal 36.5; to tip of tail 53.5; from rear 
end of 2nd dorsal to tip of tail 3.4. 

Interspace. 1st and 2nd dorsals 3.8. 

Disc 1.38 times as broad as long; maximum angle in front of 
spiracles 99°; anterior rays of pectorals extending 50 per cent of 
distance from level of orbits toward tip of snout, the latter pro- 
jecting; rostral process firm, extending to tip of snout; anterior 
margins of disc slightly convex in front of spiracles, thence concave 
and straight toward outer corners which are sharply rounded, the 
width of disc across anterior edge of orbits 25 per cent of total 
length of specimen; posterior margins and corners and inner 



6 BREVIORA No. 281 

margins all rounded. Axis of greatest breadth 71 per cent of 
distance back from tip of snout to axils of pectorals. Tail rather 
slender, the lateral folds low down, narrow, originating a little in 
advance of tips of pelvics, reaching tip of tail; length of tail from 
center of cloaca to origin of first dorsal 0.78 times as great, and to 
its tip 1.15 times as great as distance from center of cloaca to tip 
of snout. 

A row of 9 thorns along anterior and inner margins of one orbit, 
10 along the other, and 3 minute thorns opposite inner margin of 
each spiracle with several small ones over tip of rostrum. Three 
prominent thorns along midline in the nuchal region, followed by a 
space and the next thorn, a tiny one, opposite the axils of pectorals, 
with 6 more to opposite tip of anterior pelvic lobe, then 22 thorns 
alternating large and small in an unbroken series to the first dorsal 
fin; 2 thorns in front of second dorsal. A row of 21 thorns low 
down each side of the midrow on tail, more or less alternating in 
size, beginning a little in advance of tips of posterior pelvic lobe and 
reaching to opposite origin of first dorsal. Malar thorns prominent, 
sharp, backward pointing, each patch consisting of 3 to 4 rows 40 
mm long and 16 mm wide. Alar thorns well developed, each patch 
about 60 mm long and 13 mm at widest, the rows irregular, as 
many as 5 rows anteriorly reducing to a single thorn posteriorly. A 
band of fine prickles along edge of disc from opposite posterior 
malars to outer angle. Upper surface otherwise smooth. Lower 
surface with a patch of prickles on end of snout merging with a 
narrow band of prickles along edge of disc extending to about 
opposite mouth; otherwise smooth below. 

The claspers are massive, extending 75 mm beyond the tips of 
pelvics. 

Color. Upper surface plain medium brown. A prominent ocellus 
on each side of disc, situated a little posterior to the greatest axis of 
disc, its center from the midline of disc a distance about equal to 
that from tip of snout to center of orbit; distance between centers of 
ocelli 133 mm, and between centers of ocelli and centers of orbits 
127 mm. The ocelli have a pale center surrounded by a roundish 
area of solid dark brown the diameter of which is about 9 to 1 1 
mm, thence a pale area surrounded by a narrow, somewhat broken 
dark brown circle about 21 mm in diameter and finally with an 
outside narrow pale circle. Lower surface of disc and pelvics 
margined with a band of pale grayish, the tail with brownish 
blotches. 



1968 ADDITIONAL BATOID FISHES 7 

The 144 mm male has a disc 1.45 times as broad as long; 
maximum angle in front of spiracles 109°; contour of disc about 
the same as the 506 mm male, except that width across anterior 
edge of orbits is 34.7 per cent of total length of specimen; axis of 
greatest breadth 72.6 per cent; length of tail to origin of first dorsal 
0.72 times as great and to its tip 1.08 times as great as distance 
from center of cloaca to tip of snout. Two thorns along anterior 
margin and 1 at inner posterior margin of each orbit; 1 nuchal 
thorn; a row of 16 large, sharp, thorns in midline, evenly spaced, 
from opposite axils of pectorals to first dorsal and 2 between dor- 
sals; a row low down each side of midrow, from opposite tip of 
pelvics to interdorsal space, 11 on one side 12 on the other, in size 
about the same as those in midrow; otherwise smooth above. 
Smooth below except for a single row of 6 to 7 very small spines 
along edge of disc between tip of snout and nostrils. Claspers not 
reaching tip of pelvics. The distance of the center of each ocellus 
from the midline of disc is equal to that from tip of snout to an- 
terior margin of orbit; distance between centers of ocelli 40 mm 
and between centers of ocelli and centers of orbits 35.5 mm. The 
ocelli have a dark brown center and an outside solid dark brown 
circle 6 mm in diameter, with a pale area between and another 
surrounding the colored areas. The lower surface is marked with 
a broad outer band of gray from opposite mouth to axils of pec- 
torals, its inner margin irregular, and with a pair of irregular grayish 
blotches at the origin of the anterior pelvic lobes and on the tips 
of the posterior lobes, the tail with two vague grayish areas pos- 
teriorly. 

The 418 mm female agrees closely with the small male in pro- 
portional dimensions. The ocular and nuchal thorns are of the 
same number as on the 506 mm male but the midline thorns are 
more numerous, there being 6 very small ones from opposite the 
ocelli to the axils of pectorals followed by 30, alternating large and 
small, to the first dorsal, with 2 small thorns and 1 large between 
the dorsals. The side row contains about 35 thorns extending from 
near axils of pectorals to opposite the second dorsal, more or less 
alternating in size, but the posterior 5 or 6 very small. The prickles 
along the edge of disc anteriorly, above and below, are similar to 
those of the large male. The distance between the centers of the 
ocelli is 1 17 mm and between centers of ocelli and centers of orbits 
98 mm. Color above and below similar to the large male. The 
tooth count on the 7 specimens thus far examined ranges from 

3 2 '■^ 4 0* 



8 BREVIORA No. 281 

Raja clarkii Bigelow and Schroeder 1958 

Our previous records of R. clarkii (1958, 1962, 1905) include 
a total of 4 males 228-665 mm long, and 4 females of 176-747 mm. 
Four were trawled in the northern part of the Gulf of Mexico, 3 
off the coast of Nicaragua and 1 off Panama, in depths ranging 
from 200 to 300 fathoms (366-549 m). 

The present collection includes 3 males of 330-710 mm from the 
north central part of the Gulf of Mexico, "Oregon" stations 4701, 
4703, 4716; 6 males of 265-525 mm and 2 females of 256-370 
mm from off Colombia, stations 4841, 4882, 5689, 5690, 5722; 
and a male of 351 mm and 2 females of 250-275 mm off Venezuela, 
stations 4438, 4456. These stations range in depth from 220 to 
500 fathoms (400-915 m). 

This species is characterized among western Atlantic rajids by the 
presence of a band of formidable and very sharp thorns extending 
along the margin of the lower surface from the tip of the snout 
almost to the outer corners of the disc. An amplification of pre- 
vious descriptions follows. 

The median row of thorns, which is continuous from the nuchal 
region to the first dorsal fin, number from 30 to 43 of which 4 or 5 
are in the nuchal-scapular area followed by about 7 to opposite 
axils of the pectorals. The total number of thorns does not nec- 
essarily increase with the size or age of the skate, for a male and 
a female, 256 and 324 mm long, respectively, have 43 thorns 
while a male and a female of 710 and 747 mm have only 37-38 in 
the median row. There is a triangular patch of thorns in the nuchal- 
scapular area which may have but 1 thorn on each side of the mid- 
row or as many as 4 or 5. In arrangement, the orbital thorns on 
most of our specimens, including the largest, consist of 2 anterior, 
1 posterior, and from 1 to 3 aside the spiracles. 

The color of the upper surface of all the specimens we have seen 
is brown with darker punctulations scattered over the disc, pelvics, 
and more or less on the tail. Some have conspicuous white spots 
while others lack them. Thus 5 specimens, 250-747 mm long, have 
3 pairs of spots, 8 of 265-580 mm have 1 pair, and 8 of 176-525 
mm lack them. One specimen of 710 mm has 1 pair anteriorly and 
1 spot on one side posteriorly. It is noteworthy that the spots are 
located in the same parts of the disc. Thus the anterior pair is 
opposite the spiracles, the median pair, slightly closer together, 
opposite the scapular region, and the posterior pair a little in ad- 
vance of axils of pectorals and about in line with the anterior pair. 
Where but one pair is present it is located anteriorly. In shape the 



1968 ADDITIONAL BATOID FISHES 9 

spots may be barlike, oval, or spherical, and in size, if roundish, 
from less than to about equal to the area of the orbit, or, if barlike, 
about as long or longer than the diameter of the orbit. The lower 
surface is white with a broad grayish band, its inner margin ir- 
regular (Bigelow and Schroeder, 1958, fig. 9; 1965, fig. 1) ex- 
tending on most specimens from a little in advance of the outer 
angles of disc to the axils of the pectorals and along the rear margin 
of the pelvics, but on several this band originates near the tip of 
the snout. Also, a few small gray spots or blotches may be present 
here and there on the disc, including one each side of the cloaca. 

The largest male, 710 mm in length, is approaching maturity as 
the claspers are well developed, extending 56 mm beyond the tips of 
the pelvics, and a few of the alar thorns are exposed. 

The present known range of this species is from the northern 
part of the Gulf of Mexico to the offing of Colombia (12°30'N, 
72^08'W) in 200-500 fathoms (366-915 m). 

Raja floridana Bigelow and Schroeder 1962 

The original account of this species includes 27 specimens 158- 
448 mm long, and 1, the holotype of 772 mm, trawled between 
Cape Lookout, North Carolina, and Dry Tortugas, Florida, 34° 21' 
to24°17'N, in 170-225 fathoms (31 1-412 m). '' 

The present collection includes 32 specimens, 145-585 mm long, 
males and females about equally divided, taken at 12 "Oregon" 
stations (5097, 5098, 5100, 5101. 5102, 5106, 5107, 5113, 5119, 
5234, 5295, 5482) and "Silver Bay" station 5454. These fall 
within a small area between the offings of Cape Kennedy and St. 
Augustine, Florida, 28°00'-29°58'N, 79'51'-80°10'W, in 160- 
213 fathoms (293-390 m). 

In general appearance floridana resembles R. laevis Mitchill, 
1815, but it differs from laevis in a proportionately longer snout, 
wider disc, usually a shorter tail, and smaller dorsal fins. Thus, on 
6 males and 3 females of floridana 298-772 mm in total length, the 
distance from the tip of snout to the mouth is 19.5-24.9 per cent, 
the width of the disc 73.2-81.2 per cent, the length of the tail from 
center of cloaca 43.8-47.1 per cent, and the distance from origin of 
first dorsal fin to the rear end of base of second dorsal 8.4-12.1 
per cent of total length. These proportions on 2 males and 1 fe- 
male of laevis, 506-542 mm long, are 17.0-18.0 per cent, 68.0-71.0 
per cent, 49.7-51.7 per cent and 14.5-14.8 per cent, respectively. 
On 7 floridana 158-240 mm long the distance from snout to mouth 
is 18.0-21.1, length of tail 46.2-50.5 and there are 8 to 10 tail 



10 BREVIORA No. 281 

thorns, whereas on 5 laevis of 177-195 mm these proportions are 
14.1-17.8 and 54.2-58.7 and the tail thorns 13 to 18. 

We are indebted to George C. Miller of the U. S. Fish and Wild- 
life Service for pertinent data on a collection of 17 specimens, 155- 
252 mm long, and 1 of 465 mm, all taken between the offing of 
South Carolina and the Straits of Florida in 180-220 fathoms (329- 
402 m), except one from 29°04'N, 88°31'W, "Oregon" station 
3724, the latter extending the range to the northern part of the 
Gulf of Mexico. Four of these specimens have tail lengths of 50.9- 
53.8 per cent of total length, a little greater than our 50.5. The 
tail thorns number 8 to 12 except on the one of 465 mm which has 
15. Three have 2 anterior orbital thorns and 1 posterior, the others 
1 anterior and 1 posterior. 

The armature of the upper surface, at least on sizes up to 585 
mm, is limited to the orbital thorns, a single midrow of rather in- 
conspicuous thorns from somewhat in advance of the axils of 
pectorals to the first dorsal fin, and to 1 or 2 or no thorns between 
the dorsals. But there is not necessarily a progressive increase in 
the number of thorns with the size of the skate. Up to a length of 
about 400 mm most of our specimens have 1 anterior and 1 pos- 
terior orbital thorn, but a few, including the smallest, have 2 
anterior and 1 posterior. One of 585 mm has 10 or 11, and in 
addition 2 aside each spiracle, while another of 488 mm has only 
5 or 6 thorns along the margin of each orbit with 1 aside each 
spiracle. 

The thorns in the midrow usually number from 8 to 14 on 
specimens up to a length of about 400 mm, but two of 432-435 mm 
have only 3 (there is evidence that some thorns were shed and 
others still embedded), while one of 585 mm has 23, the anterior 
13 being smaller than the posterior 10. The lower surface is 
smooth. 

The immature 772 mm holotype has 12 small thorns along the 
anterior and inner margins of each orbit with 3 very small ones in 
advance of these while the midrow contains 25 small thorns of 
assorted sizes, with a few rather large ones. The lower surface is 
densely prickly in advance of axis through the mouth, except for a 
nearly smooth area immediately in front of mouth. A band of 
prickles extends along the margin of the disc from the snout to 
about opposite the 4th or 5th gill openings, the remaining lower 
surface, including the tail, being virtually smooth. 

The maximum angle of the disc in front of spiracles ranges from 
81° to 94°, except 74° on the holotype. The teeth range in number 



1968 ADDITIONAL BATOID FISHES 11 

from 27 to 38 in each jaw, and have a low triangular cusp, even 
on the smallest specimens of both sexes. In color, the smallest 
specimens are light brown above, with vague dark brown irregular 
spots about half as large to as large as the orbit, and the tail has 5 
dark bars about equally spaced. The lower surface is whitish with 
a narrow dusky margin on the outer angles of disc. Our larger 
specimens lack the spots, and below are usually plain grayish white, 
some of them partly brownish. All, small to large, have dark-ringed 
mucous pores on the under surface, most numerous anterior to the 
axis through the mouth and immediately below the lower jaw. 

It is of interest that all of the many specimens so far collected 
have been taken within the rather limited depth range of 160-225 
fathoms (293-412 m), although many trawl hauls, both shoaler 
and deeper, have been made within the known latitudinal range 
of this species. 

Raja fuliginea Bigelow and Schroeder 1954 

UntU recently this species had been known only from the holo- 
type, a juvenile male 306 mm long, USNM No. 163367, taken at 
"Oregon" station 534, in the northwestern part of the Gulf of 
Mexico, 27°32'N, 93°02'W, in 400-450 fathoms (732-823 m). 
A second specimen, a female of 330 mm, was trawled at "Oregon" 
station 4147, about 45 miles southwest of Dry Tortugas, Florida, 
24°12'N, 83°32'W, in 500 fathoms (915 m) (Bigelow and 
Schroeder, 1965). 

We now have 14 more specimens, as follows: 9 from the western 
and northern parts of the Gulf of Mexico, "Oregon" stations 4701, 
4711, 4713, 4802; 2 off Isla del Rosario, Colombia, station 4883; 
1 from about 40 miles northwest of Caracas, Venezuela, station 
4449, and 2 off Curagao, station 4416, males and females evenly 
divided, total lengths 224-445 mm. The range in latitude is from 
27°42' to 10°15.5'N, in longitude from 96°00' to 67°38'W. All 
were taken in 500-600 fathoms (915-1098 m). 

In our original account (1954), fuliginea was compared with R. 
bathyphila Holt and Byrne 1908 which it most closely resembles 
in general arrangement of thorns and prickles, in proportional 
dimensions and in the very dark coloration of the lower surface of 
disc and tail. But fuliginea differs from bathyphila in a more obtuse 
anterior contour and especially in the fact that the entire lower 
surface of the tail, apart from a very narrow median stripe, is 
densely prickled from base to tip, whereas it is naked in bathyphila. 



12 BREVIORA No. 281 

Also, the prickles on the upper surface of disc and tail are coarser 
than on bathyphila and there are no naked areas. 

Proportional dimensions in per cent of total length of all 14 
specimens: 

Disc. Extreme breadth 43.7-51.8; breadth across anterior 
margin of orbits 24.6-28.6; length 37.5-49.0. 

Snout length. In front of orbits 9.1-10.3; in front of mouth 
10.0-13.2. 

Orbits. Horizontal diameter 3.4-4.6; distance between 2.7-3.6. 

Spiracles. Length 2.1-2.9; distance between 5.7-7.3. 

Mouth. Breadth 5.3-7.8. 

Exposed nostrils. Distance between inner ends 6.2-7.3. 

Gill openings. Length, 1st 1.4-1.8; 3rd 1.4-1.8; 5th 1.1-1.5; 
distance between inner ends, 1st 10.7-14.8; 5th 5.8-8.5. 

First dorsal fin. Height 2.5-3.6; length of base 4.6-6. 1 . 

Second dorsal fin. Height 2.5-3.5; length of base 4.5-6.3. 

Pelvics. Anterior margin 10.1-13.4; origin to tip 13.4-17.5. 

Distance. From tip of snout to center of cloaca 35.2-44.0; from 
center of cloaca to 1st dorsal 42.4-48.8; to tip of tail 56.0-64.8; 
from rear end of 2nd dorsal to tip of tail 2.7-4.5. Interspace: 1st 
and 2nd dorsals 0.0. 

In armature, the upper surface of the disc, tail, and posterior 
lobe of pelvics, is densely covered with small sharp prickles. The 
dorsals, caudal fin, and the skin over eyes also are prickly. The 
orbital thorns, in various combinations, range from 2 at the inner 
margin of each orbit to 5 or 6 at the anterior and inner margins. 
There may be, in addition, a thorn aside one or both spiracles, or 
this thorn may be lacking at both. All these thorns are usually 
rather prominent but are very small on several specimens. The 
thorns on the nuchal-scapular region are in most cases arranged in 
a triangular patch, with 2 to 4 in the midrow and 1 or 2 on each 
side of these, the exceptions being 3 on each side, or none, only 1 
thorn in the midrow and, on a male of 445 mm, all the nuchal- 
scapular thorns are lacking. The thorns in the midrow, including 
the nuchal region, number from 15 to 29, the fewest (15 and 19) 
being on the larger specimens, a male of 445 mm and a female of 
412 mm in total length, and the most (28 and 29) on a male of 
224 mm and a female of 293 mm. They may continue from the 
scapular region with no interruption, or after a short space, or 
resume as far rearward as opposite the axils of the pelvics, and 
extend to the dorsal fin or fall short of the dorsal a distance as much 
as that between the spiracles. In addition to the midrow there is 



1968 ADDITIONAL BATOID FISHES 13 

a siderow of thorns, usually irregular, beginning opposite the axils 
of the pectorals or as far rearward as about the tips of the pelvics 
and ending, when complete or nearly so, about opposite the origin 
of the first dorsal. But on some specimens the row runs out well in 
advance of the dorsal, and on one, a male of 428 mm, the row is 
virtually nonexistent, there being only 2 or 3 thorns on each side 
of the midrow. The thorns in the side row may be of the same size 
as those in the midrow, or somewhat larger or smaller. 

The lower surface of disc and pelvics is smooth but the tail is 
densely set with prickles or thornlets, similar to those of the upper 
surface, but on some specimens these are lacking or sparse along 
a very narrow median stripe. 

The color of the disc and pelvics is plain grayish above, ranging 
from light to rather dark. The tail may be of the same hue or some- 
what paler. Below, the disc and pelvics are dark gray (3 speci- 
mens), dark brown (7 specimens), or blackish brown (4 speci- 
mens) some with pale areas around the entrance to cloaca and/or 
on end of snout, around the jaws, and at tips of pelvics. The tail is 
of the same shade, or slightly darker than it is above. 

Raja lentiginosa Bigelow and Schroeder 195 1 

Many specimens of this skate, 80 to 435 mm in total length, have 
been trawled by the "Oregon" in the Gulf of Mexico, where it is 
widespread, southward to the offing of Nicaragua (11°27'N, 
83°11'W) within depths of 29-305 fathoms (53-558 m). The 
present collection includes 6 more of 226-418 mm, from the north- 
ern Gulf taken in 100-240 fathoms (183-439 m), "Oregon" 
stations 4614, 4616, 4696, 4705. 

There is a band of thorns along the median zone of back and 
tail, in 3 rows anteriorly, thence merging to 5 rows, and again in 3 
rows as the tail narrows. These thorns have been described as 
originating in the nuchal region but we have since seen many speci- 
mens on which the band begins posterior to the nuchal region as 
much as the distance from tip of snout to rear margin of orbits. In 
some instances the median row originates slightly in advance of the 
side rows, but usually it begins posterior to them. 

The upper surface everywhere is densely freckled with light to 
dark brownish and whitish spots, including the tail, pelvics and 
claspers, distinguishing lentiginosa from R. gannani (Whitley, 
1939) which it closely resembles, the latter having fewer spots 
which are grouped mostly in a distinct rosette pattern. 



14 BREVIORA No. 281 

The lower surface is white, variously marked with grayish which 
may be in the form of a group of blotches, mostly fused, on each 
pectoral, or a broad band with an irregular inner margin extending 
from opposite the first pair of gill openings to as far as the inner 
margin of the pectorals. Some have an elongate blotch on the 
claspers and a blotch on the anterior and/or posterior pelvic lobes. 
Tail white. 

In the original account of lentiginosa the second dorsal and the 
caudal fin are shown as confluent, but we have found a few speci- 
mens with a short space between these fins. 

Raja olseni Bigelow and Schroeder 1951 

Previous accounts (1951b, 1953, 1962) of this species include 

15 specimens, 151-568 mm long, trawled in 53-130 fathoms (97- 
238 m) at 9 stations by the "Oregon" and at 1 by the "Silver Bay." 
All were taken in the northern and northwestern part of the Gulf 
of Mexico. The present collection includes two males of 161-196 
mm, and a female of 335 mm, from "Oregon" station 4695, 
26°16'N, 90°13'W, in 50-53 fathoms (92-97 m), in the same 
region as above. We include this record to emphasize how restricted 
is the present known range of this species in spite of the many 
bottom trawl hauls made by the "Oregon" and other vessels in 
other parts of the Gulf and along the outer coast between North 
Carolina and the offing of the Amazon. 

Raja teevani Bigelow and Schroeder 1951 

Accounts of this species (1951b; 1965) include 6 males and 11 
females, 175-635 mm long, of which 4 were trawled off Pensacola, 
Florida (3 "Oregon" stations), 1 at the northwest edge of Great 
Bahama Bank, and 1 in Santaren Channel ("Silver Bay" stations), 
and 1 1 off the coasts of Honduras and Nicaragua, the latter between 
16°43' and 12°25'N (9 "Oregon" stations), in depths of 240 to 
400 fathoms (439-732 m). We now have 5 more, all males, 306- 
840 mm long, of which 4 were taken in the northwestern Gulf of 
Mexico ("Oregon" stations 4703, 4709, 4729, 4800) in 300-400 
fathoms (549-732 m) and 1 of 500 mm from off Riohacha, 
Colombia, at ir50'N, 73°05'W (station 4911), in 175-190 
fathoms (320-348 m). 

Teevani closely resembles R. floridana from which it differs in 
having fewer orbital thorns when about half grown and larger. 
Thus, on sizes up to about 600 mm there are, with few exceptions, 



1968 ADDITIONAL BATOID FISHES 15 

1 or 2 anterior thorns and 1 posterior, and on the largest, of 840 
mm, only 5 thorns along the margin of one orbit and 3 at the other. 
Whereas young floridana up to about 300 mm have 1 anterior and 
1 posterior thorn; at 450 mm there may be as many as 8 or 9 at 
each orbit and on one of 772 mm, the largest seen, there are 12 
along the anterior and inner margins and 3 more in advance of 
these. 

Minute prickles are present on the upper surface of the disc on 
teevcmi of all sizes, on some anteriorly only, but on others scattered 
all over the disc. They are lacking on our floridana, except for the 
large one of 772 mm on which the end of the snout is densely 
covered with coarse prickles and small thorns blending into minute 
prickles to about opposite the spiracles, the rest of disc smooth. 
On the lower surface of teevani prickles are present anterior to the 
mouth and in a band along the edge of disc from near the end of 
snout to about opposite the first pair of gill openings, on small in- 
dividuals as well as large, but floridana is perfectly smooth below, 
up to a length of at least 450 mm, though prickles are present 
anteriorly on the 772 mm specimen, 

Teevani and floridana v/ere in no case taken at the same station. 
The known range of the latter is from the offing of Cape Lookout, 
North Carolina to Dry Tortugas, Florida, in 160-225 fathoms 
(293-412 m). 

Breviraja PLUTONiA (Carman) 1881 

Many specimens of this species have been recorded (Bigelow 
and Schroeder, 1953, 1962) from the offing of Cape Lookout, 
North Carolina, and southward to an area centering about 30 miles 
southwest of Dry Tortugas, Florida, in depths of 160-400 fathoms 
(293-732 m), in size from 62 to 270 mm. The present collection 
includes 100 specimens 76-257 mm long, trawled between St. 
Augustine and Vero Beach, Florida, in 188-215 fathoms (344- 
393 m), "Oregon" stations 5075, 5077, 5089, 5097, 5115, 5231, 
5233, 5483, 5484, and 7 specimens west to southwest of Dry 
Tortugas in 190-310 fathoms (348-567 m), stations 4335, 4356, 
4558. One of the latter, 275 mm, is the largest we have seen. 

Breviraja sinusmexicanus Bigelow and Schroeder 1950 

All the previous captures of this skate have been recorded (1950, 
1953, 1962, 1965) from the northern part of the Gulf of Mexico, 
except for one station in the Gulf of Campeche. There are 26 



16 BREVIORA No. 281 

specimens in the present collection, also from this region and from 
the western part of the Gulf off the Mexican coast. These were 
trawled at "Oregon" stations 3681, 4580, 4581, 4583, 4606, 4614, 
4616, 4697, 4703, 4709, 4729, 4730, 4776, 4800, 4808, 4809, 
4816. The range in length, including specimens taken on all 
cruises, is from 78 to 360 mm, and in depth of capture from 170 to 
500 fathoms (311 to 915m), except for one station at 100 fathoms 
(183 m). 

Breviraja spinosa Bigelow and Schroeder 1950 

Previous accounts of this species include several specimens 
taken by the "Albatross" during the years 1885-1886 off Delaware 
Bay and North Carolina, stations 2624 and 2730, in 258 and 727 
fathoms (472-1330 m), respectively. These were in very poor 
condition, their identity as spinosa was doubtful, and they cannot 
now be found for re-examination. 

Published reports (1950, 1953, 1962) record a large series of 
spinosa ranging from the offings of Charleston, South Carolina 
(32°58'N), to the Guianas (07°05'N), in length 77-424 mm, 
trawled in 150-400 fathoms (274-732 m). Eighteen more have 
been taken at 7 "Oregon" stations (4428, 4854, 4912, 5039, 5689, 
5690, 5722), from off Colombia, from south of Curagao, and 
southwest of Grenada, 115-528 mm long, from depths of 257-400 
fathoms (470-732 m). Following is a description of a mature male 
of 528 mm, much larger than any male previously seen (332 mm). 

Proportional dimensions in per cent of total length. 

Disc. Extreme breadth 58.5; length 46.2. 

Snout length. In front of orbits 8.7; in front of mouth 10.4. 

Orbits. Horizontal diameter 4.6; distance between 3.5. 

Spiracles. Length 4. 1 ; distance between 6. 1 . 

Mouth. Breadth 7.4. 

Exposed nostrils. Distance between inner ends 7.0. 

Gill openings. Length 1st 1.3; 3rd 1.3; 5th 1.1; distance between 
inner ends 1st 15.5; 5th 7.6. 

First dorsal fin. Height 2.6; length of base 4.2. 

Second dorsal fin. Height 2.8; length of base 4.6. 

Pelvics. Anterior margin 13.6. 

Distance. From tip of snout to center of cloaca 42.7; from center 
of cloaca to 1st dorsal 44.7; to tip of tail 57.3; from rear end of 2nd 
dorsal base to tip of tail 3.8. Interspace 1st and 2nd dorsals 0.0. 

Disc obtusely rounded in front, thence concave toward outer 
corners which are broadly rounded; posterior margin rounded; 



1968 ADDITIONAL BATOID FISHES 17 

maximum angle in front of spiracles 135°; width across anterior 
edge of orbits 28.5 per cent of total length of specimen. Axis of 
greatest breadth 75 per cent of distance rearward from tip of 
snout to axils of pectorals. 

Thorns in the nuchal-scapular region in a triangular patch, con- 
tinuing in a band of 5 rows along back, without interruption, and 
on to tail, where they are somewhat larger and in 5-6 rows an- 
teriorly, thence narrowing to 2-3 rows, ending at first dorsal; 
anterior and inner margins of orbits with a row of 7 or 8 thorns, 
also a pair opposite each spiracle; a patch of prominent and sharp 
malar thorns opposite the spiracles and orbits, the length of patch 
about 70 mm and greatest width 20 mm; alars in as many as 4 
rows, the longest row 70 mm; disc and tail covered with fine close- 
set prickles, coarsened in advance of the short triangular rostral 
process; prickles present on skin over eyes and on dorsal and caudal 
fins, also a few on posterior lobes of pelvics. Lower surface of 
disc and tail smooth. 

The claspers extend from the axil of pelvics a distance a little 
greater than from tip of snout to rear margin of orbits. 

Jaws strongly arched centrally. Teeth ^ , uppers in center of 
jaw crowded, with slender to narrowly triangular cusps, those to- 
ward corners of jaw arranged in parallel rows, not in quincunx; 
lowers similar, but those in median sector larger than correspond- 
ing upper teeth. 

Color above plain medium brown, dorsals and caudal fins 
blackish; below, black except pale around jaws, on nasal flap, 
edges of gill openings, in areas near end of snout, and at tip of 
anterior pelvic lobes. 

Cruriraja rugosa Bigelow and Schroeder 1958 

This skate appears to be relatively common within its known 
geographic and depth ranges. We have previously recorded (1958, 
1962, 1965) 103 specimens from numerous localities extending 
from the northern part of the Gulf of Mexico to the northern coast 
of Panama, the latter at 9^03'N, 81°18'VV. These ranged in length 
from 90 to 485 mm and were trawled in depths of 200-500 fathoms 
(366-915 m). 

The present collection includes 19 rugosa of 100-404 mm, from 
250-550 fathoms (457-1007 m), taken at "Oregon" stations 4414, 
4662, 4663, 4730, 4731, 4816, 4883, 4902, 5039, and "Silver 
Bay" station 5142. Most are from within the geographic range 



18 BREVIORA No. 281 

given above, but included are four additional localities: (1) the 
north coast of Haiti, (2) the coast of Colombia, (3) southwest of 
Curagao, and (4) southwest of Grenada at 1 1°40'N, 62°33'W. 

Family PSEUDORAJIDAE 
PsEUDORAJA ATLANTICA Bigclow and Schrocdcr 1962 

Many specimens of this species have been recorded (1962, 
1965) from off the Atlantic coast of Nicaragua, 13°31'N, to the 
offing of the Amazon River, 01°45'N, in depths of 135-350 
fathoms (247-640 m), in length 86-481 mm. The present col- 
lection includes 32 males and 22 females trawled at 25 "Oregon" 
stations from off the coast of Panama 9°47'N, 79°25'W to the 
vicinity of Tobago, ll°3rN, 60°5rW, in depths of 200-340 
fathoms (366-622 m), in length 90-518 mm. 

Mature males have a patch of malar thorns along the outer part 
of the pectorals from end of snout to a little in advance of a line 
through the orbits. These prominent thorns may have sharp tips 
or be covered by skin. A few may be present on individuals as 
small as 350 mm but in sizes upward of about 420 mm, they are 
well developed. 

The alar thorns usually start to appear at about 400 mm. On a 
specimen of 425 mm they are in 2 to 3 rows, on one of 447 mm up 
to 6 rows, but on the largest male of 491 mm there are only 3 to 4 
rows of thorns. The size of the patch of thorns ranges from about 
40 to 48 mm in length and 9 to 12 mm in width, usually with but 
1 or 2 thorns at the anterior and posterior ends. 

The claspers on a 230 mm specimen are minute and do not 
reach the tip of the posterior pelvic lobe; at 340-400 mm they reach 
a little beyond the pelvic lobe and at maturity, when they have 
become rigid, they extend for a considerable distance. 

Family ANACANTHOBATIDAE 

Anacanthobatis americanus Bigelow and Schroeder 1962 

Published accounts (1962, 1965) of this species include 51 
specimens taken at 12 "Oregon" stations from off the coasts of 
British Guiana, Venezuela, Panama, and Honduras, 7° 34' to 
16°35'N, in 100-400 fathoms (183-732 m), ranging in length from 
95 to 350 mm. The present collection includes 42 specimens taken 
at 15 "Oregon" stations (4412, 4413, 4415, 4416, 4424, 4841, 
4842, 4854, 4855, 4902, 4912, 4925, 5039, 5689, 5690) all from 
off the coasts of Colombia and Venezuela between 09°02'-12°30'N 



1968 ADDITIONAL BATOID FISHES 19 

and 76°31'-69°18'W, and "Silver Bay" station 5142 off the north 
coast of Haiti at 19°52'N, 71°58'W, in depths of 205-500 fathoms 
(375-915 m), in length 114-362 mm. 

The size at which males reach maturity varies. Thus, on one of 
270 mm the claspers are minute, and while one of 295 mm is 
mature, with rigid claspers, hooks exposed, pelvics and pectorals 
separated, another of 320 mm is immature, with flexible claspers 
and the pelvics and pectorals still united. The alars at maturity 
are in up to 5 rows, the patch of thorns on the largest male, of 350 
mm, being 29 mm long and 10 mm wide. 

Anacanthobatis longirostris Bigelow and Schroeder 1962 

Three specimens of this bizarre skate have been recorded (1962, 
1965) : a female of 507 mm, the holotype, from off the Mississippi 
Delta, 29°09'N, 87°53'W, in 500-575 fathoms (915-1052 m), 
"Oregon" station 2823; a male of 135 mm from Santaren Channel. 
23°59'N, 79°43'W, in 350 fathoms (640 m), "Combat" station 
450; and an immature male of 483 mm from the same locality, 
23°40'N, 79°13'W, in 290 fathoms (530 m), "Silver Bay" station 
2458. 

Two more have been trawled, both females, of 745 and 630 mm 
respectively, off the north coast of Haiti at 19°55'N, 72°00'W, in 
470-500 fathoms (860-915 m), "Silver Bay" station 5146. Pro- 
portional dimensions in per cent of total length (exclusive of fila- 
ment at tip of snout) of both specimens follow; 

Disc. Extreme breadth 61.2, 56.3; length 74.3, 64.7. 

Snout length. In front of orbits 33.0, 30.1; in front of mouth 
35.5,33.1. 

Orbits. Horizontal diameter 2.8, 2.7; distance between 3.8, 2.9. 

Spiracles. Length 1.6, 1.0; distance between 6.1, 5.7. 

Mouth. Breadth 5.3, 5.1. 

Exposed nostrils. Distance between inner ends 5.5, 4.9. 

Gill openings. Length, 1st 1.0, 1.0; 3rd 1.0, 1.0; 5th 0.7, 0.8; 
distance between inner ends, 1st 1 1.6, 10.8; 5th 8.2, 7.3. 

Pelvics. Length of anterior limb 15.3, 12.0; distance, origin of 
anterior limb to tip of posterior lobe 13.3, 13.2. 

Distance. From tip of snout (from base of filament) to center of 
cloaca 61.7, 57.1; from center of cloaca to tip of tail 38.3, 42.9. 

The axis of greatest breadth of the disc is about two-thirds the 
distance back from tip of snout to axils of pectorals. The upper 
caudal fin is vestigial on the 745 mm specimen and the lower caudal 
appears to be missing, while on the one of 630 mm the tip of tail is 



20 BREVIORA No. 281 

lacking, total length having been estimated on the basis of the disc 
width. The proportional dimensions of the latter specimen agree 
closely with the holotype but the one of 745 mm has a longer snout, 
33.0 per cent of total length of specimen; vertical length of disc, 
74.3 per cent; wider disc, 61.2 per cent; and a greater distance from 
tip of snout to center of cloaca, 61.7 per cent, these proportions on 
the holotype being 29.8, 64.7, 56.8 and 57.4 per cent, respectively. 

Springeria folirostris Bigelow and Schroeder 1951 

Many specimens of this species have been recorded (Springer 
and Bullis, 1956; Bigelow and Schroeder, 1951a, 1965), all of 
them from the northern part of the Gulf of Mexico, ranging in 
latitude from 26°46' to 29^30'N, in longitude from 85°09' to 
96°20'W, from depths of 164 to 280 fathoms (300-512 m), in 
length from 125 to 620 mm. 

In our last account of S. folirostris a description was given of a 
male, of 576 mm, the first mature specimen to come to our at- 
tention. This showed that on males the posterior lobe of the pelvics 
ceases to be adnate to the pectorals when that stage of growth is 
attained, as occurs with Anaccmthobatis americanus. 

The present collection includes three females 140-330 mm long 
and a male of 195 mm, from 27°56'N, 90°36'W, in 220 fathoms 
(402 m), "Oregon" station 4704, also a nearly mature male of 530 
mm from 27°45'N, 93°56'W, in 200 fathoms (366 m), station 
4606. 

Proportional dimensions in per cent of total length of the 530 
mm male. 

Disc. Extreme breadth 52.0; length 54.7. 

Snout length. In front of orbits 21.0; in front of mouth 22.5. 

Orbits. Horizontal diameter 3.4; distance between 2.2. 

Spiracles. Length 1.0; distance between 5.5. 

Mouth. Breadth 4.7. 

Exposed nostrils. Distance between inner ends 4.5. 

Gill openings. Length 1st 0.76; 3rd 0.76; 5th 0.66; distance 
between inner ends, 1st 9.5; 5th 4.9. 

Caudal fin. Length of base, upper 4.7; lower 4.2. 

Pelvics. Length of anterior limb 13.0; origin of anterior limb to 
tip of posterior lobe 1 1 .0. 

Distance. From tip of snout (from base of tentacle) to center of 
cloaca 47.5; from center of cloaca to tip of tail 52.5. 

These proportions agree closely with those of the holotype. a 
male 400 mm long. 



1968 ADDITIONAL BATOID FISHES 21 

The pectorals and pelvics are free, as on a fully mature male. The 
claspers, which are semi-flexible and with the hooks not yet ex- 
posed, extend beyond the tips of the pelvics a distance equal to the 
space between the outer margins of the spiracles. The alar thorns 
have broken through the skin but are not fully developed, the 
greatest width and length of the patch, on each side, being 12 mm 
and 25 mm, respectively, with the thorns arranged in as many as 4 
rows. The jaws are rather strongly arched. Teeth f^ , those in 
center of jaw the smallest and more crowded, with a triangular 
blunted cusp, those toward corners of jaw with a flat crown. 

Grayish brown above, whitish below, except for some brownish 
blotches on the posterior segment of the pelvic limb and a broad 
band of pale gray along the outer margin of the disc. 

ADDENDUM 

Illustrations are included in the following publications by Bigelow 
and Schroeder, all listed in the references cited. 

Torpedo nobiliana 1953, figs. 22, 23 

Diplobatis pictus 1962, pi. 1 

Raja bullisi 1962, fig. 1 

Raja cervigoni 1964, fig. 1 

Raja clarkii 1958, fig. 8; 1965, fig. 1 

Raja fioridana 1962, figs. 2, 3 

Raja fuliginea 1954, fig. 4 

Raja lentiginosa 1951b, fig. 1; 1953, fig. 48a 

Raja olseni 1951b, fig. 2; 1953, fig. 54a 

Raja teevani 1951b, fig. 3; 1962, fig. 3 

Breviraja pliitonia 1953, figs. 67, 68 

Breviraja simismexicanus .... 1950, pi. 5; 1953, fig. 69 

Breviraja spinosa 1950, pi. 6; 1953, fig. 71 

Cruriraja rugosa 1958, figs. 10, 1 1; 1962, fig. 10 

Pseudoraja atlantica 1962, figs. 11, 12, 13; 1965, fig. 6 

Anacanthobatis americamis . . 1962, figs. 14, 15, 16; 1965, fig. 7 

Anacanthobatis longirostris . . 1962, figs. 17, 18; 1965, fig. 8 

Springeria jolirostris 1951a, fig. 1; 1953, fig. 78a; 1965, 

fig. 9 



22 BREVIORA No. 281 

REFERENCES CITED 

BiGELOw, H. B. and W. C. Schroeder 

1950. New and little known cartilaginous fishes from the Atlantic. 

Bull. Mus. Comp. Zool., vol. 103, no. 7, pp. 385-408, 7 pis. 

1951a. A new genus and species of anacanthobatid skate from the Gulf 

of Mexico. Jour. Washington Acad. Sci., vol. 41, no. 3, pp. 

110-113, 1 fig. 

1951b. Three new skates and a new chimaerid fish from the Gulf of 
Mexico. Jour. Washington Acad. Sci., vol. 41, no. 12, pp. 
383-392, 4 figs. 

1953. Fishes of the western North Atlantic. Mem. Sears Found. Mar. 
Res., No. 1, part 2, x + 588 pp., 127 figs. 

1954. A new family, a new genus, and two new species of batoid 
fishes from the Gulf of Mexico. Breviora, Mus. Comp. Zool., 
No. 24, 16 pp., 4 figs. 

1958. Four new rajids from the Gulf of Mexico. Bull. Mus. Comp. 
Zool., vol. 119, no. 2, pp. 201-233, 11 figs. 

1962. New and little known batoid fishes from the western Atlantic. 
Bull. Mus. Comp. Zool., vol. 128, no. 4, pp. 159-244, 23 figs., 
Ipl. 

1964. A new skate. Raja cervigoni, from Venezuela and the Guianas. 
Breviora, Mus. Comp. Zool., No. 209, 4 pp., 1 pi. 

1965. A further account of batoid fishes from the western Atlantic. 
Bull. Mus. Comp. Zool., vol. 132, no. 5, pp. 443-477, 9 figs., 
2 pis. 

Bonaparte, C. L. 

1832- Iconografia della fauna Itahca — , vol. 3, Pesci, 78 pis. (not 
1841. numbered) and accompanying text (pages not numbered). 
[For dates of publication see Salvadori, Boll. R. univ. Mus. 
Zool. Anat. Comp., Turin, vol. 3, no. 48, 1888.] 

Garman, S. 

1881. Reports on the results of dredging under the supervision of 
Alexander Agassiz, along the Atlantic Coast of the United 
States. Report on the selachians. Bull. Mus. Comp. Zool., vol. 
8, no. 11, pp. 231-238. 

Holt, E. W. L. and L. W. Byrne 

1908. Second report on the fishes of the Irish Atlantic Slope. Fish- 
eries, Ireland, Sci. Invest. (1906), No. 5, 63 pp., 5 pis. 

Martin, S., Felipe 

1957. Una nueva especie de Torpedinidae del Golfo de Cariaco, Edo. 
Sucre, Venezuela. Nov. Cientif., Contr. Ocas. Mus. Hist. Nat. 
La Salle, Ser. Zool., No. 21, 4 pp., 3 figs. 



1968 ADDITIONAL BATOID FISHES 23 

MiTCHILL, S. L. 

1815. The fishes of New York, described and arranged. Trans. Lit. 
Philos. Soc. New York, vol. 1, pp. 355-492, pis. 1-6. 

Palmer, G. 

1950. A new species of electric ray of the genus Diplobatis from 
British Guiana. Ann. Mag. Nat. Hist., ser. 12, vol. 3, pp. 480- 
484, 3 figs. 

Springer, S. and H. R. Bullis, Jr. 

1956. Collections by the "Oregon"' in the Gulf of Mexico. U. S. Fish 
and Wildlife Service. Spec. Sci. Rept.-Fisheries No. 196, 134 pp. 

Whitley, G. P. 

1939. Taxonomic notes on sharks and rays. Australian Zoologist, 
vol. 9, pp. 227-262, 18 figs. 



BREVIORA 



Muiseiiim of Comparative Zoology 

Cambridge, Mass. 5 April, 1968 Number 282 

THE EXTINCT BABOON, PARAPAPIO JONESh 
IN THE EARLY PLEISTOCENE OF NORTHWESTERN KENYA 

Bryan Patterson 



Among the fossils collected during 1966 in the early Pleistocene 
sediments of the Kanapoi area, southeastern Turkana ( Patterson, 
1966), is a specimen of a small baboon. The surface find of a 
fragment of a right horizontal ramus with M-.;; prompted excava- 
tion at the spot and sifting of the slope debris. No parts were found 
in situ but a number of fragments were recovered from the slope 
and several of these fitted together to form a partial mandible. 
Good contacts are present from the symphysis back to the base of 
the ascending ramus on the right side. The bases of the incisors 
and canines and the anterior root of the right P-, are preserved in 
the symphysis; complete or nearly complete teeth present are the 
left P^, Ml and M;., and the right M^.:,. The specimen bears the field 
number 122-66K, and was found by Mr. Roger C. Wood in the 
drainage of the Kikimon, a dry wash east of the Kanapoi and. like 
it, draining into the Kakuryo. All fragments were found within a 
small area, which, together with the gentle nature of the slope, 
suggests that the fossil may not have moved far from its burial 
place. 

As is shown by the sizes of the incisors and canines relative to the 
cheek teeth and by the marked overlap of the anterior root of P.-. 
along the posterolateral face of the canine, the specimen is cer- 
tainly a male. It agrees so closely with specimens of Panipapio 
jonesi Broom from South Africa that, if Freedman's (1957, 1960) 
specific distinctions are accepted, there can be no doubt as to the 
identification. 

Frecdman has discussed the structure of this and other species of 
Panipapio in considerable detail, which makes it unnecessary to do 
more than comment on a few points. Curiously enough, the new 
specimen has the most complete symphysis of any male individual 



BREVIORA 



No. 282 



of P. jonesi yet collected. The anterior surface is not steep, form- 
ing an angle of approximately 45' with the lower border of 
the horizontal ramus. The two ridges that converge upward toward 
each of the median incisors are not very prominent; they enclose a 
shallow, median depressed area that extends dorsally from the 
foramen symphyseosum. Between the posterior portion of the 
ridge, below, and the anterior alveolus of P;., above, is a small, 




Fig. 1. Parapapio jonesi Broom. Field no. 122-66K. Dorsal view of 
incomplete mandible of male. X 1- 



rather rugose depression. Apart from this, the anterior surface is 
only slightly roughened. The incisal shelf slopes very gently down- 
ward from the incisor alveoli to a point opposite about the middle 
of P;j. The symphysis terminates at the level of the anterior end of 
P4; there is no mental spine. The ascending ramus arises a little 



1968 



EXTINCT BABOON FROM KENYA 



3 



behind M.;. The lateral face of the horizontal ramus is very slightly 
concave beneath M^. and the anterior half of M;.,. but neither here 
nor in the portion of the bone beneath the left P4 — Mi is there 
any delimitation of a mandibular fossa as such. The teeth pre- 
served call for no special comment. 

Measurements (in mm) 



Length (mesio-distal) 

Width 

Width, anterior 

Width, posterior 

Width, hypoconulid 

Depth of ramus posterior to Ms 



c 


P. 


Mx 


M. 


Mc 


6.3 


6.4 


8.2 


10.4 


12.7 


10.0 


6.2+ 












7.3 


8.8+ 


9.6 






7.1 


9.1 


8.6 
5.0 



29.1 



All measurements have been taken in accordance with Freed- 
man's methods. Dimensions of the Kanapoi specimen fall. or. in 
the case of the widths of P4 and M^. presumably fell, within the 
observed ranges for males of his sample of P. jonesi (1957. tables 
6b, 6d) in all cases but three. These, the anterior and posterior 
widths of M] and the length of M.-.,. fall within the observed ranges 
for females. As he stresses, the ranges of the two sexes overlap 
broadly for P4 — M;., in the genus. His samples for males, further- 
more, are small, numbering only 5. 4 and 3 in these cases. 

Parapapio is characteristic of the older part of the South African 
Pleistocene sequence. P. jonesi is well represented at Sterkfontein 




Figure 2. Parapapio jonesi Broom. Field no. 122-66K. Right lateral 
view of incomplete mandible of male. The ramus fragment with Pi — Mi 
has been reversed from the left side. X 1. 



4 BREVIORA No. 282 

and Makapan, by 24 and 15 specimens, respectively, and one in- 
dividual from Taung has been referred to it. The only specimens 
of Panipapio recorded from Swartkrans and Kromdraai are five 
individuals, 3 and 2, respectively, of a small form, and these are 
fragmentary. Freedman placed them with some hesitation in P. 
jonesi, pointing out that better material might in future require their 
separation. Two partial female mandibles from Swartkrans have 
"fairly large and quite deep mandibular fossae," a feature con- 
spicuously lacking in those from the earlier deposits. The Kanapoi 
specimen agrees with the earlier South African material and adds 
one more to the small list of species in common between the early 
Pleistocene of eastern and southern Africa. 

The field work was supported by National Science Foundation 
Grant no. G.A. 425. The drawings are the work of Mr. Arnold 
D. Clapman. 

REFERENCES 

Freedman, L. 

1957. The fossil Cercopithecoidea of South Africa. Ann. Transvaal 

Mus.. 23: 121-262. 
1960. Some new cercopithecoid specimens from Makapansgat, South 
Africa. Palaeont. Afric, 7: 7-45. 
Patterson, B. 

1966. A new locality for early Pleistocene fossils in northwestern 
Kenya. Nature', 212: 577-581. 

(Received 19 June 1967.) 



BREVIORA 

MmseiLiiimi of Comparative Zoology 

Cambridge, Mass. 5 April, 1968 Number 283 



SCYTHIAN AMMONOIDS FROM TIMOR 

Bernhard Kummel 

One of the more important areas of ammonitiferous Scythian 
(Lower Triassic) strata is the island of Timor. The monograph by 
Welter (1922) on the Lower Triassic ammonoids of Timor is the 
standard reference. However, an earlier paper by Wanner (1911) 
did describe four species of Lower Triassic ammonoids. A number 
of additional comments and descriptions on these Timor am- 
monoids were contributed by Spath (1934). In the course of a 
general study of all Scythian ammonoids I have had the opportunity 
to examine many collections from Timor including the type speci- 
mens of the Wanner (1911) and Welter ( 1922) studies. This note 
was prepared to make some additions and corrections to our 
knowledge of Scythian ammonoids of Timor. 

Prosphingitcs austini Hyatt and Smith is recorded for the first 
time from Timor. This is a common species of Oweuiles Zone age 
in the circum-Pacific and Arctic regions. 

Prenkites simdaicus Welter is established as the type of a new 
genus, Vickohlerites. Previous problems in the interpretation of 
this species were due to the inaccurate suture illustrated by Welter 
(1922). 

The genus Parowcnitcs Spath (1934) was established for an 
owenitid with a presumed goniatitic suture as illustrated by Welter 
(1922, pi. 169 (15). figs.^5, 8). Examination of Welter's types 
clearly shows that the goniatitic suture is the result of excessive 
grinding and polishing of the specimens. Preparation of other 
paratypes shows that the suture is ceratitic; hence the genus Par- 
owenites Spath is a synonym of Owenites Hyatt and Smith. 

The genus Metadagnoceras Tozer is recognized for the first time 
from the bed with manganese coated fossils at Nifoekoko. This 
specimen had previously been noted by Spath ( 1934, p. 269, foot- 
note) as being "very close to Dagnoceras terbunicuin." 



2 BREVIORA No. 283 

For the loan of specimens used in this study I am very grateful 
to Dr. H.K. Erben of Bonn University. Dr. M.K. Howarth of the 
British Museum (Natural History), and Dr. H.J. MacGillavry of 
the Geological Institute. Amsterdam University. The laboratory 
aspects of this study were supported by N.S.F. grant GB-5 109X. 

SYSTEMA lie PALEONTOLOGY 

Class CEPHALOPODA Cuvier, 1797 

Subclass AMMONOIDEAZittel. 1884 

Family PROPTYCHITIDAE Waagen, 1895 

Genus Owenites Hyatt and Smith, 1905 

Type species, Oweniws koeneni Hyatt and Smith, 1905 

Owenites simplex Welter 

Plate 1. figures 1-9 

Owenites simplex Welter. 1922: 153, pi. 169(15), figs. 1-8; Kutassy, 1933: 

606. 
Paiowenites simplex, — Spath. 1934: 187, fig. 5S; Kummel in Arkell et at., 

1957: L138, figs. 171-la,b; Kummel and Steele. 1962: 647. 
Owenites kwangsiensis Chao. 1959: 83, 250, pi. 22. figs. 1-6, text-fig. 26b. 
Owenites plicatiis Chao, 1959: 85. 251, pi. 22. figs. 19-21, text-fig. 26e. 
Owenites aff. plicatus Chao. 1959: pi. 22, figs. 24. 25. 
Owenites costatus Chao. 1959: 83. 249. pi. 22, figs. 10-18. 22, 23. text-fig. 

26c. 
Owenites eostatns var. lenticiiUiris Chao, 1959: 84. 249, pi. 22, figs. 7-9, 

text-fig. 26d. 
Spath (1934: 187) established the genus Parowenites primarily 
on the basis of the goniatitic suture of Owenites simplex as repro- 
duced by Welter (1922: pi. 169(15). figs. 5, 8). Examination 
of Welter's type specimens plus a number of paratypes shows that 
the sutures reproduced by Welter were exposed by grinding and 
polishing. The specimen which yielded the suture of Welter's figure 
8 is apparently lost, as it is not in the collections of the Paleontolo- 
gical Institute. Bonn University.^ However, the paratype of Wel- 
ter's plate 169( 15). figures 3-5 is available, and this shows clearly 
that excessive grinding of the lateral area resulted in the simple, 
goniatitic suture. One of the unfigured paratypes of Welter (Pi. 1, 



1 Abbreviations in this paper: MCZ = Museum of Comparative Zoology; 
GPlBo =: Geologisch-Palaontologisches Institut. Bonn; BMNH = British 
Museum (Natural History). 



1968 SCYTHIAN AMMONOIDS FROM TIMOR 3 

figs. 8, 9 of this report) yielded the suture of Figure 3E. As can 
readily be seen, the two lateral lobes are ceratitic and the whole 
aspect of the suture is owenitid. 

The measurements of 27 specimens of this species are given on 
Table I. and plotted on the graph of Figure 1. There is consider- 
able variation in the diameter of the umbilicus which interestingly 
enough is not obvious on casual inspection of the specimens. What 
is more interesting is that the diameter of the umbilicus can vary 
from one side of the conch to the other. For instance, one of the 
paralectotypes (PI. 1, figs. 3-5) has an umbilical diameter of 3.7 
mm on one side and 4.6 mm on the other side of the conch. 

H 

16 

15 

14 I- 

13 

12 

11 

10 

9 

8 

7 

6 

5 

4 

3 

2 

1 

5 10 15 20 25 30 35 40 

DIAMETER 

Figure 1. Variation in whorl height (H) and whorl width (W) of 
Oweuites simplex from limestone blocks with Owoiitcs egrediens, Bahati, 
Timor. 



w 









X 



xk 



4 BREVIORA No. 283 

The form described by Chao (1959) as Owenites kwangsiensis 
from the Owenites Zone of Kwangsi, China, I consider to be iden- 
tical to O. simplex Welter (Figs. 3E, H). From the same horizon 
and locality which yielded Owenites kwangsiensis. Chao (1959) 
described O. costatus and O. plicatiis. Of the first of these species 
Chao had 1 7 specimens which he compared mainly with Owenites 
pakiingensis Chao; the latter I consider a synonym of Owenites 
koeneni (Kummel and Erben. 1968). Owenites costatus does 
have constrictions and ribs but these are stated to be developed on 
an irregular pattern, thus differentiated from Owenites kwangsien- 
sis. Examination of the 31 specimens from Timor of Owenites 
simplex, representing all growth stages, shows that there is great 
variation in the pattern of ornamentation. Every one of the speci- 
mens of Owenites costatus illustrated by Chao ( 1959: pi. 22, figs. 
10-18, 22, 23) can be directly compared to one of the Timor speci- 
mens before me. Owenites plicatus Chao was established on two 
fragmentary specimens. On the basis of the description and illus- 
trations of these two specimens I can see no reason for separating 
them from O. simplex. The apparently more simplified suture 
(fig. 21) was noted by Chao as most probably reflecting imma- 
turity and not a distinctive new pattern. Owemtids of the type of 
O. simplex are only known from Timor and Kwangsi. China. 

Occurrence — Dark red limestone blocks with Owenites egre- 
diens of Bahati, Timor. 

Repository — All specimens are in the Paleontological Institute, 
Bonn University. 

Family PARANANNITIDAE Spath. 1930 

Genus Prosphingites Mojsisovics, 1886 

Type species, Prosphingites czekanowskii Mojsisovics, 1886 

Prosphingites austini Hyatt and Smith 

Plate 2, figures 1-16 

Prosphingites austini Hyatt and Smith, 1905: 72. pi. 7. figs. 1-4; Freeh. 1908, 

pi. 63. fig. 5: Krattt and Diener. 1909: 160; Diener. 1915: 233; Smith. 

1932: 98, pi. 7, figs. 1-4; Spath, 1934: 195-196; Kummel and Steele, 

1962: 683; Kummel 1965: 544. 
Prosphingites sp. indet. Spath. 1921: 298, 301. 
Prosphingites spathi Freboid. 1930: 20. pi. 4, figs. 2. 3. 3a; Spath, 1934: 

195, pi. 13, figs. 1, 2; Tozer. 1961: 58, pi. 13. figs. 1. 2; Kummel and 

Steele, 1962: 687; Kummel, 1965: 544. 



1968 SCYTHIAN AMMONOIDS FROM TIMOR 5 

Prusphiiigites sinensis Chao, 1959: 122. 297, pi. 25. figs. 1-5, pi. 27, figs. 
1-17. text-figs. 40a-c; Kummel. 1965: 544. 

Prosphingites radians Chao. 1959: 123. 298. pi. 28. figs. 12-16. text-fig. 39d; 
Kummel, 1965: 544. 

Prosphingites oralis Kiparisova. I960: 137, pi. 33, figs. 2-4; Kiparisova, 
1961: 115.pl. 25. figs. 6-9. 

Prosphingites orientalis Kiparisova. 1961: 117. pi. 26. figs. 1, 2. 

Prosphingites slossi Kummel and Steele. 1962: 683. pi. 101. figs. 8-17; Kum- 
mel. 1965: 544. 

None of the three previous studies on Scythian amnionoids from 
Timor (Wanner, 1911; Wehcr, 1922; Spath, 1934) contains any 
mention of the genus Prosphingites. I have available five specimens 
that can with confidence be assigned to Prosphingites austini Hyatt 
and Smith. These specimens are from Fatoe Kosat and are part 
of the Jonker Collection of the Geological Institute, Amsterdam 
University. Measurements (in mm) on four of these species are 
as follows: 



D 


W 


H 


U 


34.5 


21.7? 


13.3 


11.8 


30.8 


16.7 


13.6 


8.4 


27.8 


? 


12.8 


7.8 


27.2? 


17.0 


11.3 


8.2 


26.0 


18.5 


12.3 


8.2 



The similarity of these Timor specimens with Prosphingites slossi 
Kummel and Steele (1962: 683. pi. 101, figs. 8-17) and other mid- 
Scythian prosphingitids is indeed remarkable (PI. 2). Documenta- 
tion of species of this genus has been slow. The genotype (P. 
czekanou'skii Mojsisovics, 1886) is from northern Siberia and of 
late Scythian age. Hyatt and Smith (1905) then described Pros- 
phingites ciustini from the Meekoceras beds of the Inyo Range, 
southeastern California. Frebold (1930) and Spath (1934) re- 
corded the Spitsbergen species — Prosphingites spathi — which 
they considered late Scythian in age. It was not until the post-war 
period that descriptions of a number of new species from middle 
and late Scythian horizons began to appear. These were from 
Scythian faunas of the Primorye Region (Kiparisova. 1961). 
southern China (Chao, 1959). New Zealand (Kummel. 1965). 
and Nevada (Kummel and Steele, 1962). From the mid-Scythian, 
Owenites Zone, the following species of Prosphingites are known: 
Prosphingites austini Hyatt and Smith, P. oralis Kiparisova, P. 



6 BREVIORA No. 283 

orientalis Kiparisova, P. sinensis Chao, P. involutus Chao, P. 
kwangsianus Chao, P. radians Chao, P. spathi Frebold, and 
P. slossi Kummel and Steele. Most of these species were introduced 
over a short period of time (1959-1962) and each author was 
generally without knowledge of the others' activities. We thus have 
an accumulation of species in which very little comparative analysis 
accompanied their original descriptions. 

When I introduced the species Prosphingites slossi (Kummel 
and Steele, 1962) I was fully cognizant of its close relationship 
and possible identity to P. austini Hyatt and Smith (PI. 2, figs. 1-3) ; 
however, on the argument that Prosphingites austini was known 
only from a single and not a very well preserved specimen, I con- 
sidered it best to ignore the species. However, subsequently, on 
study of all new Scythian species that had been introduced, and 
through restudy of P. austini, I am now convinced that P. slossi is 
a synonym of P. austini. Kummel and Steele (1962) presented 
measurements on 49 specimens of P. slossi from the Meekoceras 
beds at Crittenden Spring, Nevada. These data show that at least 
within that population there is considerable variability in conch 
width and umbilical diameter. This collection also clearly showed 
that there was considerable variation in the pattern of constrictions. 

Of the Timor specimens recorded here, only four yielded com- 
plete measurements. The range of umbilical diameters in these 
four specimens falls well within the range of umbilical diameters 
in the Crittenden Spring fauna of Prosphingites slossi. The whorl 
width tends, however, to be greater in the Timor forms than in the 
Nevada fauna. This, however, considering the small size of the 
sample and the identity or close similarity of the other conch 
features, is not sufficient reason to even suggest specific distinction. 
The pattern of ornament, and variations in the nature of the um- 
bilical shoulder and wall are the same. The sutures can be said to 
be identical (Fig. 2). 

Most of the other species placed here in the synonymy of Pros- 
phingites austini are known from very few specimens. Large num- 
bers of specimens of P. spathi are in the British Museum ( Natural 
History), but only a few measurements are available and nearly all 
of this Spitsbergen collection consists of small phragmocones or 
juvenile specimens. It is uncertain from the text but it appears that 
the fauna of Prosphingites spathi from Ellesmere Island studied by 
Tozer (1961) consists of very few specimens, perhaps only three. 
Both Spath (1934) and Tozer (1961) have emphasized the varia- 
bility of this species, including its suture. I can find no criteria by 



1968 



SCYTHIAN AMMONOIDS FROM TIMOR 









Figure 2. Diagrammiitic representation of the suture of: A, Prosphiiij^itcs 
aiistini Hyatt and Smitii ( 1905: pi. 7, fig. 4) from Meekoceras beds. Union 
Wash. Inyo County, California, USNM 75256. at a diameter of 20 mm; B, 
Prosphingites slossi Kummel and Steele, from Meekoceras beds, Crittenden 
Spring, Nevada, paratype. at diameter of 18 mm; C. Prosphingites slossi 
Kummel and Steele, from Meekoceras beds. Crittenden Spring, Nevada, para- 
type. at a diameter of 30 mm; D, Prosphingites spathi Frebold, from Tozer 
(1961: pi. 13. fig. Ic) from Meekoceras beds. Blind Fiord Formation, 
Ellesmere Island, at a diameter of 16 mm; E, Prosphingites spathi Frebold, 
from Spath (1934: pi. 13, fig. le) from Owenites Zone, Spitsbergen; F, 
Prosphingites aiistini Hyatt and Smith, specimen in Yonker collection, from 
Timor, Department of Geology, Amsterdam University, at a diameter of 
22 mm; G, Prosphingites oralis Kiparisova (1961: fig. 79) from Owenites 
Zone, Primorye Region, at a whorl height of 5 mm; H, Prosphingites sinensis 
Chao (1959: fig. 40a) from Olenites Zone, Kwangsi, China, at a diameter 
of 20 mm. 



which one can separate P. spathi from P. austini. In regards to the 
age of the Spitsbergen forms it has been pointed out by Kummel 
( 1961) and Tozer (1961) that the beds containing P. spathi are 
of mid-Scythian Owenites Zone age rather than late Scythian as be- 
lieved by Spath. 

The mid-Scythian, Owenites Zone, strata of the Primorye Region 
contain, according to L. D. Kiparisova (1961), two species of 



8 BREVIORA No. 283 

Prosphingites, namely, P. ovalis Kiparisova and P. orientalis Kipari- 
sova. The first of these species, P. oralis, was said to difi'er from P. 
spathi in its larger shell, oval coiling, more denticulated lobes, and 
narrower ventral saddle. On close examination none of these 
criteria are valid. The Arctic P. spathi so far is known from very 
few specimens on Ellesmere Island and several but mainly juvenile 
forms from Spitsbergen. Even so, size is hardly a parameter for 
species distinction, especially in a case like this. In regards to the 
suture, an inspection of Figure 2 demonstrates considerable varia- 
bility in this respect. The Crittenden Spring fauna also shows 
considerable variability in details of the suture. The same varia- 
bility has been noted for P. spathi by Spath (1934) and Tozer 
(1961). Authors continue to place too high a degree of im- 
portance on minor variations in suture patterns. Time and trouble 
generally dictate the representation in a description of only one 
suture pattern. My own experience has amply shown that the 
suture is as variable as any other conch feature in most if not all 
Scythian ammonoids. 

From the same horizon and locality in the Primorye Region, 
Kiparisova (1961) has described Prosphingites orientalis. From 
the two illustrations given by Kiparisova (1961 : pi. 26, figs 1,2). 
P. orientalis differs from P. oralis primarily in the more acute 
rounding of the umbilical shoulder. The contrast in the nature of 
the umbilical shoulder of these two species is precisely the same 
as seen in the fauna of Prospliingites slossi from Crittenden Spring, 
Nevada (Kummel and Steele, 1962: pi. 104, figs. 8-17). The 
supposed ovate coiling of P. oralis is suspect, as not all of the illus- 
trated specimens show this. Even so, variants showing slightly 
ovate coiling are common in Scythian ammonoids. Again, I can 
see nothing in the suture, conch shape or pattern of constrictions 
in these Primorye forms to separate them from P. austini, P. slossi, 
or P. spathi. 

Among the very large Scythian fauna of Kwangsi, China, Chao 
(1959) recognized a number of prosphingitids. Among these, 
Prosphingites radians must be considered a synonym of P. austini 
and the other species discussed above. The conch form, pattern 
of constrictions and suture (Fig. 2) are well within the range of 
variation of what is here defined as P. austini. There is, in addition, 
Prosphingites sinensis, of which I am not as confident in placing in 
the synonymy of P. austini. The description and illustrations of this 
species leave much to be desired. However, on the basis of avail- 
able data, I consider it most probable that this species is the same 



1968 SCYTHIAN AMMONOIDS FROM TIMOR 9 

as P. austini. Even the unusual suture, in terms of the other forms 
combined with P. austini. may not be definitive (Fig. 2H). 

Prosphingites kwangsicmus Chao { 1959: 296, pi. 28, figs. 17-22) 
is reported to be a smooth form lacking constrictions. The species, 
however, is based on two incomplete specimens of modest preserva- 
tion. More satisfactory documentation of this species I suspect 
would show it to be conspecific with all the forms placed here in 
Prosphingites austini. 

Occurrence. — The Timor specimens recorded here are from 
red limestones at Fatoe Kosat, Timor. 

Repository. — Collection of Lower Triassic ammonoids made 
by Dr. H. G. Jonker during his Timor expedition in 1915-1916. 
Specimens are deposited in the Timor collection of the Geological 
Institute, University of Amsterdam. 

Genus Vickohlerites n.g. 
Type species, Prenkites sundaicus Welter, 1922 

Conch evolute, with depressed whorl section, broad rounded 
venter, conch smooth except for growth lines. Suture consists of 
two denticulated lateral lobes and an auxiliary lobe on the umbili- 
cal wall. The type species is from an unknown horizon on Timor. 
Conspecific specimens are present in the Subcolumbites fauna of 
Chios and an indeterminant species (but not conspecific) is present 
in the Subcolunibites fauna of Kotal-e-Tera, Afghanistan (Kummel, 
1968). 

Vickohlerites sundaicus (Welter) 
Plate 1, figures 10. 11 

Piciikites sundaicus Welter, 1922: 150, pi. 168 (14), figs. 18-21; Kutassy, 
1933: 621; C. Renz, 1945: 301; Renz and Renz. 1947: 60; Renz and 
Renz, 1948: 29. pi. 12. fig. 1; Chao. 1959: 306. 

"Prenkites" sundaicus. — Spalh, 1930: 77; Spath. 1934: 188, 209. 

Welter (1922) described this species on the basis of a single 
specimen from Noel Niti, Timor, and was quite positive as to the 
close relationship of his specimen to Prenkites malsorensis Arthaber 
from the Subcolumbites fauna of Albania. This close relationship 
is difficult to see. Prenkites is a more involute form, with depressed 
whorls which contract on the adoral quarter volution. The umbili- 
cal shoulders are subangular and bear fine nodes. The Timor speci- 
men has a diameter of 40.7 mm, an adoral whorl of 21.5 mm, a 
height of the adoral whorl of 13.2 mm and an umbilical diameter 



10 BREVIORA No. 283 

of 20.3 mm. The conch is evolute, the umbilicus comprising ap- 
proximately 50 per cent of the conch diameter. The whorls are 
depressed with the maximum width at the umbilical shoulder. 
The venter is arched and grades with no perceptible ventral 
shoulder to the umbilical shoulder which is acutely rounded. The 
umbilical wall is steep but not vertical. The last half volution of 
the specimens is body chamber and shows traces of delicate growth 
lines. The penultimate half volution bears a series of weak forward 
projecting ridges which are most prominent on the center part of 
the venter and disappear completely midway between the venter 
and the umbilical shoulder. 

It is in Welter's (1922: pi. 14, fig. 21) representation of the 
suture that one can pinpoint the uncertainty in interpretation of 
this form. Welter's drawing of the suture covers only the portion 
from the venter to the umbilical shoulder, but implied that is was 
a complete suture. A new drawing of the suture is given on Figure 
3J. As can be seen, there are two prominent lateral lobes but the 
umbilical wall bears a good portion of a fairly large auxiliary saddle 
and a small but very distinct denticulated lateral lobe. The first 
author to comment on Prenkites simdaicus after Welter was Spath 
(1930: 77) who remarked *' 'Prenkites' simdaicus Welter, in whorl 
shape resembles Columbites, but in suture line it is closer to Siib- 
columbites . . ." In terms of Welter's representation of the suture 
this statement is correct. In Columbites the second lateral lobe is 
generally very small consisting of a single prong, whereas in Sub- 
columbites the second lateral lobe, though much smaller than the 
first, is more highly developed in terms of its breadth and pattern of 
denticulation. On the basis of suture alone, Prenkites sundaicus 
can not be attached to either Columbites or Subcolumbites. 

There is a general similarity in conch shape of Prenkites sundai- 
cus with some groups of Subcolumbites. Among the subcolumbitids 
three distinct groups can be recognized. There is first of all the 
perrinismit/ii group with a tendency for carination of the venter; 
secondly, the dusmcmi group with a more marked development of 
the reticulate ornamentation, and a compressed whorl section, but 
lacking the tendency toward carination; finally, there is the robus- 
tus-multijormis group characterized mainly by their depressed 
whorl section. It is to this last group that Vickohlerites sundaicus 
has great resemblance in conch form. Within the two subfamilies 
of the Paranannitidae those genera assigned to the Columbitinae 
tend to have sutures lacking an auxiliary lobe, whereas within the 



1968 SCYTHIAN AMMONOIDS FROM TIMOR 11 

Paranannitinae an auxiliary lobe is commonplace, as in Prosphing- 
ites, Zenoites, Chiotites, etc. It is within this subfamily that Vic- 
kohlerites sundaicus belongs. 

Renz and Renz (1948-' 24, pi. 12, fig. 1) have described and 
illustrated a single specimen from the Siibcoliimbites fauna of 
Chios as a representative of this species. The general conch form 
of their specimen is the same as that of the type specimens from 
Timor. This Chios specimen measures 55.5 mm in diameter, 21.4 
mm for the width of the adoral whorl, 19 mm for the height, and 
23.5 mm for the width of the umbilicus. The dimensions of the 
whorl height and umbilical diameter in the two specimens are rea- 
sonably similar. However, the Timor specimen has a broader whorl 
than the Chios specimen by approximately 14 per cent. This dif- 
ference in whorl width is difficult to evaluate, as each locality has 
yielded only a single specimen. In addition, the Chios specimen, 
apparently, does not show the suture. The overall similarity of 
the Chios specimen to that from Timor is such that, in spite of the 
differences and lack of data mentioned above, the two specimens 
should be considered as conspecific. 

An undescribed specimen from a Subcolutnbites fauna at Kotal- 
e-Tera, Afghanistan, is clearly congeneric with the Timor and 
Chios specimens discussed above but is not conspecific. There 
are significant differences in whorl cross-section and in the suture. 

Occurrence. — The holotype (PI. 1, figs. 10. 11) is from Noel 
Niti, Timor. Welter (1922: 85, 86) considered this specimen to 
come from his lowest Triassic horizon. He lists (on p. 150) the 
forms associated with this specimen as Meekoceras sp. indet. The 
available evidence does not allow any precise determination of the 
age, but the biological affinities of this species suggest that it is late 
Scythian in age. 

Repository.  — The holotype is in the Paleontological Institute 
of Bonn University; the specimen from Chios is in the Natural 
History Museum, Basel J 13576. 

Family MEEKOCERATIDAE Waagen, 1895 

Genus Metadagnoceras Tozer, 1965 

Type species, Metadagnoceras pulcher Tozer 

Metadagnoceras freemani n. sp. 

Plate 1, figures 12, 13. 

This species is proposed for the specimen from Timor mentioned 
by Spath ( 1934: 269, footnote) as "an isolated example of a new 



12 BREVIORA No. 283 

species from Timor . . . which is very close to Dagnoceras terbiini- 
cum." The specimen measures 47.8 mm in diameter. 14.7 mm for 
the width of the last whorl, 21.7 mm for the height of the last whorl, 
and 10.4 mm for the diameter of the umbilicus. The conch is com- 
pressed with a low, arched venter and rounded ventral and umbili- 
cal shoulders. The suture (Fig. 3D) has a very large first lateral 
lobe with denticulations extending all along the vertical side to a 
narrow irregular first lateral saddle. There is a small denticulated 
second lateral lobe on the umbilical shoulder and wall. 

This species does not resemble Dagnoceras terbimicum in conch 
shape; the differences are centered mainly in the character of the 
venter. In the suture, the first lateral saddle and the first lateral 
lobe of the two species are quite similar but the remainder of the 
suture is quite different. In general conch morphology Metadagno- 
ceras jreenumi is quite similar to the type species, M. pulcher 
Tozer. The sutures are similar in basic plan but differ in significant 
details ( Figs. 3 A, D). The suture is quite similar to an undescribed 
species from the Tobin Formation, Nevada, but the whorls are of 
quite different proportions and the conch much more evolute 
(Fig. 3B). 

Occurrence. — Nifoekoko, Timor, from bed with manganese 
coated fossils, including Alhanites, Prohungarites, etc. 

Repository. — British Museum (Natural History) C33701, 
holotype. 



1968 



SCYTHIAN AMMONOIDS FROM TIMOR 



13 





H 





Figure 3. Diagrammatic representation of the suture of: A, Metadagno- 
ceras piilclier Tozer (1965: fig. 9). from British Columbia, at a whorl 
height of approximately 20 mm; B, undescribed new species from Tobin 
Formation, Nevada, at a whorl height of 11 mm, MCZ 9638; C, undescribed 
species of Mctaclagnoceras collected by N. J. Silberling from Star Peak 
Formation, Nevada, at a whorl height of 22 mm; D, Metadagnoceras frce- 
imini n. sp., holotype. from Nifoekoko. Timor, at a whorl height of 14 mm, 
BMNH C33701; E, Olenites simplex Welter, original drawing based on 
one of Welter's unfigured paratypes (PI. 1. figs. 8, 9 of this report) at a 
diameter of 13 mm; F, Oneiiites costatus Chao ( 1959: fig. 26c) at a diam- 
eter of 20 mm; G Owenites costatus var. lentkularis Chao (1959: fig. 26d ) 
at a diameter of 12 mm; H, Olenites kwungsiensis Chao (1959: fig. 26b) 
at a diameter of 25 mm; I, Owenites plicatiis Chao (1959: fig. 26e) at a 
diameter of 30 mm; J, Vickohlerites siindaiciis (Welter), at a diameter of 
30 mm, holotype, GPIBo 231. 



14 BREVIORA No. 283 

TABLE 1 
Measureiiiciits of Owenites simplex Welter from Timor 



N 


0. D 


W 


H 


U 


W/D 


H/D 


U/D 


1 


39.8 


10.3 


16.4 


1 0.0 


25.9 


41.2 


25.1 


2 


39.6 


10.3 


15.6 


7.2 


26.0 


39.3 


18.2 


3 


32.8 


9.5? 


13.0 


13.4 


28.9? 


39.6 


40.8 


4 


28.9 


8.9 


12.2 


7.2 


30.8 


42.2 


24.9 


5 


25.9 


7.8 


11.2 


6.3? 


30.1 


43.2 


24.3? 


6 


25.4 


8.4 


10.0 


8.0 


33.1 


39.4 


31.4 


7 


25.1 


•) 


10.9 


6.3 


'7 


43.4 


25.1 


8 


24.2 


7.3 


10.5 


5.8 


30.2 


43.4 


23.9 


9 


22.6 


7.4 


9.3 


5.1 


32.7 


41.2 


22.6 


10 


21.2 


8.0 


9.1 


5.6 


37.7 


42.9 


26.4 


1 1 


21.1 


6.9 


9.7? 


4.9? 


32.7 


45.9? 


23.2? 


i: 


19.7 


7.1 


7.5 


6.2 


36.0 


38.1 


31.5 


13 


18.8 


7.0 


8.0 


4.0 


37.2 


42.6 


21.3 


14 


17.4 


•) 


8.4 


3.1? 


•) 


48.3 


17.8? 


15 


16.1 


6.1 


7.0 


3.8 


37.9 


43.5 


23.6 


16 


16.0 


5.9 


6.8 


4.6 


36.9 


42.5 


28.8 


17 


15.6 


6.2? 


6.8 


3.8 


39.7? 


43.6 


24.4 


18 


15.5 


6.1 


7.8 


4.0 


39.4 


50.3 


25.8 


19 


15.1 


6.1 


6.4 


3.7 


40.4 


42.4 


24.5 


20 


13.3 


5.3 


6.2 


2.7 


39.8 


46.6 


20.3 


21 


13.3 


5.1 


5.4 


3.3 


38.3 


40.6 


24.8 


-) ■» 


14.6 


5.6? 


6.3 


3.7 


38.4? 


43.2 


25.3 


23 


12.4 


5.3 


5.4 


•) 


42.7 


43.5 


•) 


24 


10.4 


4.3 


4.8 


1 .6? 


41.3 


46.2 


15.4? 


25 


10.1 


4.7 


4.7 


1.5? 


46.5 


46.5 


14.9? 


26 


8.0 


3.7 


3.7 


9 


46.3 


46.3 





27 


7.3 


3.8 


3.3 


1.1? 


52.1 


45.2 


15.1? 



D = diameter; W = width of adoral whorl; H - height of adoral whorl; U = 
diameter of umbilicus. 

No. 2. Lectotype. Welter (1922: pi. 169(15), figs. 1. 2: PI. 1. figs. 1, 2 of 
this report) GPIBo 234a. 

No. 8. Paralectotype, Welter (1922: pi. 169(15). figs. 3-5; PI. 1. figs 3-5 
of this report) GPIBo 234h. 

No. 10. Paralectotype. Welter ( 1922: pi. 169( 15), figs. 6, 7; PI. 1, figs. 6, 7 
of this report) GPIBo 234c. 

No. 20. Suture specimen from Bahati, Timor (PI. 1, figs. 8, 9, Te.\t-fig. 3E of 
this report) GPIBo. 



1968 SCYTHIAN AMMONOIDS FROM TIMOR 15 

REFERENCES 

Arkell. W. i. et ul. 

1957. Treatise on Invertebrate Paleontology, Ammonoldea. R. C. 
Moore ed., Pt. L, Mollusca. 4, 490 pp. 

Chao, Kingkoo 

1959. Lower Triassic ammonoids from western Kwangsi, China. 
Palaeont. Sinica, 145: 1-355. 

DiENER, Carl 

1915. Fossiliiim Catalogus. I. Animalia. Pt. 8, Cephalopoda Triadica. 
Berlin, 369 pp. 

Frebold, H. 

1930. Die Altersstellung des Fischhorizontes des Grippianiveaiis iind 
des unteren Saurierhorizontes in Spitzbergen. Skr. Svalb. og 
Ishavet, 28: 1-36. 

Frech, F. 

1903- Lethaea Geognostica. Das Mesozoicum. I, Trias. Stuttgart, 

1908. 623 pp. 

Hyatt, A. and J. P. Smith 

1905. The Triassic cephalopod genera of America. Prof. Pap. U.S. 
Geol. Surv., 40: 1-394. 

Kiparisova, L. D. 

1960. Novye vidy drennikh rastenii i besopzvonochnykh SSSR, 
Chast' 2. Novye rannetriasovye nautilus i prosfingit Yuzhnogo 
Primorya. Vsesoiuznyi nauchno-issIedovateTskii geologicheskii 
institut (VSEGEI). [New species of ancient plants and in- 
vertebrates of the U.S.S.R., Pt. 2. New early Triassic nautiloids 
and prosphingitids from the Primorye Region. All Union Sci. 
Res. Geol. Insl.. pp. 136-139.] 

1961. Paleontologicheskoe obosnovanie stratigrafii triasovykh otloz- 
henii Primorskogo kraia. Chast'l. Golovonogie moUiuski. 
Vsesoiuznyi nauchno-issledovatel'skii geologicheskii institut 
(VSEGEI). trudy. [Paleontological foundation for the strati- 
graphy of the Triassic deposits of the Primorye Region. Pt. I, 
Cephalopoda. Trans. All Union Sci. Res. Geol. Inst.. N. Ser., 
48: 1-278.) 

Krafft, a. v. and Carl Diener 

1909. Himalayan Fossils. Lower Triassic Cephalopoda from Spiti, 
Malla Johar, and Byans. Mem. Geol. Surv. India, Palaeont. 
Indica, Ser. 15,6(1): 1-186. 

Kummel, B. 

1961. The Spitsbergen arctoceratids. Bull. Mus. Comp. Zool., 123: 
499-532. 



16 BREVIORA No. 283 

1965. New Lower Triassic ammonoids from New Zealand. New 

Zealand J. Geol. Geophys.. 8: 537-547. 
1968. Additional Scythian ammonoids from Afghanistan. Bull. Mus. 

Comp. Zool.. in press. 

KuMMEL. B. and H. K. Erben 

1968. Lower and Middle Triassic cephalopods from Afghanistan, 
Palaeontographica, in press. 

KuMMEL, B. and G. Steele 

1962. Ammonites from the Meekoceras gracilitatis zone of Crittenden 
Spring. Elko County. Nevada. J. Paleont.. 36: 638-703. 

KUTASSY, A. 

1933. Fossilium Catalogus. I. Animalia. Pt. 56, Cephalopoda Triadica 
II. Berlin, pp. 371-832. 

Mojsisovics, E. V. 

1886. Arktische Triasfaunen. Beitriige zur palaeontologischen Char- 
akteristik der Arktisch-Pacifischen Triasprovinz. Mem. Acad. 
Imp. Sci. St. Petersburg, Ser. 7, 33: 1-159. 

Renz. Carl 

1945. Beitriige zur Slratigraphie und Paliiontologie des ost-mediter- 
ranen Jungpaliiozoikum und dessen Einordnung im griechischen 
Gebirgssystem, Teil I and 11. Eclogae Geol. Helvetiae, 38: 211- 
313. 

Renz, Carl and Otto Renz 

1947. Uhersicht ijber eine untertriadische Ammonitenfauna von der 
Insel Chios (Griechenland ). Verhandl. Naturforsch. Gesell. 
Basel, 58: 58-79. 

1948. Eine untertriadische Ammonitenfauna von der griechischen 
Insel Chios. Abhandl. Schweiz. Palaont., 66: 1-98. 

Smith, J. P. 

1932. Lower Triassic ammonoids ox North America. Prof. Pap. U.S. 
Geol. Surv., 167: 1-199. 

Spath, L. F. 

1921. On ammonites from Spitsbergen. Geol. Mag.. 58: 297-305, 

347-356. 
1930. The Eo-Triassic invertebrate fauna of East Greenland. Medd. 

omGronland, 83: 1-90. 

1934. Catalogue of the fossil Cephalopoda in the British Museum 
(Natural History). Part IV, The Ammonoidea of the Trias. 
London, pp. 1-521. 

Tozer, E. T. 

1961. Triassic stratigraphy and faunas, Queen Elizabeth Islands, 
Arctic Archipelago. Mem. Geo!. Surv. Canada, 316: 1-116. 



1968 SCYTHIAN AMMONOIDS FROM TIMOR 17 

1965. Latest Lower Triassic ammonoids from Ellesmere Island and 
northwestern British Columbia. Bull. Geol. Surv. Canada, 
123: 1-45. 

Waagen, W. 

1895. Salt Range fossils. Fossils from the Ceratite Formation. Pt. I, 
Pisces-Ammonoidea. Mem. Geol. Surv. India, Palaeont. Indica, 
Ser. 13, 2: 1-323. 

Wanner, J. 

1911. Triascephalopoden von Timor und Roti. N. Jahrb. Mineral. 
Geol. Palaont.. Beil.-Bd.. 32: 177-196. 

Welter, O. A. 

1922. Die Ammoniten der unteren Trias von Timor. Palaont. Timor, 
11(9): 83-154. 

(Received 11 July 1967.) 



18 BREVIORA No. 283 



EXPLANATION OF PLATE 1 

Figs. 1-9. Olenites simplex Welter from Timor. Figs. 1, 2, lectotype 
(Welter, 1922: pi. 169(15), figs. 1. 2) GPIBo 234a, XI; 3-5 paralectotype 
(Welter. 1922: pi. 169(15), figs. 3-5) GPlBo 234b, XI. 5; 6. 7, paralecto- 
type (Welter, 1922: pi. 169(15). figs. 6, 7) GPIBo 234c. XI. 5: 8, 9, 
paralectotype from which suture of Text-figure 3E was taken, GPIBo, X3. 

Figs. 10. 11. I'ickohlcrites siindaiciis (Welter) from unknown horizon 
on Timor. Holotype GPIBo 231. XI. 

Figs. 12, 13. Metadii^iiocerus frcenuini n. sp. from beds with managanese 
coated fossils, Nifoekoko. Timor. Holotype. BMNH C337()l. XI. 



1968 



SCYTHIAN AMMONOIDS FROM TIMOR 



19 










^1 






10 




13 






;!-* 



- w. *, 



11 



"t>!. 







8 



\ 




> 



20 BREVIORA No. 283 



EXPLANATION OF PLATE 2 

Figs. 1-16. Prosphin^itcs aiistini Hyatt and Smith. Figs. 1-3, from 
Meekoceras beds Union Wash, Inyo Range. California, holotype USNM 
75256, XI. 5: 3, 4. Prosphlngites slossi Kummel and Steele, from Meeko- 
ceras beds. Crittenden Spring, Nevada, paratype MCZ 5781, XI; 6. 7, 
Prosphini^ites slossi Kummel and Steele from Meekoceras beds, Crittenden 
Spring, Nevada, paratype MCZ 5616. Xl; 8, Prospliingites slossi Kummel and 
Steele from Meekoceras beds, Crittenden Spring, Nevada, paratype MCZ 
5615 XI; 9. 10. Prosphingites slossi Kummel and Steele, from Meekoceras 
beds, Crittenden Spring. Nevada, holotype MCZ 5614. XI; 11-16. Pros- 
phingites aiistini. Jonker Collection, Timor, Geology Department, University 
of Amsterdam, the suture of Text-figure 2F was taken from the specimen 
illustrated here on figures 15, 16. 



1968 



SCYTHIAN AMMONCIDS FROM TIMOR 



21 




Z^r^ 



^^^ 





/ 




4 



V- 



Vi 



 4, 



*»»H 



,,_-<^«£^H^'''^?^W"f 




.#; 



5^- 

8 







12 



11 






5 

V 




14 




t4,'«» 




/^- 



i'- 



^^: 




BREVIORA 

Museiuiini of Comparative Zoology 

Cambridge, Mass. 5 April. 1968 Number 284 



THE RELATIONSHIPS OF ANOLIS OF THE ROQUET 
SPECIES GROUP (SAURIA:IGUANIDAE) — III. 
COMPARATIVE STUDY OF DISPLAY BEHAVIOR 

George C. Gorman ^ 

INTRODUCTION 

In separate papers on chromosome cytology and electrophoresis 
of blood proteins (Gorman and Dessauer. 1966; Gorman and 
Atkins, 1967), the relationships of the roquet group of Anolis 
have been discussed in detail. The present paper is an attempt to 
use comparative ethology as an additional index of relationship 
and evolutionary history of the group. No attempt is made here 
to provide an "ethogram"' of any of the forms. Instead, some 
stereotyped action patterns that vary from island population to 
island population are compared as taxonomic characters. 

Taxonomy and Distribution 

Eight forms are considered in this paper. The species of the 
roquet group are found on all the islands of the Lesser Antilles 
from Martinique south to Grenada and west to Bonaire. To the 
north of Martinique are anoles of the hinuiculatus group; to the 
south, on Trinidad (a continental island), and to the west, on 
Curacao and Aruba, are members of the chrysolepis group (classi- 
fication into species groups follows Etheridge, 1960). These 
species groups seem to exclude each other, and the only recorded 
overlaps" (on Trinidad, St. Lucia, and Guayana) appear to be recent 
introductions (Gorman and Dessauer, 1965, 1966). Figure 1 
summarizes the distribution of the group. 



1 Present address: Museum of Vertebrate Zoology. University of Cali- 
fornia, Berkeley, California. 



BREVIORA No. 284 





66° 


64° 




! 1 

62° 60° 

OCCULATUS ^ Dominica 

ROQUET ' -A Wl3'''i"iq"e 




— 14° 








LUCIAE ^/j sj. Lucia 

TRINITATIS S. .Vincent 
GRISEUS EXTRE- 
■'Bequia MUS 


14 — 

Barbados 


— 12° 


LINEATUS 
Curacao ^^ 


BONAIRENSIS 
^ Bonaire i 


AENEUS 
RICHARDI ,' 

^j Grenada 


12'- 


— 




1 

68 '_ 


J 


RICHARDI^ -* Tobacjo 

CHRYSOLEPIS ,-V<S5?^ 

'^^^: Trinidad 




— 10° 








,.:<^^^ 


10" — 




66° 

1 


64° 

1 




62' 60° 

1 1 





Figure 1. The distribution of Anolis of the roquet species group (un- 
hatched islands). Names of forms considered, in capitals. Names of 
islands, in capital and small letters. The anoles on the islands indicated 
by oblique lines are not in the roquet group. 



By the use of chromosomes and blood proteins, the species 
group has been divided into several subgroups and species desig- 
nated. A basic dichotomy exists in chromosome number. Three 
forms have 2n=34, the other five have 2n=36. The latter is the 
presumably primitive number for the family Iguanidae (Gorman. 
Atkins, and Holzinger, 1967). 

Those anoles with reduced diploid number have always been 
considered very closely related on morphological evidence. Reduc- 
tion in chromosome number is rare and generally an advanced 
character, and this is taken as additional evidence that indeed 
these three anoles form a natural subgrouping. For reasons docu- 
mented elsewhere (Gorman and Dessauer, 1966), similarity in 



1968 DISPLAY BEHAVIOR OF SOME ANOLES 3 

hemoglobins was used to define species. Thus the group could be 
divided in the following manner: 

/ liiclae i rocjiict roquet ) 

\ trinitatis 2n = 34 3 roquet extremus ) co"i"i«n Hb 

2n = 36 >, hoiiaireiisis i aeueus 

J ricluirdi richardi 

I , ,. . common Hb 

( runarcli i^ri.seus 

Tn the 2n=36 group (primitive) A. r. ric/uirdi and A. r. gn'seus 
form a natural subunit characterized not only by proteins but by 
giant size. 

An hypothesis about the biogeographic movements of the group 
is being presented in a separate paper (Gorman and Atkins, 
1968). The defense of this hypothesis will not be repeated here. 
However, it is presented as a working model with which the be- 
havorial results may be compared. 

A. luciae is considered the most primitive member of the group. 
Reasons include chromosome number, size (intermediate between 
the giants and the other five forms), and geographic position (more 
or less in the center of the group.) From St. Lucia, St. Vincent 
was twice colonized, leading to two species, trinitatis (small) and 
gri.sen.s (giant). From St. Vincent, the two reached Grenada and 
through further differentiation have become aeneiis (small) and 
richardi (giant). A. ricluirdi has reached Tobago without further 
difi"erentiation. The small Grenada species colonized Barbados 
(extremus) and from there Martinique {roquet). Or. alternatively, 
the small St. Vincent form (trinitatis) colonized Barbados directly, 
followed by a double radiation from Barbados to Grenada and 
Martinique. 

Behavior in Anolis 

Because of the elaborate displays of these strongly territorial 
lizards, the Anolis species have been of long-standing ethological 
interest. Greenberg and Noble (1944) presented the most com- 
prehensive discussion of a single species, A. carolinensis. This 
report was based upon four years of observation of approximately 
6(X) lizards. Included in their paper are sections on aggression, 
courtship and mating, egg laying, dominance, subordination and 
hierarchies, territory, and sex recognition. Other studies, such as 
Kastle's ( 1963) on Norops f= Anolis) auratus. Carpenter's (1965) 
on A. townsendi, and Evans' (1938) on A. sagrei, provide valuable 
descriptions of displays and social behavior in single species. 



4 BREVIORA No. 284 

The only attempt at a broad comparative survey of Anolis be- 
havior was by Ruibal (1967), who described aggressive displays 
in twelve species of Greater Antillean Anolis. 

The genus is divided into two major groups (alpha and beta), 
based on an osteological difference (Etheridge, 1960), and sup- 
ported independently by chromosome differences (Gorman, 1965). 
Ruibal (1967) was interested in determining if there was any 
parallel between the alpha-beta dichotomy and behavior. Although 
Ruibal found species-specific differences, he wrote, "the analysis 
of the initial response in these anoles demonstrates that the behav- 
ioral patterns are of limited value in determining relationships 
within the West Indian species of Anolis." 

My own study, presented here, indicates the value of behavior 
as a systematic character within at least one species group. 

METHODS 

In 1963-64, I spent 10 months in the Caribbean area, based at 
the University of the West Indies in Trinidad. Subsequently, several 
additional brief visits were made to the islands where the roquet 
group occurs, as well as to neighboring islands. At first, the lizards 
were simply observed in the field and extensive notes on ecology 
and behavior were prepared. Later, they were studied in cages at 
the University of the West Indies. In general, two adult males were 
placed on opposite sides of an opaque partitioned cage. Several 
hours to days later, the partition would be removed and social 
interactions described. To gather quantifiable data the lizards were 
photographed on 16 mm motion picture film. The bulk of the data 
presented are from analysis of films made in the field. In some 
cases the films are of natural interaction, but high intensity terri- 
torial disputes are not frequent in natural situations; they are easily 
elicited, however, by tethering an "intruder" to a long pole and 
placing him within the territory of a resident. Films were analyzed 
on a Weinberg-Watson single frame analyst. This allows frame-by- 
frame analysis of the temporal sequence of displays. Amplitudes of 
movements were not measured precisely but were estimated. No 
great emphasis has been placed upon amplitude differences in 
display. 

Elicitation of Display 

Iguanids respond to visual cues. The introduction of a mirror 
into a cage elicited highly stereotyped displays identical to those 



1968 DISPLAY BEHAVIOR OF SOME ANOLES 5 

seen in the field in natural encounters. Carpenter (1962) has re- 
ported results of mirror experiments in Sceloporus, and a rather 
accurate description of an A. roquet extremiis display to a mirror 
was written for a popular audience by Dr. Maurice Burton (London 
Illustrated News, 29 August 1964). 

Neither the opponent's dewlap color nor the dewlap per se seems 
to be of great importance in eliciting a response from a resident. 
Greenberg and Noble (1944) found that restraining the dewlap 
by means of a coUodian coating did not affect the social status of 
the male. Dominant animals were still able to hold territory and 
win combats with normal rivals. In addition, normal males dis- 
played to males unable to extend the dewlap. 

I have found that members of the roquet group in captivity will 
display to other species if these are in the same size range. 

A planned experiment to study releasers by use of models pre- 
sented to residents in the field was thwarted at the outset. A. trini- 
tcitis are very abundant and quite willing to display to other anoles 
in the Botanic Garden in St. Vincent. A life-size wooden model of 
a male triniiati.s was constructed and. I thought, accurately painted. 
A slit was made in the throat region for insertion of paper dewlaps 
of the "right" or "wrong" color. If males responded aggressively 
to the model, then variations in size and color were planned to 
elucidate exactly which parameters were important to a resident 
territorial male. However, responses to the dummy were uniformly 
ncizative. 

Experiments involving a decapitated intruder were somewhat 
more successful. The decapitated head was pinned to a piece of 
sponge, cut approximately in the shape of a lizard torso. Several 
resident males attacked and/or displayed actively to this model, 
and even to the decapitated head alone, when dangled freely. This 
experiment was terminated when a griseus ran down from the 
crown of a tree and grabbed and ate the trinitatis head. 

In a reciprocal trial, a head-sized piece of sponge was pinned to 
the body of the decapitated lizard. This did not elicit aggressive 
display. Some males were painted bright red, others bright white. 
Results in several trials were equivocal. Some residents displayed 
or attacked, others fled from the intruder. Thus, something in the 
configuration of the head probably acts as a releaser. 

Another apparently important component in elicitation of dis- 
play is movement on the part of the intruder. A tethered male will 
usually flatten himself on the substratum in a classically submissive 



6 BREVIORA No. 284 

posture.^ The resident will watch. Slight jiggling of the intruder is 
sufficient to elicit immediately a high intensity response (biting at- 
tack or display). 

Description of Display 

Carpenter (1962) has standardized his descriptions of iguanid 
behavior by describing eight characteristics of display — site, posi- 
tion, posture, movement type, parts moved, units of movement, 
sequence, and cadence. 

Ruibal listed and defined the following movements and postures 
to describe aggressive encounters in Anolis. 

1 . Dewlapping — the characteristic erection of the large gular 
appendage. 

2. Bobbins — the vertical motion of the head. 

3. Pushups — an extension of the limbs to raise the body off 
the ground. 

4. Tail wags — vertical or horizontal movement of the tail. 

5. Tail lashes — a very exaggerated movement of the tail in 
a broad whipping manner. 

6. Tongue — the tongue may be everted and protruded from 
the mouth and in some species may become purple with en- 
gorged blood. 

7. Nuchal and dorsal crests — the dorsal regional of the neck 
and the middorsal region of the body are usually raised as 
thin fins. 

Application of these descriptions to the roqiict species group 
would be as follows: 
A . The eight categories of Carpenter. 

1 . Site: All the anoles of the roquet group are strongly arboreal. 
Most daytime observations are made of animals on tree trunks 
four to seven feet above the ground. Most displays are given when 
the lizards are on tree trunks; however, during a battle they will 
display from the ground. When they display they try to find a 
slightly elevated site, such as a knot on a tree. 

2. Position: Orientation is usually perpendicular to the recipient 
of the display. If the opponents are parallel, then the heads are 
facing each other. The position chosen maximizes the opponent's 
view. 



1 Darwin (1872) pointed out that submissive postures were often the 
antitheses of aggressive ones. Thus, in the anoles in submissive posture the 
crests are down, dewlap in. body low. 



1968 DISPLAY BEHAVIOR OF SOME ANOLES 7 

3. Posture: In aggressive encounters, nuchal and dorsal crests 
are generally raised. As noted by Greenberg and Noble (1944) 
for A. carolinensis. "The initial hostile reaction is a puffing or 
gorging of the throat followed almost immediately by lateral flat- 
tening so that the male appears much larger from side view." 
This serves as an excellent description for the roquet group and 
shall be referred to as "engorged head." 

These three categories are not of further use in this comparative 
study because they are constant for the species group. 

4-6. Movement type, parts moved, and units of movement are 
encompassed by Ruibal's categories, and are discussed below. 

7. Sequence: The order of appearance of units of movement 
will be exemplified by quotations from notes on actual encounters. 

These categories again are consistent for the species group. 

8. The measured time for the performance of each unit of 
movement. This is the important category employed in comparing 
displays within the species group. 

B. The seven movements of Ruibal. 

1. Dewlapping: The dewlap of the roquet group and all its 
near neighbors is well developed. The cadence of extension differs 
between the roquet group and the anoles that geographically sur- 
round it. 

2. Bobbing: In the roquet group all eight forms bob the head 
in high intensity display, and each form shows a temporal se- 
quence that is unique. I also refer to nodding, in which the head 
is raised and lowered very slowly. This is a less stereotyped move- 
ment and seems to occur at lower intensity. 

3. Pushups: Again, all forms in the group rise up on all limbs 
in high intensity display. Some have also been observed to include 
"rhythmic movements" as a part of the challenge display. This is 
a series of movements at very precise intervals — it may simply be 
head bobs, or pushups, or pulsating locomotion. 

4. Tail wags: Horizontal movement of the tail is seen under 
situations of stress. It occurs when two anoles are actually light- 
ing and is not seen in the clearly dominant animal. It also occurs 
when an anole is captured by a herpetologist and is held in the 
hand. 

5. Tail lashes: All species of the roquet group may use the 
tail in a challenge display. The frequency and amplitude of tail 
lashes is an important character in tracing relationships. 

6. Tongue: Several members of the roquet group have been 
seen to protrude the tongue during display. I do not know how 
frequent or how widespread this movement is. The tongue is 



8 BREVIORA No. 284 

engorged with blood and quite bright in color. At the termina- 
tion of the display, while the dewlap is being retracted, the mouth 
is opened and closed several times as though a swallowing move- 
ment were occurring. This has been recorded for A. r. roquet, A. 
tiinitatis and A. bonairensis and may be present in the other 
species. 

7. Nuchal and dorsal crests: Present and raised in all species 
during aggressive encounters. 

THE DISPLAY CATEGORIES OF THE ROQUET GROUP OF ANOLIS 

There are three obvious categories of display in this species 
group: activity bobs, courtship, and aggressive display. Only the 
last was studied in detail. 

(1 ) The activity bob corresponds to Carpenter's (1962) "As- 
sertion Display." For example, in changing position a lizard may 
run a few feet, stop, bob, run a few more feet, stop, bob, etc. 
This bob is also commonly seen after feeding. I have reason to 
believe that it is stereotyped and may be species-specific. However, 
I do not have enoutih data to discuss it further. This bob is most 
commonly given by females. 

(2) Courtship was difficult to study under field conditions, as 
most displays directed by males to females were followed by rapid 
departure by the female. There are no obvious differences among 
the species in courtship display, and my observations are con- 
cordant with those of Greenberg and Noble (1944) on the fol- 
lowing points: 

a) There is no sign ot aggression; i.e., engorged head, crest 
erection, and tail jerks are all absent. 

b) The approach pattern differs from the aggressive encounter 
in that there are numerous rapid low amplitude bobs and an irregu- 
lar, jerking walk. 

(3) Aggressive display. 

The most elaborate of the displays are also the most stereotyped, 
hence the most useful for analysis of evolutionary relationships. 
Intensity of display obviously varies, as has been pointed out by 
Kasde (1963) and Ruibal (1967). Ruibal limited his study to 
what he called the "initial response," which he defined as "the 
immediate and first response of the test animal to the decoy." 
Later he states that it is comparable but not identical to the chal- 
lenge display of Carpenter ( 1962) — "the initial response actually 
represents the first part of the challenge display." 



1968 DISPLAY BEHAVIOR OF SOME ANGLES 9 

I agree completely with Ruibal that there is a predictable, stereo- 
typed portion of lizard display behavior. However, the term 
"initial" in RuibaFs phraseology is unfortunate, because the lit- 
erally immediate response of a resident male to an intruder is 
quite unpredictable: the resident may flee, go into the engorged 
head posture and wait, attack by biting, or display at varying in- 
tensities. I prefer to use the term challenge display. 

All eight members of the roquet group have challenge displays 
which follow a definite sequence of actions: (1) extend dewlap; 
(2) rise high on all four hmbs; (3) raise the tail, then lower it; 
(4) lower the body; (5) give rapid head bob; (6) retract dewlap. 
The following excerpt from my field notes will exemplify this 
pattern: 

A. r. roquet — Martinique — 2 July 1965: 

A tethered male intruder is placed about 12 inches above a resident 
male (R) perched on a tree. The intruder starts running away from 
resident. R slowly moves up several inches, pauses. Intruder jiggled. 
R backs away. R turns perpendicular to intruder. Intruder moves 
and R follows, stopping a foot above. Head is in engorged position 
R moves to two feet above intruder; extends dewlap fully; rises 
high on all four limbs; 5 tail lashes; lowers body; dewlap half re- 
tracted. R pauses for several seconds. Extends dewlap fully while 
body rising high on all four limbs. Tail raised high and then lowered 
slowly. Bobs head. Retracts dewlap. Display is repeated again. 
Note that the sequence of movements is the same in the two 
full displays, but in the first instance head bobbing did not follow 
the tail movement. 

The six sequential steps of the challenge display are present or 
absent in independent fashion, but when they do appear they are 
evidently never out of order. That is, the dewlap might not be 
extended but the resident male still lashes his tail and then bobs; 
or he may omit the bobs, or he may not fully retract the dewlap 
at the end of the sequence. However, he will never bob, then 
lash the tail, and then extend the dewlap. 

Some of the steps are better correlated than others. The tail 
movement almost always occurs with the dewlap fully extended. 
And, if there be a rapid bob with dewlap extended, the dewlap is 
almost always retracted at the end of the sequence unless the dis- 
playing lizard immediately launches into a new sequence. 

The high intensity challenge display is interpreted as occurring 
when there is an intermingling of attack and withdrawal stimuli. 
The elaborate display is often preceded by backing away from the 
intruder. Indeed, the displays often follow unsuccessful direct 



10 HREVIORA No. 284 

attacks. The following encounters may serve as illustrations. 

1. A. trinitatis — St. Vincent — 8 July 1966: 

Intruder is placed three feet below resident who is 10 feet up in a 
coconut tree. R runs down, bites at intruder. Bites again. They 
lock jaws. R breaks away and runs back up. Bobs rapidly. Attacks 
again several times, biting at the head of intruder. R backs away. 
Extends dewlap half-way. Rapid bob. R attacks again. Grab jaws. R 
flipped to ground. 

2. A. r. roquet — Martinique — 3 July 1965: 

Tethered male intruder placed four feet below resident who turns 
immediately, runs down and stops one foot above intruder. Pauses. 
Then rushes at intruder and bites at head; extends dewlap, bobs. 
Runs jerkily above intruder; full dewlap extension, head bob, dewlap 
slowly retracted. This is repeated again. Then R moves toward in- 
truder and bites at him. K then backs away and moves to one foot 
above intruder. Dewlap fully extended, head bobs, then dewlap 
retracted. Repeated several times. Each time there is a slight move- 
ment of the intruder, there has been a full dewlap extension followed 
by head bobbing. Resident then runs to one foot above intruder. 
Faces down and is approximately perpendicular. Rises high on all 
four limbs, tail comes up slowly, full dewlap extension; body lowered, 
head bob, dewlap retracted. Then R moves up and away from 
intruder. 
In Trinidad. I filmed a long sequence of a male A. aeueus dis- 
playing to a tethered intruder in which the resident gave the full 
display, then ran up and away from the intruder. I then moved the 
intruder up a comparable distance, the resident again quickly dis- 
played and moved away again. This was repeated several more 
times. 

These examples serve to support the hypothesis that the elab- 
orate challenge display is not simply a ritualization to avoid physi- 
cal fighting, but rather a display given in a situation with conflicting 
stimuli of attack and flee. Under natural field conditions these 
displays have been seen only when the opponents were extremely 
evenly matched. Usually the engorged head of a dominant resident 
is enough to stimulate flight in a subordinate or intruding male. 

Combat is by no means always the initial high intensity response, 
as the following notes indicate. 

A. r. roquet — Martinique — 3 July 1965: 

Intruder placed near male resident who immediately goes into 

engorged head posture and moves down toward intruder. R extends 

dewlap fully and rises up on all four limbs; three tail lashes; dewlap 

slowly retracted. 

Some encounters are settled without actual biting. I once 

witnessed a very lengthy dispute between two male A. trinitatis in 



1968 DISPLAY BEHAVIOR OF SOME ANGLES H 

the Botanic Gardens of St. Vincent. Both were on a broad tree. 
At the start, one was about two feet above the other. After some 
fifteen minutes with dozens of high intensity displays, they slowly 
moved towards one another and, when contact was made, they 
butted heads, snouts pointing downwards. Then one backed off 
and climbed up and around the tree. 

Although all these encounters describe males, females also dis- 
play to females. Although the secondary sexual characters of 
dewlap and crests are much reduced, and the females are also 
much smaller than adult males, the postures and motions appear 
identical to those of males. I never saw a natural interaction be- 
tween females, but did observe and film female display by use of 
a tethered intruder. 

SYSTEMATIC SECTION 
I. Definition of the roquet species group on the basis of 

BEHAVIOR 

Although each of the eight forms has a display which is quite 
distinct because of variation in cadence, all members of the species 
group have basically a very similar display. In the use of the 
dewlap, the roquet group differs from its near neighbors in the 
chrysolepis and bimaculatus groups. Thus, behavior provides fur- 
ther evidence that the group is a natural evolutionary unit. 

In all eight forms, the dewlap is extended and held out for the 
entire display sequence, and sometimes is not retracted at the end 
of the sequence. The dewlap is never pumped in and out in any 
rhythmic manner. This contrasts sharply with the other Lesser 
Antillean anoles. Figure 2 illustrates the challenge display of 
A . roquet extremus. 

In place of the activity bob. members of the binioculatus group 
seem to have a true assertion display sensu Carpenter. I have 
watched and filmed A. oculatus on Dominica pumping the dewlap 
in and out in a slow rhythmic manner — even when no recipient of 
the display was apparently present. In high intensity challenge, 
dewlap pumping is an integral part of a complex display which 
also involves bobbing and tail movements. 

A. lineatus (chrxsolepis series of beta anoles) of Curacao also 
pumps the dewlap. This species is quite unrelated to the bimacula- 
tus group (alpha anoles), and serves to illustrate the problem 
Ruibal (1967) encountered in a search for a major diagnostic 
character to split the genus by means of behavior. 



12 



BREVIORA 



No. 284 




Figure 2. Challenge display of A. roijuet extrciniis. Dewlap fully ex- 
tended, tail up, and body high off the perch. 



1968 



DISPLAY BEHAVIOR OF SOME ANOLES 



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14 BREVIORA No. 284 

A. clirysolepis of Trinidad and the South American mainland 
displays by flashing the dewlap extremely rapidly and in a stereo- 
typed manner. The entire display appears to consist of dewlapping. 
The head is held high but is not bobbed. Figure 3 show.v clirysolepis 
in challenge positions. 

Because dewlapping is widespread in both the alpha and beta 
anoles, it is interpreted as a primitive character which has been 
secondarily lost in the roquet group. Further evidence for this will 
be presented below. 

IT. Description of the challenge display of A. luciae 

A. luciae is considered the most primitive member of the species 
group. It is thus worth analyzing its display in some detail, to 
serve as a basis for comparison. 

Descriptions of interactions 

1. Forestiere. St. Lucia — 6 July 1965 

Resident male is high in a tree. Tethered intruder placed four feet below 
him. R rises up on all limbs, extends dewlap, tail goes up once, then 
down. Bobs head. There are four distinct series of bobs while the dewlap 
is held out. Dewlap slowly retracted. 

2. Vigie Beach. St. Lucia — 6 July 1965 

Resident male is in a breadfruit tree. Intruder placed one foot below him. 
R immediately orients perpendicularly to intruder, head facing head. Resi- 
dent rises up high. Extends dewlap and bobs head. The head bobs are 
in four distinct series, with a partial dewlap retraction following each series. 
Intruder turns, and R turns, maintaining perpendicular orientation. Repeats 
display, again with four series on a single dewlap. 

The display of luciae may be characteriezd as distinct from other 
members of the roquet group by the following: 

1 ) Tail — the tail is infrequently used in display sequence. 
When it is used, it is raised once slowly, then lowered. 

2 ) Head bobs — luciae has a head bob sequence that is ex- 
tremely long in duration and that is clearly divided into several 
series. That is, a series of bobs is followed by a pause, followed by 
a series of bobs, followed by a pause, etc. (Figs. 4L and 5L ). 

3) Partial dewlap retraction — during the long pauses, the 
dewlap is very slightly retracted, then re-extended. A. luciae 
is the only member of this species group to move the dewlap at all 



1968 DISPLAY BEHAVIOR OF SOME ANGLES 15 

in mid-sequence. This movement might be a vestige of the dewlap 
pumping common to most Anolis. 

The following quantitative data were analyzed: 

1) Number of peaks preceding a pause. A pause was arbi- 
trarily defined as 6/16 of a second (N.B. all films were made at 
16 frames/second, and time units discussed are in 16th of a second 
or frame) with no head movement. A peak was defined as the 
highest point the head reached before the next downward move- 
ment. Thus sub-peaks were not counted. The distribution in num- 
ber of peaks per bobbing series proved to be quite variable, with 
a range from 1-18. However, the mean is 5.2 and 54 per cent 
of the sequences had 4, 5 or 6 peaks. 

2) The time interval of bobbing series was also measured. 
Again the range is very broad, 2-47 frames. The mean is 18. 

3) The pause lengths between bobbing series were also meas- 
ured and the distribution of these time intervals is shown in Figure 
7. Note that the first category is also the largest. This is probably 
an artifact — i.e. a pause was defined as 6 or more frames. This 
does not mean that it is necessarily a "real'" pause to the animal. 
If 10 or more frames were chosen as the arbitrary limit, there would 
be a gradual rise in frequency to the 30-33 frame class, and then a 
drop. No relationship could be demonstrated between the number 
of peaks in a sequence and the length of the following pause. 

Two other categories also easily lent themselves to quantitative 
analysis: number of bobbing series in a single display sequence, and 
total time elapsed in the full bobbing sequence. Because of the 
repetitious nature of the long displays and the high cost of motion 
picture film, few full sequences were filmed. Some examples are 
tabulated below: 



. bobbing 


series 


Total time 


Full 


sequence filmed 


5 




10.5 sec. 






no 


3 




8.5 






yes 


5 




19.3 






yes 


5 




12.5 






yes 


4 




12.5 






no 


3 




6.7 






yes 



The total elapsed time for a full bobbing sequence in A. luciae 
is far greater than for any other member of the species group. 



16 



BREVIORA 



No. 284 





1968 



DISPLAY BEHAVIOR OF SOME ANOLES 



17 




Figures 4-6. Head bobbing sequences in roquet group AnoUs. Time 
intervals are one-half second between successive dots on upper and lower 
borders of each figure. Long pauses are indicated by a break, with the 
time interval in seconds written in. 

Figure 4 

L. A. Incicie 

B. A . honiiirensis 

T. A . irini talis 



Figure 5 



L. A. Iiicicic 

G. A. ricliardi griseiis 

RI. A. r. ricliardi 



Figure 6 

E. A. roquet e.xtrenuis 
RO. A. r. roquet 
A. A. (iciieiis 



18 



BREVIORA 



No. 284 





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PAUSE LENGTHS (FRAMES) 



34-37 



> 37 



Figure 7. The frequency distribution of pause lengths which precede 
subsequent bobbing series in A. Iiiciae. One frame is equivalent to one- 
sixteenth of a second. 



ITT. Presumed derivatives of A. luciae 

A. luciae, for reasons stated earlier, is presumed to be the primi- 
tive member of the species group. Following the model presented 
in the zoogeographic study of the roquet group, three forms are 
believed to be derived directly from luciae: double colonization of 
St. Vincent from St. Lucia gave rise to trinitatis and griseu.s, and 
colonization of Bonaire gave rise to bonaireusis. In this section we 
shall compare the displays of these three forms to those of A . luciae. 
a) A. bonairensis. 

The relationships of this species to the remainder of the species 
group are probably the least clear. Bonaire is far west of the 
Lesser Antillean chain (see Figure 1), but the anoles on the adja- 
cent islands just west of Bonaire are certainly unrelated to bonair- 
ensis. When the species was described (Ruthven, 1923), it was 
compared with roquet and aeneus. The osteological study of 
Etheridge placed this form very close to the other members of the 



1968 DISPLAY BEHAVIOR OF SOME ANOLES 19 

roquet group, and this was confirmed by blood proteins and 
chromosomes. In our paper on blood proteins (Gorman and Des- 
sauer, 1966), it wa* pointed out that although boiiaircusis was 
clearly a member of the roquet group, it was not particularly close 
to any one form. We speculated that it might be derived from ae- 
neus, since Bonaire is due west of Grenada'and the general current 
flow is westerly. 

However, when we examined chromosomes (Gorman and At- 
kins, 1967), we found that bonairensis belonged to the more 
primitive group (2n=36), making it highly improbable that it was 
derived from aeneus. The geographically intermediate form A. 
bonairensis blanquillanus has been unavailable for study. 

It is, of course, possible that Bonaire was colonized independ- 
ently by a mainland South American Anolis. However, that is not 
likely. Although the roquet group forms are clearly allied to South 
American alpha anoles, they share a unique complex of osteological 
and biochemical characters which most likely evolved in the Lesser 
Antilles. 

Thus our hypothetical ancestor would be a ///c/ac-like or trini- 
tatis-Wkt anole. J. D. Lazell, Jr., who is studying the external mor- 
phology of Lesser Antillean anoles, relates bonairensis to luciae 
(personal communication). 

The display of bonairensis is basically similar to other members 
of the species group. It may include tail lashes and head bobs. 

Description of an interaction 

4 May 1964 

Intruder placed on a tree. Resident male comes running across a fallen 
stump to stop 2 feet from intruder. He runs at least 15 feet. Head is in 
engorged posture. 1 jiggled intruder. R immediately extends dewlap fully, 
rises high on all four limbs, tail is raised once, dropped, body lowered. 
Bobs head and retracts dewlap. 

The head bob sequence of bonairensis is unique in the large 
number of peaks and the great length of time. The bobbing may or 
may not be broken up by pauses of 6 frames or longer. In 20 filmed 
sequences, 14 are without pauses. These sequences have a range 
of 12-23 peaks (mean 17.7) and a time range of 43-95 frames 
(mean 62.2). Of six sequences with pauses, five have only a 
single pause and one sequence has two pauses. This last case may 
be an artifact, for the first pause follows a single slow head rise, 
which is probably not part of the bobbing sequence. As opposed 
to luciae, which has pause lengths of as long as 40+ frames, the 



20 BREVIORA No. 284 

pauses of honairensis are all short, ranging from 6-10 (mean 8.3) 
frames. 

Those sequences which have pauses range in total peak number 
from 13-24 (mean 17.7), which is almost identical to the non- 
pause displays. The total time intervals range from 47-91 frames 
(mean 69.0), which is within the range of non-pause displays. 

Thus the head bobbing sequence of A. bonairensis is best 
described as a long single sequence with many peaks, occasionally 
broken into two series. The display of honairensis could be derived 
from the lucicie display with but one minor alteration: elimination 
of the pauses which break the bobbing sequence into distinct series. 

Bonairensis also uses tail lashing in high intensity displays. In 
this respect the species resembles other members of the roquet 
group more than luciae. 

b) A. trini talis. 

A. trinitatis is the widespread small anole on St. Vincent. It also 
occurs on Trinidad, where it was probably introduced by man. On 
Trinidad it hybridizes with aeneus. A detailed analysis of hybridiza- 
tion is being prepared. The data on trinitatis presented in this 
paper include St. Vincent and Trinidad specimens, as no discernible 
difference could be found in their displays. 

The challenge display of trinitatis consists of very rapid tail 
lashes followed by rapid bobs. Long pauses are infrequent. A slow 
tail movement has not been observed. Figure 8 compares the 
characteristic rapid tail movement of trinitatis with that of ex- 
tremiis, which is slow. Descriptions of encounters appeared in the 
introductory part of this paper. 

The variation in 30 rapid bob sequences of trinitatis is: 

Rapid bob time interval No. peaks per bobbing sequence 

range (sixteenths of a sec.) 12-31 2-10 

mean 23.0 5.7 

56% are 4. 5, or 6 

A typical rapid head bob of trinitatis is presented in Figure 4T. 

The full bobbing sequence in trinitatis resembles a single series 
of bobs within a sequence in A. luciae. The display of trinitatis 
differs from that of luciae in the rapid tail lash and in the shorter, 
non-paused head-bob sequence. The latter, however, could easily 
be derived from a foreshortening of the luciae display. 

c) A. ricliardi griseus. 

Although this is the largest, it is paradoxically the shyest mem- 
ber of the roquet species group; that is, A. r. griseus is quick to 



1968 DISPLAY BEHAVIOR OF SOME ANGLES 21 

flee at one's approach. As a result, I have few data. No natural 
aggressive interactions were seen. Two males did display to tethered 
intruders, and their several bobbing sequences were recorded on 
film. These two males were filmed in the summers of 1964 and 
1966, respectively, and on opposite ends of St. Vincent. The display 
action patterns are quite similar. Thus, although lacking details on 
the total range of variation display, the data are probably valid 
from the broadly comparative viewpoint. 

High intensity display consists of dewlap extension, rapid tail 
lashes, head bobs, and dewlap retraction. 

The head bobs are in series, as in /I. luciae. Figure 5G illustrates 
a typical bobbing sequence of griseus. 

I do not have enough pauses measured to compare in detail, but 
here too the distribution seems similar to luciae. Of 1 1 measured 
pauses, 5 are between 6-9 frames, 5 are 20 or more frames, and 
only one falls between these two intervals (compare with Figure 7). 

There does seem to be a reduced number of peaks per series; 
in 18 measured series the mean is 3.3, and 61 per cent of the 
sequences have 1, 2, or 3 peaks. In luciae the mean is 5.2, and 54 
per cent of the series have 4, 5, or 6 peaks. However, the mean 
time interval per bobbing series ( 16.6 frames) is very close to that 
of luciae (18.0). This implies that the head bobs of griseus are 
broader (slower) than those of luciae. Such a trend can be seen 
by comparing Figures 5L and 5G. 

The total time of a bobbing sequence, adding all series and 
pauses, may also be shorter than in luciae. (Compare these data 
with those on p. 15.) 



No. bobbinij 


; series 


Total time 


Ful 


1 sequence filmed 


2 




5.0 sec. 




yes 


2 




4.8 




yes 


5 




8.4 




yes 


4 




7.3 




yes 



In summary, the displays of griseus are quite similar to those of 
luciae. In head bobbing there seems to be a tendency for fewer 
peaks per series without a significant decrease in the time of the 
series, whereas the total time of a bobbing sequence is reduced. 

Also, griseus may use several vigorous tail lashes in the challenge 
display, whereas luciae does not use the tail frequently, and does 
not use multiple lashes when it does use the tail. 



22 BRHVIORA No. 284 

IV. Presumed derivatives of griseus and trinitatis 

a) A. r. richardi. 

A. r. richardi is very closely related to griseus. On the basis of 
blood proteins (Gorman and Dessauer, 1966), the two forms were 
considered conspecific and distinct from all other members of the 
species group. 

Unlike griseus, richardi is not shy, and numerous displays have 
been observed in Grenada and in Bequia, the northernmost of the 
Grenadines. No displays were studied in Tobago. 

Descriptions of encounters 

1. St. George's, Grenada - 17 July 1965 

A male resident is seven feet up in a mango tree. Intruder placed two 
feet helow. R immediately into engorged head posture watching intruder. 
Intruder runs up. R extends dewlap fully, rises up high on all limbs, tail 
up a bit, then down slowly. Again up and then down slowly, dewlap still 
out. Intruder removed. 

This is interpreted as a low intensity response. Male richardi may 
rise up slowly and the head is sometimes slowly nodded. The fol- 
lowing is an example of a high intensity response, the type of 
response that was analyzed. 

2. Beausejour. Grenada - 17 July 1965 

Intruder placed near resident male. Dewlap extended. No tail motion. 
Bobs vigorously about 7 times. Dewlap partially retracted. Intruder 
jiggled. R rises up, lashes tail four times, then a fifth time with lower 
amplitude. Dewlap is held out. R looking at intruder. Intruder removed. 

The high intensity display of richardi is characterized by vigorous 
tail lashing, followed by head bobbing. 

Unlike griseus, there are not multiple series, but rather a single 
series, in a bobbing sequence. Figure 5 RI shows an example of 
richardi head bobs. These bobs are broad, i.e. the rise and fall are 
relatively slow and the peak is flatter than in griseus. The display 
appears much more stereotyped than that of either hiciae or griseus, 
both in number of peaks (range 4-7. mean 5.8) and in total elapsed 
time in the bobbing sequence (range 39-58, mean 47.3 frames). 
Although there are only eight filmed displays, numerous other 
displays observed in the field seemed to conform to these ranges. 

b) A. aeneus, extremus and roquet. 

These three forms are very closely related and were considered 
conspecific by Underwood (1959), although he later changed his 
mind (1962). They were first studied biochemically by Gorman 



1968 



DISPLAY BEHAVIOR OF SOME ANGLES 



23 



and Dessauer (1965), who found that on the basis of electro- 
phoresis of serum proteins they were closely related, and a linear 
chain of relationship could be established: aeneiis and extrcmus 
differ in hemoglobin mobility, extremus and roquet in transferrin 
mobility, and twneus and roquet differ in both these characters. 
In a more detailed study of blood proteins encompassing the entire 
species group (Gorman and Dessauer, 1966), extremus was rec- 
ognized as a subspecies of roquet, and aeneus was recognized as a 




Figure 8. Comparison of the slow tail movement of A. roquet extreiniis 
(upper) with the rapid tail movement of A. trinitatis (lower). Numbers at 
the tip of the tail indicate frames elapsed from start of display (one frame 
equal to 1/16 second). 

A. Tail rise in extrcmus. The tail is slowly arched up. A full two 
seconds elapse before peak is reached. 

B. Tail lowering (same display). The tail is dropped more quickly 
than raised. Multiple dashes at 36 indicate tail was in rapid motion and 
film was blurred on that frame. Total time interval for sequence is 40 
frames {IVi seconds). 

C. Tail rise in trinitatis. The peak is reached in 3 frames. 

D. Tail lowering (same display). The tail is down one frame later. 
Total time interval. 4 frames ( Vi second). The drawings were made by 
superimposing images from a single frame projector. 



24 BREVIORA No. 284 

species. The three forms are obviously very close to trinitatis. The 
chromosome study showed that trinitatis was a likely ancestral form, 
as it had the primitive chromosome number. 

In display behavior, there are great similarities among the three 
in gross movements. That is, the sequences of movements are 
identical, all have a variable tail movement, slow to moderately 
fast — but always slower than trinitatis (Figure 8) — and all bob 
the head. But here there are consistent differences. 

A. r. extremus has a head bobbing pattern that appears to be the 
most rigidly stereotyped. There are several rapid peaks and the 
display is over. There are never any long pauses. Figure 6E illus- 
trates an example of this head bob. Thirty-two head bobbing 
sequences were filmed, the number of peaks per sequence ranged 
from 2-5 (mean 4), and the time interval in frames ranged from 
6-14 (mean 10.6). 

Although in blood proteins c.xtrenuis seemed to be intermediate 
in this series of three forms, in head bobbing, roquet and aeneus 
are more similar to each other than to extremus. Both forms have 
displays characterized by a long pause; thus the total time interval 
of the bobbing sequence is longer than that of extremus. A. r. 
roquet tends to have more peaks than does aeneus, while aeneus 
tends to have longer pauses and thus a longer bobbing sequence. 
However, there is considerable overlap, as tabulated: 

Total time of bob sequence 
Rapid bob No. peaks (sixteenths of a sec. ) 

no. range mean range mean 

aeneus (25) 2-4 2.8 27-62 44.6 

roquet (28) 2-9 4.3 18-47 33.3 

Figures 6A and RO compare rapid bobs in the two forms. 

The examples depicted accurately show the typical differences in 
the head bobbing sequences between the two forms. Not only are 
there fewer peaks and a longer pause in aeneus, but it also may be 
seen that in the typical display of aeneus a head rise follows the 
drop after the long pause, whereas in roquet, following the long 
pause the head is dropped and the bobbing sequence is terminated. 

A. trinitatis is the presumed ancestor of this group. Chromosome 
loss being rare, it is highly probable that trinitatis gave rise to only 
one of these forms. Two alternatives are plausible. 

1. Colonization of Barbados, loss of chromosome pair, giving 
rise to extremus. A. r. extremus then sends out two radiations, one 
to Martinique where a transferrin difference evolves (roquet), the 



J968 DISPLAY BEHAVIOR OF SOME ANGLES 25 

Other to Grenada where a new hemoglobin evolves {aeneits). The 
display of extremus is not very different from that of trinitatis. Both 
have head-bob sequences consisting of several peaks in a short time 
interval. The sequence of trinitatis is more variable, in both number 
of peaks and time interval and is occasionally punctuated by pauses. 
The similarity in display of roquet and aencus would have to be 
accounted for by convergence. 

2. Colonization of Grenada, loss of a chromosome pair, giving 
rise to aeneus. Evolution of the aeneus type display (increase in 
tendency for long pause, decrease in number of peaks) is followed 
by colonization of Barbados {extremus), and from there coloniza- 
tion of Martinique {roquet). There would be minor change in the 
head-bob display of roquet, and evolution of a new head-bob 
display in extremus. 

There is one other bit of behavioral evidence that can be 
examined to assess these two alternatives. Members of the roquet 
group have a movement which I termed "rhythmic motion." This 
may consist of bobbing, or walking, or climbing in a very jerky 
manner. The animal will move, pause, move, pause, etc. The pause 
is long and the movement is very sudden and exaggerated. The 
actual context of the display is not fully understood by me, but it 
appears to occur at a somewhat lower intensity than the tail move- 
ments and the rapid head bob in male-male territorial encounters. 

This movement was seen most frequently in extremus and aeneus, 
but was also filmed occasionally in trinitatis and roquet. 

In A. r. extremus, in eleven different displays, 100 intervals- 
between-bobs were measured. The range in frames (one frame = 
1/16 sec.) was 5-12 (5-9 in 97 per cent of the intervals) and the 
mean interval between bobs is 7.27. 

In A. trinitatis in four displays (from same individual) 30 inter- 
vals range from 7-1 I with a mean of 8.46 frames. 

In A. r. roquet only one sequence was filmed; there are 21 
pauses ranging from 7-9 frames with a mean interval of 7.75. 

In A. aeneus in ten displays 103 intervals were measured ranging 
from 9-19 frames with a mean of 1 1.9. 

If this character has any meaning in terms of relationships, it 
implies that aeneus is the most divergent member of the series and 
that trinitatis is closer to extremus. This supports the first alternative 
above. 

Table 1 (p. 29) summarizes some of the data for the roquet 
group. 



26 BREVIORA No. 284 

DISCUSSION 

At the level of the species group, the display behavior is useful 
as an additional taxonomic character. The roquet group can be 
defined as distinct from its geographic near neighbors by the absence 
of dewlap pumping within a display sequence. Furthermore, dif- 
ferences can be delineated which distinguish the eight forms of the 
species group. 

Why stereotyped displays in island races of lizards? Why inter- 
island differences? Do they relate to isolating mechanisms? Among 
some well-established cases of vocalization in birds, island races 
may have much more variable, less stereotyped, and less complex 
calls than the corresponding mainland races (Marler, 1959). The 
assumption here is that there are fewer sympatric congeners, and 
that there is therefore reduced selection for specificity. 

The few studies on island lizards, however, do not show a similar 
pattern. The display of Anolis townsendi, a species restricted to 
Cocos Island some hundreds of miles west of the Central American 
mainland, is quite stereotyped (Carpenter, 1965). Furthermore, 
Carpenter ( 1966) found, in a study similar to this one, that 1 1 of 
12 Galapagos islands have iguanids of the genus Tropidurus with 
species-specific (or island-specific) display patterns. Each island 
has only one Tropidurus. These differences were attributed to 
genetic drift. 

In the study by Ruibal ( 1967) on comparative anole behavior, 
island races were discussed, but the species studied were from 
Greater Antillean islands where each species might have a half 
dozen sympatric congeners at the same locality. Yet the displays 
described are no more stereotyped than the ones described here. 

It could be argued that some of the sympatric roquet group 
anoles evolved behavioral differences as additional isolating mech- 
anisms. For example, there was double colonization of St. Vincent 
from St. Lucia, which led to trinitatis and griseus. Perhaps some of 
the differences in the displays as seen today reflect selection for 
species-specific differences. 

Why are the displays no less stereotyped where there is only one 
form on an island? Probably the least variable display in the roquet 
group is that of extremus from Barbados, an island which has and 
probably has always had only one Anolis. 

I have attempted to emphasize that certain components in the 
display of each form characterize that form, yet there is some 
variation in the temporal sequence of displays. Since each island 
was colonized by a presumably small number of arrivals from 



1968 DISPLAY BEHAVIOR OF SOME ANOLES 27 

another island, population differences in display are not at all 
surprising. There is sampling error to begin with. Then, as the new 
population adjusts to the new island, there is reconstruction of 
the gene pool. Displays are almost certainly polygenic, virtually all 
genes have pleiotropic effects, and thus differences in challenge 
displays in anoles on isolated islands are the expected result. 

But selection, too, could be playing a very strong role. When we 
look at the display of A. chrysolepis and compare it with the 
roquet group, we find few similarities. A. chrysolepis hardly moves 
its body, but flashes its cherry-red dewlap quickly; in the roquet 
group the tail is waved, the head is bobbed, and there are vigorous 
push-ups. 

When we look at the ecology of these anoles, this behavior makes 
perfect sense. A. c/ir\solepis is a small brown forest anole living in 
deep shade. Body movements would not be conspicuous, the wind 
might be rustling leaves. But a sudden flash of red color appearing 
from "nowhere" is quite striking. Anoles in the roquet group are 
larger and display from more open sites. Thus head and body move- 
ments are conspicuous sign stimuli. 

At a subtler level, this line of reasoning could account for display 
differences within the roquet group. The islands are by no means 
identical. Bonaire, for example, is sparsely vegetated. St. Lucia has 
more characteristic lush tropical flora. Could differences in the 
configuration of the habitat act to select for display differences? In 
afl probability they could. Perhaps it is advantageous for luciae to 
repeat himseif by giving multiple head-bobbing series while a single 
long series suflRces in the more open terrain for A. bonairensis. 

In cases of sympatry in the Lesser Antilles, display differences 
may be very important in sexual selection, even though these dis- 
plays are "territorial," not "courtship," displays. In a study of two 
sympatric species of grackles, Selander and Ciller (1961) found 
that males hold mutually exclusive territories and solicit nesting 
females of either species. It is the females that must select con- 
specific mates based upon a display difference. Possibly, in AnoUs, 
females associate only with males giving the "proper" display. 
Hunsaker (1962) showed that female Sceloporus preferentially 
associated with mechanical models which gave the species-specific 
head-bob. Because of the importance of display in the social struc- 
ture of the population, even on a one-anole island we must expect 
limited variation, i.e. stereotypy. 

Despite differences in displays of all the forms considered, it 
should be emphasized that the basic display patterns are quite 



28 BREVIORA No. 284 

similar and tiiat this may be used as a valuable taxonomic charac- 
ter. Quite possibly, in the Greater Antilles where 20 or more species 
of Anolis may be found on a single island, there might be very 
strong selection for behavioral differences (including challenge 
display) as isolating mechanisms in closely related forms. This 
would render the patterns taxonomically useless, except to show 
that different species had different displays: it would mask, rather 
than elucidate patterns of relationship. 

In general, the behavioral data support my proposed phylogeny 
of the roquet group based upon chromosomes and proteins. The 
presumed prototype, A. luciae, has a display which could well be 
the basis for further elaborations and foreshortenings, leading to 
the displays of the other forms. It is interesting that luciae and 
griseus have very similar displays, which differ from ricluirdi. An 
entirely different approach, marshalling biochemical evidence, has 
led me to the conclusion that ric/hirdi had to be derived from griseus 
and not vice-versa (Gorman and Atkins, 1968). 

The relationship between luciae and bonairensis is less clear. If 
blanquillanus, the presumed intermediate, had been available for 
study, this perhaps could have been clarified. 

Relating trinitatis and luciae closely is not a novel idea. Under- 
wood (1962) considered them conspecific, although his reasons 
were not particularly clear. The two forms differ considerably in 
blood proteins; however, the similarity in karyotype and certain 
similarities in display behavior tend to support the idea of close 
relationship. 

To compare the different island forms I have presented the data 
in an oversimplified manner. That is, I implied that a display once 
established is static. Thus, I call luciae ancestral to various other 
forms and compare the displays of today's luciae to the presumed 
derived forms. Obviously, the displays of all forms living in isola- 
tion are subject to change. The minor use of the tail in luciae may 
well be a secondary loss rather than a primitive character that was 
later elaborated in all the other members of the species group. 

Despite the oversimplification, however, comparative behavioral 
data when combined with other information are valuable in tracing 
the evolution and relationships of the roquet group. 

ACKNOWLEDGMENTS 

I am indebted to Dr. Garth Underwood, formerly of the Uni- 
versity of the West Indies in Trinidad, for sparking my interest in 
West Indian Anolis, and for providing space and facilities in the 



1968 DISPLAY BEHAVIOR OF SOME ANOLES 29 

Zoology Department of the University of the West Indies. I have 
been aided by the kindnesses of many people living in the West 
Indies, in particular: Dr. J. L. Price, Dr. J. Kenny, Mr. H. Boos, 
and Mr. J. Boos of Trinidad; Dr. J. L. Groome of Grenada; Mr. 
and Mrs. R. Milligan of Barbados; Dr. E. Kirby and Mr. F. Prescod 
of St. Vincent; Mr. E. Long of St. Lucia; and Dr. A. Krumperman 
of Bonaire. Field assistance has been provided by Miss C. Raybin. 
Miss A. Swidler, Mr. A. R. Kiester, and Mr. R. Ross. Illustrations 
used in this paper were prepared by Mrs. P. Kerfoot. 

In 1963-1964 I held a travelling fellowship from the University 
of California. Further field work has been supported by NSF Grant 
GB-2444 to Dr. E. E. Williams, and grants from the Explorers 
Club, Sigma Xi, and the Evolutionary Biology Committee of 
Harvard University. In addition to my travelling fellowship, my 
graduate studies have been supported by an NSF Predoctoral 
Fellowship (1964-1966) and a Turtox Fellowship (1966-67). 

Dr. E. O. Wilson provided valuable criticism of the manuscript. 
The frequent consultations with Dr. E. E. Williams, and his con- 
stant interest in this study, have been instrumental in its fruition. 

TABLE 1 
Head bob patterns of roquet group anoles 

Time Interval in 16thsof Sec. 



p 


eaks, 


/Bobbing 


Sequence 






No. 


filmed 






Form 


sequences 


Range 


Mean 


liiciae 1 




83 


1-18 


5.2 


honairensis - 




14 


12-23 


17.7 


honairensis'-^ 




6 


13-24 


17.7 


trinitatis 




30 


2-10 


5.7 


griseits i 




18 


1-8 


3.3 


ricluirdi 




8 


4-7 


5-8 


extrcnuis 




32 


2-5 


4.0 


aeiieiis 




25 


2-4 


2.8 


roquet 




28 


2-9 


4.3 



Range 


Mean 


2-47 


18 


43-95 


62.2 


47-91 


69.0 


12-31 


23.0 


1-41 


16.6 


39-58 


47.3 


6-14 


10.6 


27-62 


44.6 


18-47 


33.3 



1 In liiciae and gri\c'tis. a head bobbing sequence consists of several 
series of bobs, with pauses between the series. These data are for the 
individual series. 

- In bomiirensis there are occasionally paused displays, as in hiciae and 
griseiis. Here are tabulated non-paused displays. 

•■■ This tabulates individual series of the paused displays. 



30 BREVIORA No. 284 

LITERATURE CITED 

Carpenter, C. C. 

1962. Patterns of behavior in two Oklahoma lizards. Amer. Midi. 
Natur. 67: 132-151. 

1965. The display of the Cocos Island anole. Herpetologica 21: 
256-260. 

1966. Comparative behavior of the Galapagos lava lizards (Tropi- 
diinis). In: The Galapagos. Proceedings of the Galapagos 
International Scientific Project, R. I. Bowman, editor (pp. 269- 
273). Berkeley and Los Angeles, Univ. of California Press. 

Darwin, Charles 

1872. The Expression of Emotions in Man and Animals. London, 
Murray, 374 pp. 

Etheridge, R. 

1960. The relationships of the anoles (Reptilia:Sauria:Iguanidae). An 
interpretation based on skeletal morphology. Ph.D. thesis. 
University of Michigan. (University Microfilms. Inc., Ann 
Arbor.) 

Evans, L. T. 

1938. Cuban field studies of territorial behavior of Anulis sagiei 
J. Comp. Psychology 25: 97-125. 

Gorman, G. C. 

1965. Interspecific karyotypic variation as a systematic character in 
the genus AnoUs (SauriaTguanidae ). Nature (London) 208: 
95-97. 

Gorman, G. C. and L. Atkins 

1967. The relationships of Anulis of the loqiict species group (Sauria: 
Iguanidae). IL Comparative chromosome cytology. Systematic 
Zoology 16: 137-143. 

1968. The zoogeography of Lesser Antillean Anolis (Sauria:lguani- 
dae). An interpretation based on chromosomes and lactic 
dehydrogenases. In press. 

Gorman, G. C, L. Atkins, and T. Holzinger 

1967. New karyotypic data on 15 genera of lizards in the family 
Iguanidae, with a discussion of taxonomic and cytological im- 
plications. Cytogenetics 6: 286-299. 

Gorman, G. C. and H. C. Dessauer 

1965. Hemoglobin and transferrin electrophoresis and relationships of 
island populations of Anolis lizards. Science 150: 1454-1455. 



1968 DISPLAY BEHAVIOR OF SOME ANOLES 31 

1966. The relationships of AnoUs of the roquet species group (Sauria: 
Iguanidae). I. Electrophoretic comparison of blood proteins. 
Comp. Biochem. and Physiol. 19: 845-853. 

Grhenbhrg. B. and G. K. Noble 

1944. Social behavior of the American chameleon {Aiioli.s caroliiicn- 
sis). Physiol. Zool. 17: 392-439. 

HUNSAKER, D. 

1962. Ethological isolating mechanisms in the Sceloporus toiquatiis 
group of lizards. Evolution 16: 62-74. 

Kastle, W. 

1963. Zur Ethologie des Grasanolis (Noyops aiiratiis). Z. Tierpsych. 
20(1): 16-33. 

Marler, p. 

1959. Developments in the study of animal communication. //;. 
Darwin's Biological Work, P. R. Bell, editor, (pp. 150-206). 
Cambridge, Cambridge University Press. 

RUIBAL, RODOLFO 

1967. Evolution and behavior in West Indian anoles. //;.• Lizard 
Ecology, A Symposium, W. W. Milstead. editor, pp. 1 16-140. 
Univ. of Missouri Press, Columbia. 

RUTHVHN, A. G. 

1923. The reptiles of the Dutch Leeward Islands. Occ. Pap. Mus. 
Zool. Univ. Michigan No. 143: 1-10. 

Selander, R. K. and D. R. Giller 

1961. Analysis of sympatry of the great-tailed and boat-tailed grackles. 
Condor 63: 29-86. 

Underwood, G. 

1959. Anoles of the eastern Caribbean. Part 111. Revisionary notes. 
Bull. Mus. Comp. Zool., 121: 191-226. 

1962. Reptiles of the eastern Caribbean. Caribbean Affairs, N.S. 
1: 1-192. 

(Received 19 July 1967.) 





E V I 






Mmsemiii of Comparative Zoology 

Cambridge, Mass. 5 April, 1968 Number 285 



THE GENERA OF PUFFBIRDS (BUCCONIDAE) 
G. W. Cottrell 

In 1882, in the last part of A monograph of the jacamars and 
puff-birds, pages xxxiv-xli, P. L. Sclater divided the family of 
puffbirds (Bucconidae) into seven genera: Bitcco, Malacoptila, 
Micromonacha, Nonnula, Hapaloptila, Monacha (Sclater's emenda- 
tion of Monasa Vieillot 1816), and Chelidoptera. The large 
genus Bucco, containing twenty of the forty-three species enum- 
erated for the family, he grouped in two subgenera (following 
VVagler 1827, Sy sterna avhmi): Bucco, "rostro rotundato ad basin 
dilatato" (fifteen species), and Nystalus, "rostro compresso et 
magis elevato" (five species). He saw no grounds for division into 
subfamilies. 

By 1914 the trend of the times had swelled the number of genera 
to thirteen, as set forth by Ridgway in The buds of North and 
Middle America 6: 371-373, the increment all taking place within 
the limits of Sclater's genus Bucco. The roster in Ridgway runs 
Notharchus,^ Argicus, Nystactes, Bucco, Hypnelus, Ecchaunornis, 



1 Misprinted "NortharcJius" in the key to the genera, "Notharchiis" else- 
where. This name, with two of its trivial adjuncts, has had an ill-starred 
orthographic career. In 1958 Eisenmann, Auk 75: 101, called attention 
to the misprinting in Peters 1948, Check-list 6: 10, of the subspecific 
hyperrhynchus (Sclater 1856) as "hypeirynchus." And Ridgway's "Nortlwr- 
chiis" has just been noted. But there is worse to come. In recent decades 
(first observed in Griscom 1932, The distribution of bird-life in Guatemala), 
Notharchus has frequently been printed "Notliarcus," contrary to the original 
publication, with indicated derivation, of Cabanis and Heine 1863, Museum 
Heineaniim 4, 1: 149 {Northarchus formed from vwdrjs, sluggish, slothful, 
and dpxos, leader, chief — though the latter incorrectly accented apxos 
by Cabanis and Heine). Of less consequence than the generic misspelling 
is the shift (dating back at least to Wagler 1827, Sy sterna avium) of the 
ending of the specific name macrorhynchos as first published by Gmelin 
1788, Systema naturae 1, 1: 406, from -os to -us, latterly perhaps by 
attraction to Sclater's hyperrhynchus, originally a specific designation but a 
subspecies of macrorhynchos since 1939 (Sassi, Temminckia 4: 161). 



2 BREVIORA No. 285 

Nystalus, Malacoptila, Micronwnacha, Hapaloptila, Monasa, Non- 
nula, Chelidoptera. The first seven genera (comprising Sclater's 
Biicco) are grouped as the subfamily Bucconinae, the remaining 
six as the subfamily Malacoptilinae. Cory 1919, Catalogue of birds 
of the Americas 2, 2: 390-410, has the same thirteen genera as 
Ridgway, but without division into subfamilies. 

Thirty years later the emphasis in classification had shifted from 
dissimilarity toward similarity: Peters 1948, Check-list 6: 10-24, 
reduces the seven genera equated by Ridgway with Sclater's Bucco 
to four, Argicus and Nystactes being merged with Bucco, and 
Ecchaunornis with Nystalus. The other six genera remain un- 
changed. No subfamilies or subgenera are recognized. Peters' 
arrangement has been regularly followed down to the present (e.g., 
de Schauensee 1966, The species of birds of South America). 

A new review of the family has led to the conclusion that a full 
return to Sclater's generic system should be made by merging 
Notharchus, Nystalus, and Hypnelus, maintained by Peters, with 
Bucco.'- Since the Check-list does not justify its systematics, one 
must revert to Ridgway for criteria of differentiation. Ridgway 
follows Sclater in dividing his subfamily Bucconinae (Sclater's 
genus Bucco) into two major groups: one with thick bill, "its 
width at nostrils nearly to quite equal to its depth at same point" 
(Sclater's subgenus Bucco), the other with compressed bill, "its 
width at nostrils much less than its depth at same point" (Sclater's 
subgenus Nystalus). In the thick-billed group he included his 
genera Notharchus, Argicus, Nystactes, and Bucco, in the com- 
pressed-billed group the genera Hypnelus, Ecchaunornis, and 
Nystalus, paralleling Sclater's division except with respect to the 
species included in Hypnelus, which were placed by Sclater in the 
thick-billed group. An actual check of specimens shows the 
amount of compression to be only very slightly greater in Sclater's 
subgenus Nystalus (most in maculatus) . 

Lesser criteria of differentiation in Ridgway relate to variations 
in bill shape and color, tarsus length, length of rictal bristles, 
length of remiges or rectrices, plumage markings, as set out in his 



2 This same merging of genera was proposed and followed by Charles 
G. Sibley in a seminar syllabus on the Piciformes, Cornell University, ca. 
1956, but was not published. Its actual priority is hereby duly recorded. 



1968 GENERA OF PUFFBIRDS 3 

key."^ Thus, Notharchus is distinguished from Argicus, Nystactes, 
and Biicco by tarsus shorter than outer anterior toe without claw, 
upper parts mostly plain blackish, forehead sometimes white, as 
opposed to tarsus equal to or longer than outer anterior toe with- 
out claw, upper parts mostly brown or rufous-tawny, forehead 
never white. We have seen that Peters included Argicus and 
Nystactes in Bucco. Within his compressed-billed group, Ridgway 
distinguishes Hypneliis from Ecchaunornis and Nystalus by a 
strongly bifid maxilla (see below), longer rictal bristles, longest 
primaries decidedly longer than secondaries, indistinct spotting of 
upper parts, black pectoral band, plain (unbarred) tail, wholly 
black (not red. orange, or horn) bill. Peters merged Ecchaunornis 
with Nystalus. 

Such morphological criteria, as employed by Ridgway for his 
seven genera, and tacitly maintained by Peters for the four genera 
recognized by him, do not seem adequate for separation at the 
generic level. With respect to ethology, there are no recorded 
aspects that militate against the proposed merging — though 
information is in general scanty and fragmentary (also repetitious), 
and in some species almost totally lacking. 

The genus Bucco, now re-extended to Sclater's limits (and 
equivalent to Ridgway's subfamily Bucconinae), is marked by a 
relatively stout and straight bill, much broadened at the base, with 
strongly uncinate tip to the maxilla and terminally ascending more 
or less convex gonys. A striking feature, found to greater or less 
degree in all but one of the species, is the bifid tip to the maxilla. 
An examination of specimens shows that this peculiar process, part 
of the rhamphotheca and not of the underlying bone, is particularly 
prominent in tamatia and ruficollis (including bicinctus, Phelps and 
Phelps 1958, Proc. Biol. Soc. Wash. 71: 122), pronounced in 
macrorhynchos, pectoralis, ordii, tectus, noanamae, striolatus. 



^ Ridgway 's key cannot be relied on even within its own frame of refer- 
ence. There is, for example, no subdivision "bb" to match "b" under "aa," 
which renders subsequent allocations indeterminate. The genus Hapaloptila 
appears under the subdivision "aa," characterized as having the tip of the 
maxilla not uncinate, whereas in fact the tip in Hapaloptila is strongly 
hooked. According to the key the bill in Monasa, Nonniila, and Chelidoptera 
is red, orange, or yellow, true in reality of Monasa only. Again, Chelidoptera 
is keyed as not "tawny" beneath, though actually with cinnamon-chestnut 
belly. 



4 BREVIORA No. 285 

maculatus, fairly slight in capensis, slight in radiatus, very slight in 
chacuru, and not distinguishable at all in macrodactylus. In a 
study of this bifid process, Boetticher 1936, Festschrift ziim 60. 
Geburtstage von Professor Dr. Embrik Strand 1 : 74-11, suggested 
that it might serve as a sort of vise for holding the heads of cap- 
tured flying insects (especially butterflies) while the wings were 
sheared off by the mutually adapted tomia. He admitted that there 
were no specific observational data to support his hypothesis; nor 
have any been reported since. 

The treatment of this character in Ridgway is inconsistent and 
in part inaccurate (notably with regard to extralimital taxa). In 
the key there is the general statement, applicable to all seven 
genera, that the tip of the maxilla is sometimes cleft medially, 
but no discrimination between individual genera (and hence 
species) until the compressed-billed group, comprising Hypnelus, 
Ecchaunornis, and Nystcdus, is reached. Here Hypnelus (consist- 
ing of the currently recognized ruficollis) is correctly signalized as 
having the tip of the maxilla strongly bifid, and Ecdmunornis 
(consisting of radiatus and chacuru) equally correctly as having 
the tip not distinctly if at all bifid. But Nystahis (consisting of 
striolatus and maculatus) is united in this respect with Ecchau- 
nornis, whereas in actuality the medial cleft is pronounced in its 
component species. The subsequent generic description for Nothar- 
chus (consisting of macrorhynchos, pectoralis, ordii, and tectus) 
includes a more or less distinctly bifid tip to the maxifla, but there 
is no later indication as to the situation in Argicus {macrodacty- 
lus), Nystactes (tamatia), or Bucco (noanamae, capensis), de- 
scriptions being omitted for extralimital genera. 

A review of the remaining six bucconine genera as found in 
Sclater, Ridgway, and Peters reconfirms their distinctness. They all 
have a less stout, more compressed, less basally broadened bill 
than the reconstituted Bucco, with the gonys more or less de- 
curved terminally and, except in Hapaloptila, the maxifla more 
gradually decurved and non-uncinate. In none of the six does the 
bifid maxflla occur. Differences in size, in shape and proportions 
of bill, wings, and tail, and in plumage patterns serve adequately 
to separate one from another, as described and fllustrated by 
Sclater, and hence are not detaUed here. Certain ethological fea- 
tures reinforce the morphological distinctions, as may be seen in 
the social behavior of Monasa and the mode of flight and flocking of 
Chelidoptera, this latter being in many respects the most divergent 
of the taxa included within the family. 



1968 GENERA OF PUFFBIRDS 5 

Consequent upon the merging proposed above, the genera and 
species of the Bucconidae stand as follows (order from Peters, 
species as in Peters except as indicated, subspecies not shown) : 

Biicco macrorhynchos, pectoralis, ordii, tectus, macrodactylus, 
tamatia, nocmamae, capensis, radiatus, chacuru, striolatus, 
maculatiis, ruficollis (includes bicinctus of Peters, following 
Phelps and Phelps 1958, Proc. Biol Soc. Wash. 71: 122) 

Malacoptila striata, jusca, semicincta (subspecies of fusca in 
Peters; distinct species here, following Traylor 1951, Fieldi- 
ana, Zool. 31: 614), julvogularis, ruja, panamensis, mystacalis 

Micromonacha lanceolata 

Nonnula rubecula, sclateri, brunnea, mficapilla (includes jron- 
talis of Peters, following de Schauensee 1946, Not. Nat. 163: 
5), amaurocephala 

Hapaloptila castanea 

Monasa atra, nigrifrons, morphoeus, flavirostris 

Chelidoptera tenebrosa. 

(Received 26 July 1967.) 



BREVIORA 



MiiseiMii of Comparsitive Zoology 



Cambridge, Mass. 5 April, 1968 Number 286 



THE STATUS OF NESOGALAXIAS NEOCALEDONiCUS 
(WEBER AND DE BEAUFORT) 
(PISCES, GALAXIIDAE) 

R. M. McDowall, 

Museum of Comparative Zoology, Harvard University^ 
Fisheries Research Division, New Zealand Marine Department- 

ABSTRACT 

The validity of the genus Nesogalaxias, proposed by Whitley 
(1935) for Galaxias neocaledonicus Weber and de Beaufort 
(1913), is supported by its very reduced pleural ribs, a character 
which has not been observed in other galaxiids. N. neocaledonicus 
occurs in mountain lakes in southern New Caledonia and is the 
most tropical species in the family. It is thought to have dispersed 
from Australia to New Caledonia when temperatures were lower 
during the Pliocene and Pleistocene, and the post-Pleistocene 
warming seems to have confined it to a few mountain lakes. Hither- 
to recorded only from Lac en Huit, N. neocaledonicus is now also 
known to occur in Grand Lac. 



INTRODUCTION 

In 1913, Weber and de Beaufort described a small fish species 
in the genus Galaxias from a lake in the mountains of southern 
New Caledonia. Several workers, e.g. Whitley (1935) and Scott 
(1936, 1966), have listed this species or commented on its generic 
position, but none appears to have re-examined it thoroughly. 
Since its generic status is currently uncertain, a re-examination of 
the species is necessary. 



1 Present address. 

- On leave. Fisheries Research PubUcation No. 105. 



2 BREVIORA No. 286 

In 1964, Dr. P. H. J. Castle collected a small sample of this 
species from Lac en Huit, the type locaHty, and more recently 
(1966), Mr. W. Skrzynski collected further samples from this 
lake, and also from Grand Lac, apparently for the first time. 
These samples have been generously made available to me, en- 
abling an assessment of the affinities of this species to be made. 

GENERIC STATUS 

Although Whitley (1935) proposed a new genus for G. neo- 
caledonicus, he supplied no generic diagnosis, merely stating that 
"it is not a true galaxiid and may receive the new generic name 
Nesogalaxias." Scott (1966) found that "no feature in the origi- 
nal description would seem to preclude attribution to Galaxias" 
but apparently not having specimens, he took no further action. 

Nesogalaxias is correctly included in the family Galaxiidae, its 
lack of scales, fin positions, lateral line pores on the head, dentition, 
lack of a mesocoracoid, and jaw structure all conforming to the 
general galaxiid pattern. N. neocaledonicus was found to come 
within the range of variation of members of the family in New 
Zealand, in most of the characters examined. Superficially it 
difters little from some of the more generalized species of Galaxias. 
Stokell (1945) revised the generic classification of the Galaxiidae. 
He suggested that the very long head of the specimen figured by 
Whitley (1935) indicated deformity and classed the genus as "not 
recognised." 

However, Stokell's (1945) conclusion that the very long head 
in Whitley's (1935) figure is a deformity is incorrect, as the 
species does have a very long head (see Table 1). The pelvic 
fins have more posterior insertion than in most galaxiids, but this 
is a result of the great head length contained in the pre-pelvic 
dimension. In ratios of both standard length/head length and 
standard length/pre-pelvic length N. neocaledonicus represents an 
extreme for the family, but the range of values overlaps the ranges 
for these ratios in other species, e.g. Galaxias argenteus (Gmelin). 
Vertebral number (41-43) is very low, but not as low as in G. 
zebratus (Castelnau), which Jubb (1965) reported as having 
39-41 vertebrae. 

In most osteological characters, N. neocaledonicus is conven- 
tional for the family Galaxiidae. The caudal skeleton shows the 



1968 



STATUS OF NESOGALAXIAS 



irregular variability which is characteristic for the family, and is 
not distinctive. The pectoral girdle lacks the postcleithrum, but 
this bone has disappeared in several other unrelated species, e.g. 
G. divergens Stokell, G. paucispondylm Stokell. However, in the 
absence of pleural ribs from those vertebrae behind the pelvic fins, 
N. neocaledonicus is different from all other galaxiids examined. 
These bones are present on vertebrae between the head and about 
the level of the pelvic fins, but thereafter disappear quite suddenly 
from the remainder of the abdominal vertebrae. The ribs present 
anteriorly, although short, are well developed, appear to be strongly 
ossified and have well developed heads articulating with the para- 
pophyses of the vertebrae. There are no epipleural ribs. The 
habits of A', neocaledonicus are somewhat different from other 
galaxiids (see p. 5) and this species seems to have entered a 
somewhat unusual type of niche for the family. These characters 
suggest that N. neocaledonicus stands apart from the rest of the 
family Galaxiidae and is best regarded as generically distinct from 
the more generalized species. The name Nesogalaxias Whitley 
(1935) has priority and is available. 

Nesogalaxias Whitley, 1935 

Type species: Galaxias neocaledonicus Weber and de Beaufort, 
1913, by original designation. 

Diagnosis: Trunk cylindrical to a little compressed, naked, 
dorsal fin posterior, above vent. Pelvic fins present, usually seven 
rayed. Caudal fin forked with 16 principal rays. Teeth uniserial 
on jaws and entopterygoids, biserial on basihyal. Postcleithrum 
absent. Pleural ribs lacking from abdominal vertebrae behind 
the pelvic fins. 




Figure 1. Nesogalaxias neocaledonicus (Weber and de Beaufort, 1913). 
Lac en Huit, New Caledonia. Length to caudal fork 52 mm. 



4 BREVIORA No. 286 

Nesogalaxias neocaledonicus (Weber and de Beaufort, 1913) 

Galaxias neocaledoniciis 'Weher and de Beaufort, 1913: 173. 

Nesogalaxias neocaledonicus, Whitley, 1935: 42; Scott, 1936: 85, 1966: 250. 

Syn types: Zoology Museum, Amsterdam, Holland, no. ZMA 
104, 700, 2 specimens (seen). Museum National d'Histoire Natu- 
relle, Paris, France, nos. 2312-3 (Bertin and Esteve, 1951, not 
seen). 

Type locality: Recorded as "Lac en 8, sudliches Neu-Kale- 
donien" by Weber and de Beaufort. The locality for the paratypes 
in the Museum National d'Histoire Naturelle is listed as "Lac en 
Huit" by Bertin and Esteve (1951). 

Description (Fig. 1, Table 1): A small, rather slender species, 
trunk deep anteriorly but tapering to a very slender caudal peduncle 
which is much longer than deep. Greatest body depth somewhat 
behind pectoral fin bases. Head very long, slender, flattened 
ventrally, tapering rather abruptly forward dorsally to a slender, 
long snout. Eye large, projecting above head profile, interorbital 
concave, very narrow. Jaws sub-equal, long, extending to about 
anterior third of eye, gape narrow. Profile of lower jaw from 
ventral aspect deep and U-shaped. Canine teeth weakly developed 
in jaws; three to four moderately developed entopterygoidal teeth. 
Gill rakers strongly developed; two weak pyloric caeca. 

Dorsal and anal fins membranous and short based, rather high. 
Anal origin a little behind a line through dorsal origin. Pectoral 
fin inserted rather low, long in interval between pectoral and 
pelvic fin bases. Pelvic fin long in interval between pelvic base 
and anal origin, fin tapering to a point medially with middle rays 
longest. Caudal fin forked, rather short, fin tips rounded; flanges 
of caudal peduncle rather poorly developed. 

Size: Weber and de Beaufort listed a maximum size of 76 mm 
in their sample. Amongst fishes examined by the writer, the 
largest was only 56 mm long, and most adults and sub-adults 
were 40-50 mm long. 

Coloration: Trunk a uniform grayish brown, darkening dorsally 
and on head; ventrally paler, creamish gray. Pigmentation extends 
a little on to bases of fins, especially the caudal. 

Meristic variation: Dorsal fin rays (segmented) 8 (6), 9 (12) 
10 (2); anal rays 9 (2), 10 (14), 11 (4); caudal rays (principal) 
15 (1), 16 (19); pectoral rays 11 (5), 12 (14), 13 (1); pelvic 
rays 6 (1), 7 (19); branchiostegals 6 (7), 7 (13); vertebrae 41 



1968 STATUS OF NESOGALAXIAS 5 

(3), 42 (16), 43 (3); gUl rakers 3-11 (1), 3-12 (0), 3-13 (3), 
4-11 (1), 4-12 (6), 4-13 (4), 5-12 (1), 5-13 (3), 6-13 (1). 

DISTRIBUTION AND HABITAT 

N. neocaledonicus is known only from two small lakes in the 
Plaine des Lacs, New Caledonia (Fig. 2) — Lac en Huit and 
Grand Lac. Skrzynski (personal communication) has kindly sup- 
plied the following notes on the lakes and the habitats in which he 
found this species. 

The Plaine des Lacs is about 100 km from Noumea. There are 
two main lakes, Lac en Huit and Grand Lac, each at an altitude 
of about 250 m, and a few square kilometers in area. They are 
about 1 Vi km apart and separated by a ridge, their outflows merg- 
ing a few kilometers below the lakes. 

Both lakes are depressions in an iron pan, their bottoms being 
solid rock with loose rock and soil in some places. The only aquatic 
plant growth observed consisted of loose patches of reeds in a few 
areas and encrusting algae. Neither of these provide much cover 
for fishes and the only cover in which N. neocaledonicus was ob- 
served was loose rock on the lake bed and crevices in the bed 
itself. The fish were found to conceal themselves in this cover 
during the day, but at night were commonly observed swimming in 
mid-water. Small unpigmented juveniles about a centimeter long 
were found to shoal freely in mid-water during the day. 

RELATIONSHIPS AND ORIGIN 

N. neocaledonicus is the most tropical species generally recog- 
nized as a galaxiid. G. indicus Day, from India, has not been re- 
collected since it was first described in 1888, and is doubtfully a 
galaxiid. Apart from A', neocaledonicus and the doubtful G. indi- 
cus, fishes of this family are confined to the southern temperate 
zone and, like the Northern Hemisphere salmonoids, are primarily 
cold water fishes. The fact that N. neocaledonicus has found refuge 
in the mountain lakes of New Caledonia suggests that it too pre- 
fers or is limited to cold waters. If this is so, it seems probable 
that it reached New Caledonia at a time of cooling, most likely in 
the Pliocene-Pleistocene, when lowered temperatures would have 
facilitated dispersal to New Caledonia. With subsequent post- 
glacial warming occurring, the populations have become confined 
to the cooler mountain lakes. 



BREVIORA 



No. 286 



E 
in 



> 




a 

'2 
o 

u 
U 
<u 



o 

O 

u 
3 
60 



1968 STATUS OF NESOGALAXIAS 7 

The relative closeness of New Caledonia to Australia points to 
likely origin of its galaxiid populations in Australia. To what 
species or species group in Australia A^. neocaledonicus is most 
closely related is quite unknown, but it seems likely that its 
ancestor was diadromous, since oceanic dispersal would have been 
necessary for the fishes to reach New Caledonia. Analysis of the 
Australian galaxiid fauna may indicate the relationships of 
N. neocaledonicus. 

ACKNOWLEDGEMENTS 

I am grateful to Dr. Giles W. Mead and Mr. G. D. Waugh for 
reading the manuscript, to Dr. P. H. J. Castle for the loan of speci- 
mens, and to Mr. W. Skrzynski for specimens and his notes on 
localities and habitats. Thanks are also due members of the staff 
of the Institut Frangais d'Oceanie Noumea, who assisted both 
Dr. Castle and Mr. Skrzynski in making their collections. The 
financial support of a New Zealand National Research fellowship 
is gratefully acknowledged. 

LITERATURE CITED 

Bertin, L. and R. Esteve 

1951. Catalogue des types de poissons du Museum National d'Histoire 
Naturelle, 6 Partie. Haplomi, Heteromes, Catostomes. Mu- 
seum National d'Histoire Naturelle, Paris, 60 pp. 

JUBB, R. A. 

1965. Freshwater fishes of the Cape Province. Ann. Cape Province 
Museum 4: 1-72, 41 figs. 

Scott, E. O. G. 

1936. Observations on fishes of the family Galaxiidae. Pap. Roy. 
Soc. Tasmania, 1935: 85-112, 4 figs. 

1966. The genera of Galaxiidae. Austral. Zool. 13(3) : 244-258. 
Stokell, G. 

1945. The systematic arrangement of the Nev/ Zealand Galaxiidae. 
I. Generic and sub-generic classification. Trans. Roy. Soc. New 
Zealand 75(2): 124-137, 10 figs. 
Weber, M., and L. F. de Beaufort 

1913. iiber neue Fische von Neu-Kaledonien. Zool. Anz. 52: 172- 
174. 
Whitley, G. P. 

1935. Whitebait. Vict. Nat. Melbourne 52(3): 41-51, 1 pi., 2 figs. 

(Received 28 August 1967.) 



8 BREVIORA No. 286 

TABLE 1 

Morphometric variation in 20 Nesogalaxias neocaledonicus. 

Min. Mean Max. 

Length caudal fork/standard length 

Standard length/body depth at vent 

Standard length/length caudal peduncle 

Length caudal peduncle/depth caudal peduncle 

Standard length/pre-dorsal length 

Pre-anal length/pre-dorsal length 

Standard length/length dorsal fin base 

Greatest length of dorsal fin/basal length 

Standard length/length anal fin base 

Greatest length of anal fin/basal length 

Pre-pelvic length/standard length 

Pectoral-pelvic length/standard length 

Pectoral fin length/pectoral-pelvic length 

Pelvic-anal length/standard length 

Pelvic fin length/pelvic-anal length 

Standard length/head length 

Head length/head depth 

Head length/head width 

Head length/snout length 

Head length/post-orbital head length 

Head length/interorbital width 

Head length/diameter of eye 

Head length/length of upper jaw 

Head length/length mandible 

Head length/width of gape 



1.07 


1.12 


1.16 


7.86 


8.36 


9.06 


6.46 


7.11 


8.25 


1.50 


1.96 


2.25 


1.34 


1.38 


1.40 


1.03 


1.05 


1.07 


8.95 


10.36 


12.33 


1.43 


1.63 


1.93 


8.00 


9.45 


10.26 


1.36 


1.59 


1.82 


0.53 


0.56 


0.59 


0.24 


0.27 


0.28 


0.57 


0.62 


0.70 


0.20 


0.22 


0.23 


0.55 


0.66 


0.72 


3.04 


3.23 


3.39 


1.96 


2.19 


2.45 


1.96 


2.14 


2.33 


2.89 


3.25 


3.53 


1.85 


2.03 


2.15 


3.70 


4.36 


5.09 


4.45 


4.91 


5.45 


2.42 


2.65 


2.83 


2.57 


2.72 


2.88 


2.89 


3.66 


4.08 



BREVIORA 

MiLiseiulm of Comparative Zoology 



Cambridge, Mass. 5 April, 1968 Number 287 



THE SEQUENCE OF GENERA IN THE 
ESTRILDIDAE (AVES) 

Ernst Mayr 



The manuscript for the list of the Estrildidae in volume XIV of 
Peters' Check-list of Birds of the World is being prepared by three 
authors, Melvin Traylor (Africa), R. Paynter (Oriental and 
Palearctic Regions), and myself (Papuan Region and Australia). 
The task of coordinating the three accounts was assigned to me. 
Since each species, with few exceptions, is restricted to a single 
geographical region, the main problem of coordination is the recog- 
nition of genera and the choice of a sequence. To have a widely 
acceptable sequence of genera is particularly important in view of 
the fact that members of this family are used increasingly for be- 
havior studies and in physiological research. 

Research on the classification of the Estrildidae has been very 
active in the last two decades, particularly since the publication in 
1943 of Delacour's classical study. This was the first consistent 
attempt to arrange the genera in groups, and to utilize various bio- 
logical characters (courtship, palate markings of the nestlings, etc.). 
The subsequent researches of Steiner, Wolters, Immelmann, Nico- 
lai, and others, have all been endeavors to improve on the founda- 
tion laid by Delacour. 

The ranking of the taxon Estrildidae and its delimitation from 
the Viduinae and the Ploceidae have been discussed by several 
authors. Until recently the estrildids were considered a subfamily 
of the Ploceidae. Delacour (1943) derived them from the Sporo- 
pipinae, and neither Chapin (1917) nor Sushkin (1924, 1927) 
questioned the close relationship of the weavers and estrildids. The 
realization has, however, grown in recent years that not all 
"finches" are necessarily a monophyletic assemblage, and that a 
cone-shaped bill may have been acquired independently in several 
groups of seed-eating birds. There is no a priori necessity for a 
close relationship between ploceids and estrildids. Steiner (1955) 
undertook a careful comparison of the two groups and concluded 



2 BREVIORA No. 287 

that they differ in so many important characters that he questioned 
a close relationship. Some of these differences are valid only in a 
comparison of the Estrildidae and the Ploceinae (sensii stricto). 
The opinion has therefore been expressed, that the stated differ- 
ences are only subfamily differences. It is still possible that the 
Ploceidae are the nearest relatives of the Estrildidae; nevertheless, 
the differences are numerous and striking. Estrildids and ploceids 
are clearly different in courtship posture, clutch size, egg colora- 
tion, pattern in mouth of the young, and mode of begging of the 
young. In an analysis of the structure of the digestive tract, Zis- 
wiler (1967) comes to the conclusion that the Estrildidae are far 
more different from the Ploceidae than the ploceid subfamilies are 
from each other. The Viduinae, interestingly, agree in this charac- 
ter quite well with the other Ploceidae rather than with the Estril- 
didae. Africa is the home of the most primitive genera of estrildids 
and, presumably, the home of the family. It must be an old family, 
considering the subsequent rich radiation in the Australian Region. 
It is advisable for the time being to consider the estrildids a sepa- 
rate family, perhaps distantly related to the Ploceidae, but suffi- 
ciently distinct to deserve family rank. 

The Viduinae are nest parasites of the Estrildidae and have 
generally been considered to be very closely related to them. The 
similarity in the palate pattern of the nestlings was undoubtedly the 
major reason for this opinion. Delacour (1943: 73) questioned 
this assumption: "As there exists no transition between [the Estril- 
dinae and the Viduinae], it appears that their similarities are due to 
recent adaptation and I have come to the conclusion that the 
Estrildinae have probably evolved independently from the Sporo- 
pipinae, just as have the Viduinae and the Ploceinae." The fact 
that the nestlings of each species of Vidua have the palate markings 
of one particular species of waxbills clearly indicates convergence 
rather than common descent. In the meantime, Nicolai (1964) has 
demonstrated convincingly that the similarities between viduines 
and estrildids are due to nest parasitism. He has, in addition, pro- 
duced evidence to indicate a derivation of the viduines from 
Euplectes. To indicate this relationship, the Viduinae should be 
treated as a subfamily of the Polceidae rather than of the Estril- 
didae. Sibley (MS) has discovered similarities in the starch gel 
patterns of viduines and Passerinae. 

The valid name of the (sub-) family of the estrildids is still 
somewhat in doubt. The first available name for this taxon is 
apparently Spermestinae Cabanis (1847, Arch. Naturg., 13: 331, 
also 1851, Mus. Heineanum, 1, Singvogel: 172), based on the 



1968 GENERA OF ESTRILDIDAE 3 

generic name Spermestes Swainson 1837. This name has since 
been used occasionally, as in Gray, 1870 (Handlist Birds, 2: 49), 
Reichenow, 1904 (Vogel Afrikas, 3: 103), Steiner, 1960 (J. Or- 
nith., 101: 92-112), and in some of the avicultural literature. 

In the scientific ornithological literature the name Spermestinae 
has been almost completely displaced by the name Estrildidae. 
This name was apparently first proposed by Bonaparte, 1851 
(Consp. Av., 1: 450) (erroneously spelled Estreldinae), based on 
the genus Estrilda Swainson 1827. Bonaparte's name, in its cor- 
rected spelling Estrildinae, is used in virtually all those modern 
standard works in ornithology, in which the (sub-) family is recog- 
nized as different from the Ploceidae. It is the name used by Stuart 
Baker (1926, Fauna Brit. India, Birds, 3: 77), Sclater (1930, 
Syst. Av. Aethiop., p. 769), Berlioz (1950, in Grasse's Traite 
Zool., 15: 1039), Chapin (1954, Birds Belgian Congo, 4: 451), 
and Ripley (1961, Synopsis Birds India: 602). 

The name Estrildinae is likewise the name adopted by most mod- 
ern revisers of the family, for instance by Delacour (1943, Zoolog- 
ica, 28: 69), Wolters (1957, Bonn. Zool. Beitr., 8: 90-129), and 
Keast (1958, Emu, 58: 219-246). It is the name adopted in gen- 
eral classifications, such as Mayr and Amadon (1951, Amer. Mus. 
Novit., 1496: 38). 

The name Spermestinae has presumably been rejected almost 
universally because the name Spermestes Swainson 1837, on which 
it is based, is considered by most modern authors to be a synonym 
of Lonchura Sykes 1832. If the name Spermestinae were restored, 
the valid name of the type genus of this subfamily would be Lon- 
chura, an inconvenient lack of congruence. The name Spermestes 
has not even been used by recent authors for any of the subdivisions 
of the Estrildidae, the waxbills (Estrildae), the grass finches 
(Erythrurae or Poephilae), or the mannikins (Amadinae or Lon- 
churae). 

It is clear that this is a situation to which Article 23d (ii) of 
the International Code is applicable. There is strong justification 
for the name Estrildidae to be placed on the Official List and 
the name Spermestidae on the Official Index. 

Delacour's revision was followed by reviews of the family (or 
some of the species) by Desmond Morris (1954, 1958), Moynihan 
and Hall (1954), Wolters (1957), Steiner (1960), Hall (1962), 
and Immelmann ( 1962). Steiner, in particular, proposed a classifi- 
cation which differed drastically in many ways from that of 
Delacour. For the 108 (-110) species of the family, Delacour rec- 
ognized 15 genera with a total of 26 subgenera (including the 



4 BREVIORA No. 287 

nominate subgenera), while Steiner recognized 34 genera with 53 
subgenera, and Wolters 3 1 genera with 62 subgenera. 

It has become very evident that there is at present a real differ- 
ence of opinion between the taxonomists and the ethologists con- 
cerning the size of genera. In recent years many species of estrildids 
have been kept in captivity and their courtship and other aspects of 
biology have been carefully analyzed. Through these studies the 
ethologists have discovered more and more behavior differences 
even between the most closely related species, and are therefore 
enormously impressed with the distinctness of every species (except 
a few sibling species or semispecies). As a consequence, they tend 
to admit numerous genera, many of them monotypic. For instance, 
Immelmann (1965) recognizes 13 genera for the 16 native Aus- 
tralian superspecies of estrildids (2 species have 2 semispecies 
each). Such an emphasis on distinction destroys the concept and 
function of the genus as a means of expressing relationship. Ad- 
mittedly, there are some extremely distinct species which must be 
classified as monotypic genera, but for others it is much better to 
place them in the same genus with the most nearly related species. 
Recognition of subgeneric status is sufficient to bring out the proper 
level of distinctness. 

The discrepancy between the various authors with respect to the 
sequence of genera is equally disconcerting. Although Delacour's 
sequence, waxbills, grass finches, and mannikins, has been, on the 
whole, adopted by most subsequent authors, there has been great 
variation in the sequence of genera within each subgroup, and in 
the assignment of certain genera, particularly Aegintha, Amadina, 
Erythnira, and Chloebia, to the three subgroups. 

In an endeavor to achieve a reasonable consensus, I worked out 
a tentative sequence of recognized genera, and submitted it to the 
following specialists for their opinions: Collin Harrison, Klaus Im- 
melmann, Allen Keast, Desmond Morris, J. Nicolai, Melvin A. 
Traylor, and H. W. Wolters. They responded most generously and 
supplied me with abundant details bearing on the question of rela- 
tionship and the importance of various taxonomic characters. A 
revised sequence, prepared on the basis of this first set of answers, 
was circulated together with a questionnaire in which 19 specific 
questions were asked concerning sequence and the advisability of 
recognizing certain genera. 

The present paper is the result of an analysis of this correspond- 
ence. It is obvious that relationships cannot be decided by majority 
vote, and yet I feel that an arrangement that has been reached in 
consultation with the active specialists has a greater prospect for 



1968 GENERA OF ESTRILDIDAE 5 

Stability than an arrangement based on the opinions of a single 
worker. The following comments may be useful to other students 
of the Estrildidae. Publication of these notes at this time may 
stimulate further research that can still be utilized in the prepara- 
tion of the manuscript on this family for Peters' Check-list of 
Birds of the World. 

The Estrildidae appear to be unique among bird families in that 
color pattern is a relatively poor clue to relationship. On the other 
hand, courtship song (Hall, 1962) and display postures seem to 
have high taxonomic weight. As a consequence, a sorting of the 
species and genera on the basis of coloration leads to a classifica- 
tion which is flatly contradicted by various biological characteristics, 
including the palate pattern of the young. As in all weighting of 
taxonomic characters, there is an element of subjectivity in the de- 
cision to give higher weight to behavioral characters than to 
plumage pattern, yet this decision is supported in this family by a 
correlation among the various biological characters. The sequence 
here proposed is based on this correlation. 

THE WAXBILLS (ESTRILDAE) 

A glance at Delacour's diagram of relationships (1943, fig. 1) 
clearly shows the central position of Clytospiza. One might, there- 
fore, start the sequence of genera with Clytospiza and follow it 
with the several derived sequences. However, this not only results 
in several breaks in the sequence, but it also removes Clytospiza 
far from the Estrilda group which also seems to be derived from a 
Clytospiza-WkQ ancestor. Under the circumstances, it is better to 
adopt the following sequence: Parmoptila, Nigrita, Nesocharis, 
Pytilia, Mandingoa, Cryptospiza, Pirenestes, Spermophaga, and 
Clytospiza. This arrangement differs from that of Delacour in two 
significant respects: Nesocharis Shelley is considered a valid genus, 
probably related to Nigrita; Mandingoa (nitidida) is considered 
related to Cryptospiza rather than to Hypargos. All 9 genera are 
well defined and form a primitive group among the waxbills, a 
group on the whole more closely associated with bushes or forest 
than with grass. "I place Nesocharis near Nigrita to which it has 
some similarity in mouth markings and habits, but affinities with 
Nigrita are by no means certain. However, I can find no better 
place for this very distinct, tit-like genus" (Wolters in litt.). 

There is comparatively litde argument about the Estrilda group 
among the waxbills, except on the number of valid genera. Dela- 
cour placed all species in Estrilda (and Hypargos), while Steiner, 



6 BREVIORA No. 287 

Wolters, and Immelmann would like to recognize a minimum of 8 
genera, probably best arranged in the following sequence: Hypar- 
gos, Euschistopiza, Lagonosticta, Uraegintlius {^= Graiiatina), 
Estrilda, Ortygospizci, Amandava (incl. Sporaeginthus), and Stic- 
tospiza. Two pairs of these "genera" are rather close to each other, 
and it may result in more equal standards when Lagonosticta and 
Uraegintlius are combined, and also Amandava with Stictospiza. 
"Uraegintlius is closer to Lagonosticta than to Estrilda with which 
it has been merged by several authors" (Wolters in litt.). Estrilda, 
Lagonosticta, and Uraeginthus have been lumped at various times 
or split more finely. Estrilda and Lagonosticta seem sufficiently 
different in song and courtship to be maintained as two genera. 
Uraeginthus, on the other hand, is sufficiently closer to Lagonosticta 
than to Estrilda, that it might be treated only as a subgenus (Wol- 
ters, 1966). The species of this group are to be assigned as follows 
(not necessarily exactly in the listed sequence) : 

Estrilda 

melanotis, paludicola, melpoda, rhodopyga, troglodytes, as- 
trild, nonnula, atricapilla, erythronotos (-f- charmosyna) , 
thomensis, perreini, caerulescens, rufibarba, nigriloris 

Lagonosticta, subg. Lagonosticta 

larvata -\- nigricolUs + vinacea, rara, rubricata, rhodopareia 
-j- jamesoni, senegala, rufopicta, nitidula, landanae 

Lagonosticta, subg. Uraeginthus 

bengalus, angolensis, cyanocephala, granatina, ianthinogaster. 

Granatina (1850) would have one year priority over Uraegin- 
thus (1851), if it were an available name. It was, however, pub- 
lished by Bonaparte in the plural as Granatinae, and is thus not 
available according to Article llf of the Code. The subgenus has 
to be called Uraeginthus. 

"The similarities between Ortygospiza and Amandava in plumage 
pattern, mouth markings, and behavior are so numerous that it 
would be difficult to believe that they should be due to con- 
vergence" (Wolters in litt.). Ortygospiza deserves only subgeneric 
status. 

THE GRASS FINCHES (POEPHILAE) 

Aegintha {temporalis) is a problem. On first sight, the Aus- 
tralian temporalis seems very close to the African Estrilda (e.g. 
melpoda), and this is where Delacour placed it. Yet, in its display 
and some other respects it seems close to the Australian Emblema 



1968 GENERA OF ESTRILDIDAE 7 

group and is placed with the Australian genera by Steiner, Immel- 
mann, and Wolters. Others still think that the similarity to Estrilda 
is more than convergence. At present, it would seem best to place 
Aegintha in the grass finches, but closest to the waxbills. "Aegintha 
is the 'Estrilda-type'' among the Australian finches. Superficially, 
there is considerable similarity with African Estrilda, but the green 
of Aegintha is not found anywhere among the Estrildae (the^kind 
of green in E. melanotis is quite different). The eyestripe is above 
the eye, not through the eye, and of a different red. The courtship 
has rather primitive aspects as in Estrilda and Zonae ginthus, and 
does not provide decisive information. I am not at all convinced 
of any close relationship between Aegintha and Estrilda." (Immel- 
mann, 9. IV. 1966 in litt.). 

Delacour recognized only three genera, Zonaeginthus [=:Em- 
blema], Poephila, and Erythrura. All subsequent work has con- 
firmed the naturalness of these three groups, but this is where the 
agreement ends. Steiner, for instance, splits Emblema into 7 
genera {Zonaeginthus, Stagonopleura, Emblema, Aidemosyne, 
Bathilda, Neochmia, and Oreostruthus) , all but Zonaeginthus 
(2 species) monotypic. Aidemosyne, as shown by recent re- 
searches (Steinbacher and Wolters, 1953 ff.; Keast, 1958; Morris, 
1958; Immelmann, 1962), is close to Lonchura and must be shifted 
to the mannikins. The palate pattern is the only resemblance with 
the Emblema group. To split the remaining Emblema group into 6 
genera seems excessive. Recognition of three genera, Emblema 
(with Stagonopleura and Zonaeginthus), Oreostruthus (the some- 
what aberrant Papuan genus), and Neochmia (with Bathilda) 
would seem to give better balance. 

There is great confusion about the nomenclatural validity of the 
generic names in the Emblema group. The name Stagonopleura 
was proposed by Reichenbach in 1850 for a bird of which a black- 
and-white drawing of head, legs, and tail is given. Subsequent 
authors have applied the name to various groups of Australian 
estrildines, but the combination of a gray (or at least dark) crown, 
black lores and eye patch, and light (?white) unmarked chin and 
throat occurs in only one species, the Diamond Firetail Finch 
{guttata). Cabanis was therefore quite right (1851, Mus. Hein., 
1: 172) in designating Eoxia guttata Shaw as type for "Reichen- 
bach's genus." The name Stagonopleura (pleurae with drop-like 
spots) also fits this species excellently. 

Bonaparte (1851, Consp. Av., 1: 456), even though crediting 
the name to Reichenbach, inadvertently misspelled it Steganopleura 
(covered-up or roofed-over pleurae), which makes no sense for 



8 BREVIORA No. 287 

this species. This spelHng does not constitute an available name, 
since it is clearly an "incorrect subsequent spelling" and not an 
"intentional emendation" in the sense of Article 33. As stated in 
the Code: "Any change in the spelling of a name, other than an 
emendation, is an 'incorrect subsequent spelling'; it has no status in 
nomenclature and therefore does not enter into homonymy and 
cannot be used as a replacement name." 

Ignoring the unavailability of this name, Mathews (1919, Birds 
of Australia, 7: 434) designated Fringilla bichenovii Vigors and 
Horsfield as the type of Steganopleura Bonaparte and used it for 
the group of species previously known under the generic names 
Stictoptera Reichenbach or Stizoptera Oberholser. This was clearly 
an invalid action. 

For Stagonopleura of Reichenbach 1850, Mathews uses his own 
name Tavistocka 1919, while Delacour, Keast, and Immelmann use 
Zonaeginthus Cabanis 1851. Actually, as shown above, Stagono- 
pleura continues to be an available name, but there is doubt as to 
its validity (see under Emblema below) . 

Keast employs a subgenus Zonaeginthus {Emblema) , but since 
Emblema was proposed in 1842 and Zonaeginthus in 1851, it 
clearly will have to read Emblema (Zonaeginthus) . Immelmann 
recognized Emblema as a separate genus from Zonaeginthus, which 
carries splitting rather far. For the closely knit group of species 
consisting of pictus, oculatus, bellus, and guttatus Shaw we have 
thus three generic names available: Emblema Gould 1842 (picta), 
Stagonopleura Reichenbach 1850 (guttata), and Zonaeginthus 
Cabanis 1851 (bellus), and the only remaining question is whether 
the name Emblema is valid or preoccupied. 

Emblema Gould 1842 was in unchallenged use between 1842 
and 1930, when Iredale (1930, Austral. Zool., 6: 175) contended 
that it was preoccupied by Emblema Deshayes 1840 (mollusks). 
He renamed it Cayleyna. Deshayes' name was neither an erroneous 
spelling of Amblema Rafinesque nor a valid emendation. When 
one looks up the original citation of Emblema Deshayes (1840, 
D'Orbigny's Diet. Univer. Hist. Nat. 1: 334), one finds that he 
mentions Emblema merely in a purely philological comment, as the 
correct transliteration of the Greek word from which it was derived, 
but lists his discussion under "Ambleme (Amblema).'" Since he 
does not adopt Emblema, this spelling cannot qualify as an emen- 
dation in the sense of Article 33. It is a spelling without nomen- 
clatural status. It is fortunate for the stability of nomenclature that 
there are only a few authors who would use the flimsy excuse of 



1968 GENERA OF ESTRILDIDAE 9 

this philological discussion to upset the nomenclature of a well- 
known genus that had been totally stable for the preceding 88 
years! Nor have the malacologists ever employed the version 
Emblema. Since 1914, the name Amblema Rafinesque has been 
used universally for the so-called washboard clams (type, A. costata 
Rafinesque, 1820) and there is not the slightest reason for sup- 
pressing Emblema Gould for fear that it might be confused with 
the moUuskan genus Amblema. A recent Opinion of the Commis- 
sion (not yet published) deals with this name. Since writing this, I 
have discovered that Mees (1961, J. Roy. Soc. West. Austr., 44: 
123) had independently come to the same conclusions. 

The complete synonymy of the genus Emblema is thus as fol- 
lows: 

Emblema Gould, 1842, Birds Australia, pt. VII. Type, by 
monotypy, E. picta Gould. [Not preoccupied by the nomen- 
claturally unavailable Emblema Deshayes 1 840. j 

Stagonopleura Reichenbach, 1850, Av. Syst. Nat., pi. LXXV. 
Type, by subsequent designation (Cabanis, 1851), Loxia guttata 
Shaw. 

Steganopleura Bonaparte, 1851, Consp. Gen. Av., 1: 456. 
Nomenclaturally unavailable, as an "incorrect subsequent spelling." 

Zonaeginthus Cabanis, 1851, Mus. Hein., 1:171. Type, by 
subsequent designation (Gray, 1855: 76), Loxia nitida Latham =i 
L. bella Latham. 

Tavistocka Mathews, 1919, Birds Australia, VII: 434. New 
name for Stagonopleura Reichenbach. 

Cayleyna Iredale, 1930, Austr. Zool., 6: 175. New name for 
Emblema Gould. 

The Poephila group is typical for the difficulties one faces in the 
grass finches. There are three fairly distinct species groups, con- 
taining species that "are highly specialized in color pattern and 
general proportions, and no intermediates exist between them. 
That is the reason why so many monotypic, and in my judgment 
utterly unnecessary genera have been created for them" (Delacour, 
1943: 80). The behavior specialists, on the other hand, are im- 
pressed by the differences. I propose a compromise solution, in 
which two subgenera {Poephila, Taeniopygia) are recognized. The 
species bichenovii is not sufficiently different from guttata Vieillot to 
justify recognition of a third subgenus (Stizoptera). The grass 
finches sensu stricto, after the removal of Erythrura (see below), 
should be called the Poephilae, consisting of the genera Emblema, 
Oreostruthus, Neochmia, and Poephila. 



10 BREVIORA No. 287 

THE MANNIKINS (LONCHURAE) 

Gould's Finch (gouldiae), as shown by recent behavior studies 
(Wolters, 1950; Mitchell, 1958; Immelmann, 1959; and Steiner, 
1960), does not belong in Poephila. It was first thought to fit into 
Erythrura but recent studies indicate similarities also with Loncliura 
(Nicolai MS; Hall MS; Immelmann, 1962). Recognition of the 
genus Chloebia (near Erythrura) for gouldiae would seem indi- 
cated until the situation is better understood. It is also now becom- 
ing clear that Erythrura is not really a grass finch, related to 
Emblema-Poepliila , but that it is very distantly related to the Lon- 
chura group. The Erythrura group is, however, very isolated. 

"Chloebia shows a great deal of similarity with Erythrura in 
plumage color and mouth markings but reminds one in pattern also 
of some species of Lonchura (e.g. castaneothorax) . In behavior it 
occupies an equally intermediate position, only there are more simi- 
larities with Lonchura than with Erythrura. The voice of Chloebia 
is typical for Erythrura, while the display is very similar to that of 
Lonchura malacca, L. maja, and L. castaneothorax. Chloebia sings 
with the same rigid posture and a puUed-in head as these species of 
Lonchura, but has retained a number of ancestral components of 
the Estrildidae courtship which have been lost by the Lonchura spe- 
cies, such as the hopping during the courtship dance. The species 
of Erythrura which I know best (trichroa, cyanovirens, psittacea) 
not only fail to carry grass stems during their display but even the 
dance itself has completely disappeared." (Nicolai, in litt.). 

The precopulatory display of Erythrura is unique among the 
Estrildidae and resembles neither that of Lonchura nor that of any 
other genus. "Erythrura prasina is somewhat intermediate between 
typical Erythrura and Chloebia and since the latter has many simi- 
larities with Lonchura, I believe that it is best to place Chloebia 
between Erythrura and Lonchura." (Nicolai, 4 August 1965, in 
litt. ) . 

The mannikins are, on the whole, a homogeneous group. The 
only genera whose assignment to the Lonchurae can be questioned 
are Amadina and Chloebia-Erythrura. As far as the remaining spe- 
cies are concerned, there is only the question of the number of 
genera to be recognized. Are Euodice, Odontospiza, and Spennestes 
valid? Spennestes (and Odontospiza) have certain primitive traits, 
and so does Euodice (which Harrison, 1964, cannot confirm). 
Some recent authors recognize Spermestes as different from Lon- 
chura; others, like Delacour, do not. Kunkel (1965) lists the be- 
havioral differences, which to me do not seem to justify more than 



1968 GENERA OF ESTRILDIDAE H 

subgeneric rank. The same is true for Euodice. Odontospiza de- 
serves generic status even less and is best placed in the same sub- 
genus {Euodice) as malabarica-cantans. The Java Finch {oryzi- 
vora) is so distinct that it is best recognized as a separate genus 
(Padda), even though admittedly Padda juscata connects this genus 
with Lonchwa; "Heteromunia" (pectoralis) is so close to Lon- 
chura, particularly the castcmeothorax group, that it cannot be rec- 
ognized as a monotypic genus, even though it has some remarkable 
specializations (Immelmann, 1965). The suggestion that it is 
closely related to modesta (Aidemosyne) does not seem well sub- 
stantiated. 

Even though neither Spermestes nor Euodice is sufficiently dif- 
ferent from Lonchwa to justify generic separation, I have recog- 
nized them as subgenera for those who like to emphasize their dis- 
tinctiveness. 

The grouping and the sequence of species in Lonchwa (sensu 
stricto) is a problem. Certain groups are well defined, for instance 
the castaneothorax group (with flaviprymna, castaneothorax, tee- 
rinki, stygia, monticola, montana, and melaena). Another fairly 
well defined assemblage consists of punctulata, kelaavti, leucogastra, 
mohicca, leucogastroides, juscans, and striata. The two New 
Guinea species tristissima and leucosticta stand well apart. The 
difficulties are with the remaining species. L. quinticolor (Lesser 
Sunda islands) is usually placed near grandis of New Guinea, but 
the similarity may well be superficial. The species maja and pallida 
are usually placed near the castaneothorax group, supposedly con- 
nected by flaviprymna, but I consider this similarity as secondary. 
L. flaviprymna seems to have lost the typical characters of the 
castaneothorax group, perhaps as a result of life in arid Australia. 
There has been a good deal of parallelism and convergence in this 
group of mannikins, and further analysis of songs and other charac- 
ters is needed. (The sequence adopted in the Check-list attempts 
to bring close relatives together, but is surely not the last word.) 

Amadina is perhaps the most aberrant genus of the mannikins. 
It does not quite fit with anything and is best listed last. Most 
likely it is an aberrant offshoot of the Lonchwa group. Padda is 
close to Lonchura, but somewhat specialized. Lonchwa is un- 
doubtedly the central genus of this whole assemblage and the man- 
nikins should be called Lonchurae. 

It is possible that the three groups now informally designated as 
the Estrildae, Poephilae, and Lonchurae, will eventually deserve 
to be recognized as three tribes. At present, it seems however, that 
the differences are not sufficiently trenchant to justify tribal status. 



12 



BREVIORA 



No. 287 



Furthermore, there are too many genera, the placement of which 
is still doubtful. An informal classification, as first proposed by 
Delacour, would seem more appropriate for the time being. 

REVISED SEQUENCE OF GENERA OF ESTRILDIDAE 



I. 


ESTRILDAE 


II. 


POEPHILAE 


1 


Parmoptila 


15 


Aegintha 


2 


Nigrita 


16 


Emblema 


3 


Nesocharis 


17 


Oreostruthus 


4 


Pytilia 


18 


Neochmia 


5 


Mandingoa 


19 


Poephila 


6 


Cryptospiza 






7 


Pirenestes 






8 


Spermophaga 


III. 


LONCHURAE 


9 


Clytospiza 


20 


Erythrura 


10 


Hypargos 


21 


Chloebia 


11 


Euschistospiza 


22 


Aidemosyne 


12 


Lagonosticta 


23 


Lonchura 


13 


Estrilda 


24 


Padda 


14 


Amandava 


25 


Amadina 




BIBLIOGRAPHY 





Chapin, J. P 

1917. The classification of the weaver-birds. Bull. Amer. Mus. Nat. 
Hist., 37: 243-280. 

Delacour, J. A. 

1943. A revision of the subfamily Estrildinae of the family Ploceidae. 
Zoologica, 28: 69-86. 

Hall, M. F. 

1962. Evolutionary aspects of estrildid song. Zool. Soc. London. 
Symposia No. 8: 37-55. 

Harrison, C. J. O. 

1964. The taxonomic status of the African silverbill Lonchura cantans 
and Indian silverbill L. malabarica. Ibis, 106: 462-468. 

Immelmann, K. 

1959. Die Verwandtschaftsbeziehungen der Gouldsamadine. Gef. 
Welt, 83: 205-208. 



1968 GENERA OF ESTRILDIDAE 13 

1962. Beitrage zu einer vergleichenden Biologic australischer Pracht- 
finken (Spermestidae). Zool. Jb. Syst., 90: 1-196. 

1964. A critique of Mitchell's paper on the relationship of Neodinva. 
J. Ornithol., 105: 97-99. 

1965. Australian finches in bush and aviary. Angus and Robertson, 
Sydney, 196 pp. 

1966. Ecology and behavior of African and Australian grass finches 
— a comparison. Ostrich, supp. 6: 371-379. 

Keast, a. 

1958. Infraspecific variation in the Australian finches. The Emu. 
58: 395-411. 

KUNKEL, P. 

1965. Verhaltensstudien an den Kontinental-afrikanischen Elsterchen 
{Spermestes, ^vjdimson). Die Vogelwelt, 86: 161-178. 

Mitchell, I. G. 

1958. The taxonomic position of the Gouldian Finch. The Emu, 58: 
395-411. 

1962. The taxonomic position of the Crimson Finch. The Emu, 62: 
115-125. 

Morris, D. 

1954. The courtship behavior of the Cutthroat Finch. Avicultural 
Magazine, Sept./Oct.: 169-171. 

1958. The comparative ethology of grass finches (Erythrurae) and 
mannikins (Amadinae). Proc. Zool. Soc. London, 131: 389- 
439. 

MoYNiHAN, M., and M. F. Hall 

1954. Hostile, sexual and other social behavior patterns of the Spice 
Finch {Lonchiira punctulata) in captivity. Behaviour, 7: 33-76. 

NiCOLAI, J. 

1964. Der Brutparasitismus der Viduinae als ethologisches Problem. 
Zeitschr. Tierpsych., 21: 129-204. 

Steinbacher, J., and H. E. Wolters 

1953-1960. Vogel in Kafig und Voliere. Teil Prachtfinken. Aachen., 216 pp. 

Steiner, H. 

1955. Das Brutverhalten der Prachtfinken, Spermestidae, als Ausdruck 
ihres selbstandigen Familiencharakters. Acta XI Cong. Int. Orn. 
Basel: 350-355. 

1960. Die Klassifikation der Prachtfinken, Spermestidae, auf Grund 
der Rachenzeichnungen ihrer Nestlinge. J. Omith., 101: 92-112. 



14 BREVIORA No. 287 

SUSHKIN, p. p. 

1924. Outlines of classification of Fringillidae and allied groups. Bull. 

Brit. Orn. Club, 45: 36-39. 
1927. On the anatomy and classification of the weaver-birds. Bull. 

Amer. Mus. Nat. Hist., 57: 1-32. 

WOLTERS, H. E. 

1949. Die Finkenvogel. (1) Beitrage zur Gattungssystematik der 
Vogel. Goecke & Evers. Krefeld, pp. 3-17. 

1950. iiber einige Gattungen der Estrildinae. Bonn. Zool. Beitr., 
1: 33-38. 

1957. Die Klassifikation der Webefinken (Estrildinae). Bonn. Zool. 
Beitr., 8: 90-129. 

1966. On the relationships and generic limits of African Estrildidae. 
Ostrich, supp. 6: 75-81. 

ZlSWILER, V. 

1965. Zur Kenntnis des Samenoffnens und der Struktur des hornernen 
Gaumens bei kornerfressenden Oscines. J. Ornith., 106: 1-48. 

1967. Vergleichend morphologische Untersuchungen am Verdauungs- 
trakt kornerfressender Singvogel zur Abklarung ihrer syste- 
matischen Stellung. Zool. Jb. Syst., 94: 427-520. 

(Received 20 December 1967.) 



BREVIORA 



Museiuiim of Coiniparsitive Zoology 

Cambridge, Mass. 29 May, 1968 Number 288 

THE EPIDERMAL GLANDS OF LYGODACTYLUS 
(GEKKONIDAE, LACERTILIA) 

P.F.A. Maderson' 



Recent histological and ultrastructural studies have shown that 
the squamate epidermis is quite different from that of other verte- 
brates. In snakes and lizards, unique "epidermal generations" 
(Maderson, 1967) can be identified. These units, whose periodic 
appearance, maturation, and loss are reflected in a series of dif- 
fering histological pictures at various times in the sloughing cycle, 
are made up of six (and sometimes more) different cell populations 
arising sequentially from an apparently homogeneous stratum 
germinativum. Some of these populations, e.g. those that form the 
superficial Oberhaiitcheu component of the /3-layer, or the in- 
nermost "clear layer," normally comprise only a single contiguous 
cell layer over the entire body surface. Others, e.g. the cells com- 
prising the remainder of the /:i-layer, or the a-layer, are repre- 
sented by numerous cell layers, although there may often be con- 
siderable variation from one genus or species to another or even 
from one part of the body or from one scale to another. 

Although the details of the morphology of the squamate epi- 
dermis have only recently been reported (Maderson, 1965a, b, 
1966b, 1967; Maderson and Licht, 1967; Roth and Jones, 1967), 
certain specialized modifications of this organ system have been 
described. Conclusive demonstration of an associated nerve supply 
(Miller and Kasahara, 1967) has confirmed the identity of so- 
called "sense organs" in lizards, described by earlier workers 
(Schmidt, 1920;''Preiss, 1922). These studies, and those of the 
climbing organs of gekkonid and anoline lizards (Maderson, 
1964a;"l966a; Ruibal and Ernst, 1965; Ernst and Ruibal, 1966; 
Lillywhite and Maderson, 1968) have shown that although the 
piimary function of the Oberhautchen is to facilitate sloughing 

1 Research Associate, Department of Dermatology, Harvard Medical School. 



2 BREVIORA No. 288 

(Maderson, 1966b), this cell layer has undergone considerable 
adaptive specialization in various lizard groups. 

Specialization of the squamate epidermis is not restricted to the 
formation of sensory and climbing organs; gland-like structures 
have also been reported (Coie, 1966a, b). While I was assembling 
material for a study on the so-called "glandular escutcheon scales" 
(Taylor and Leonard, 1956) of the Central American gekkonid 
Gonatodes, Dr. E. E. Williams of Harvard University drew my 
attention to the recorded presence of apparently similar structures 
in the East African gekkonid Ly^odaciylus (Pasteur, 1964). Pre- 
liminary examination of a few specimens revealed fundamental 
differences in the microscopic anatomy of the organs in the two 
genera. The structure of the escutcheon scales of Gonatodes proved 
to be not only of intrinsic interest but also contributed considerably 
to our understanding of the pattern of development of a squamate 
epidermal generation (Maderson, 1967). ¥yo\w a comparative 
cytological point of view, the superficiaily similar structures in 
Lygodactylus are more of a curiosity than of fundamental sig- 
nificance, but a detailed account of their structure is warranted 
for other reasons. First, it provides another example of the re- 
markable evolutionary potential of the squamate epidermis for the 
formation of specialized structures; second, it represents a further 
contribution to our knowledge of gekkonid anatomy; and finally, 
it helps to shed some light on the problem of the homology of the 
various types of "pores' found in gekkonids (Kluge, 1967). 

MATERIAL AND METHODS 

Through the courtesy of Dr. E. E. Williams of Harvard Univer- 
sity and Dr. J. Peters of the Smithsonian Institution, I was able to 
obtain 29 male specimens of Lygodactylus spp. Loveridge's ( 1947) 
and Pasteur's (1964) statements to the effect that only males show 
the specialized glandular scales and pre-anal pores were confirmed 
by dissection and histological examination of 12 females from the 
same sources. The species were identified as L. guttiiralis, L. pic- 
turatus, L. p. keniensis and L. fisc/ieri from various known localities 
in East Africa. No significant differences in the structure of the 
glands were seen between species. 

Pieces of skin were removed from the posterior abdominal sur- 
face just anterior to (but sometimes including) the pre-anal pores, 
or from the ventral aspect of the femoral region, which shows 
similarly modified scales (see below). Histological examination re- 
vealed that there are similarly modified scales between the line of 



1968 GEKKONID SKIN GLANDS 3 

pre-anal pores and the vent, but there are no external indications of 
their presence. The history of some of the material was uncertain 
but it was probably fixed in the field in formalin and later stored in 
70 per cent alcohol. The specimens were dehydrated in alcohol, 
cleared in chloroform, embedded in 56°C paraffin, and cut at 1 ^x. 
vertical to the skin surface, either transverse or longitudinal to the 
scale axis (see below). Sections were mounted serially, a few from 
each ribbon being mounted unstained and examined by phase con- 
trast. The remainder were stained in Ehrlich's hematoxylin and 
eosin, or aniline blue-orange G. or Masson's trichrome (Gurr, 
1958). The material in the pre-anal pores proved to be exception- 
ally difficult to cut. giving confirmatory evidence of its nature (see 
below). 

OBSERVATIONS 
The Scales 

Macroscopic appearance. The posterior abdominal, ventral 
femoral, and ventral tibio-fibular scalation is quite different in the 
males (PI. 1. fig. 1, left) than in the females (PI. 1, fig. 1, right). 
In the male, 2 to 3mm anterior to the vent, there is a transverse 
row of slightly enlarged scales on whose outer surface (Maderson, 
1964b) is seen a circular opening about 0.3 mm across, in which a 
plug of material may be seen. There are 7 to 9 of these so-called 
"pre-anal"' pores depending on the species (Pasteur, 1964); not 
infrequently there may be one or more which is considerably larger 
than the others. In the male, the darker pigmentation of these re- 
gions is distinctive. The scales lying in a blunt V-shaped area 
Teading forwards from the line of pre-anal pores are the same shape 
(regularly trapezoid, the short axis lying parallel to the body axis, 
with little overlapping of successive elements) as the more anterior 
abdominal scales, but are distinctly darker. On the femoral region 
there are patches of scales of essentially similar shape (Loveridge, 
1947), which are very much darker than those on the abdomen. 
This femoral patch grades into non-specialized, lighter-colored 
scales anteriorly (PI. \ fig. 1, left) but ends sharply posteriorly. 
There is a similarly modified patch on the ventral aspect of the 
tibio-fibular surface (Loveridge, 1947). The individual scales all 
show a more lightly pigmented marginal area; this is seen in histo- 
logical sections^ to indicate the lateral margins of the specialized 
zones on each outer scale surface (Maderson, 1964b). 

Microscopic appearance. As has been described elsewhere for 
Gonatodes (Maderson, 1967), the glandular scales of Lygodactylus 



4 BREVIORA No. 288 

show a number of different histological conditions which can be 
interpreted as manifesting changes in association with periodic 
sloughing. This assumption will be followed in the descriptions 
below of four significantly different conditions which are important 
in interpreting the morphological structure of the specialized scales. 

All the specimens examined showed conditions which could be 
interpreted as belonging to the "proliferation-renewal phase" 
(Maderson, 1965a, b, 1966b. 1967; Maderson and Licht, 1967). 
That a large museum sample should show 100 per cent of the 
material in the proliferative phase is very unusual in the author's 
experience. Mr. Allen Greer of Harvard University collected the 
21 specimens examined of the Kenyan L. picturatus during an 
ecological study (Greer, 1967), and he provided me with further 
data regarding this material. The animals were caught by hand, 
or shot with dust-shot, and then fixed in formalin within at least 
four hours. Experience in collecting Australian gekkonids (Mader- 
son, unpublished data) suggests that the stress of capture initiates 
a new epidermal cycle (Maderson, 1967), with the result that ma- 
terial obtained from animals kept in captivity for three or four days 
will always show various stages in epidermal proliferation. Al- 
though Mr. Greer tells me that he rarely saw L. picturatus actually 
sloughing, on the few occasions that he did so, they were engaged 
in removing material from their body surfaces with their teeth. 
This behavior pattern has been described in a number of squamate 
species (Bustard and Maderson, 1965). In Gekko gecko (Chiu, 
Phillips and Maderson, 1967) and Auolis carolinensis (Maderson 
and Licht, 1967) the complete physical removal from the body of 
the material to be shed is accomplished in a very short time (some- 
thing less than 5 hours and often during the night [Maderson, 
unpublished data]). Although the material examined here did not 
permit an estimate of the time occupied by the various stages of 
epidermal differentiation (Maderson and Licht, 1967), the picture 
which emerges is that Lygodactxlus either sheds very frequently or 
else has a cycle in which the "resting phase"' (Maderson, 1967) is 
extremely short or absent. It would be of interest to know to what 
extent this is associated with the probable ecological and be- 
havioral significance of the glands and pores (Cole, 1966a, b). 

In median longitudinal section the individual scales are seen as 
triangular structures arising from the body surface with very little 
overlapping of adjacent elements (PI. 5, fig. 2). In transverse sec- 
tion the picture varies depending on the level along the scale axis, 
but in general one sees a rather wide-based structure, again with 
little overlap of adjacent scales. The outer scale surface (Mader- 
son, 1964b) is always distincdy concave with the deepest portion 



1968 



GEKKONID SKIN GLANDS 



towards the posterior distal region of the scale; it is in this con- 
cavity that the speciahzed epidermal material is seen. In gross and 
low-power microscopic appearance there is a very general similarity 
to the escutcheon scales of Gonatodes (Maderson, 1967). 



OUTER 
EPIDERMAL 
GENERATION 



splitting 
zone "^ 



INNER J 
EPIDERMAL 
GENERATION 




^'s^^^^^^^^mesos layer 

1 

alpha -layer 



lacunar 
tissue 

clear layer 
Oberhautchen 



"1 

BHayer 

/ 



eratinizjng 
B-cells 



mesos 



If An 



I first presumptive 
alpha cells 



stratum 
germinativum 



Figure 1 . Schematic representation of a generalized squamate epidermis 
several days before sloughing is due to take place. (Figure taken from 
Maderson [1967] and reproduced here by kind permission of the editors of 
Copeia.) 



To avoid undue repetition of references in the text of the follow- 
ing descriptions, all the terms used are those discussed and defined 
in previous works (Maderson, 1965a, 1966b, 1967; Maderson 
and Licht, 1967). This nomenclature is shown in schematic form 



in Figure 1. 



Condition One (6 specimens); PI. 1, figs. 2-4. 
Plate 1, figure 2 shows a sagittal section through a scale taken 
from the anterior margin of the specialized abdominal region where 
the gland development is smallest; an enlarged portion of the low- 
power view is shown in Plate 2, figure 4. 



6 BREVIORA No. 288 

The surface of the body is formed from a homogeneous, chromo- 
phobic tissue which shows no indications of nuclear remains or cell 
outlines; this is the /^-layer of the outer epidermal generation. The 
extreme superficial surface of the fS-layer is formed by the charac- 
teristic Oberhaiitc/wn which in gekkonids (Maderson, 1966b, 1967; 
Ruibal and Ernst. 1965) is always seen as a scries of minute ver- 
tical spinules (PI. 1, fig. 4). Towards the distal portion of the 
outer scale surface, the Oherlunitchen is separated from the normal 
/:^-material described above by a quite different material (PI. 1, 
fig. 4). The latter is basophilic in hematoxylin stained sections and 
stains a patchy orange and blue in sections stained with aniline 
blue/orange G. This suggests that the material is of a keratina- 
ceous nature, as adjudged by its affinity for orange G (Gurr, 
1958), but with a mixed complement of possible mucoprotein ma- 
terial which stains with the aniline blue. There are also copious 
deposits of melanin granules. This is here termed the /:^-gland 
material, for reasons which will be discussed later. Initial examina- 
tion posed the problem of how this could be regarded as a "gland" 
if the material was confined between the Oberluiutchen (which is a 
very strong unit in spite of its being composed of only a single cell 
layer [Maderson, 1966a]) and the rest of the /?-layer. Although 
examination of serial sections showed that often only the edges of 
the material, proximally, distally, and laterally were covered by 
the Oberhautchen, it was assumed that histological preparation had 
destroyed a contiguous central portion. The proof that this is not 
so will be presented later, and in fact there is an approximately 
circular area on the outer scale surface where the gland material is 
directly exposed to the external environment; this is due to a re- 
gional non-development of the Oberhautchen. This is the first time 
that the author has ever seen a situation where there is not a con- 
tinuous Oberhautchen over the whole epidermis so as to cover the 
entire body surface. Even on the inner scale surface and in the 
hinge region where the /^-layer is often difficult to recognize histo- 
logically, ultrastructural studies (Ernst and Ruibal, 1966) have 
shown that the /:^-layer is represented here by the Oberhautchen 
alone. 

The ^-layer invariably separates from the underlying tissues dur- 
ing histological preparation, and the artifactual gap is traversed 
by refractive strands which represent the only histological indica- 
tions of the mature mesos layer. This is often difficult to identify 
in geckos but is marked by an X in Plate 1 . figure 4. 

Beneath the mesos layer is a region of lamellar chromophilic ma- 
terial containing dense deposits of melanin, showing no indications 



1968 GEKKONID SKIN GLANDS 7 

of nuclei except for the occasional pycnotic figure seen as a pre- 
sumptive a-cell becomes incorporated into the tissue (Maderson 
and Licht. 1967) at its very base: this is the a-layer of previous 
accounts. No further changes occur in the histological appearance 
of the tissues described thus far and they will not be mentioned 
again in subsequent descriptions. 

Beneath the z-layer are two or more rows of flattened or rec- 
tangular cells. The outermost layer(s) (plto, PI. 1, fig. 4) are 
not continuous over the entire outer surface of the scale where new 
gland material is seen (see below) but are easily recognizable in 
other regions (see below). This represents the presumptive lacunar 
tissue which plays an important role in the maturation of an epi- 
dermal generation prior to shedding (Maderson. 1967; Maderson 
and Licht. 1967). The innermost layer ipclo, PI. 1, fig. 4) is 
not readily recognizable as a typical gekkonid clear layer (Mader- 
son, 1966b. 1967) at this time, and only its later development 
really reveals its identity (see below). The tissues and layers thus 
far described comprise a "complete" outer epidermal generation 
(Maderson, 1967). 

Between the presumptive clear layer and the stratum germina- 
tivum is a cell mass comprising tight-packed polygonal cells. In 
sagittal section (PI. 1, fig. 4) there is a slight indication of oblique 
orientation of the cells towards the scale apex, but in transverse 
section (PI. 1, fig. 3) their orientation is vertical away from the 
stratum germinativum. The nuclei are rounded in the early stages 
of development (PI. I, fig. 4) but tend to become ovoid later (PI. 
1, fig. 3). The cytoplasm presents a slightly filamentous appearance 
but is often floccular in the early stages of development. There is 
only a single recognizable cell type in the region at this time. This 
mass of cells represents the first indications of the new gland ma- 
terial which will eventually be exposed to the external environment 
after the next slough. As the number of cells in this region in- 
creases, the basally situated epidermal melanocytes extend dendrites 
far above the stratum germinativum and deposit melanin granules 
into the cells (PI. l,fig."3). 

The stratum germinativum consists of cuboidal cells when the 
cell mass above it first appears (PI. 1, fig. 4), but there is a gradual 
change to a more columnar condition later (PI. 1 , fig. 3 ) . 

Condition Two (8 specimens); PI. 2, figs. 1-2. 
A longitudinal section of a major glandular scale (PI. 2. fig. 
1 ) shows that the most significant change from the previous 
condition is that the new gland material is now seen to be an in- 
tegral cell mass confined distally by a region of highly flattened 



8 BREVIORA No. 288 

cells and proximally by the stratum gerniinativum. The flattened 
cells (PI. 2, fig. 1) beneath the darkly chroniophilic 3:-layer will 
eventually form a 2 to 3 layered lacunar tissue, which wiU be 
bounded basally by a clear layer, and the flattened layer beneath 
this (the one forming the outermost boundary of the mass of new 
gland material cells) will become the Oberluiutclien. Two points 
must be emphasized here. First, the cell layers which are here 
termed the "presumptive clear layer" (pclo, PI. 2, fig. 2) and the 
"presumptive Oberhautchen" (pObi, PI. 2. fig. 2) are not yet 
recognizable on the basis of their cytology, only by their topo- 
graphical position. Second, Plate 2, figure 2 shows a section 
through a lateral part of a scale where a distinct lacunar tissue and 
clear layer /Ober/uiutc/ien complex forms (see below); the picture 
in a median sagittal plane would be very different. Here the base 
of the mature a-layer would lie directly against the uppermost cells 
of the mass of new gland material. The high power micrograph of 
the lateral margin as seen in transverse section (shown as PI. 2, 
fig. 2) shows some of the cytological detail visible at this time. In 
this particular field the lacunar tissue is not developed (it is of 
variable occurrence and distribution in lacertilians [Maderson, 
1966b; Maderson and Licht, 1967]). The Oberhautchen (which is 
not as regular in shape here as over the general body surface of 
other gekkonids [Maderson, 1966b]) is just recognizable under oil 
immersion by the slightly thickened outermost cell membranes. 
The overlying clear layer is also distorted in most regions of the 
glandular scales in L\godact\lus, as compared with that of other 
gekkonids (Maderson. 1966b). 

The extraordinary variations in cell shape, distribution of ele- 
ments, etc., described above would make identification of the 
lacunar tissue, clear layer, and Oberhautchen very tenuous, were it 
not for the fact that in serial sections the various regions and layers 
can be traced along the adjacent "semi-specialized" (see later) and 
unspecialized scales. In these last locations, the histology ap- 
proaches the conditions described for the normal body epidermis 
of Gekko gecko during the early proliferative phase (Stages 2 
and 3 [Maderson, 1966b]). Certain features of these adjacent 
regions, which provide clues as to the homologies of the various cell 
types seen in the specialized scales of Lygodactylus, will be dis- 
cussed later. 

Condition Three (13 specimens); PI. 2, figs. 3, 4; PI. 3, figs. 1, 2. 
In these specimens one sees the laying down and subsequent 
maturation, i.e. keratinization, of cells which can be recognized as 



1968 GEKKONID SKIN GLANDS 9 

a ^-, a mesos, and a partially formed a-layer of the inner epidermal 
generation (Maderson, 1967). The pattern of histological changes 
occurring in the lacunar tissue and clear layer of the outer epi- 
dermal generation supports the general assumption that the gland 
scales resemble a much-modified form of the typical gekkonid body 
epidermis during the pre-sloughing period (Stages 3-5 [Maderson, 
1966b]). Apart from the gland material, all of the regions which 
are identified and discussed below are contiguous with similar re- 
gions of the epidermal surface of adjacent unmodified scales which 
resemble exactly the general body epidermis described elsewhere 
(Maderson, 1966b). 

A low-power view of a transverse section through a glandular 
scale when the first presumptive ^-cells become visible is shown in 
Plate 2. figure 3, and the region indicated by the vertical strip is 
shown in Plate 3, figure 1 . The latter shows a portion of the scale 
where the new gland material is not bounded by an Oberhautchen 
and there is no clear layer or lacunar tissue associated with the 
outer epidermal generation, so that the outermost layer of new 
gland cells abuts directly onto the base of the a-layer, which is 
shown artifactually split away in Plate 2, figure 3. 

The mass of cells termed the "new gland material" (PI. 2, fig. 3) 
shows two distinct types of cell. The most predominant form is 
a polygonal shaped cell with a slightly thickened membrane and 
relatively large central ovoid or rounded nucleus lying in a densely 
filamentous cytoplasm (PI. 3, fig. 1). These filaments are slightly 
basophilic in hematoxylin stained sections and also show a strong 
affinity for orange G. The only similar cytological picture which 
the author has ever seen is in chick scales during the later em- 
bryonic stages when the characteristic avian /:i-keratin is being laid 
down in that location (Baden and Maderson, unpublished data). 
For this reason and to distinguish the glands of Lygodactylus from 
the escutcheon scales of Gonatodes (the formative stages of which 
are totally different [Maderson, 1967]), I suggest that these ab- 
dominal glands in Lygodactylus should be termed "/^-glands." In a 
series of specimens I have observed the gradual death of these 
cells, as indicated by the increasing frequency of pycnotic figures 
(PI. 3, fig. 2; PI. 4, fig. 3). The filaments become less and less 
distinct and the entire cell eventually becomes either finely baso- 
philic (hematoxylin) or totally orange in orange G-stained material. 
The conspicuous thickening of the cell membranes which normally 
(Maderson, 1966b. 1967) characterizes the cytogenesis of the 
subjacent presumptive /^-cells is not seen here; this may be due to 



10 BREVIORA No. 288 

the fact that the gland material never becomes quite as homoge- 
neous in appearance in its mature state as does the actual /:?-layer 
of the outer epidermal generation (see p. 6). 

The second cell type seen in the mass of presumptive gland ma- 
terial has a quite different cytology. In hematoxylin and eosin sec- 
tions the cytoplasm is tloccular and is faintly eosinophilic. The 
central nucleus is about the same size and shape as that of the other 
cells. In sections stained with aniline blue the floccular appearance 
takes on the form of a dense blue mesh-work with occasional deep- 
staining blue granules. These cells are scattered randomly between 
the filament-filled cells in the main body of the new gland material 
(PI. 3, fig. 1 ; A') but tend to be concentrated in clumps around the 
lateral, proximal, and distal margins of the material (PI. 3, fig. 3; 
PI. 4, fig. 2; X). These positions correspond to portions of the 
mature material where the blue-staining (with aniline blue/orange 
G stain) is most dense. On adjacent scales (PI. 4, fig. 1 ), cells of a 
similar nature are seen, but their distribution and relative abun- 
dance are different. Here a distinct Oberhautchen with char- 
acteristic spinules is seen; beneath this are 1 to 3 layers of the 
blue-staining granular cells (PI. 4. fig. 1; gc), and beneath these, 
in turn, normal presumptive /:^-cells in various stages of maturation 
are seen. In other instances (further away from the main "gland 
area" on the abdominal and limb surfaces) occasional cells of 
this type are seen. In the latter, the cytoplasm resembles that of 
the "protruding" Oberhautchen cells seen in the epidermis of 
Gekko gecko (Maderson, 1966b, p. 43), except that whereas the 
latter definitely showed spinules on the outer membrane, in Lygo- 
dactxlus the cells contact the clear layer cells directly with no indi- 
cation of spinule development. A final point which will be of 
significance in homologizing the various cell types is the fact that 
in Lygodactyhis (and numerous other gekkonid species [Maderson, 
unpublished data] ) definhe Oberhautchen cells, i.e. cells which 
eventually form the outermost surface of the body, which develop 
in association with overlying clear layer cells, and which show 
various degrees of spinule development, can be seen to contain 
granules which stain with aniline blue; these granules are not seen 
in the underlying presumptive /y-cells. 

Material showing this condition confirms the regional absence of 
the Oberhautchen covering over much of the gland material; the 
very sharp discontinuity of this covering is seen in transverse (PI. 
3, fig. 2) and sagittal (PI. 3, fig. 3) sections (see arrows). The 
clear layer and lacunar tissue are also absent from those regions 
where there is no Oberhautchen. A slight problem is presented by 



1968 GEKKONID SKIN GLANDS 11 

the morphology of the cell layer indicated by Y in Plate 3, figure 3. 
This layer lies between the main mass of new gland cells and the 
easily recognizable presumptive /3-cells which are nearly fully kera- 
tinized. Plate 3, figure 3 shows that this cell layer is very different 
from the underlying cells and does not contain the conspicuous 
filaments seen in the gland cells. In many ways it resembles a 
typical gekkonid Oherhautchen except for a complete absence of 
any sign of spinule development. However, the total lack of any 
indication of topographic continuity between this layer and the 
Oberliautchen (of characteristic gekkonid form) finishing at the 
arrow (PI. 3, fig. 3) suggests that the cytological similarity is mis- 
leading. I suggest that there is a large area where there is no de- 
velopment of a characteristic Oherhautchen; this point is im- 
portant in assessing the homologies of the various cellular elements 
in the glandular scales. 

The morphological appearance and subsequent maturation of the 
cell layers termed the lacunar tissue and the clear layer are exactly 
the same as those described in Gekko gecko (Maderson, 1966b, 
1967). Subsequent development of the elements comprising the 
rest of the inner epidermal generation (presumptive /y-cells, pre- 
sumptive mesos layer, presumptive a-cells) is also quite typical, 
and details of their structure will be found elsewhere (Maderson, 
1966b, 1967; Maderson and Licht. 1967). The columnar appear- 
ance and relative chromophobia of the stratum germinativum cells 
as the mesos layer is laid down (PI. 4, figs. 1, 2, 4) (Maderson 
and Licht, 1967) are emphasized. 

Condition Four (2 specimens); PI. 2, fig. 5; PL 4, fig. 4; PI. 5, 

fig. 1. 
One specimen showed a very late phase in the keratinization of 
the new /^-layer of the inner epidermal generation, with subjacent 
presumptive mesos cells showing pycnotic figures, and beneath 
these numerous presumptive ct-cells (Maderson, 1966b, 1967; 
Maderson and Licht, 1967). Another (PI. 5, fig. 1) showed a 
definite "pre-sloughing" condition with the inner epidermal genera- 
tion having a mature /3-layer, a mature mesos layer showing as 
refractive strands, as described earlier, and with the first indications 
of a new a-layer beneath. Beneath the 3 to 4 layers of presumptive 
a-cells which would normally be incorporated into the a-layer 
during the post-sloughing period (Maderson, 1966b, 1967; Mader- 
son and Licht, 1967), there were some cells which resembled 
exactly the first indications of the next mass of new gland cells, 
as described earlier. In the absence of more material one hesitates 



12 BREVIORA No. 288 

to make definitive statements, but it is probable that this genus 
never shows a typical "resting phase" in the sloughing cycle. 

Sloughing, i.e. the physical removal of the outer epidermal gen- 
eration from the body surface, involves the separation of the clear 
layer of the outer generation from the Oberhautchcn of the inner 
generation (cf. Fig. 1; Maderson, 1966b, 1967; Maderson and 
Licht, 1967) as seen in Plate 5, figure 1, left. Where the clear 
layer /Ober/uuitc/K'n complex is absent (over the central portion of 
the new gland material), the separation occurs at the base of the 
old a-layer, which lies directly on top of the new gland material (PI. 
5, fig. 1, right). Cell shapes are still visible in the latter at this 
time, and the material appears to occupy a greater vertical depth 
than does the mature material; this may be due to drying of the 
material once it is exposed to the external environment. The stain- 
ing characteristics of the gland material in the pre-slough specimen 
are exactly those of the exposed material (see p. 6). 

The Pre-anal Pores 

Macroscopic appearance. The external appearance of the pore 
openings has already been described above (p. 3). Dissection of 
the integument in the pre-cloacal region reveals the inner portions 
of the pores as apparently simple sacs running directly anteriad 
from the opening for a distance of 1.2 to 1.4 mm. The sacs lie 
between the subcutaneous tissue and the ventral musculature. They 
are surrounded by fat. 

Microscopic appearance (PI. 5, figs. 2-4; PI. 6, figs. 1-4). A 
median longitudinal section through the opening of a pre-anal pore 
is shown in low power view in Plate 5, figure 2. The scales anterior 
and posterior to the scale on which the pore opening is situated 
show ^-glands in "condition three" as described above. It is note- 
worthy that the rest of the epidermis on either side of the pore 
lumen is not modified in any way (PI. 5, figs. 3 and 4) and its 
structure is that which has been described elsewhere (Maderson, 
1966b) as a stage four condition of the normal gekkonid body 
epidermis. Plate 5, figure 4 gives a clear picture of the slightly 
modified condition normally seen towards the inner scale surface 
and hinge region where there is quite extensive development of a 
lacunar tissue (Ito) in which the characteristic basophilic granules 
(keratohyalin? [see discussion, Maderson, 1966b]) are to be seen. 
Following the epidermis from the scale surface into the mouth of 
the pre-anal pore, one sees the striking discontinuity of all the 
component layers and regions seen on the normal epidermis; these 



1968 GEKKONID SKIN GLANDS 13 

are marked by arrows in Plate 5, figures 3 and 4. The most rigor- 
ous examination of the pre-anal pore walls (see below) reveals 
no homologous layers or any resemblance to them, except for a 
stratum germinativum. The major portion of the mouth of the 
pre-anal pore in Plate 5, figure 2 is empty, but numerous fragments 
of the original "plug'' of material can be seen. This material is very 
much harder to cut than either the /?-gland material, the escutcheon 
gland material of Goimtodes (Maderson, 1967a), or iguanid gland 
material (Maderson, unpublished data). It is very slightly eosi- 
nophilic and stains fairly densely with orange G. The shape of the 
individual fragments suggests that they are single, hardened cells, 
but there are no signs of nuclear remains in the mature material. 
The histological structure of the lining and of the lumen contents 
varies as one follows down the length of the gland. 

At about the region where the gland turns rather sharply forward 
(PI. 5, fig. 2). the lumen is normally filled with mature material as 
described above and the lining shows the histological picture seen 
in Plate 6, figure 1 . There is a cuboidal stratum germinativum and 
above it 2 to 3 layers of loosely arranged, slightly flattened cells. 
There is no indication that new cells, contributing to the actual pore 
material, arise from this region. 

About half-way down the length of the pore, immature glandular 
material may be seen (PI. 6, fig. 2). This stains deeply with orange 
G, individual cells can be recognized, and some contain pycnotic 
nuclei. There is no obvious separation between these cells and the 
mature material described above; they merge imperceptibly into 
one another. 

In the anterior portion of the gland the histological picture is 
quite different. Just beyond the region described above there are 
masses of tightly packed cells of irregular polygonal shape with a 
strongly heterogeneous fibrillar cytoplasm (PI. 6, fig. 3). The 
nuclei are all viable and the cytoplasm appears to contain tightly 
packed fibrillar elements lying in an effectively floccular matrix. 
The fibrils stain intensely with orange G, suggesting a keratinaceous 
nature, while the background is patchy blue, indicating an uptake 
of aniline blue. There is no indication of thickening of the cell 
membranes. The figure suggests that intercellular spaces are com- 
mon; this is an artifact of sectioning, since certain regions show 
this appearance, but in other parts the cells are closely packed. 

At the base of the pore are seen cells which can reasonably be 
interpreted as "mother cells." Their very specific appearance gives 
a clear indication of how the cells described above are formed. 
Lines of columnar cells reach into the pore lumen from the stratum 



14 BREVIORA No. 288 

germinativum. Those nearest the germinal layer (PI. 6, fig. 4) 
show a distinct division into three parts. Nearest the germinal cells, 
the relatively large, ovoid nucleus can be seen. Next, there is a 
region of cytoplasm with a slightly amorphous appearance, some- 
times suggesting irregular droplets or granules; this region stains 
conspicuously with orange G. The distal portion of the cell con- 
tains a large droplet (PI. 6, fig. 4; arrow); this is slightly eosinophi- 
lic but stains particularly intensely with aniline blue. As the 
cells move out into the lumen, and begin to move towards the 
pore mouth, the orientation of the three portions is lost, and the 
distinct droplet soon disappears. One assumes that the patchy blue 
staining visible in cells further up the lumen results from a disinte- 
gration of the large droplets and a spreading of the material 
throughout the cell cytoplasm. 

All the various stages described above were visible at various 
levels wherever a portion of pre-anal gland material had been 
sampled with the overlying scales; there was no indication of any 
cycle of activity. 

DISCUSSION 

There have been comparatively few studies of specialized 
"glandular structures" in lizards (Cole, 1966b). and those works 
which are available are for the most part concerned with follicular 
glands, such as femoral pores (Cole, 1966a). 

Taylor and Leonard (1956) considered the "escutcheon scales" 
of Gonatodes, and I have recently re-examined the problem in the 
light of our present knowledge of the squamate epidermis (Mader- 
son, 1967). The specialized scales of Gonatodes and of Lygodac- 
tyliis are similar in that their pattern of formation is intimately 
associated with the periodic sloughing cycle of the epidermis, but 
here the similarity ends. 

The major differences between the mature and developing gland 
material in the two genera can best be expressed in the following 
table: 

Gonatodes (Maderson, 1967) Lygodactyhis 

1. Specialized scales restricted to 1. Specialized scales seen on pos- 

a V-shape anterior to cloaca. terior abdomen, immediately 

(N. B. In other sphaerodactyline anterior to the cloaca, and on 

forms [Sphaerodactyliis, Thorn- the ventral aspects of the hind 

as and Schwartz, 1966] special- limbs, 
ized scales may be found on the 
femoral region also.) 



1968 



GEKKONID SKIN GLANDS 



15 



Gland material is borne on the 
surface of the /i-layer of the out- 
er epidemal generation and has 
an incomplete Oherliaiitchen 



running beneatli it. 



3. Gland material is derived from 
the basal portion of the epi- 
dermal generation which was 
lost at the previous slough. Thus 
in the "resting phase" there are 
tissues in the epidermis derived 
from two different epidermal 
generations. 



4. Single cell type seen in develop- 
ing gland material. 

5. Staining of keratinaceous ele- 
ments of the gland material sug- 
gests an affinity with the a-layer 

6. At sloughing, splitting zone over 
glandular material is at base 
of the oc -layer of the outer epi- 
dermal generation, leaving a 
portion of the outer epidermal 
generation behind. 



2. Gland material is found //; the 
/i-layer of outer epidermal gen- 
eration with an incomplete 
Oberhaiitcheii partially covering 
it. 

3. Gland material is derived from 
the superficial portions of the 
epidermal generation upon 
which it is borne. As there is 
some doubt that a true "resting 
phase" ever exists, there may 
always be tissues from two dif- 
ferent generations present in the 
epidermis, but for quite diflferent 
reasons (see text, p. II). 

4. Two distinct cell types seen in 
developing gland material. 

5. Staining of keratinaceous ele- 
ments of gland material suggests 
an affinity with the /i-layer. 

6. At sloughing, entire outer epi- 
dermal generation is lost. Al- 
though the splitting zone does 
occur at the base of the a-layer 
of the outer generation over the 
gland material, this is due to a 
partial non-development of the 
lacunar tissue and clear layer of 
the outer generation. 



These differences are so fundamental that I suggest the special- 
ized scales in the two genera are not in any way homologous. 

The problem of the relationship of the specialized scales to the 
pre-anal pores is slightly complicated by the possibility that this 
particular genus has no true resting phase in its sloughing cycle. 
Cole (1966a) suggests that there is a continuation of the normal 
epidermis (referred to in his paper, pp. 125-126, as the "stratum 
corneum") into the femoral gland mouth in Crotaphytus, which 
splits the "plug" into an outer portion which is lost at shedding and 
an inner portion which takes the place of the latter. This would 
imply that glandular activity is directly correlated with the epider- 
mal sloughing cycle. Cole (1966a) also suggests that there is some 
evidence that there is annual cyclical activity of the iguanid femoral 



16 BREVIORA No. 288 

pores. In a study of Gonatodes (Maderson, 1967), I have indi- 
cated that gekkonids typically shed very frequently throughout 
the year. // there is indeed annual cyclical activity of the gland (? 
in association with the reproductive cycle), then one would have 
expected that at some stage in the evolution of true pores or glands, 
their pattern of cyclical protein synthesis would have had to become 
independent of the rest of the epidermis. In Gonatodes (Maderson, 
1967) there are morphological indications of how this might have 
occurred. As far as one can tell in Lygodactylus there is no indica- 
tion of a separation between the completely mature pre-anal pore 
material and the cells still undergoing differentiation, so that the 
pre-anal pores in this genus resemble a tube of tooth-paste from 
which material is slowly squeezed out. It is unprofitable to specu- 
late on this point in the absence of material collected throughout 
the year in an attempt to demonstrate a true annual cycle of activ- 
ity in the pre-anal pores. 

Apart from the problem of the relative activity of the pre-anal 
pores and the /:^-glands, there is the problem of the relationship 
between the two types of structure. Here I would unhesitatingly 
suggest that the /:^-glands represent an ancestral condition of the pre- 
anal pores in this genus (see below). The striking discontinuity of 
the Oberluiutchen has been emphasized. If, in association with gland 
development on the scales, one can see the absence of a superficial 
layer, there is no logical reason to deny the possibility that 
subjacent layers could also be reduced and eventually lost during 
evolution. In fact I have demonstrated that this process has already 
begun in the absence of the lacunar tissue and clear layer from the 
central portion of the /:;-glands. Thus, reference to Plate 5, figure 2 
shows that one only has to imagine that the rest of the epidermal 
generation beneath the developing gland material (presumptive- 
/:^-cells. presumptive mesos cells and presumptive a-cells) could dis- 
appear and one would be left with a "shallow pre-anal pore."' 
Furthermore, the figure shows that if a particular portion of the 
individual scale, i.e. the portion over which the gland material 
forms, invaginated deeper and deeper into the subcutaneous tissues, 
a pre-anal pore type of structure would be developed. The epider- 
mis of the scale upon which the pore opens is notably different 
from the adjacent scales showing no indication of normal gland 
development; this would be homologous with the extreme unmodi- 
fied margins of the /:?-gland scales which show a quite typical 
"normal body epidermis" structure (Maderson, 1966b). A con- 
sideration of probable intergeneric homologies of scales and pores 



1968 GEKKONID SKIN GLANDS 17 

depends on the interpretation of the various component cell types 
which will be discussed next. 

The presence of two distinct cell types in the ^-glands has been 
emphasized. Although the chemistry of epidermal proteins is a 
highly complex field, and consideration thereof is not pertinent to 
an article of this kind, it is permissible to suggest that those cells 
which show a definite affinity for orange G are synthesizing a kera- 
tin, while those showing an affinity for aniline blue are synthesizing 
a mucoprotein of some description. The vertebrate epidermis has 
the capacity to form either keratins or mucins, as evidenced by 
studies on the structure of the amphibian epidermis (Parakkal and 
Matoltsy, 1964), and under certain experimental conditions (New, 
1963). It seems that in Lygodcictylus yS-glands the two functions 
are carried on within different cells. However, there is evidence 
from adjacent scales in this and other genera which suggests that 
Oberluiutchen cells (or at least, cells belonging to the same portion 
of the epidermal generation as the Oberhautchen or presumptive 
;ff-cells) are capable of both activities. Whether these modified 
cells occurring either singly or in a double or triple epithelium 
(PI. 4, fig. 1) should be regarded as "Oberhautchen" cells is a 
semantic problem; one would prefer to retain the term only for 
those cells which show the characteristic development of spinules 
or setae on the outermost membrane (Ruibal and Ernst, 1965; 
Ernst and Ruibal. 1966). Within the pre-anal pores, there is no 
indication of two distinct cell types. There is however evidence of 
localization of mucoprotein synthesis and keratin synthesis within 
a single cell. This then should be regarded as a specific specializa- 
tion of the tendency seen in Oberhautchen cells on the non- 
specialized scales of this and other genera. In the femoral pores of 
Gekko gecko (Maderson, unpublished data) such "mixed-func- 
tion" cells are not seen; here there are cells which only stain 
positively for keratin and cells which only stain positively for 
mucoprotein. A "non-association" with normal epidermal prolif- 
eration is also seen in Gekko gecko, as indicated by the disconti- 
nuity of all the normal layers and regions of the inner epidermal 
generation at the femoral pore mouth, exactly as has been described 
here in Lygodactyhis. Although there is a need for detailed studies 
of the basic anatomy and possible cyclical activity in glands from a 
variety of gekkonid types, it is suggested that the problem of direct 
homology between the pre-anal pores of Lygodactyhis and the 
femoral pores of Gekko gecko will be one of semantics. Granted 
that there are situations where "single-function" and "mixed-func- 
tion" cells may be identified, the fundamental relationship of the 



18 BREVIORA No. 288 

pore cells to the normal epidermal generation is more important. 
Thus I suggest that there is some evidence that lygodactyline ^- 
glands are definitely homologous with the pre-anal pores of that 
genus and with the femoral pores of Gekko gecko — all materials 
deriving from the outermost portions of the epidermal generation 
— but there is no evidence of homology with the type of glandular 
scale seen in the sphaerodactyline gekkonids, where the glandular 
material derives from the innermost portions of an epidermal gen- 
eration. The absence of pre-anal pores from sphaerodactylines 
(Kluge, 1967) makes it impossible to prove or disprove Kluge's 
(1967) statement that the escutcheon scales in these forms are 
"almost certainly modified pre-anal organs" (p. 18). Taylor and 
Leonard (1956) suggested that the escutcheon scales of sphaero- 
dactylines represented structures from which pre-anal organs in 
other groups were derived; the analysis of the cellular components 
in Gomitodes (Maderson. 1967) and the evidence presented here 
suggest that this is unlikely. In the present state of knowledge one 
can only surmise that the specialized scales thus far described in 
sphaerodactylines appear to be the result of an independent evolu- 
tionary pathway from those found in Lygodactylus and Gekko 
gecko. Further comment on the phylogenetic implications of scale, 
pore, and gland structure must await detailed analysis of material 
from a variety of forms. 

SUMMARY 

1. The histological structure of the scalation of the abdominal, 
femoral and tibio-fibular surfaces and pre-anal pores of 29 male 
specimens of Lygodactylus spp. has been studied. 

2. The epidermis of the scales in these regions shows a number 
of histological conditions which can be interpreted as representing 
different stages in the histogenesis of the integument in association 
with periodic sloughing. All the characteristic elements of a typical 
squamate "epidermal generation" are represented. There is pres- 
ent an additional region consisting of a keratinaceous material with 
a mucoprotein component; this material develops in association 
with the /:^-layer of the epidermal generation and lies between the 
Oberluiutchen and the subjacent /i-layer. In the mature state the 
material is partially exposed to the external environment by a 
unique regional non-development of the Oberhautchen. 

3. The pre-anal pores are simple invaginations of single scales 
running forwards in the ventral body wall musculature. There is no 
obvious indication of any cyclical activity, and it would appear that 



1968 GEKKONID SKIN GLANDS 19 

the activity of these glands is quite independent of the sloughing 
cycle of the rest of the body covering. 

4. Comparison of the structure and development of the modi- 
fied scales with what is known of superficially similar "escutcheon 
scales" in sphaerodactyline gekkonids suggests that the two organ 
systems are of quite independent evolutionary origin, there being 
no evidence of any homology between them. To prevent further 
confusion in this context, it is suggested that the organs described 
here in Lygodactylus should henceforth be termed "/:?-glands." 

5. Comparison of the structure of the /^-glands with the pre- 
anal pores suggests that the former present a definite possible an- 
cestral form of the latter in this genus. 

6. The problem of the evolutionary relationships of gekkonid 
glands and pores is briefly discussed. The small amount of avail- 
able evidence suggests the possibility that whereas there is a broad 
basis for assuming an homology between the pre-anal pores in 
Lygodactylus and Gekko gecko, no homology of any description 
can be assumed with the escutcheon scales of sphaerodactylines. 
The basis for assessing homologies of these epidermal structures in 
gekkonid lizards should depend on a consideration of the funda- 
mental relationship of the "gland" material to the epidermal gen- 
eration. 



ACKNOWLEDGMENTS 

I wish to thank Dr. E. E. Williams of the Museum of Compara- 
tive Zoology for introducing me to the problem, for supplying me 
with material, and for reading the draft manuscript. Thanks are 
also due to Mr. A. Greer of Harvard for information regarding the 
life of the animals and the method of collection. Dr. J. Peters of 
the Smithsonian Institution supplied other material. Thanks are 
also due to Miss Joyce Stanganelli for typing the manuscript. Fi- 
nancial assistance for this work was provided by a National Cancer 
Institute Grant No. 5 ROl CA 5401-07 and a Damon Runyon 
Foundation Grant No. DRG947. 



20 BREVIORA No. 288 

LITERATURE CITED 

Bustard. H. B. and P. F. A. Maderson 

1965. Skin-eating in squamate reptiles. Herpetologica, 21: 306-308. 
Chiu, K. W.. J. G. Phillips and P. F. A. Maderson 

1967. Some observations on tlie thyroid control of the sloughing cycle 
in Gekko gecko. J. Endocrinol, 39: 463-472. 
Cole. C. J. 

1966a. Femoral glands of the lizard Crouiphyiii.s collinis. J. Morph. 

118: 119-136. 
1966b. Femoral glands in lizards: A review. Herpetologica, 22: 199- 
206. 
Ernst, V. and R. Ruibal 

1966. The structure and development of the digital lamellae of lizards. 
J. Morph., 120: 233-266. 

Greer, A. 

1967. Observations on the behavior and ecology of two sympatric 
Lygocldctxiiis geckos. Breviora, No. 268: 1-19. 

GURR, E. 

1958. Methods of analytical histology and histochemistry. Leonard 
Hill, London. 327 pp. 
Kluge, a. 

1967. Higher taxonomic categories or gekkonid lizards and their 
evolution. Bull. Amer. Mus. Nat. Hist., 135: 1-60. 

Lillywhite, H. B. and P. F. A. Maderson 

1968. Histological changes in the epidermis of the sub-digital lamellae 
of Anoli.s ciiroHncnsis (Iguanidae) during the sloughing cycle. 
J. Morph. i In press. ) 

LOVERIDGE, A. 

1947. Revision of the African lizards of the family Gekkonidae. Bull. 
Mus. Comp. Zool., 98: 3-469. 
Maderson, P. F. A. 

1964a. Keratinized epidermal derivatives as an aid to climbing in 
gekkonid lizards. Nature (London). 203: 780-781. 

1964b. The skin of snakes and lizards. Brit. J. Herp., 3 (6): 151-154. 

1965a. Histological changes in the epidermis of snakes during the 
sloughing cycle. J. Zool., 146: 98-113. 

1965b. The structure and development of the squamate epidermis. Pp. 
129-153. //;.• The biology of the skin and hair growth. Pro- 
ceedings of a Symposium held in Canberra, Australia, August 
1964. (Eds. A. G. Lyne and B. F. Short), Anus and Robertson, 
Sydney. 

1966a. Some macroscopic and microscopic observations on the foot- 
pads of the tokay {Gekko gecko). Mem. Hong Kong Nat. Hist. 
Soc. 7: 6-10. 

1966b. Histological changes in the epidermis of the tokay {Gekko 
gecko) during the sloughing cycle. J. Morph., 119: 39-50. 



1968 GEKKONID SKIN GLANDS 21 

1967. The histology of the escutcheon scales of Gonatodes, with a 
comment on the squamate sloughing cycle. Copeia, 1967: 
743-752. 
Maderson, p. F. a. and P. Light 

1967. The epidermal morphology and sloughing frequency in normal 
and prolactin injected Anolis caroliiwnsis. J. Morph., 123: 
157-172. 
Miller, M. and M. Kasahara 

1967. Studies on the cutaneous innervation of lizards. Proc. California 
Acad.Sci., 34: 549-568. 
New, D. a. 

1963. Effects of excess vitamin A on cultures of skin and buccal epi- 
thelium of embryonic rat and mouse. Brit. J. Derm., 75: 320- 
325. 

Parakkal, p. F. and A. G. Matoltsy 

1964. A study of the fine structure of the epidermis of Rcina pipiens. 
J. Cell Biol., 20: 85-94. 

Pasteur, G. 

1964. Recherches sur revolution des lygodactyles lezards Afro-mal- 
gaches actuels. Trav. Inst. Scient. Cherif. (Zool.), 29: 5-132. 

Preiss, F. 

1922. Uber Sinnesorgane in der Haut einiger Agamiden. Zugleich ein 
Beitrag zur Phylogenie der Saugetierhaare. Jena. Z. Naturwiss., 
58: 25-76. 
Roth, S. I. and W. A. Jones 

1967. The uitrastructure and enzymatic activity of the boa constrictor 
(Constrictor constrictor) skin during the resting stage. J. Ultr. 
Res., 18: 304-323. 
RuiBAL, R. and V. Ernst 

1965. The structure of the digital setae of lizards. J. Morph., 117: 
271-294. 

Schmidt, W. J. 

1920. Einiges iiber die Hautsinnesorgane der Agamiden, insbesondere 
von Ccilotes, nebst Bermerkungen iiber diese Organe bei Gek- 
koniden und Iguaniden. Anat. Anz., 53: 113-139. 
Taylor, E. H. and A. B. Leonard 

1956. Concerning the relationship of certain neotropical gekkonid 
lizard genera, with comments on the microscopic structure of 
their glandular scales. Univ. Kansas Sci. Bull., 38: 1019-1029. 
Thomas, R. and A. Schwartz 

1966. Sphaerodactylus (Gekkonidae) in the greater Puerto Rico re- 
gion. Bull. Florida State Mus., 10: 193-260. 

(Received 19 July 1967.) 



22 



BREVIORA 



No. 288 



ISO 

OLO 

do 

clo/Obi 

f 

GM 
H 

ISS 

Ito 

m 

mi 

mo 

NGM 

Obi 

Oho 

OSS 

Plii 

py.i 

pclo 

plto 

pmi 
pObi 



ABBREVIATIONS USED IN PLATES 

/;?-layer of inner epidermal generation 
IdAayQV of outer epidermal generation 
a-layer of outer epidermal generation 
Clear layer of outer epidermal generation 
Clear laytv/Ober/iautc/ien complex 
Filaments 
Granular cells 
Exposed glandular material 
Hinge region 
Inner scale surface 

Lacunar tissue of outer epidermal generation 
Melanin granules 

Mesos layer of inner epidermal generation 
Mesos layer of outer epidermal generation 
New glandular material 
Oberhautchen of inner epidermal generation 
Oberhautchen of outer epidermal generation 
Outer scale surface 

Presumptive /:?-layer of inner epidermal generation 
Presumptive a-layer of inner epidermal generation 
Presumptive clear layer of outer epidermal generation 
Presumptive lacunar tissue of outer epidermal genera- 
tion 
Presumptive mesos layer of inner epidermal generation 

Presumptive Oberhautchen of inner epidermal genera- 
tion 

Stratum germinativum 



PLATES 



24 BREVIORA No. 288 



PLATE 1 

Figure 1. Posterior abdominal and sub-caudal surfaces of Lygodcictyliis 
picturatiis male (left) and female (right). Note the darker scales on the 
abdominal, femoral and tibio-fibular surfaces of the male as compared with 
the female. The small square of tissue missing from the abdominal surface 
of the male is seen in figs. 2-4. Scale divisions in mm. 

Figure 2. Low-power view of sagittal section through two successive scales 
from the region shown in fig. 1 . The portion outlined is shown in fig. 4. This 
specimen shows the simplest form of "Condition One" (text pages 5-7). 
Hematoxylin and eosin stain. 

Figure 3. High-power view of transverse section through a more typical 
"jQ-gland" from the abdominal surface showing a later "Condition One" than 
in figs. 2 and 4. The /i-layer and the exposed gland material are absent. 
Hematoxylin and eosin stain. 

Figure 4. Oil immersion montage of region outlined in fig. 2. 



1968 



GEKKONID SKIN GLANDS 



25 




PLATE 1 



26 BREVIORA No. 288 



PLATE 2 

Figure 1. Sagittal section through epidermis of a specialized scale from 
the femoral region showing "Condition Two." Masson's trichrome stain. 

Figure 2. Extreme lateral margin of a specialized scale from the femoral 
region showing "Condition Two" as seen in transverse section. This shows 
the layers of cells which will later be recognizable as the lacunar tissue, the 
clear layer and the Ohcrhaittchen. The /i-layer of the outer epidermal genera- 
tion (represented only by the Oberhaittclien in this region [Ernst and Ruibal, 
1966]) is not shown. Hematoxylin and eosin stain. 

Figure 3. Transverse section through a specialized scale from the abdom- 
inal surface showing an early "Condition Three." The /i-layer of the outer 
epidermal generation and the exposed gland material are not shown. The 
region enclosed by the rectangle is shown in PI. 3, fig. 1. Hematoxylin and 
eosin stain. 

Figure 4. Transverse section through a specialized scale from the abdom- 
inal surface showing a mid-late "Condition Three." The /i-layer of the outer 
epidermal generation is not shown. The region enclosed by rectangle marked 
A' is shown in PI. 3, fig. 2, and that marked Z is shown in PI. 4, fig. 3. Aniline 
blue-orange G stain. 

Figure 5. Median sagittal section through a specialized scale from the 
abdominal surface showing a very late "Condition Three" or early "Condi- 
tion Four." The /:i-layer of the outer epidermal generation is not shown. A 
region comparable to that enclosed by the rectangle is show in Pi. 3, fig. 3. 
Aniline blue-oran<je G stain. 



1968 



GEKKONID SKIN GLANDS 



27 







PLATE 



28 BREVIORA No. 288 



PLATE 3 

Figure 1. Oil montage of region indicated by rectangle in PI. 2, fig. 3. 
Note the conspicuous filaments (/) in the more numerous cell type in the 
new gland material {NGM) and the occasional cell with granules (X). The 
outermost portion of the new gland material abuts directly onto the base of 
the a-layer of the outer epidermal generation (not shown). 

Figure 2. Region comparable to that indicated by rectangle marked X in 
PI. 2, fig. 4. Note the pycnotic nuclei visible in the new gland material and 
the sharp discontinuity of the Oberhautchen of the inner epidermal genera- 
tion marked by an arrow. 

Figure 3. Region comparable to that indicated by a rectangle in PI. 2, 
fig. 5. The clear layer of the outer epidermal generation (do) is recogniz- 
able by pycnotic nuclei. The sharp discontinuity of the new Oberhautchen 
is indicated by an arrow. Compare with PI. 4, fig. 2. 



1968 



GEKKONID SKIN GLANDS 



29 




MGM 



X 



^ 







PLATE 3 



30 BREVIORA No. 288 



PLATE 4 

Figure 1. Micrograph of epidermis from an adjacent "semi-specialized 
scale" showing a modification of a "Stage Four-Five" (Maderson, 1966b) 
with the addition of two layers of granular cells (gc) between the Oberhaiit- 
cheii {Obi) and the rest of the presumptive /i-layer {p^i). Aniline blue- 
orange G stain. 

Figure 2. Similar region of a specialized scale, as seen in sagittal section, 
to that shown in PI. 3, fig. 3. This specimen shows a slightly more advanced 
stage of histogenesis of the new /i-Iayer. It is readily comparable to the 
condition shown in figure 1 above. Note that it is possible to see where 
the spinules of the new Oherliaiiiclien have separated from the keratinized 
clear layer of the outer epidermal generation. Aniline blue-orange G stain. 

Figure 3. Similar region to that indicated by rectangle marked Z in PI. 2, 
fig. 4. Hematoxylin and eosin stain. 

Figure 4. Similar region to that shown in fig. 2 in an early "Condition 
Four." Note the chromophobic appearance of the stratum germinativum 
in this figure and figs. 1 and 2; this feature is characteristic of the period 
before, during, and after the new mesos layer is laid down. Hematoxylin 
and eosin stain.. 



1968 



GEKKONID SKIN GLANDS 



31 








.i 








PLATE 4 



32 BREVIORA No. 288 



PLATE 5 

Figure 1. Extreme lateral margin of specialized scale from abdominal 
region, seen in transverse section, showing an immediate pre-slough condi- 
tion (late "Condition Four"). The /;-i-layer of the outer epidermal genera- 
tion is absent from the photograph. The new Ohcrluiiitchen separates from 
the clear layer to facilitate sloughing, to the left of the picture. In this 
region there is a lacunar tissue (Ito) above the clear layer. To the right 
of the picture, there is no development of the lacunar tissue, clear layer, or 
Oherluutlchen so that the a-layer of the outer epidermal generation (ao) 
separates directly from the new inner epidermal generation. This material 
will be exposed when sloughing occurs. The mesos layer (ino) of the inner 
epidermal generation has now keratinized and resembles the similar com- 
ponent of the outer generation. The stratum germinativum cells are regain- 
ing their chromophilic properties as the first presumptive a-cells laid down 
ipcci). Hemato.xylin and eosin stain. 

Figure 2. Plan view of a sagittal section through the scale on which the 
pre-anal pore opens. The rectangles marked a, h, c, d are shown in figs. 3 
and 4 and PI. 6, figs. 1 and 2. respectively. Note that the scales anterior 
and posterior to the scale on which the pore opens show a late "Condition 
Three" of the /i-glands, but the scale itself is unspecialized (see figs. 3 and 
4). H indicates direction toward head, and T indicates direction towards 
tail. Hematoxylin and eosin stain. 

Figure 3. Region shown in rectangle a in fig. 2. This shows the epidermis 
in a typical Stage Four condition (Maderson, 1966b). Note the sharp dis- 
continuity of the development of the complex epidermal generation in the 
mouth of the pore as shown by the arrow. The fine vertical striations 
indicating the old (Oho) and new (Obi) Oherhautclien spinules can just be 
seen. Several fragments of the pre-anal pore material can be seen lying in 
displaced positions. The lumen of the pore in this and fig. 4 (below) is 
indicated by L. 

Figure 4. Region shown in rectangle b in fig. 2. The epidermis here 
shows the modified Stage Four condition (Maderson, 1966b) with a greatly 
reduced jS-layer which characterizes the inner scale surface. Note the 
much-enlarged clear layer cells and the enclosed granules which may be 
keratohyalin (see discussion, Maderson, 1966b). Again the sharp discon- 
tinuity of the epidermal generation in the pore mouth can be seen (arrow). 



1968 



GEKKONID SKIN GLANDS 



33 







^^^JO^ 




^*C 'il4 



PLATE 5 



34 BREVIORA No. 288 



PLATE 6 

Figure 1. Region indicated by rectangle c in PI. 5, fig. 2. L indicates the 
lumen of the pore and D the dermis. Note the complete absence of any 
indication of "epidermal generation" structure. 

Figure 2. Region indicated by rectangle (/ in PI. 5, fig. 2. This shows 
the cellular elements comprising the pre-anal pore secretion which are al- 
most fully mature and keratinized. Two pycnotic nuclei are indicated by 
arrows. 

Figure 3. Cells in the anterior region of the forward-running pre-anal 
pore where the cells are immature. Note the extremely conspicuous intra- 
cellular fibrils. Aniline blue-orange G stain. 

Figure 4. Cells in the extreme anterior portion of the pre-anal pore; this 
region is apparently the "germinal region." the site of origin of all the pore 
secretion. As the cells arise from the germinal layer (sg) they show three 
conspicuous features. The nucleus lies towards the germinal layer. There 
is a central region where fibrils are just visible (/) which stain intensely 
with orange G. There is a distal region which is occupied by a large 
droplet (Dr) which stains with aniline blue. As the cells move away from 
the anterior part of the gland, the droplet disintegrates, giving the picture 
seen in fig. 3. Aniline blue-orange G stain. 



1968 



GEKKONID SKIN GLANDS 



35 



© 




"^f D 



1^' 



y 




PLATE 6 



BREVIORA 



Musemm of Comparative Zoology 



Cambridge, Mass. 29 May. 1968 Number 289 

THE GENUS MIAGRAMMOPES (ARANEAE, ULOBORIDAE) 
IN PANAMA AND THE WEST INDIES 

Arthur M. Chickering 



The collection of members of this genus of spiders has been of in- 
terest to me for many years. As a result, I have accumulated a 
considerable number of specimens of both sexes from Panama and 
the West Indies. This seems to be a convenient time to bring to- 
gether the results of this field work extending over a period of more 
than thirty-five years. 

Up to the present time, so far as I have been able to determine, 
only one species of the genus has been definitely known from Pan- 
ama. Mr. Banks (1929) reported the collection of M. alboguttatus 
O. P. - Cambridge on Barro Colorado Island, Canal Zone, but the 
specimens so identified plainly belong to M. simiis Chamberlin and 
Ivie. 1936. At present I am obliged to recognize six species of this 
genus from Panama. Among these are four kinds of males and four 
kinds of females. Although it is difficult to match males and females, 
I seem to be able to do this for two of the species, but four species 
will remain known only from a single sex until the genus can be 
studied more intensively. 

In the West Indies, up to the present time, four species belonging 
to this genus have been definitely recorded. Franganillo reported 
M. thwaitesii O. P. - Cambridge from Cuba in 1936 and earlier but 
this identification has been shown to have been an error. For this 
reason this species, described originally from Ceylon, will not be 
further considered in this paper. At the present time I am obliged 
to recognize a total of nine species from the West Indies. Five are 
described as new to science; three of these are known only from 
males and the remaining two only from females. 

All types, together with my entire collection of this genus, are 
being deposited in the Museum of Comparative Zoology, Harvard 
University. 



2 BREVIORA No. 289 

Grant No GB-1801 from the National Science Foundation made 
it possible for me to spend seven months collecting spiders in the 
West Indies and Panama during the latter part of 1963 and the first 
five months of 1964. Grant GB-5013 from the NSF made it possible 
for me to spend nearly four months during 1966 collecting on nine 
diflferent West Indian islands. During this time, however, I did not 
take a single mature specimen of Miagrammopcs. As I have fre- 
quently stated in my published papers, I am deeply grateful for 
financial help from several foundations, together with help and 
encouragement from members of the staff of the Museum of Com- 
parative Zoology at Harvard University over a period of many years. 
Special acknowledgements should be extended to Dr. Ernst Mayr, 
Director; Dr. P. J. Darlington, Jr., Alexander Agassiz Professor of 
Zoology; Dr. Herbert W. Levi, Associate Curator of Arachnology; 
Miss Nelda Wright, Editor of Publications; and Dr. Frank M. 
Carpenter, Alexander Agassiz Professor of Zoology and Editor of 
Psyche. My thanks are extended to Dr. G. Owen Evans and Mr. D. 
J. Clark, British Museum (Natural History), for the loan of very 
helpful specimens of M. scoparius Simon, 

Genus Miagrammopes O. P. - Cambridge, 1870 

The type species of the genus is Miagrammopes thwaitesii O. 
P. -Cambridge, 1870. The holotype is a female and is now defi- 
nitely known only from Ceylon and, possibly, from India. The male 
remains unknown. Apparently, the P. - Cambridges had only fe- 
males for the nine species described by them. The genus is now 
known to be widely distributed in Asia, Africa, Australia, and es- 
pecially in the Neotropical region. Roewer, 1954, lists twenty-one 
species from the Neotropical region. Twelve of these are known 
only from females, two only from males, six from both sexes, and 
one from an immature specimen. 

In addition to the general family characteristics, generic features 
common to the species from Panama and the West Indies, studied in 
the preparation of this paper, may be stated as follows: Cephalo- 
thorax in general rather low; carapace often with marked grooves 
and pits in the cephalic area; sternum often conspicuously sub- 
divided and laterally extended between second and third coxae. 
Eyes: small; only four in number; with laterals on more or less 
prominent tubercles. Chelicerae of modest size, probably typically 
with few true teeth but with cusps and associated long hairs or 
bristles along margins of fang groove. Legs: 1423 in order of length 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 3 

with first legs very long and quite robust; in females the first meta- 
tarsi and, to some extent, the first tibiae bear conspicuous fringes of 
hair; in males the first tibiae have more or less prominent batteries 
of robust spines; in females the fourth metatarsi and tarsi bear 
robust spines; the calamistrum in females extends from two-thirds 
to three-quarters of entire length of fourth metatarsi; in both sexes 
the femora appear to be well supplied with long trichobothria. In 
females the palpal tarsal claw is conspicuously toothed; in males the 
palpi are short and simple, with exception of the tarsus, which is 
moderately complicated. In females the abdomen is usually more 
or less swollen dorsally; the epigynum is often obscurely patterned. 
The complete list of species of the genus Miagrammopes from 
Panama, as they are now recognized, may be given as follows: 
Miagrammopes aspinatiis sp. nov.; M. intempiis sp. nov.; M. larun- 
dus sp. nov.; M. licinus sp. nov.; M. simiis Chamberlin and Ivie; M. 
unipus sp. nov. The complete list of species of this genus from the 
West Indies, as they are now recognized, may be given as follows: 
M. animotus sp. nov. from Puerto Rico; M. ciliatus Petrunkevitch 
from St. Thomas and. possibly, also from Puerto Rico; M. cubanus 
Banks from Cuba; M. latens Bryant from Cuba and Dominican Re- 
public; M. molitus sp. nov. from Jamaica; M. obliicus sp. nov. from 
Jamaica; M. pinopiis sp. nov. from St. John; M. scoparius Simon 
from St. Vincent; M. tonatiis sp. nov. from Jamaica. 

Key to tlie males of known species of Miagrammopes from 
Panama and the West Indies 

la. Species in which the carapace is nearly or quite as wide as long 
(aspinatiis, simiis, unipus) 2 

lb. Species in which the carapace is clearly longer than wide (ani- 
motus, latens, licinus, molitus, scoparius, tonatus) 4 

2a. Palpal tarsus with three prominent, terminal apophyses (Fig. 
10) M. aspinatiis 

2b. Palpal tarsus lacking the three prominent terminal apophyses 
(simus, unipus) 3 

3a. First tibia armed with a battery of about 21-22 robust, erect spines 
extending nearly the entire length of the segment (Fig. 43). M. simus 

3 b. First tibia with fewer and weaker spines extending through about 
the distal two-thirds of the segment (Fig. 50) M. unipus 

4a. Carapace about twice as wide at level of lateral eyes as at posterior 
end where is it squarely truncate (Fig. 24) M. molitus 

4b. Carapace less than twice as wide at level of lateral eyes as at pos- 
terior end (animotus, latens, licinus, scoparius, tonatus) 5 

5a. Palpal tibia concave dorsally; first tibia with very few spines as 
seen in retrolateral view (Figs. 20-23) M. licinus 



4 BREVIORA No. 289 

5b. Species lacking such a combination of features as given above 
(animotus, latens, scoparius, tonatiis) 6 

6a. Carapace nearly rectangular in outline in posterior two-thirds; 
first tibia with few long spines showing in retrolateral view (Figs. 
45, 46) M. tonatiis 

6b. Species without combination of features given above {animotus, 

latens, scoparius) 7 

7a. Carapace with a broad, white, median stripe throughout; palpal 
tarsus with a pair of robust, distal apophyses; first tibia with nu- 
merous dorsal spines (Figs. 1, 2, 5) M. animotus 

7b. Species lacking such a combination of features as given above 
{latens, scoparius) 8 

8a. Palpal tarsus terminating in a single robust apophysis together with 
a series of small, sharply pointed spines (Bryant, 1936, fig. 4) 
M. latens 

8b. Palpal tarsus with a distal apophysis divided into a robust hook and 
a slender, sickle-shaped extension (Fig. 36) M. scoparius 

MlAGRAMMOPES ANIMOTUS Sp. nOV. 

Figures 1-5 

Holotype. The male is from Mayaguez, Puerto Rico, W. I., in a 
wooded area near Nuclear Center, January 27, 1964. The name of 
the species is an arbitrary combination of letters. 

Description. Total length from anterior border of carapace to 
posterior end of spinnerets about 2.8 mm (body curled). Carapace 
0.97 mm long; 0.71 mm wide at level of lateral ocular tubercles 
where it is widest; with little hair; only a slight depression to repre- 
sent the median thoracic fovea; grooves and depressions in ocular 
region, so conspicuous in some species, only slightly developed 
here; lateral ocular tubercles also weakly developed (Fig. 1 ). Eyes: 
only four, in a gently recurved row as seen from above; ratio of eyes 
ME : LE = about 6:7; ME separated from one another by about 
4.6 times their diameter, from LE by a little less than three times 
their diameter; with very little black pigment in ocular region. 
Chelicerae, maxillae, and lip difficult to observe closely but ap- 
parently typical of males of the genus. Sternum: very much swollen 
in middle and greatly expanded between second and third coxae; in 
general, typical of males of the genus. Legs: tibial index of first leg 
12, of fourth leg 10; first leg with a conspicuous battery of stout 
spines on tibia and metatarsus (Figs. 2-3); first leg somewhat more 
than twice as long as second leg and a little more than three times 
as long as third leg. Calamistrum lacking. Palp: essential features 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 




i/mmi 




Figures 1-5. Miagrammopes animotiis sp. nov. Fig. 1. Carapace; from 
above. Fig. 2. Right first tibia; retrolateral view. Fig. 3. Right first meta- 
tarsus; dorsal view (several spines restored from scars). Figs. 4-5. Male 
palp; retrolateral and prolateral views, respectively. 



shown in Figures 4-5. Abdomen without special features; appar- 
ently quite typical of males of the genus. Color in alcohol: carapace 
with a broad, median, white stripe covering the median third from 
clypeus to posterior border; lateral thirds of the carapace brownish 
gray with black, irregular, short lines; sternum yellowish with ir- 
regular grayish spots; legs generally yellowish but first legs are 
brown dorsally; abdomen with nearly the whole dorsum light 
yellowish with white flecks; a pair of minute black dots occurs on 
the dorsum a little in front of the middle and another pair of black 
dots occurs a little behind the middle of the dorsum; a black, lateral 
stripe on each side extends the whole length of the abdomen; the 
central part of the venter is very light brownish and the remainder is 
nearly the same but with many whitish flecks. 

Records. Two immature specimens taken on the same day as the 
holotype and in the same locality are tentatively placed in this 
species, but their identification must remain uncertain. 



6 BREVIORA No. 289 

MlAGRAMMOPES ASPINATUS Sp. nOV. 

Figures 6-10 

Holotype. The male is from Barro Colorado Island, Panama 
Canal Zone, June, 1950. The name of the species is an arbitrary 
combination of letters. 

Description. Total length from anterior border of carapace to 
posterior end of extended spinnerets about 3.82 mm. Carapace 
about 1.29 mm long; 1.21 mm wide at level of lateral ocular 
tubercles where it is widest; about 0.57 mm tall; quite hairy; with 
moderately well developed median thoracic fovea, pits and grooves 
in ocular regions and lateral ocular tubercles (Fig. 6). Eyes: only 
four, as usual in the genus; with the row slightly recurved as seen 
from above; ratio of eyes ME : LE = 6 : 7.5; ME separated from 
one another by about nine times their diameter, from LE by about 
five times their diameter. Chelicerae, maxillae and lip apparently 
typical of males of the genus but details not clearly observed be- 
cause of fragility of holotype and closely coiled appendages. Ster- 
num: quite typical in general; quite convex in middle but less so 
than in some species; much narrowed at posterior end and, appar- 
ently, continued by an extra, very narrow sclerite between fourth 
coxae, which are separated by slightly more than one-fifth of their 
width. Legs: tibial index of first leg 11, of fourth leg 12; first leg 
nearly twice as long as second leg; nearly three times as long as 
third leg; first tibia with a conspicuous battery of robust spines 
much as in M. simus Chamberlin and Ivie; fourth tibia also with 
numerous long, stout spines (Figs. 7-8). Palp: essential features 
shown in Figures 9-10; tibia with quite distinctive form. Abdomen: 
typical of males of the genus. Color in alcohol: yellowish through- 
out with some variation; only a moderate amount of black pigment 
in ocular region; abdomen with three pairs of minute dots on dor- 
sum equally spaced from the base; legs with few gray areas. 

Records. The female is unknown. Another specimen of the 
genus was taken at the same time and in the same locality as the 
holotype, but it is immature and its identification is very uncertain. 

MlAGRAMMOPES ciLiATUS Petruukevitch 

Miagrammopes ciliatus Petrunkevitch, 1926: 47, fig. 7. The present loca- 
tion of the holotype is quite uncertain. Petrunkevitch, 1930: 232; 
Roewer, 1954: 1349; Bonnet, 1957: 2830. 
The holotype of this species was a female from St. Thomas, U. S. 
Virgin Islands, collected in July, 1915. No description of the epigy- 
num was given. Dr. Petrunkevitch later had several specimens of 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 7 

what he assumed to be the same species from Puerto Rico. Among 
these were three females regarded as mature and several immature 
specimens of both sexes. I think it unlikely that the Puerto Rican 
specimens belong to M. ciliatus Petrunkevitch but I cannot be posi- 
tive at this time. In my collection from Puerto Rico, made in 
January and February, 1964, I have a single mature male and sev- 
eral immature specimens but not a single mature female. The 
mature male is being described as a new species elsewhere in this 
paper. 




Figures 6-10. Miagrammopes aspinatus sp. nov. Fig. 6. Carapace; from 
above. Fig. 7. Right first tibia; dorso-retrolateral view. Fig. 8. Right fourth 
tibia; prolateral view. Figs. 9-10. Male palp; prolateral and retrolateral 
views, respectively. 



8 BREVIORA No. 289 

MiAGRAMMOPES CUBANUS Banks 

Miagrammopes cuhaniis Banks. 1909: 159. The holotype is in the Mu- 
seum of Comparative Zoology. Bryant, 1940: 329; Roewer, 1954: 
1349; Bonnet, 1957: 2830. 
The holotype was very briefly described and no figures were 
published. The holotype is immature and its status cannot be deter- 
mined until careful collecting is pursued in the type locality. Miss 
Bryant (1940) stated that M. latens Bryant might prove to be a 
synonym for M. ciibanus Banks. A few specimens assigned to M. 
cubanus Banks since the holotype was described are plainly M. 
latens Bryant. 

Miagrammopes intempus sp. nov. 
Figures 11-13 

Holotype. The female is from Porto Bello, Panama, August 11, 
1936. The name of the species is an arbitrary combination of 
letters. 

Description. Total length from anterior border of porrect cheli- 
cerae to posterior end of spinnerets 6.24 mm; length from anterior 
border of carapace to posterior end of anal tubercle 5.85 mm. Cara- 
pace 1.54 mm long; 1.08 mm wide at level of lateral ocular tu- 
bercles where it is widest; grooved behind ME in somewhat dis- 
tinctive manner; with a poorly defined median thoracic fovea (Fig. 
11); about 0.55 mm tall; lateral ocular tubercles less prominent 
than in some other species; with many hairs. Eyes: four as usual, in 
a slightly recurved row as seen from above; ratio of eyes ME : LE 
^ 6.5 : 8; ME separated from one another by about eight times 
their diameter, from LE by nearly 3.5 times their diameter. 
Chelicerae, maxillae and lip apparently quite typical of females 
of the genus. Sternum: conspicuously distinctive; anterior por- 
tion sharply narrowed from median portion which is greatly 
widened and extended between second and third coxae thus 
forcing first and second legs forward and third and fourth legs back- 
ward; between third and fourth coxae there is a transverse shelf and 
behind this a greatly depressed and narrowed portion (Fig. 12); in 
front of third coxae are loose, membranous structures and just be- 
hind second coxae are smaller, similar structures; fourth coxae 
separated by slightly less than one-fourth their width. Legs: tibial 
index of first leg 13, of fourth leg 10; first leg with copious hair 
fringes on tibia and metatarsus; first leg nearly twice as long as 
second leg and about 2.5 times as long as third leg; fourth tibia a 
little longer than first tibia; second and third coxae very widely 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 




Figures 11-13. Miagrammopes intempus sp. nov. Fig. 11. Carapace; 
dorsal view. Fig. 12. Sternum, seen from below. Fig. 13. Epigynum, seen 
from below. 



separated as already noted; only occasional slender spines on palps 
and legs with exception of fourth tarsi and metatarsi where special, 
short, robust spines occur similar to those figured for M. simiis 
Chamberlin and Ivie. Black palpal claw quite conspicuous. Cal- 
amistrum about three-fourths as long as fourth metatarsus. Abdo- 
men with a conspicuous dorsal hump a little in front of the middle; 
otherwise apparently quite typical of females of the genus. Epigy- 
num: essentials shown in Figure 13. Color in alcohol: carapace 
light brownish in general but with many darker streaks; lighter 
through the center and darker along lateral sides; sternum light 
yellowish brown; legs brownish in general with first pair darkest on 
dorsal surfaces and light brownish below; abdomen with a dark 
median stripe widest in anterior third then narrowed at top of dorsal 
hump and widened again at posterior end; each side of the central 
dark stripe there is a lighter, finely reticulated area; these reticulated 
areas continue laterally and nearly throughout the venter; a grayish 
area on each lateral side of the abdomen extends backward from the 
base to nearly opposite the dorsal hump. 

Records. The male is not known and there are no female para- 
types. 



10 



BREVIORA 



No. 289 



MlAGRAMMOPES LARUNDUS Sp. nOV. 

Figures 14-17 

Holotype. The female is from Barro Colorado Island, Panama 
Canal Zone, August, 1950. The name of the species is an arbitrary 
combination of letters. 

Description. Total length from anterior border of extended cheli- 
cerae to posterior end of spinnerets 4.92 mm. Carapace 1.23 mm 
long; 1.32 mm wide at level of lateral ocular tubercles where it is 
widest; 0.45 mm tall at level of eyes where it is tallest; with grooves 
and depressions well developed (Fig. 14); depressions medial to 
ME deep and conspicuous; division and course of grooves around 
ocular regions somewhat complicated; median thoracic fovea hardly 
discernible; with a fairly well developed coat of short hair directed 
forward. Eyes: four as usual, in a slightly procurved row as seen 
from above; ratio of eyes ME : LE = 10 : 11; ME separated from 




Mlfc ' |:f ill 
0§^' Mill .v^i^>'i 



f' 



mmiux 




Figures. 14-17. Miograinmopes lariindiis sp. nov. Fig. 14. Carapace, 
dorsal view. Fig. 15. Maxillae and lip. Fig. 16. Sternum, seen from be- 
low. Fig. 17. Epigynum, seen from below. Fig. 18. Miagrammopes latens 
Bryant. Epigynum, seen from below. 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 11 

one another by nearly nine times their diameter; separated from LE 
by about 1.3 times their diameter; with black pigment nearly cover- 
ing the ocular tubercles. Chelicerae: apparently typical of females 
of the genus. Maxillae, lip and sternum with features as shown in 
Figures 15-16. Legs: tibial index of first leg 10, of fourth leg 9; 
calamistrum as usual in females of the genus; fourth metatarsus and 
tarsus with the short, stout spines similar to those figured for M. 
simus Chamberlin and Ivie; first leg nearly twice as long as second 
leg and nearly three times as long as third leg. Abdomen without a 
dorsal, medial swelling such as frequently occurs in females of the 
genus; otherwise typical of females of the genus and without note- 
worthy features. Epigynum: obscurely distinctive; observable 
features shown in Figure 17. Color in alcohol: carapace with most 
of dorsum light yellowish brown but with numerous gray streaks 
and spots; lateral areas gray with considerable black pigment in 
ocular regions; sternum light grayish with a yellowish median 
stripe; mouth parts generally yellowish with a few grayish spots; 
legs generally brownish. Abdomen with a conglomerate color pat- 
tern; anterior two-thirds of dorsum yellowish, irregularly reticulated 
and with two pairs of irregularly shaped darker, oblique spots; pos- 
terior end of abdomen irregularly brown; lateral sides with several 
long, narrow, alternating, irregular light and dark lines; venter 
brownish with a series of stripes leading from genital groove to near 
the posterior end; there is a narrow, indefinhe, brownish, central 
stripe, a pair of pale, narrow stripes, a pair of broader brownish 
stripes, and a pair of light lines most lateral of all. 

Records. A female collected on May 5, 1964, in the type local- 
ity appears to be a paratype; an immature male and an immature 
female also collected in the type locality at about the same time 
probably belong to this species. 

MiAGRAMMOPES LATENS Bryant 
Figure 18 

Miagrammopes latens Bryant, 1936: 326, pi. 23, fig. 4. The holotype male 

is in the Museum of Comparative Zoology; it was taken in Cuba, 

Sierra del Cobre, Loma del Gato. Bryant, 1940: 330; 1948: 393; 

Roewer, 1954: 1350; Bonnet, 1957: 2831. 

The holotype male is from the mountains of Cuba; the palp is 

still in good condition and well represented by Miss Bryant's plate 

23, figure 4. I have not been able to find the specimen regarded by 

Miss Bryant as the allotype but I have studied a female from the 

same locahty and, apparently, taken with the allotype. My Figure 



12 BREVIORA No. 289 

18 is taken from this specimen. I see the epigynum somewhat dif- 
ferently than represented by Miss Bryant's figure 98 (1940) ; I have 
noted, however, considerable variation in the appearance of this 
organ among available specimens assigned to this species. The 
matter will probably not be clarified until numerous specimens are 
available for study following extensive collecting in the regions 
where the species occurs. 

MlAGRAMMOPES LICINUS Sp. nOV. 

Figures 19-23 

Holotype. The male is from El Volcan, Panama, August 14, 
1950. The name of the species is an arbitrary combination of 
letters. 

Description. Total length from anterior border of slightly porrect 
chelicerae to posterior end of spinnerets 3.32 mm. Carapace 1.19 
mm long; 0.84 mm wide at level of ocular tubercles; 0.48 mm tall; 
ocular tubercles quite well delineated by grooves; with a fairly well 
defined median thoracic fovea; with a pair of depressions shortly 
behind ME; with a fairly well developed coat of hair which appears 
whitish in the light used for description (Fig. 19). Eyes: four as 
usual, in a gently recurved row as seen from above; ME separated 
from one another by seven times their diameter, from LE by slight- 
ly more than four times their diameter; ratio of eyes ME : LE = 
5 : 7. Chelicerae, maxillae and lip: all apparently typical of males 
of the genus. Sternum: extremely convex in the middle; fourth 
coxae contiguous. Legs: tibial index of first leg 12, of fourth leg 
13; first tibia with few spines (Fig. 20) and with different numbers 
on right and left; first metatarsus with spines as shown in Figure 20; 
first leg slightly more than twice as long as second leg and nearly 
three times as long as third leg; the calamistrum is lacking but the 
fourth metatarsus and tarsus bear several robust, ventral spines 
similar to those which occur on females; tarsal claws as usual. Palp: 
general features shown in Figures 21-23; the tibia is very distinctive 
with its dorsal expansion and concavity; copiously supplied with 
slender spines; tarsus complicated with its parts difficult to dis- 
tinguish. Abdomen: without special features; apparently typical of 
males of the genus. Color in alcohol; carapace with a yellowish. 
median, longitudinal stripe about one-third as wide as this part of 
the body; this yellowish stripe contains a narrow, grayish stripe 
which widens near the middle into an irregular diamond-shaped 
figure; a broad dark gray stripe covers the remainder of the cara- 
pace on each side; sternum yellowish; abdomen with a fairly broad, 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 



13 



grayish, central, dorsal stripe indented along two-thirds of its length 
and then irregularly broken into a series of five or six somewhat 
separate spots; on each side of this central stripe there is a yellowish 
stripe; lateral sides of abdomen irregularly dark grayish; the venter 
has a grayish central stripe and on each side a narrow, light yellow- 
ish stripe and a narrow, darker stripe; all five of these ventral stripes 
are quite inconspicuous; legs in general yellowish; first pair with 
considerable dark gray coloration on dorsal and prolateral surfaces. 
Records. The female is unknown and there are no paratypes. 




Figures 19-23. Miagrammopes licinus sp. nov. Fig. 19. Carapace, 
dorsal view. Fig. 20. Right first tibia and metatarsus; retrolateral view. 
Figs. 21-22. Left palp; prolateral and retrolateral views, respectively. 
Fig. 23. Left palpal tibia; dorsal view. 



14 



BREVIORA 



No. 289 



MlAGRAMMOPES MOLITUS Sp. nOV. 

Figures 24-28 

Holotype. The male is from Jamaica, W. I., Portland Parish, 
Hardwar Gap, October 2, 1957. The name of the species is an 
arbitrary combination of letters. 

Description. Total length about 3.15 mm (body curled closely). 
Carapace 1.17 mm long; 0.81 mm wide at level of lateral ocular 
tubercles; about 0.44 mm tall at level of median eyes where it is 
tallest; with median thoracic fovea barely indicated; with grooves in 
ocular region very obscure (Fig. 24). Eyes: four as usual with row 
gently recurved; ratio of eyes ME : LE ■=6:1; ME separated from 




Figures 24-28. Miagrammopes molitiis sp. nov. Fig. 24. Carapace, 
dorsal view. Fig. 25. Left first tibia and patella; dorsal view. Fig. 26. 
Left first metatarsus; prolateral view. Figs. 27-28. Left male palp: pro- 
lateral and retrolateral views, respectively. 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 15 

one another by slightly more than five times their diameter, from 
LE by about 3.3 times their diameter; with only a moderate amount 
of black pigment around eyes. Chelicerae, maxillae, lip and sternum 
impossible to observe clearly because of sharply curled position of 
holotype and its fragility. Legs: tibial index of first leg 11, of fourth 
leg 10; first leg with a series of spines on tibia and metatarsus as 
shown in Figures 25-26; these spines are less robust and have a 
different placement than in other species treated in this study; other 
legs with few or no spines but with a copious hair covering; no true 
fringes observed; first legs considerably more than twice as long as 
second legs, three times as long as third legs and 1.7 times as long 
as fourth legs. Palp: complicated; parts difficult to observe because 
of transparency; form essentially as shown in Figures 27-28. Abdo- 
men: apparently quite typical of males of the genus. Color in al- 
cohol; carapace with a broad, yellowish, median stripe extending 
the whole length of this part of the body and occupying the middle 
third; lateral thirds of the carapace grayish brown; sternum yellow- 
ish; legs in general yellowish but first legs somewhat grayish brown 
along dorsal surfaces; abdomen with a median, dorsal, yellowish, 
longitudinal stripe and a narrower grayish stripe on each side; re- 
mainder of the abdomen yellowish with variations. 

Records. The female is unknown; there are no male paratypes; 
an immature specimen was taken in the same locality and on the 
same day as the holotype but its identification is uncertain. 

MlAGRAMMOPES OBLUCUS Sp. nOV. 

Figures 29-30 

Holotype. The female is from Jamaica, Portland Parish, Hard- 
war Gap, November 22, 1957. The name of the species is an 
arbitrary combination of letters. 

Description. Total length from anterior border of somewhat ex- 
tended chelicerae to posterior end of spinnerets about 5.57 mm 
(body considerably curled). Carapace 1.58 mm long; 1.12 mm 
wide at level of lateral ocular tubercles where it is widest; about 
0.55 mm tall; median thoracic fovea barely discernible; with a 
moderately developed coat of hair (Fig. 29). Eyes: four as usual, in 
a gently recurved row, seen from above; ratio of eyes ME : LE = 
8:10; ME separated from one another by nearly 6.25 times their 
diameter, from LE by about 3.5 times their diameter. Chelicerae, 
maxillae and lip apparently quite typical of females of the genus. 
Sternum: essentially as described for M. simus Chamberlin and 
Ivie except that there is an angular distortion at the swollen and 



16 



BREVIORA 



No. 289 




^li'!!# 





33 




Figures 29-30. Miagrammopes obliicus sp. nov. Fig. 29. Carapace; 
from above. Fig. 30. Epigynum; seen from below. Figs. 31-33. Miagram- 
mopes pinopus sp. nov. Fig. 31. Carapace; dorsal view. Fig. 32. Epigynum; 
seen from below. Fig. 33. Abdomen; lateral view, right side. 



expanded median part; posterior end not clearly visible but appears 
to terminate in a narrow, rounded point. Legs: tibial index of first 
leg 12; of fourth leg 8; first leg slightly more than twice as long as 
second leg, nearly three times as long as third leg and about 1.4 
times as long as fourth leg; calamistrum extends over about five- 
sixths of the length of the fourth metatarsus; first metatarsus with a 
definite dorsal fringe; fourth metatarsus and tarsus each with a row 
of short spines on ventrolateral surface essentially as in M. simus 
Chamberlin and I vie (Fig. 40). Abdomen: robust, with a slight 
swelling a little in front of the middle; otherwise quite typical of 
females of the genus as observed in this study. Epigynum: some- 
what obscure; essential observed features shown in Figure 30. Color 
in alcohol: carapace generally grayish brown; with a lighter, median, 
irregular stripe extending the entire length of this part of the body; 
the lateral thirds darker and with a narrow, marginal white stripe. 
Sternum light yellowish brown. Mouth parts yellowish with varia- 
tions; the lip is largely grayish. Legs: yellowish in general; first and 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 17 

second legs with dorsal and dorsolateral surfaces dark grayish 
brown; fourth metatarsus dark dorsally and prolaterally. Abdomen: 
with a central, dorsal, very irregular brownish gray stripe flanked on 
each side by a lighter, irregular stripe; a pair of black dots is present 
on the dorsum somewhat in front of the middle, and another pair of 
black dots occurs just behind the middle; lateral sides irregularly 
brownish; the venter is a kind of light, reddish brown with a narrow, 
central stripe flanked on each side by a narrow lighter stripe; two 
pairs of black dots appear in the posterior half of the venter. 

Records. The male is unknown. One female was taken on the 
same day and in the same general locality as the holotype and is 
tentatively regarded as a paratype, but there are differences which 
may be due to immaturity. 

MlAGRAMMOPES PINOPUS Sp. UOV. 

Figures 31-33 

Holotype. The female is from St. John, U. S. Virgin Islands. 
Centerline Road, 4 miles east of Cruz Bay, March 4, 1964. The 
name of the species is an arbitrary combination of letters. 

Description. Total length from anterior border of extended cheli- 
cerae to posterior end of extended abdomen 6.9 mm. Carapace: 
1.76 mm long; 1.07 mm wide at level of lateral ocular tubercles; 
about 0.55 mm tall; with grooves in ocular region only moderately 
developed; with median thoracic fovea shallow and inconspicuous 
(Fig. 31). Eyes: four as usual, in a gently recurved row; ratio of 
eyes ME : LE = 6.5 : 7; ME separated from one another by about 
seven times their diameter, from LE by slightly more than half that 
distance. Chelicerae and maxillae apparently typical of females of 
the genus. Lip: long, slender, convex; widest just before the middle; 
distal end a fine point. Sternum: long, slender, except for excessive 
width between second and third coxae; narrowest between first and 
second coxae; posterior end very hairy, somewhat tuberculate, 
rounded and ending just opposite bases of fourth coxae, which are 
separated by a little more than one-third of their width. Legs: tibial 
index of first leg 1 1, of fourth leg 7; first leg slightly more than twice 
as long as second leg, somewhat less than three times as long as 
third leg, and about 1.4 times as long as fourth leg; all tarsi with 
claws as usual; calamistrum as usual in females of the genus; fourth 
metatarsus and tarsus with spines essentially as shown in Figure 40 
for M. simiis Chamberlin and Ivie; other spines seem to be lacking 
on legs. Palp: with well developed claw and with about five long, 
slender teeth. Abdomen: moderately raised dorsally; with posterior 



18 BREVIORA No. 289 

end continued into a definite caudal extension above anal tubercle 
(Fig. 33) and spinnerets; otherwise typical of females of the genus. 
Epigynum: essential features shown in Figure 32; quite distinctive; 
resembles that of M. latens Bryant. Color in alcohol: carapace 
grayish brown with many narrow, darker, irregular lines and spots 
and with lateral sides somewhat lighter; sternum yellowish brown; 
legs generally yellowish with first pair gray on dorsal surfaces; 
mouth parts yellowish. Abdomen: light yellowish in general; very 
reticulate; with a grayish brown, broken stripe along the median 
dorsal region and with lateral regions bearing irregular and poorly 
defined, darker, oblique bands; venter highly reticulate and with a 
pair of very narrow, dark stripes extending from epigynum to the 
vicinity of the cribellum. 

Records. The male is unknown; there are no female paratypes. 

MlAGRAMMOPES SCOPARIUS SimOU 

Figures 34-36 

Miagrammopes scopariiis Simon, 1891: 555. The types, male and female, 
are in the British Museum (Natural History) and were taken on St. 
Vincent, B.W.I. Simon, 1892: 220; Petrunkevitch, 1911: 97; 1930: 
232; Lutz, 1915: 79; Roewer, 1954: 1350; Bonnet, 1957: 2831. 

Lutz (1915) reported this species from Puerto Rico, but this is 
now regarded as an error in identification and, apparently, the 
species is known only from the type locality. No figures were pub- 
lished with the original description, but in 1902 Simon published a 
figure of the male palp (fig. 164). Through the courtesy of the 
Department of Zoology, British Museum (Natural History), I have 
been able to examine eight specimens from St. Vincent. Three of 
these are males and one of them has been selected for brief de- 
scription as given below. The five females all appear to be in 
various stages of immaturity and, for this reason, no detailed de- 
scription has been prepared. 

Male. Total length about 3.2 mm (body somewhat curled). 
Carapace 1.1 mm long; 0.79 mm wide at level of LE; general ap- 
pearance as shown in Figure 34. Simon gave length of male as 4.5 
mm. Eyes: four as usual, in a recurved row as seen from above; 
ME separated from one another by 4.6 times their diameter, sep- 
arated from LE by about 2.25 times their diameter; ratio of eyes 
ME : LE r= 7 : 8. Chelicerae, maxillae, lip and sternum appar- 
ently all typical of males of the genus. Legs: 1423 in order of 
length; first leg more than twice as long as second leg, more than 
three times as long as third leg but a little less than twice as long as 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 19 




Figures 34-36. Miagrammopes scopariits Simon. Fig. 34. Carapace of 
male from above. Fig. 35. First right tibia; retrolateral view. Fig. 36. 
Left palpus of male; prolateral view. 



fourth leg. Tibial index of first leg 11, of fourth leg 9. First tibia 
with a conspicuous battery of robust spines (Fig. 35). Calamistrum 
lacking; present in immature male. Palp: essential features shown 
in Figure 36. Abdomen: typical of the genus; without unusual 
features. Color in alcohol: carapace light yellowish brown in gen- 
eral, somewhat lighter in middle of anterior half; whole area ir- 
regularly reticulated with fine dark lines; legs yellowish in general 
with variations; first leg grayish dorsally; abdomen yellowish through 
the center and grayish along lateral sides with venter yellowish. 

Females. As already indicated, the five females accompanying 
the males seem to be immature with no developed epigyna. Their 
general appearance agrees quite well with that of the males. The 
largest female has a total length of about 4.22 mm (Simon gave 
length of female as 6 mm). 

Miagrammopes simus Chamberlin and Ivie 
Figures 37-44 

Miagrammopes simus Chamberlin and Ivie, 1936: 12, pi. 2, figs. 11-12. 

The female holotype from Barro Colorado Island, Panama Canal 

Zone, is probably in the University of Utah. Roewer, 1954: 1350; 

Bonnet, 1957: 2832. 
Miagrammopes alboguttatus, — Banks, 1929: 89. (Not M. alboguttatus 

F. P. -Cambridge.) 



20 BREVIORA No. 289 

I have examined the specimens identified as M. alboguttatus F. 
P. -Cambridge by Mr. Banks and am convinced that they are M. 
simus. Because the original description of the holotype female is so 
brief I have thought it worth while to offer a more detailed descrip- 
tion of a selected female from the locality where I collected the 
original specimen and loaned it to the senior author. It is difficult 
to match males and females in this genus but I believe I have done 
so in this case. A male has also been selected and described in the 
following pages. Males and females are by far the most numerous 
of any species of the genus in my collection. 

Female. Total length from anterior border of somewhat porrect 
chelicerae to posterior end of posterior spinnerets 6.63 mm. Cara- 
pace about 1.76 mm long; 1.41 mm wide at level of lateral ocular 
tubercles where it is widest; with a conspicuous pair of depressions 
slightly behind and somewhat medial to ME; with these depressions 
continued to the lateral margins by grooves in a slightly diagonal 
direction; with width only slightly narrowed to posterior margin; 
medial thoracic fovea short and slightly procurved. Eyes: four as 
usual, in a slightly procurved row, viewed from above; ratio of eyes 
ME : LE = 10 : 13; ME separated from one another by slightly 
less than nine times their diameter, from LE by about 1.6 times 
their diameter. Chelicerae: removal of these organs seems to show 
that the promargin of the fang groove has a single tooth and several 
minute cusps each with a stout, long hair; the retromargin of the 
fang groove appears to have a series of minute cusps each also with 
a stiff hair. Maxillae and lip: apparently quite typical of females of 
the genus (Fig. 37). Palpal tarsus with a robust claw having four 
or five very slender teeth (Fig. 38). Sternum: very angular; swol- 
len near middle where a conspicuous shelf crosses from interval 
between second and third coxae; slopes sharply from high region 
both anteriorly and posteriorly; a second less conspicuous shelf 
crosses the sternum between third and fourth coxae; posterior end 
obscure but not extended between fourth coxae which are separated 
by about one-third of their width (Fig. 39). Legs: tibial index of 
first and fourth legs 10; all tarsi short; first tarsus slightly more than 
one-fifth as long as first metatarsus; all legs with fine hair; first tibia 
with a fringe of fairly long hair; apparently three true claws with 
several spurious claws on each tarsus; spines on fourth metatarsus 
and tarsus as shown in Figure 40; trichobothria appear to be present 
on femora, tibiae and metatarsi; the calamistrum extends over about 
seven-twelfths of length of fourth metatarsus; first leg nearly 3.5 
times as long as third leg. Abdomen: essentially as shown in dorsal 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 21 




Figures 37-44. Miagrcunmopes simiis Chamberlin and Ivie. Fig. 37. 
Female maxillae and lip. Fig. 38. Female left palpal tarsus; prolateral 
view (with aid from dissected specimen). Fig. 39. Female sternum; seen 
from below. Fig. 40. Right fourth metatarsus and tarsus of female; re 
trolateral view. Fig. 41. Epigynum from below. Fig. 42. Carapace of male 
from above. Fig. 43. First tibia of male; retrolateral view (a few spines 
restored from scars). Fig. 44. Left palp of male; retrolateral view. 



22 BREVIORA No. 289 

view in figure 11 of Chamberlin and Ivie, 1936; cribellum entire; 
spinnerets six as usual. Epigynum: somewhat obscured by hair; 
essential features shown in Figure 41 (variations in appearance 
have been noted among numerous specimens assigned to this 
species). Color in alcohol: legs generally brown with variations; 
first pair darkest of all but with tarsus light yellowish brown and 
metatarsus somewhat darker; carapace light yellowish brown 
throughout the dorsal region and with lateral sides dark brown; 
sternum yellowish brown with variations. Abdomen: with a semi- 
transparent cardiac area and white spotted area posterior to cardiac 
region; a pair of irregular, white stripes extend along the dorsum 
nearly the entire length; lateral regions brownish with an irregular 
white spot a little in front of the middle; venter light yellowish brown 
with some variation. Considerable variation in color pattern has 
been noted among the numerous specimens of this species now in 
the collection. 

Male. Total length from anterior border of slightly porrect cheli- 
cerae to posterior end of abdomen about 3.05 mm (body curled 
and difficult to measure accurately). Carapace 1.06 mm long; 0.99 
mm wide at level of lateral ocular tubercles; narrowed to 0.84 mm 
just behind ocular tubercles; pits and diagonal grooves, so con- 
spicuous in the female, are here much less prominent; general ap- 
pearance shown in Figure 42. Eyes: four as in female, in a straight 
row as seen from above; ME separated from one another by 8.5 
times their diameter, from LE by about twice their diameter; ratio 
of eyes ME : LE = 7 : 10; lateral eyes directed somewhat ven- 
trally. Chelicerae, maxillae and lip apparently as in female except 
for great reduction in size; details hidden by curled position and 
interlaced legs which are too fragile for disentanglement. Sternum: 
greatly swollen in middle but lacks the conspicuous transverse folds 
seen so clearly in the female; fourth coxae almost contiguous. Legs: 
tibial index of first leg 12, of fourth leg 11; calamistrum apparently 
lacking; first leg more than twice as long as second leg and nearly 
three times as long as third leg; first tibia with a battery of nearly 
two dozen robust spines on dorsal and dorsolateral sides essentially 
as shown in Figure 43; few spines observed elsewhere and the most 
conspicuous of these are on the first metatarsus. Palp: with femur, 
patella and tibia short and without special features; tarsus compli- 
cated and distinctive (Fig. 44). Abdomen: without noteworthy 
features; typical of males of the genus. Color in alcohol: in general 
light yellowish with variations; carapace with a somewhat angular, 
nearly white, central dorsal spot, with remainder of dorsum yellow- 
ish and with lateral sides grayish and with considerable black pig- 
ment in ocular region; sternum grayish; abdomen with dorsum light 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 23 

yellowish with numerous white flecks; a broken, grayish stripe ex- 
tends along each side of abdomen and the venter is irregularly 
grayish. Legs are yellowish in general but with variations; first legs 
with femora grayish retrolaterally; first tibiae, metatarsi and tarsi 
are all yellowish brown; other legs with grayish spots. 

Records. Specimens assigned to this species have been collected 
on Barro Colorado Island, Panama Canal Zone, during each of my 
several periods of field work there beginning in 1934; I have a few 
records of the species from other localities in the Canal Zone and in 
Panama proper. The described female is from Barro Colorado 
Island, Canal Zone, May 18, 1964; the described male is from the 
same locality, August 2, 1954. 

MlAGRAMMOPES TONATUS Sp. nOV. 

Figures 45-49 

Holotype. The male is from Jamaica, W. I., St. Andrew Parish, 
Mt. James at Plaintain River, October 24, 1957. The name of the 
species is an arbitrary combination of letters. 

Description. Total length 3.45 mm. Carapace 1.08 mm long; 
0.79 mm wide at level of lateral ocular tubercles; not conspicuously 
grooved as in some species in the genus; depressions just posterior 
to ME only moderately developed; median thoracic fovea only 
moderately depressed (Fig. 45). Eyes: four as usual, in a mod- 
erately recurved row, seen from above; ratio of eyes ME : LE r= 
5 : 6; ME separated from one another by about 6.5 times their 
diameter, from LE by slightly more than four times their diameter; 
lateral ocular tubercles only moderately developed. Chelicerae, 
maxillae and lip apparently quite typical of males of the genus. 
Sternum: with the usual form in this genus; greatly swollen, widened 
and extended between second and third coxae. Legs: tibial index 
of first leg 11, of fourth leg 8; calamistrum apparently lacking; first 
tibiae and metatarsi with few conspicuous, erect spines, others hori- 
zontal and hidden by hairs (Fig. 46); first leg more than twice as 
long as second leg, nearly twice as long as fourth leg and about 
three times as long as third leg; tarsal claws as usual; several tri- 
chobothria observed on tibiae and other segments. Palp: very short 
as usual; essential features shown in Figures 47-49. Abdomen ap- 
parently typical of males of the genus. Color in alcohol: carapace 
generally a dark gray with lighter spots and darker, irregular lines 
together with a dull, yellowish, ventral margin; sternum dark gray, 
lighter at anterior end; first pair of legs fight yellowish brown ventral- 
ly, darker gray elsewhere except for tarsus which is yellowish brown; 



24 



BREVIORA 



No. 289 




47 



Figures 45-49. Miagrammopes tonatus sp. nov. Fig. 45. Carapace of 
male from above. Fig. 46. First right tibia; retrolateral view. Figs. 47-48. 
Male palp; retrolateral and prolateral views, respectively. Fig. 49. Tip 
of tarsal bulb; nearly dorsal view. 



legs 2-4 much as first but less conspicuously so; mouth parts 
yellowish with grayish streaks. Abdomen: with a somewhat in- 
dented, dark gray, median stripe extending from base for about 
three-fourths of length of abdomen and then joining a darker area 
which covers the posterior quarter of the dorsum; lateral to this 
median region are lighter areas with irregular, yellowish white spots; 
venter with a central, light, reticular stripe extending from genital 
groove to posterior end; on each side of this central stripe there is a 
narrow, irregular, semitransparent stripe and further laterally are 
reticulated areas; entire color pattern indefinite and difficult to 
describe adequately. 

Records. An immature male taken with the holotype appears to 
belong to this species. The female is unknown. 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 



25 



MlAGRAMMOPES UNIPUS Sp. nOV. 

Figures 50-55 

Holotype. The male is from Barro Colorado Island, Panama 
Canal Zone. June, 1950. The name of the species is an arbitrary 
combination of letters. 

Description. Total length from anterior border of chelicerae to 
posterior end of spinnerets 3.58 mm. Carapace 1.06 mm long; 1.0 
mm wide at level of lateral ocular tubercles; about 0.4 mm tall; 
with general form essentially as in M. simus Chamberlin and Ivie 





50 



54 




55 




53 




52 



Figures 50-55. Miagrammopes unipus sp. nov. Fig. 50. Right first 
tibia of male; retrolateral view. Figs. 51-52. Left palpal tarsus of male; 
prolateral and retrolateral views, respectively. Fig. 53. Left palpal femur, 
patella and tibia; retrolateral view (from male paratype). Fig. 54. Ster- 
num of described female paratype; seen from below. Fig. 55. Epigynum; 
seen from below. 



26 BREVIORA No. 289 

with depressions medial to ME moderately conspicuous but grooves 
leading to lateral border almost lacking. Eyes: four as usual, in a 
gently procurved row as seen from above; ME separated from one 
another by about eight times their diameter, from LE by slightly 
more than their diameter; ratio of eyes ME : LE = 8 : 10.5. 
Chelicerae: short and largely hidden by elongated maxillae which 
are twice as long as lip; all of these parts hidden by fragile sur- 
rounding structures. Sternum: elongated, narrow, quite convex 
opposite second coxae; continued laterally between second and 
third coxae; obtusely terminated between bases of fourth coxae but 
not continued between the latter, which are separated by about one- 
fifth of their width. Legs: tibial index of first leg 10, of fourth leg 7; 
first leg nearly three times as long as third leg; second leg only a 
little shorter than fourth leg; first tibia with a set of robust spines 
(Fig. 50); first metatarsus with fewer and less robust spines; tarsal 
claws as usual in the genus. Palp: distinctive features shown in 
Figures 51-53; femur of moderate length; patella and tibia extreme- 
ly short. Abdomen: considerably flattened dorsally; otherwise 
typical of males of the genus. Color in alcohol: carapace with a 
narrow, central, somewhat broken grayish stripe; with considerable 
black pigment in ocular area; with a broad, lateral, grayish stripe on 
each side from lateral eyes to posterior border; the lower part of this 
gray stripe becomes lighter, thus adding another stripe to the lateral 
sides and, finally, the ventral margin is darker gray, thus making a 
total of seven distinguishable stripes on this part of the body; ster- 
num yellowish with fairly broad, grayish margins. Abdomen: 
yellowish dorsally with two pairs of narrow, elongated, grayish spots 
dorsolateral in position and a series of small grayish spots at pos- 
terior end above spinnerets; laterally there are several short, gray- 
ish stripes; the venter has a narrow, median, grayish stripe with a 
narrow yellowish stripe on each side, and further laterally is a 
broader, light grayish stripe on each side; shortly in front of the re- 
duced cribellum there is a narrow, grayish, transverse bar. Legs 
yellowish in general with several grayish spots and whole segments; 
fourth legs with most conspicuous grayish parts; fourth patellae 
dark gray; distal parts of fourth tibiae the same and nearly the entire 
metatarsi are gray. 

Female paratype. The described female paratype measures 5.2 
mm from anterior border of carapace to posterior end of spinnerets; 
length from anterior border of somewhat extended chelicerae to 
posterior end of spinnerets about 5.59 mm. Carapace 1.3 mm long; 
1.32 mm wide at level of lateral ocular tubercles; medial to ME 
there is a fairly conspicuous depression in communication with a 



1968 MIAGRAMMOPES IN PANAMA AND THE WEST INDIES 27 

narrow groove which extends along the border of the ocular tubercle 
to the lateral border of the carapace; another narrow groove in front 
of the ocular tubercle passes to the lateral border of the carapace 
and is connected to a short, narrow groove extending about half way 
through the tubercle between ME and LE; otherwise apparently 
typical of females of the genus. Chelicerae hidden and not clearly 
visible. Maxillae long, parallel, with rounded lateral borders. Lip: 
about three-fifths as long as maxillae; central portion swollen; 
sternal suture may not be a movable joint. Sternum: only moder- 
ately convex; not conspicuously swollen as in most females of the 
genus seen in this study; not extended between fourth coxae which 
are separated by nearly one-fourth of their width (Fig. 54). Legs: 
tibial index of first leg 10, of fourth leg 9; first leg twice as long as 
second leg and more than three times as long as third leg; fourth 
leg about 0.7 as long as first leg; spines appear to be lacking except 
for the moderately developed set of robust spines on fourth meta- 
tarsus and tarsus similar to those shown in Figure 40; legs with a 
fairly well developed coat of hair and first metatarsus has a mod- 
erately well developed ventral fringe; calamistrum appears typical 
of females of the genus. Abdomen: with a pronounced dorsal 
medial swelling; quite typical of females of the genus. Epigynum: 
obscure and difficult to observe clearly; observed essentials shown 
in Figure 55; a dorsal injury to the abdomen has permitted eggs to 
extrude, thus indicating that the female is nearly or quite mature. 
Color in alcohol: essentially as in male holotype except that the 
gray parts in that sex are here all considerably extended and 
blackened. 

Records. The described female paratype was taken on Barro 
Colorado Island, Panama Canal Zone, August 22, 1939. Several 
male paratypes are in the collection from the type locality and taken 
as follows: August, 1936, 1950; July, 1939, 1950, 1954; February, 
1958. One male paratype was taken at Summit, Panama Canal 
Zone, July, 1950. Three additional females are, with some un- 
certainty, placed in this species and were taken as follows: July, 
1934; August, 1939; May, 1964, all from Barro Colorado Island, 
Canal Zone. 

REFERENCES CITED 

Banks, Nathan 

1909. Arachnida of Cuba. Sec. Rept. Centr. Exper. Stat. Cuba. Pp. 

150-174, pi. 45. 
1929. Spiders from Panama. Bull. Mus. Comp. Zool., 69: 53-96, 
4 pis. 



28 BREVIORA No. 289 

Bonnet, Pierre 

1957. Bibliographia Araneorum. Toulouse. Vol. 2 (3). 
Bryant, Elizabeth 

1936. Descriptions of some new species of Cuban spiders. Mem. 

Soc. Cubana Hist. Nat., 10: 325-332, pi. 23. 
1940. Cuban spiders in the Museum of Comparative Zoology. Bull. 

Mus. Comp. Zool., 86(7) : 249-532. 22 pis. 
1948. The spiders of Hispaniola. Bull. Mus. Comp. Zool., 100(4): 
331-447, 12 pis. 

Cambridge, O. P. - 

1870. Descriptions and sketches of two new species of Araneidea with 
characters of a new genus. Jour. Linn. Soc. London, Zool., 
10: 401, figs. 1-12. 

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

1889- Arachnida-Araneida. Vols. I-IL ///.• Biologia Centrali-Ameri- 

1905. cana. Dulau & Co., London. 
Chamberlin, Ralph V., and Wilton Ivie 

1936. New spiders from Mexico and Panama. Bull. Univ. Utah, 
27 (5) (Biol, series, 3, No. 5): 3-103, 17 pis. 

Franganillo, Pelegrin Balboa, S. J. 

1936. Los Aracnidos de Cuba hasta 1936. Privately printed in 
Havana, 183 pp. 

LuTZ, Frank E. 

1915. List of Greater Antillean spiders, with notes on their distri- 
bution. Ann. New York Acad. Sci., 26: 71-148. 

Petrunkevitch, Alexander 

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

South America, etc. Bull. Amer. Mus. Nat. Hist., 29: 1-809. 
1926. Spiders from the Virgin Islands. Trans. Conn. Acad. Arts 

Sci., 28: 21-78. 
1930. The spiders of Porto Rico. Pt. 2. Trans. Conn. Acad. Arts 

Sci., 30: 159-355, 240 figs. 

ROEWER, C. Fr. 

1954. Katalog der Araneae. Brussels, Vol. 2b: 927-1751. 

Simon, E. 

1891. On the spiders of the Island of St. Vincent. Pt. I. Proc. Zool. 

Soc. London 1891: 549-575, pi. 42. 
1892- Histoire naturelle des araignees. Deuxieme edition. 2 Vols. 
1903. Librairie Encyclopedique de Roret, Paris. 

(Received 28 August 1967.) 



BREVIORA 



Meseuinii of Comparative Zoology 

Cambridge, Mass. 29 May, 1968 Number 290 

GEOGRAPHIC VARIATION IN THE HISPANIOLAN FROG 
ELEUTHERODACTYLUS WETMOREI COCHRAN 

Albert Schwartz' 



Eleutherodactylus wetmorei Cochran is a brightly colored lepto- 
dactylid frog which was described from a series of four specimens 
collected by Dr. Alexander Wetmore at Fond des Negres, Departe- 
ment du Sud, Haiti. The holotype and paratypes were taken from 
a communal nest of the Palm Chat {Dulus dominicus Linnaeus) 
along with two Hyla dominicensis Tschudi (Cochran, 1932:191). 
Later, Cochran (1941:76-77) reported 11 additional specimens 
of E. wetmorei, collected by W. L. Abbott, from Moron, Departe- 
ment du Sud, Haiti, near the extreme western tip of the Tiburon 
Peninsula. Both the typical and Abbott specimens were especially 
poorly preserved, but the frog is a very distinctive one in pattern 
(and in coloration in life). Two populations of E. wetmorei have 
the concealed surfaces of the hindlimbs with dark patterns on the 
brightly colored (orange to yellow) ground color; Cochran pointed 
out, however, that the Moron frogs seemed to lack this distinctive 
thigh pattern, and Shreve and Williams (1963:324) commented 
that two of the Moron series they examined lacked thigh markings, 
in contrast to the single individual they had from the Petionville 
area. 

The two-note voice of E. wetmorei is prominent in nocturnal 
choruses throughout southern Haiti; the frogs, however, are arbo- 
real (as the situation for the type series intimates), and collecting 
series of E. wetmorei is extremely difficult. Through the efforts 
of Dr. Ernest E. Williams, several excellent lots of E. wetmorei 
are now available from some Haitian localities, including the south- 
ern slope of the Massif de la Selle in the vicinity of Thiotte, and 
from the area between Jeremie and Dame-Marie at the extreme tip 
of the Tiburon Peninsula. The latter specimens amply confirm the 



^Dept. of Biology, Miami-Dade Junior College, Miami, Fla. 33167. 



2 BREVIORA No. 290 

distinctness of the distal peninsular population of E. wetmorei, and 
the Thiotte specimens likewise demonstrate characters (which also 
occur elsewhere along the Dominico-Haitian border) distinctive to 
the populations in that region. Remarkably, there is now more 
material of these two peripheral populations than there is of the 
nominate subspecies, although I have taken specimens of the latter 
at Camp Perrin on the southern foothills of the Massif de la Hotte. 
Aside from my own material in the Albert Schwartz Field Series 
(ASFS), I have examined specimens in the American Museum of 
Natural History (AMNH), Carnegie Museum (CM), Museum of 
Comparative Zoology (MCZ), and United States National Museum 
(USNM); for the loan of these frogs I am grateful to Charles M. 
Bogert and George W. Foley, Neil D. Richmond and Clarence J. 
McCoy, Jr., Ernest E. Williams, and Doris M. Cochran. In the 
field I have had the capable assistance of Ronald F. Klinikowski, 
David C. Leber, and Richard Thomas. Messrs. Leber and Thomas 
were successful in securing the first specimen of E. wetmorei from 
the Republica Dominicana; it was this frog, strikingly different 
from material I had seen in life at Camp Perrin, which prompted 
the present study. The excellence of the illustrations is once more 
due to the work of Mr. Leber. 

The collections in Haiti made for the Museum of Comparative 
Zoology were financed by NSF Grant 1 6066 to Dr. Ernest E. Wil- 
liams. 

SYSTEMATIC ACCOUNTS 
Eleutherodactylus wetmorei Cochran, 1932 

Eleutherodactyliis wetmorei Cochran, 1932, Proc. Biol. Soc. Washington, 45: 

191. 
Eleutherodactylus auriculatus wetmorei, — Cochran, 1941, Bull. U. S. Natl. 

Mus., 177:74. 
Eleutherodactylus varians wetmorei, — Schwartz, 1960, Sci. Publ. Reading 

Public Mus. and Art Gallery, 11:6. 
Eleutherodactylus wetmorei, — Schwartz [in press]. Studies Fauna Curasao 

and other Caribbean Islands. 

Remarks: Although E. wetmorei was originally named as a dis- 
tinct species, Cochran later (1941 : 75) considered wetmorei closely 
related to E. auriculatus Cope (= E. varians Gundlach and Pet- 
ers). Later still, I (Schwartz, 1960:6) suggested that wetmorei 
might indeed be correctly regarded as a subspecies of E. varians, 
since the two forms have several structural and pattern features 
in common (although chromatically they differ strikingly). The 



1968 ELEUTHERODACTYLUS WETMOREI 3 

calls of E. varians and E. wetmorei are, however, very different 
(Schwartz, in press), the call of E. wetmorei resembling that of the 
Cuban E. eileenae Dunn rather than E. varians. Accordingly, I 
have once more separated E. wetmorei nomenclatorially from E. 
varians; the former species is limited to Hispaniola, the latter to 
Cuba and the Isla de Pinos. 

ELEUTHERODACTYLUS WETMOREI WETMOREI Cochrau 

Type locality: Fond des Negres, Dept. du Sud, Haiti; holotype, 
USNM 72617. 

Definition: A subspecies of E. wetmorei characterized by anterior 
and posterior faces of thighs reticulate black to dark brown on 
an orange ground, and groin orange with similarly colored reticu- 
lum (Fig. 2A). 

Remarks: There are only ten specimens which I assign to the 
nominate subspecies; of these, the two paratypes (USNM 72618- 
19) are in extremely poor condition, and three (ASFS X2993-95) 
are juveniles with snout-vent lengths between 11.5 and 18.2 (all 
measurements in millimeters). Mensural data for a single male 
(ASFS X2684) are: snout-vent length 28.7, head length 11.3, 
head width 12.8, tympanum 2.0, eye 4.0, naris to eye 3.8, femur 
12.8, tibia 15.2. fourth toe 11.5; measurements of four females 
(extremes and means) are: snout-vent length 33.5-35.1 (34.2); 
head length 12.0-13.5 (12.9); head width 13.3-14.7 (14.1); tym- 
panum 2.2-2.5 (2.3); eye 3.8-4.8 (4.2); naris to eye 3.7-4.5 (4.2); 
femur 14.1-15.2 (14.8); tibia 17.1-17.8 (17.4); fourth toe 13.4- 
14.3 (13.8). 

The condition of the topotypes of E. w. wetmorei precludes ab- 
solute assurance that the material from Camp Perrin is identical in 
pattern to them. Cochran's figure (1941:75, fig. 24) however, 
shows quite clearly that the anterior and posterior faces of the thigh 
are darkly reticulate, and these markings plus the pale interocular 
bar are still barely visible on the holotype. Although Fond des 
Negres lies in the extreme northeastern foothills of the Massif de 
la Hotte and Camp Perrin lies in the southwestern foothills of the 
same range, the Camp Perrin specimens agree fairly closely in 
thigh design with the Fond des Negres material. Accordingly I con- 
sider the Camp Perrin specimens to be E. w. wetmorei, although 
there is a possibility that, with the acquisition of fresh topotypical 
material, the Camp Perrin population will be found to differ from 
that at Fond des Negres. 



4 BREVIORA No. 290 

Fresh specimens of E. w. wetmorei from Camp Perrin are tan to 
brownish tan in hfe, with a very faint pair of dorsolateral pale lines 
(ASFS X2684) present in one individual. The posterior and an- 
terior faces of the thigh, the groin, and the upper surface of the pes 
were orange (pi. 2 D 12; all color designations from Maerz and 
Paul, 1950) in life, and the entire ventral surface was yellow (pi. 
9 L 5). Both faces of the thighs, as well as the groin, are overlaid 
with a dark brown to greenish black reticulum; reticulum remnants 
more or less outline the orange groin patch and separate the bright 
color from the more drab dorsal tan. The sides are at times vaguely 
marked with a brown reticulum which extends, in a diluted fashion, 
onto the chest as an area of brownish dots or flecks. The throat 
may be flecked with brown, and there is a pair of yellow-orange 
glands on the posterior portion of the throat; inguinal glands are 
absent in the species. The underside of the hindlimbs is usually 
marked with brown continuations of the anterior and posterior thigh 
pattern, and the crus also shows some ventral brown reticulations. 
The ventral crural reticulum is a continuation of the anterior and 
posterior crural reticulum, occupying the concealed surfaces of the 
crus. All specimens have a pale (tan to buffy) broad interocular 
bar, with one exception (MCZ 35199) which now appears to lack 
this feature. In the juveniles, the entire thigh was yellow in life, but 
the thigh pattern is identical to that of aduhs. In adults, the iris is 
golden above and below, whereas in juveniles the iris is metaUic 
buffy. 

At Camp Perrin, E. w. wetmorei was abundant in trees about 
the settlement, and the ringing two-note call made up a prominent 
portion of each night's choruses. Specimens, on the other hand, 
were difficult to secure; the three juveniles were taken by Mr. Leber 
from arboreal bromeliads, whereas the adult male (ASFS V2684) 
was obtained while calling on a Sansevieha leaf only one foot off 
the ground. Like the equally arboreal Cuban E. varians, E. wet- 
morei occasionally can be collected when it calls from low vocaliz- 
ing sites. 

As pointed out above, all known localities for E. w. wetmorei are 
associated with the lower slopes of the Massif de la Hotte. The 
locality near Miragoane is unlocatable on any current map, so it 
is possible that this specimen came from a lower elevation than 
the other E. w. wetmorei; the elevation at Camp Perrin is 1000 feet 
(305 meters) and that at Fond des Negres is 730 feet (240 
meters). Figure 1 shows localities for specimens of E. w. wetmorei. 



1968 



ELEUTHERODACTYLUS WETMOREI 



Specimens examined: Haiti, Dept. du Sud, Camp Perrin, 6 
(ASFS X2684, X2897, X2962, X2993-95); Fond des Negres, 3 
(USNM 72617 — holotype; USNM 72618-19 — paratypes); Bu- 
tete, nr. Miragoane, 1 (MCZ 35199). 




Fig. 1. Map of southwestern Haiti, showing the Tiburon Peninsula. 
Localities for E. wetmorei wetmorei, E. w. williamsi, and E. w. ceraemeriis 
are shown as solid circles. 



ELEUTHERODACTYLUS WETMOREI CERAEMERUS 

new subspecies 

Holotype: MCZ 36101, an adult female, from Thiotte, Dept. de 
rOuest, Haiti, one of a series collected April 1962 by G. Whiteman. 

Paratypes: MCZ 36097-100, 36102-05, USNM 146614, same 
data as holotype; MCZ 37216-18, same locality as holotype, sum- 
mer 1962, G. Whiteman; CM 38550-55. same locality as holotype, 
June 1962, G. Whiteman; MCZ 36107-11, Tete a I'Eau, near Sal- 
trou, Dept. de I'Ouest, Haiti, April 1962, G. Whiteman; CM 
37786, MCZ 34501. Marbial, 21 km NE Jacmel, Dept. de I'Ouest, 
Haiti, 20-21 April 1961, L. Whiteman; AMNH 44034, ridge of 
Massif de la Selle, just south of Savane Zombi, on Saltrou road, 
"4500 feet," Dept. de TOuest, Haiti, 6 April 1935, W. G. Hassler; 
MCZ 34503, AMNH 44050, Colombia (= Colombier), Dept. de 
rOuest, Haiti, 6 April 1935, W. G. Hassler; MCZ 34502, AMNH 
44036-37, La Mahot. near Colombia {— Colombier), Dept. de 
rOuest, Haiti. 7 April 1935, W. G. Hassler; MCZ 31733, Boutil- 
lier Road, Dept. de TOuest. Haiti, 9 August 1959, E. E. Williams 



BREVIORA 



No. 290 



and A. S. Rand; MCZ 37141, La Boule, Dept. de TOuest, Haiti, 
18 April 1960, J. A. Rivero; ASFS V271 1, 7 km SE Los Arroyos, 
2200 feet (720 meters), Pedernales Province, Republica Domini- 
cana, 29 June 1964, R. Thomas. 

Definition: A subspecies of E. wetmorei characterized by anterior 
and posterior faces of thigh bright orange with a pattern of black 
spots rather than a reticulum, and groin orange with scattered black 
dots and outlined by black (Fig. 2B). 






Fig. 2 Three subspecies of E. wetmorei, as follows: A, E. w. wetmorei, 
ASFS X2962, Camp Perrin, Dept. du Sud, Haiti; B, E. w. ceraemeriis, MCZ 
36101, holotype, Thiotte, Dept. de I'Ouest, Haiti; C, E. w. williamsi, MCZ 
31151 , holotype, Marfranc, Dept. du Sud, Haiti. 



Description of holotype: An adult female with the following 
measurements: snout-vent length 35.6, head length 12.7, head 
width 14.5, longitudinal diameter of tympanum 2.5, longitudinal 
diameter of eye 4.5, naris to eye 4.0, femur 14.7, tibia 17.5, fourth 
toe 13.8. Head distinctly broader than long; snout truncate, with 
nares conspicuous at anterior end of canthus rostralis; diameter of 
eye slightly longer than distance from naris to anterior corner of 
eye; diameter of tympanum much less than diameter of eye, dis- 
tance from tympanum to eye slightly less than diameter of tym- 
panum. Interorbital distance 5.4, greater than diameter of eye. 
Digital discs present, large, those on digits 2, 3, and 4 only slightly 
smaller than area of tympanum. Fingers short, basally with a very 
small web, 3-4-2-1 in order of decreasing length; subarticular 
tubercles pale, concolor with palmar surface of hand. Toes moder- 
ately long, oiily very slightly webbed basally, 4-3-5-2-1 in order of 
decreasing length; subarticular tubercles prominent, concolor with 
plantar surface. Heels overlap strongly when femora are held at 



1968 ELEUTHERODACTYLUS WETMOREI 7 

right angles to body axis. Inguinal glands absent. Dorsum smooth. 
Throat and chest smooth, belly coarsely granular. Dorsal surface 
of fore- and hindlimbs smooth. Posterior and ventral faces of thighs 
covered with moderately sized juxtaposed flattened granules. 
Tongue large, free and notched behind, its greatest width equal to 
about two-thirds of that of floor of mouth. Vomerine teeth in two 
small, almost diagonal patches, enclosed within the median margins 
of the choanae, separated from them by a distance equal to the 
diameter of a choana, the two patches separated medially by a dis- 
tance equal to about half the length of a single vomerine patch. 

Coloration and pattern of Jiolotype: Dorsum (as preserved) 
brown and without pattern except for a vaguely discernible paler 
interocular bar and a very faint middorsal hairline; hindlimbs con- 
color with dorsum; concealed surfaces of thigh, crus, and pes paler 
(presumably orange in life) with scattered dark brown dots and 
outlined with a faintly darker brown line, punctate with dark brown 
dots along its length; groin spot pale (presumably orange in life) 
with a few scattered dark dots on the left side (right side immacu- 
late), the groin spot outlined by a vague darker line and sharply set 
off from the dorsal color; forelimbs dark brown, concolor with the 
dorsum but with a sharply distinct pale mark on the wrist and an- 
other at the forelimb insertion. Venter pale (presumably orange in 
life), heavily suftused with brown stippling on the sides anteriorly 
(anterior to the pale groin spot); throat densely stippled with dark 
brown; underside of forelimbs dark brown on antebrachium, pale 
on brachium. A pair of moderately prominent glandular areas on 
the posterior throat anterior to the level of the forelimb insertions. 

Variation: The series of nine male E. w. ceraemerus has the fol- 
lowing measurements (extremes and means): snout- vent length 
28.5-33.4 (30.4), head length 9.8-11.8 (10.8), head width 10.8- 
12.6 (12.2), tympanum 1.7-2.0 (1.8), eye 3.4-4.0 (3.7), naris to 
eye 2.9-3.9 (3.5), femur 11.6-14.2 (13.2), tibia 14.5-15.9 (15.2), 
fourth toe 11.2-12.3 (11.7). Twenty females have the following 
measurements: snout-vent length 26.7-38.3 (34.4), head length 
9.6-13.3 (11.9), head width 10.8-15.0 (13.5), tympanum 1.6-2.5 
(2.1), eye 3.1-4.7 (4.1), naris to eye 3.2-4.5 (3.8), femur 11.9- 
16.8 (14.6), tibia 14.1-18.9 (17.2), fourth toe 10.4-14.3 (13.2). 

As preserved, the paratypes agree well in color with the holotype, 
although some specimens are paler (tan rather than brown). In 
coloration, the concealed hindlimb surfaces and the groin spot are 
always sharply and conspicuously set off from the remainder of the 
hindlimb. and the pattern of these areas is regularly one of dots or 



8 BREVIORA No. 290 

spots, never a reticulum as in E. w. wetmorei. At times the spots or 
dots are somewhat fused (MCZ 34501), but the condition never 
approaches that of the nominate subspecies. The groin spot may 
have some dark flecking within it, or it may be immaculate; in either 
case it is outlined dorsally by a dark and punctate line as in the 
holotype. Some specimens show a pale interocular bar (MCZ 
36105), but this is not the regular condition in ceraemerus. A very 
fine median hairline is barely discernible in some specimens 
(USNM 146614), as it is in the holotype, but most frogs lack this 
pattern element. Other specimens (MCZ 36102) have some fine 
and vague stippling along the lower sides above the venter, but this 
condition likewise occurs only sporadically. Only those frogs which 
are dorsally as dark as the holotype have the sides of the venter 
anterior to the groin spot as darkly pigmented as does the holotype. 

There are only two specimens from north of the Massif de la 
Selle, those from La Boule and Boutillier Road; the balance of the 
series is from the southern side of that massif. I do not know if the 
ranges of these two populations are continuously directly across the 
massif, but it does not seem likely, since my impression is that E. 
wetmorei is generally a frog of low to moderate elevations in 
forested situations. In any event, the two frogs from north of the 
La Selle agree in thigh pattern details with those from the southern 
slope except that the degree of fusion between the spots is greater 
than is customary for southern specimens. It seems quite likely 
that another subspecies will be found to inhabit the northern slopes 
of the La Selle and its associated ranges above Port-au-Prince; the 
thigh pattern of these northern specimens does not approach the 
reticulum of E. w. wetmorei. 

Structurally, the paratypes agree with the holotype except that the 
buccal cavity of the latter is somewhat aberrant. In most specimens 
of E. w. ceraemerus, the vomerine patches are small, compact, and 
enclosed well within the inner margins of the choanae, separated 
from the choanae by a distance equal to that of one vomerine series 
and separated from each other by a similar distance. In the holo- 
type, the vomerine patches are closer to one another than in the 
balance of the series. 

Comparisons: Comparison of the subspecies wetmorei and 
ceraemerus is hardly necessary; the two forms can be easily dis- 
tinguished by the pattern of the concealed surfaces — reticulate in 
wetmorei, dotted in ceraemerus. Size differences are difficult to 
assess, since the series of E. w. wetmorei is very small. E. w. 
ceraemerus seems the larger of the two forms. 



1968 ELEUTHERODACTYLUS WETMOREI 9 

Although there are no color data on most specimens of E. w. 
cememerus, Richard Thomas recorded the color in life of the 
Dominican specimen as follows: dorsum grayish tan (= putty 
colored), venter bright orange. Forearm insertions, pre- and post- 
femoral surfaces, and groin bright orange with black dotting. 
Undersides of legs and dorsal and ventral surfaces of feet bright 
orange; dark flecking roughly outlining region of contact between 
orange and tan on both limbs and groin. 

Remarks: The abundance of specimens of E. w. ceraemerus, in 
contrast to the paucity of those of the nominate subspecies, is 
puzzling, although it is possible that the frog is more abundant or 
more easily secured on the southern slope of the Massif de la Selle 
than it is farther west. The Dominican specimen was collected while 
it was calling about ten feet above the ground in a coffee tree; 
others were heard calling in the same general region, but most were 
vocalizing high in the trees and thus were inaccessible. The scarcity 
of specimens from the Morne I'Hopital above Petionville is equally 
puzzling; the calls of this frog were commonly heard between 
Petionville and Fermate, but there are only two specimens from this 
entire region. The species apparently is absent from the Sierra de 
Baoruco, where the dominant, moderately sized, more or less arbo- 
real Eleutherodactylus is E. armstrongi Noble and Hassler. This 
species also occurs south of Port-au-Prince on the Montagnes Noires 
at Furcy, an area where E. wetmorei is absent. The Dominican 
specimen is from the extreme edge of the Republica Dominicana in 
what may well be considered the eastern extreme of the Massif de 
la Selle. 

The altitudinal limits of E. w. ceraemerus vary from a low eleva- 
tion of about 600 feet (200 meters) at Marbial to high elevations 
of about 4270 feet (1400 meters) near Savane Zombi. These 
elevations are greater than those of the nominate subspecies to the 
west. The name ceraemerus is derived from the Greek "keraia," a 
dot, and "meros," thigh, an illusion to the dotted concealed surfaces 
of the hindlimbs in this subspecies. 

Eleutherodactylus wetmorei williamsi new subspecies 

Holotype: MCZ 37757, an adult female, from Marfranc, Dept. 
du Sud, Haiti, one of a series collected 26-27 December 1962 by 
Hill and Vuilleumier. 

Paratypes: MCZ 37751-56, 37758-63, same data as holotype; 
MCZ 37587, Perine, near Jeremie, Dept. du Sud, Haiti, December 
1962, G. Whiteman; MCZ 37586, Carrefour Sanon, near Jeremie, 



10 BREVIORA No. 290 

Dept. du Sud, Haiti, December 1961, G. Whiteman; USNM 60627- 
35, Moron, Dept. du Sud, Haiti, 20 December 1917, W. L. Abbott. 
MCZ 28599-600, Moron, Dept. du Sud, Haiti, 24 December 1917, 
W. L. Abbott. 

Definition: A subspecies of E. wetmorei characterized by com- 
plete absence of pattern on anterior and posterior faces of thigh and 
on groin spot (Fig. 2C). 

Description of holotype: An adult female with the following 
measurements: snout-vent length 33.8, head length 11.8, head 
width 13.0, longitudinal diameter of tympanum 2.0, longitudinal 
diameter of eye 4.2, naris to eye 4.1, femur 14.2, tibia 16.9, fourth 
toe 13.1. Head distinctly broader than long; snout truncate, with 
nares conspicuous at anterior end of canthus rostralis; diameter of 
eye about equal to distance between naris and anterior corner of 
eye; diameter of tympanum much less than diameter of eye, dis- 
tance from tympanum to eye equal to about one-half the diameter 
of tympanum. Interorbital distance 4.0, slightly less than diameter 
of eye. Digital discs present, large, those on digits 2, 3, and 4 only 
slightly smaller than area of tympanum. Fingers short, basally with 
a very small web, 3-4-2-1 in order of decreasing length; subarticular 
tubercles pale, concolor with palmar surface of hand. Toes moder- 
ately long, only very slightly webbed basally, 4-3-5-2-1 in order of 
decreasing length; subarticular tubercles prominent, concolor with 
plantar surface. Heels overlap strongly when femora held at right 
angles to body axis. Inguinal glands absent. Dorsum smooth. 
Throat and chest smooth, belly coarsely granular. Dorsal sur- 
face of fore- and hindlimbs smooth. Posterior and ventral faces 
of thighs covered with moderately sized, juxtaposed, flattened gran- 
ules. Tongue elongate and narrow, free and notched behind, its 
greatest width equal to about one-half of that of floor of mouth. 
Vomerine teeth in two very small patches, well enclosed within the 
median margins of the choanae, separated from them by a distance 
equal to the length of one vomerine series, the two patches separated 
by a distance equal to the length of one vomerine patch. 

Coloration and pattern of holotype: Dorsum (as preserved) pale 
tan, heavily stippled with brown on the trunk; head less densely 
stippled and with remnants of a pale interocular bar, outlined an- 
teriorly and posteriorly by remnants of brown lines; upper surfaces 
of hindlimbs concolor with dorsum; concealed surfaces of thigh, 
crus and pes paler (presumably brightly colored in life) without any 
dark markings or outlining in darker; groin spot also paler and 
without dark markings or outlining; forelimbs tan, concolor with 
dorsum. Venter pale (presumably brightly colored in life), without 



1968 ELEUTHERODACTYLUS WETMOREI 11 

heavy stippling; throat unstippled but with a pair of darker glan- 
dular areas anterior to the forelimb insertions. 

Variation: The single male E. w. williamsi has the following 
measurements: snout-vent length 32.7, head length 11.5, head 
width 13.1, tympanum 1.6, eye 4.0, naris to eye 3.9, femur 14.6, 
tibia 16.4, fourth toe 12.9; the series of 21 females has the following 
mensural data (extremes and means): snout-vent length 25.2-36.4 
(32.7), head length 9.5-12.2 (1 1.5), head width 10.7-14.0 (13.1), 
tympanum 1.2-2.3 (1.8), eye 3.0-4.7 (3.9), naris to eye 3.1-4.5 
(3.5), femur 11.2-15.2 (13.5), tibia 14.0-18.0 (15.7), fourth toe 
9.1-14.0 (11.9). 

The paratypic series includes both long-preserved and freshly 
collected material; the former is the lot from Moron and the latter 
material is from the type locality and from near Jeremie. Many 
recent specimens have the more or less speckled dorsal aspect of 
the type, but this style of pattern does not occur in the Moron 
material, a fact which I assume is due to the length of time in 
preservation. The presence of the interocular bar is variable, but it 
is demonstrated in both old and new specimens; many frogs show 
this feature more diagrammatically than does the holotype. A 
median dorsal hairline occurs in some individuals (MCZ 37759, 
USNM 60629). There is never any indication of dark pattern on 
the concealed surfaces nor in the groin, these areas being quite pale; 
the freshly taken specimens still show a weak pinkish-orange wash 
on the thigh, so that I assume that these areas were quite brightly 
colored in life, presumably some shade of yellow or orange like the 
two other subspecies of E. wetmorei. Structurally, the paratypes 
agree very closely with the description of the holotype. 

Comparisons: E. \v. williamsi is easily distinguished from the 
nominate subspecies and E. w. ceraemerus since it lacks any indica- 
tion of thigh or groin pattern. As preserved, the specimens of 
williamsi are regularly paler than those of the other subspecies, and 
it seems possible that there is a dorsal chromatic difference between 
williamsi and the other subspecies. E. w. williamsi does not appear 
to reach so large a size as does ceraemerus but exceeds slightly the 
maximum size recorded for E. w. wetmorei. The absence of detailed 
color data in life for E. w. williamsi hampers more detailed 
comparisons. 

Remarks: E. w. williamsi occupies the extreme northwestern 
portion of the Tiburon Peninsula in Haiti. Its range, as now known, 
is circumscribed, with all localities being associated with the Monts 
Cartaches to the west of Jeremie. Altitudinally, E. w. williamsi 



12 BREVIORA No. 290 

occurs somewhat lower than the two more eastern subspecies, from 
a low elevation of about 130 feet (40 meters) at Moron to 1440 
feet (440 meters) at Carrefour Sanon; Ferine is not locatable. 

The distinct patternless condition of the western population was 
first recognized by Cochran (1941:76), who pointed out that the 
series from Moron was light in color; later, Shreve and Williams 
(1963:324) again commented that the Moron series lacked limb 
and groin markings, in contrast to their specimen from Boutillier 
Road {ceraemerus) . The new series in the Harvard collection 
amply confirms these differences and demonstrates that they are 
not due to fading or the age of the previously known material from 
this region. I take great pleasure in naming the far western sub- 
species of E. wetmorei for Dr. Ernest E. Williams, whose interest in 
Hispaniolan herpetology, sponsorship of field work on that Antillean 
island, and generous cooperation with others involved with His- 
paniolan herpetological problems can be acknowledged only in a 
token fashion by the use of his name as a patronymic designation 
for this distinctive subspecies. 

DISCUSSION 

E. wetmorei is now known to be widely distributed along the 
length of the Tiburon Peninsula in Haiti; the altitudinal limits of the 
species indicate that it is an inhabitant of low to moderate elevations 
associated with the north and south flanks of the massifs de La 
Selle and de La Hotte. There is but one record from the Republica 
Dominicana, along the Dominico-Haitian border in the extreme 
eastern ranges of the Massif de La Selle. The species remains un- 
known from the Sierra de Baoruco proper. Doubtless many areas 
in Haiti whence E. wetmorei remains unknown will ultimately be 
found to be inhabited by this species; since the frog is predomi- 
nantly arboreal, it is difficult to collect, and often the species has 
been heard vocalizing in areas where specimens were impossible 
to secure. 

As in several other south island (sensu Williams, 1961) am- 
phibians and reptiles, E. wetmorei has differentiated into a series of 
three distinct populations which are easily recognizable. Although 
there is really too little material to be certain, it is interesting that 
the specimens from Fond des Negres agree with those from Camp 
Perrin on the southern slopes of the Massif de La Hotte farther to 
the west. Such a similarity is remarkably like that shown in 
Sphaerodactylus copei Steindachner, wherein material from the 
vicinity of Miragoane (and thus near the Fond des Negres) is like 



1968 ELEUTHERODACTYLUS WETMOREl 13 

5. c. cataplexis Schwartz and Thomas (at Camp Perrin) rather than 
like S. c. picturatus Carman (at Jeremie). Apparently the low and 
forested pass across the Massif de La Hotte in the Aquin-Fond des 
Negres-Miragoane region has allowed passage of some southern 
subspecies to the north coast, thereby more or less separating the 
two north coast populations {S. c. picturatus and S. c. copei sep- 
arated by S. c. cataplexis, just as E. w. williamsi and E. w. cerae- 
merus are separated by E. w. wetmorei; see Schwartz and Thomas, 
1965, for details of gecko distributions in this region). Such 
similarities seem hardly due to chance. 

LITERATURE CITED 

Cochran, Doris M. 

1932. A new frog, Eleiitherodactylus wetmorei, from the republic of 

Haiti. Proc. Biol. Soc. Washington, 45: 191-193. 
1941. The herpetology of Hispaniola. Bull. U. S. Natl. Mus., 177:i-vii, 
1-398, 120 figs, 12 pis. 

Maerz, a., and M. Rea Paul 

1950. A dictionary of color. McGraw-Hill Book Co., pp. i-vii, 1-23, 
137-208, 56 pis. 

Schwartz, Albert 

1960. Nine new Cuban frogs of the genus Eleiitherodactylus. Reading 
Public Mus. and Art Gallery, Sci. Publ. 11:1 -50, 6 figs. 

(IN PRESS). The Antillean Eleiitherodactylus of the auriculatus group. 

Studies Fauna Curagao and other Caribbean Islands. 

and Richard Thomas 

1965. Subspeciation in Sphaerodactyliis copei. Quart. Jour. Florida 

Acad. Sci., 27(4) :316-332, 1 pi. 

Shreve, Benjamin, and Ernest E. Williams 

1963. The herpetology of the Port-au-Prince region and Gonave Island, 
Haiti. Pt. II. The frogs. Bull. Mus. Comp. Zool., 129(5) : 302-342, 
5 pis. 

Williams, Ernest E. 

1961. The evolution and relationships of the Anolis semilineatus 
group. Breviora, Mus. Comp. Zool., No. 136:1-7, map. 

(Received 9 November, 1967.) 



BREVIORA 



MitaseiLam of Comparative Zoology 



Cambridge, Mass. 29 May. 1968 Number 291 

A NEW SPECIES OF TRIBOLONOTUS (LACERTILIA: 

SCINCIDAE) FROM BOUGAINVILLE AND BUKA, 

SOLOMON ISLANDS, WITH COMMENTS ON THE 

BIOLOGY OF THE GENUS 

Allen E. Greer and Fred Parker' 

INTRODUCTION 

The skinks of the genus Tribolonotus are undoubtedly one of the 
most bizzare taxa of lizards. Indeed, many of their anatomical fea- 
tures such as abdominal glands (Parker, 1940). volar pores (Roux, 
1930 and 1934; Parker, 1940), and single right functional ovi- 
duct (discussed below) are as yet unreported in other lizards. 
Among skinks, these features, as well as their anomalous head and 
body squamation, make them one of the most easily distinguished 
groups in an otherwise taxonomically difficult family. 

In this paper we describe an unnamed species of the genus and 
discuss various aspects of the ecology, reproductive biology, and 
behavior of several species. 

DESCRIPTION OF A NEW SPECIES OF TRIBOLONOTUS 

During a recent period of residence (1962-1963) on Bougain- 
ville Island and a subsequent return trip (1966) to the northern 
Solomons. Parker collected a large number of specimens of a 
hitherto unnamed species of Tribolonotus on the islands of Bou- 
gainville and Buka. The new species is very similar in many details 
of squamation to T. ponceleti Kinghorn, 1937, which occurs on 
Bougainville and Shortland islands, and may therefore be known as 



' Department of District Administration. Kundiawa. Territory of New 
Guinea, 



2 BREVIORA No. 291 

Tribolonotus PSEUDOPONCELETii new species 

Holotype: Museum of Comparative Zoology (MCZ) 1291 A, an 
adult male collected by Fred Parker at Kunua, Bougainville, on 
30 December 1962. 

Paratypes (610 specimens) : BOUGAINVILLE: KUNUA: MCZ 
12161-12119, 72781-72785, 72790-72791, 72793-72805, 72807- 
72813, 72815-72829, 72831-72863, 72865-72866, 72868-72869. 
72871-72873. 72875-72900, 72910-72912. 76148, 76155-76178, 
76181-76191, 76193-76201. 76412-76469. 78094-78282. + 4 
untagged specimens; American Museum of Natural History 
(AMNH) 92026-92058, + 10 untagged specimens; Australian 
Museum (AM) R 26603-26607; British^Museum (BM) 1963.578- 
1963.584; Field Museum of Natural History (FMNH) 141982- 
141985, 152596; University of Kansas Natural History Museum 
(UKNHM) 98510-98511; Zoological Museum of Berlin (ZMB) 
39155 (2 specimens). BOKU: MCZ 65878-65880, 67723-67726; 
AMNH 89433. KIETA: MCZ 65875-65877, 67249-67250. 
MATSIOGU area: MCZ 92423-92426, 92428. 92468. 92492- 
92495. MELILUP: MCZ 92429-92433. MUTAHI: MCZ 
87615-87623. 88782-88794, 91467-91472, 92394-92422, 92427, 
92434-92440. Above POPHEIARAI: MCZ 76179. RAMAZON 
RIVER (1600-2400 feet): MCZ 92502-92503. 

BUKA ISLAND: MCZ 67706-67716, 73850-73861; AMNH 
89434. KUBA: MCZ 92491. 

Diagnosis: Pseiidoponceleti differs from blancliardi of the Sol- 
omons in having two rows of enlarged vertebral scales instead of 
a single row as in bkmchardi. Sclmiidti of the Solomons also has 
two rows of enlarged vertebral scales, but whereas these scales 
extend anteriorly to the single large parietal scale in schmidti, they 
only extend to a point just anterior to the level of the insertion of 
the forelegs in pseiidoponceleti. Size is the easiest way to dis- 
tinguish ponceleti from pseiidoponceleti. The three specimens of 



^ The similarity in certain points of squamation between T. ponceleti and 
the new species, as well as their supposed sympatry (but see Ecology sec- 
tion of paper below), has caused the new species to be confused with 
ponceleti both in the field and in museum collections. Fortunately, this 
confusion has been carried over into the literature only once (Zweifel, 
1966). To help clarify the taxonomic confusion, it seems best to take the 
path of least nomenclatural confusion and call the new species pseiido- 
ponceleti. 



1968 NEW SPECIES OF TRIBOLONOTUS 3 

poucelcti known in collections range from 116-125 mm in snout- 
vent length, whereas the largest of 611 pseudoponceleti examined 
is only 73 mm in snout-vent length. 

Gracilis and novaeguineae of New Guinea can be immediately 
distinguished from pseudoponceleti by means of the greatly en- 
larged spines on the two vertebral and paravertebral scale rows, 
and the head casque which, posteriorly, is raised above the level 
of the nape. 

Pseudoponceleti differs from annectens of New Britain (known 
from the holotype and more recently from a second specimen in 
the Zoologisk Museum, Copenhagen i) in several scale characters 
which are summarized here from Zweifel (1966; see Fig. 1). "In 
both species there is a prominent row of enlarged scales paralleling 
the much larger vertebrals, but where at midbody in pseudopon- 
celeti there is one of these smaller scales for each vertebral, an- 
nectens has only one for every two vertebrals. There are two 
primary temporal scales in pseudoponceleti and three in annectens." 
The first infralabial in pseudoponceleti is long and thin and extends 
posteriorly almost to exclude the second infralabial from the edge 
of the lip, or in a very few cases ( 1 specimen in 46 examined by 
Zweifel, 1966) the first labial does just exclude the entire second 
infralabial from the edge of the lip. In annectens the first infra- 
labial extends posteriorly to exclude the second (in the type) or 
the second and third (in the Copenhagen specimen) infralabials 
trom the edge of the lip.- 

Description (Fig. 2; also see fig. 1 in Zweifel, 1966): T. 
pseudoponceleti measures between 26 and 73 mm in snout-vent 
length, with the tail approximately 59 to 65 per cent of the total 
length. The pentadactyl limbs are well developed and overlap when 
adpressed to the body. The head is deep and rather triangular in 
shape when viewed from above. The longitudinal striations on the 
head, the keeled vertebral and caudal scales, and the enlarged 
scales on the sides and limbs give the animal a rather rugose 
appearance. 



'The locality data for this second known specimen of annectens are as 
follows: Yalom, 35 km S.E. of Cape Lambert. Gazelle Peninsula, New 
Britain. Elevation 1000 meters above sea level. 

- As these infralabials are excluded from the edge of the lip, they might 
be called sublabials as Zweifel (1966) notes. We choose to call them in- 
fralabials here, however, to emphasize the proper homologous relationships. 



4 BREVIORA No. 291 

The rostral is approximately 2'/2 times as wide as deep. A very 
large frontonasal covers the dorsal surface of the snout, and pre- 
frontals are absent. The frontal is slightly shorter than the fronto- 
nasal and touches this scale throughout its entire anterior length. 
The frontal is in contact with the 2 anterior supraoculars laterally 
and the frontoparietals posteriorly. There are 4 supraoculars. The 
frontoparietals are distinct or fused to varying degrees. A distinct 
interparietal is lacking. The single large parietal, within which lies 
the parietal eye, is bounded on both sides by a smaller anterior and 
posterior parietal scale.' 

The nostril is contained in a single nasal. A large, roughly rec- 
tangular loreal extends from the anterior corner of the eye to the 
nasal and first supralabial. There are 5 supralabials; the first is 
extremely long and thin and extends posteriorly to the level of the 
anterior half of the eye, thereby excluding the anterior 2/5 of the 
large, rectangular second supralabial from the margin of the lip. 
A large 3rd and smaller 4th and 5th supralabial follow. Both the 
second and third supralabials border the smaller scales of the en- 
tirely scaly eyelid. There is a small scale above the 4th supralabial 
and two superposed scales are above the 5th supralabial. Posterior 
to these last 3 scales are the two temporal scales, of which the 
lower is about twice the size of the upper. 

The mental is followed on either side by 5 or 6 infralabials. The 
first of these is long and thin and extends posteriorly about as far 
as does the first supralabial above. The first infralabial excludes 
approximately Vi-Va of the second infralabial from the margin of 
the lip. A single large postmental is followed by a pair of very 
large chin scales which form a suture along the midline and are 
followed by a second part of chin scales of slightly smaller size. 
These chin scales are separated by 4 of the smaller gular scales. 

All the large head scales bear longitudinal ridges or keels. 

The tympanum is almost level with the surface of the head. 
Auricular lobes are lacking. 



^ Zweifel's (1966) interpretation of the single large median parietal as 
possibly resulting from the fusion of the interparietal and parietals seems 
perfectly plausible to us. In all other lygosomine skinks the parietal eye 
occurs in an interparietal which would argue for the interparietal's being 
part of the single median parietal of Triholonotiis. Zweifel's observation 
of the incomplete posterior median suture in the large parietal can cer- 
tainly be seen in many pscitdoponceleti and again argues for the paired 
lygosomine parietals having become part of the single shield. 



1968 NEW SPECIES OF TRIBOLONOTUS 5 

There are 21-26 pairs of enlarged, keeled, vertebral scales be- 
tween the base of the tail and a point just anterior to the insertion 
of the forelimbs. Between each of these pairs of enlarged ver- 
tebrals there is usually a single small median scale. 

The side of the body between the large vertebral scales and the 
large ventral scales is covered with many small granular scales and 
fewer enlarged, almost tubercle-like scales di^sposed in roughly 
oblique rows. Granular scales predominate in an area adjacent to 
and as wide as the vertebral rows, except for a row of slightly 
enlarged, keeled scales that alternate with the vertebrals. 

The tubercle-like scales extend from the side of the body forward 
onto the side of the neck and dorsally onto the nape between the 
enlarged post-parietal scales and the anteriormost pair of enlarged 
vertebrals. 

The ventral scales are large and at the level of the abdominal 
glands are roughly disposed in tranverse rows 8 scales in breadth. 
Each ventral scale, except for those covering the abdominal glands, 
bears a small median keel and is drawn out to a point posteriorly. 
The scales covering the abdominal glands are smooth and lack the 
keel and posterior point. There are two large, keeled, preanal 
scales. 

The scales of the limbs are rather strongly keeled, the keels being 
drawn up into almost tubercle-like projections on the scales of the 
upper and outer surfaces of the hind leg. 

The scales of the tail are keeled and arranged in annuli. 

The digits are covered above by single scales throughout most of 
their entire length, although at their base there may be 1 or 2 paired 
scales (Group I of Brongersma, 1942). There are 20-29 (17 on 
one apparently normally developed toe; see Table I) subdigital 
lamellae on the 4th (longest) toe. The lamellae on the basal third 
of the toe consist of paired scales, one scale being large and cover- 
ing most of the subdigital surface and the other scale being small 
and laterally situated. The lamellae on the distal % of the toe are 
smooth and consist of single scales. 

Adult males have both palmar and plantar pores. The palmar 
pores are few in number (2-4), and are distributed in an arc on 
the anterior edge of the palm. The plantar pores (3-10) are dis- 
tributed in two rows across the sole: a longer series from the ex- 
treme base of the 4th toe and a shorter series from the base of the 
3rd toe. In larger males there may be one or two pores on the base 
of the 5th toe. 

Females always lack palmar pores, and whereas all females have 
large plantar scales similar to those bearing pores in males, the 



6 BREVIORA No. 291 

pores are developed in only some females. When present in females 
the plantar pores are less well developed and less numerous (1-6) 
than in males. 

There is a pair of abdominal glands which are covered by 2 
pairs of midventral scales lacking keels and the posterior median 
point. 

Color (Fig. 2): In preservative, specimens are generally brown 
above and on the sides and light yellowish brown below. Most 
individuals are uniformly dark above, but some show various 
degrees of light tan to whitish mottling on the nape. back, and 
upper surface of the base of the tail. The lighter mottled pattern 
tends to be bilaterally symmetrical on the nape and back and is 
often expressed as 1 or 2 transverse, posteriorly projecting chevrons 
on the dorsal base of the tail. 

Variation: There is a slioht difference in the dorsal color of the 
Bougainville and Buka specimens (Fig. 2). Pseudoponceleti from 
Bougainville tend to be darker brown dorsally with less light 
mottling than is found in Buka specimens. In Buka pseudoponce- 
leti the light dorsal color tends to be more pronounced, with a few 
individuals being predominately creamy tan on the body with some 
brown mottling. 

The general differences in color pattern between the Buka Island 
and Bougainville populations are correlated with differences in the 
distribution of certain scale counts between the two populations 
(Table 1). Using the coefficient of difference (CD.) statistic 
(Mayr, et ai, 1953) to determine the degree of joint non-overlap 
between the Bougainville and Buka populations in regard to these 
differences, the CD. was found to be lower in each case than the 
minimum CD. of 1.28 and 90 per cent joint non-overlap conven- 
tion commonly accepted for giving subspecific recognition to two 
populations. These differences do not. therefore, seem to warrant 
subspecific recognition for the Bougainville and Buka populations. 

Distribution (Fig. 1): Tribolonotus pseudoponceleti has been 
collected at several localities on Bougainville as well as on Buka, 
just north of Bougainville (Fig. 1). Altitudinally, the species is 
known from the coastal lowlands up to an elevation of about 4000 
feet (in an area 7-8 miles east of Kunua. Bougainville). None of 
these skinks has ever been taken by Parker or his native collectors 
on the Buin Plain at the south end of Bougainville, however. In 
this region the species was only found from about 2000 feet and 
above in the mountains. 



1968 



NEW SPECIES OF TRIBOLONOTUS 



6°S 



7°S 



155° E 



BUKA I 



156° E 



30 miles 




6°S 



' o ° 



SHORTLAND I S ^^Lofung 



I55°E 



MONO I cC^ 



156° E 



Figure 1. Map of Bougainville and surrounding islands showing the 
known collecting localities for Triholonotiis pseudoponceleti (circles) and 
T. poiiceleti (triangles). 



8 BREVIORA No. 291 

It is curious that the species was not found on Shortiand in the 
Bougainville Straits, approximately 6.1 miles south of Bougain- 
ville. This smaller island is separated from Bougainville by a sea 
passage not more than 25 fathoms in depth and was probably 
connected with the mainland during the Pleistocene. 

Comparisons: As Zweifel (1966) has provided a detailed com- 
parison between cmnectens and pseudoponceleti (under the name 
ponceleti) as well as between cmnectens and other species of the 
genus, the comparisons in this section are limited to cmnectens and 
ponceleti and are intended to complement Zweifel's recent dis- 
cussion. 

In his diagnosis of cmnectens, Zweifel (1966) noted that the 
"ventral scales number 40 in T. cmnectens; 44 to 54 (mean 48.6) 
in 45 specimens of T. ponceleti." This particular difference be- 
tween cmnectens and pseudoponceleti (Zweifel's ponceleti) is, 
however, less clear cut with the additional data now at hand. There 
are, indeed, generally more gular and ventral scales' in Bougain- 
ville (whence came all but one of the specimens of pseudoponceleti 
(42-54, avg. = 48.4) than in annectens (40 in the type, 46 in the 
Copenhagen specimen; avg. = 43.0), but the Buka Island popula- 
tion of pseudoponceleti has a somewhat intermediate number of 
gular and ventral scales (42-49, avg. = 45.0). 

Until now ponceleti was known in the literature only from the 
single individual of the original description. Parker has, however, 
recently obtained two more specimens for the Museum of Com- 
parative Zoology, which have allowed us to make direct compari- 
sons between ponceleti and pseudoponceleti. 

Other than size (see Diagnosis) there are several subtle differ- 
ences in squamation between pseudoponceleti and ponceleti. In 
pseudoponceleti the first supralabial and infralabial, which are 
long and thin, extend posteriorly to at least the level of the anterior 
corner of eye, whereas in ponceleti the first supralabial and infra- 
labial extend posteriorly only to a point about half way between 
the snout and the anterior corner of the eye. The position of the 
first supralabial thus permits the second supralabial to form a 
suture with the nasal in ponceleti, but in pseudoponceleti the first 



^ Counted along the ventral midline from the enlarged, paired chin 
scales to, but not including, the large, paired preanal scales (Zweifel's 
1966 standardization). Presumably Zweifel's use of "ventral scales" in 
the preceding quote of this paragraph is meant to include the gular 
scales also. 



1968 NEW SPECIES OF TRIBOLONOTUS 9 

supralabial meets the loreal, separating the second supralabial from 
the nasal. In pseudoponceleti the enlarged vertebral scales become 
poorly defined just anterior to the level of the insertion of the fore- 
limbs, but in ponceleti the enlarged vertebral scales maintain their 
continuity farther on to the nape. The keels of the vertebral scales 
and two rows of tubercles just lateral to the vertebral scales are 
relatively larger and more pronounced in ponceleti than in pseud- 
oponceleti. The one or two scales situated in the dorsal midline 
between a preceding and following pair of enlarged vertebrals are 
also relatively larger in ponceleti than in pseudoponceleti. There 
are two rows of plantar pores in male pseudoponceleti, one extend- 
ing from the base of the 4th toe onto the sole and the other extending 
from the base of the 3rd toe onto the sole. In male ponceleti there 
is only one row of pores extending from the base of the 4th toe. 

Breeding: T. pseudoponceleti is oviparous and lays but one ob- 
long, leathery, shelled egg in a clutch. Furthermore, the left ovi- 
duct is reduced to a small, vestigial appendage of the cloaca leaving 
the right as the functioning oviduct. In only two of 101 females 
with oviducal and/or ovarian eggs collected at Kunua, Bougain- 
ville, was there a small yolky egg in the reduced left oviduct. These 
2 eggs, however, did not appear to be viable. 

It is noteworthy that the left ovary is still functional. Ten of the 
101 females with oviducal and/or ovarian eggs had a single large 
yolky egg in the left ovary, but in all 1 females the left ovary had 
shifted to the right side, where, presumably, the ovulated eggs 
would pass into the right oviduct, i.e., the only functional oviduct. 
In no case was a large ovarian egg found in the left ovary when it 
was in its normal position on the left side of the body cavity. These 
data would indicate that as yolking proceeds in a follicle of the left 
ovary, the ovary is displaced to the right side, perhaps as a result 
of "crowding" with other internal organs, e.g., the stomach which 
tends to lie more on the left side of the body. 

It would appear that ovulation of a second egg may occur just 
prior to or just after depositon of a preceding egg. This supposi- 
tion is based on observations of single oviducal eggs the same size 
as the largest ovarian egg in the right oviduct just anterior to a 
full term egg in the same oviduct. In other cases there is simply a 
large ovarian egg and no oviducal egg. Such a situation would per- 
tain if a full term oviducal egg had been laid just before ovulation 
of the second egg. 

The 101 females with large ovarian eggs and/or oviducal eggs 
ranged in snout-vent length from 44-58 mm (avg. = 51.8 mm). 



10 BREVIORA No. 291 

The smallest female contained a large, shelled oviducal egg. Given 
the large total sample size, it would appear that 44 mm is near the 
snout-vent length at which females first become capable of repro- 
ducing. 

Ecology: T. pseudoponceleti is sympatric with T. blanc/iardi on 
Bougainville, although the two species occur in rather different 
ecological situations. 

T. blanchardi is a montane species, and is probably to be found 
throughout the mountain ranges in the center of Bougainville. 
Specimens have been collected at 2000-3000 feet in the ranges 
north of Buin. Other specimens from the ranges east of Kunua 
were collected in river valleys from about 500-3000 feet. The 
habitat varies somewhat with altitude. In the lower altitudes where 
the ridges are still warm and comparatively dry, the species lives 
only in the steep sided and heavily shaded small creek valleys. At 
higher altitudes the species occurs farther from the waterways, but 
is still easier to collect along the creeks. The lizard does not live 
in the open in any environment, but prefers tall, shaded, primary 
forest and abundant moisture. In river and creek beds individuals 
are found under stones, logs and leaf rubbish at, and just above, 
water level, but only very occasionally may one be found moving 
on the ground. It often is found in company with Sphenomorphus 
cranei and S. concinnatus. 

T. pseudoponceleti has a greater altitudinal range (coast to above 
4000 feet) than T. blanchardi, but it is found more frequently at 
lower altitudes. T. pseudoponceleti is terrestrial and very secretive, 
living in and under decayed logs in all its types of habitat. It is com- 
mon in swampy areas, and in some places is common on the plains 
where it is found in primary forest and in secondary growth. In 
addition, newly cleared gardens usually contain numbers of these 
skinks. The species likes moist conditions but not the very wet 
conditions preferred by blanchardi. 

In the mountains there is little syntopy between the two species, 
because while blanchardi is confined to creek beds initially, then 
the shaded slopes of the steep ridges, pseudoponceleti is to be found 
only on the sharp ridges where the scrub is more open and the 
ground drier and warmer. This restriction to the ridges has been 
confirmed by extensive collecting east of Kunua from 2000-4000 
feet. A number of other coastal species which range to about 2000 
feet in the mountains are also confined to the drier ridge tops. 

In the Boku area the natives believe pseudoponceleti prefers the 
hairy base of a large ground fern as a dwelling place, but this was 
not borne out by the breaking up of a number of these ferns in a 



1968 NEW SPECIES OF TRIBOLONOTUS 



11 



hunt for the species. The habitat in this area was decaying wood 
on the ground. 

Whether T. ponceleti also occurs on Bougainville along with 
pseudoponceleti and blanchardi is uncertain. The single specimen 
from which the species was described (Kinghorn, 1937) was said 
to have come from "Buin, east Bougainville Island, Solomon Is- 
lands." However, there are no ponceleti among several hundred 
reptiles that Parker and his native collectors obtained in southern 
Bougainville near Buin. 

Upon inquiry, Mr. Harold Cogger of the Australian Museum 
found the notes that accompanied Father J. B. Poncelet's zoological 
collection of which the single specimen of ponceleti was a part. 
The notes consist mainly of a list of the common English and native 
names for each of the specimens and are prefaced by an unequivo- 
cal statement regarding the provenance of the collection — "All 
those {sic) specimens have been collected at Buin (South Bougain- 
ville), 10 to 15 miles in the interior, during the months of Septem- 
ber and October 1934, by Father J. B. Poncelet S.M., of the Buin 
Catholic Mission." 

Native collectors have obtained the only other two known ponce- 
leti at Lofung on Shortland Island (Fig. 1), but whether this large 
Tribolonotus also occurs on Bougainville is still an open question. 

The presence of ponceleti and the absence of pseudoponceleti on 
Shortland Island seems well established. It is also fairly clear 
that pseudoponceleti is absent from the Buin Plain. It is not certain 
whether ponceleti occurs on the Buin plain, although it is fairly 
well established that ponceleti does not occur elsewhere on Bougain- 
ville where large collections have been made (Fig. 1 ). Reasons for 
the absence of pseudoponceleti from the Buin plain and Shortland 
Island are obscure as are the reasons for the absence of ponceleti 
from the rest of Bougainville, if this species occurs on Bougainville 
at all. There seem to be few ecological differences between the 
Buin plain, the other lowland areas of Bougainville where pseudo- 
ponceleti has been collected, and Shortland Island (Parker, per- 
sonal observation). 

As no noticeable ecological differences are known to be corre- 
lated with the ranges of ponceleti and pseudoponceleti, the only 
likely remaining explanation for their allopatry is possibly compe- 
titive exclusion — perhaps of a very dynamic nature. But this facile 
explanation seems implausible, as the great size difference between 
the two species would certainly suggest a difference in the utilization 
of the habitat by the two species. 



12 BREVIORA No. 291 

On the basis of our present information there is Httle more that 
can be said beyond this description of the problem. The ecologic 
and geographic relationships of ponceleti and pseucloponcelcti will 
be interesting field problems for future workers on Bougainville. 

Little seems to have been published on the ecology or habits of 
the closely related species T. gracilis and T. novaeguineae, but Mr. 
Harold Cogger has written to us about his personal experience with 
T. gracilis. Mr. Cogger says, "I am familiar with this species only 
on Karkar Island, where I obtained approximately 90 specimens. 
All specimens collected by me, or in my presence, were found under 
decaying logs and other vegetation usually on the banks of small 
non perennial streams, up to an altitude of about 2,000 feet, usually 
in fairly dense primary or secondary forest." 

Behavior: T. pseudoponceleti is cryptic in its habits and never 
moves about in the open. No lizards were ever collected or seen in 
the open, and none was found moving about at night. In captivity 
they will come into the open parts of the cage to eat if there is little 
or no light. When found in its usual habitat under logs pseudopon- 
celeti makes no attempt to escape but may move away slowly if 
given time. When an individual is uncovered there seems to be an 
instinctive reaction to flatten on the ground to avoid being noticed. 

Usually two adults, a male and a female, are found together, and 
occasionally, if the log is large enough, another pair or two or three 
partly grown specimens may be close by. Rarely is a lone individ- 
ual found — a close search into the ground and log nearby usually 
reveals a second individual. 

T. pseudoponceleti is not strictly speaking a burrower but moves 
through insect burrows and hollows in and under logs. In captivity 
the species will eat termites. The lizards seem to be keen sighted 
and can detect movements in a dimly lit room some 10-15 feet away 
from the cage containing captive skinks. Other species of skinks 
of different genera seemed to be able to detect similar movements 
only when within 5-6 feet of the cage. 

T. blanchardi is very similar to pseudoponceleti in its behavior. 
It moves slowly, although when being chased it will make short 
bursts of speed to escape. In captivity it is not given to the quick, 
nervous movements found in other skinks, e.g., Emoia and Spheno- 
niorphus species, either when frightened or feeding. When moving, 
the skink holds the whole body and sometimes the tail off the 
ground. 

Virtually nothing is known of the ecology or habits of ponceleti. 
The two specimens of ponceleti collected for Parker by the natives 
at Lofuno, Shortland, had dirt in the crevices between the scales 



1968 NEW SPECIES OF TRIBOLONOTUS 



13 



which may indicate that the species is something of a burrower. 
Many pseudoponceleti also have dirt between their scales when col- 
lected and this species is known to spend much of its time in close 
contact with the soil in secretive situations. 

VOLAR PORES AND ABDOMINAL GLANDS 
IN THE GENUS IRIBOLONOJUS 

Among lizards, volar pores and the dermal abdominal glands are 
unique to the genus Tiibolonotus. The presence of volar pores was 
first pointed out by Roux (1930) in male T. schmidti when he used 
them as a character diagnostic of a new genus — Pediporus. Roux 
(1934) later rectified this mistake when he found similar pores on 

some of the toes and sole of a male T. novacguineac the type 

species of the genus Tribolonotus. 

Parker (1940) extended Roux's observations on volar pores of 
gracilis and provided additional information on their distribution in 
schmidti and novaeguineae. Parker also provided the first account 
of abdominal glands in the genus, having noticed them in male 
gracilis and novaeguineae. 

Function: The function of both volar pores and abdominal glands 
is speculative. Secretion seems to be the primary purpose of both 
structures, and from the distribution of the glands on the body of 
the animal — belly and volar surfaces — it would seem as though 
the secretion would be placed in a new part of the microenviron- 
ment with every step taken by an animal. What, then, is the value 
of such a secretion? 

The only information we have on the ecology of Tribolonotus is 
that given above for blanchardi, pseudoponceleti, and gracilis. All 
three species appear to be highly secretive and spend most of their 
time under objects such as stones, leaf rubbish, and rotten logs — 
microhabitats that are both dark and relatively moist, and if the 
secretions are volatile and odiferous, they might be important olfac- 
tory signals. 

Furthermore, the fact that both structures are generallv better 
developed in adults than juveniles and in males rather than females 
suggests that their function may have some sexual significance. 
For instance, the secretions provided by the pores and glands may 
be olfactory markers of territory. 

Distribution of volar pores in Tribolonotus (Table 2): The two 
known specimens of annectens are females and both lack volar 
pores. 



14 BREVIORA No. 291 

Plantar pores occur in both male and female pseudoponceleti 
where they are found in a series behind both the 3rd and 4th toe. 
In some males, but never in females, there may be an additional 
pore or two on the basal third of the 5th toe. Palmar pores are 
present in males but not in females. 

The three known specimens of ponceleti are males and all have 
a single series of plantar pores directly in line with the 4th toe 
(Fig. 3). Palmar pores are absent. 

Both gracilis and novaeguineae females apparently lack volar 
pores. Males of both species have pores on the basal half of the 
3rd and 4th toes and on the sole in a line directly behind the 3rd 
toe. Male gracilis possess palmar pores (personal communication, 
Harold Cogger re Karkar Island gracilis), but whether palmar pores 
are present or absent in male novaeguineae is unclear as yet. 

Female blanchardi also lack volar pores. Males have both 
palmar and plantar pores which are distributed mainly on the palm 
and sole, although occasionally a pore is found on the basal third 
of one of the digits. 

In sclunidti, females lack volar pores, but males have both palmar 
and plantar pores, the latter being distributed in two series: one 
behind the 4th toe and a second, shorter, series behind the 3rd toe. 

Distribution of abdominal glands in Tribolonotus (Table 2): In 
the Copenhagen specimen of the female annectens which we have 
examined, the glands are covered by four pairs of median ventral 
scales. 

Both sexes of pseudoponceleti have a single pair of unlobed ab- 
dominal glands lying beneath two pairs of midventral scales. The 
glands are slightly smaller in females than in males. 

In ponceleti, which is at present known from only 3 males, the 
abdominal glands are bilobed and covered by one or two pairs of 
ventral scales (Fig. 3). 

Female gracilis and novaeguineae lack abdominal glands, but in 
males of both species the glands are paired and unlobed and extend 
a distance of three or sometimes four ventral scales along the mid- 
line. 

Female blanchardi also lack abdominal glands. Males have four 
pairs of bilobed abdominal glands which extend the length of four 
or five ventral scales. 

The glands are very small and minutely lobed in female schmidti, 
but the ventral scales covering the glands are not visibly different 
from the other ventral scales. In males the two pairs of ventral 
scales covering the glands are characteristically smooth along their 
posterior edges rather than mucronate as are the surrounding en- 
larsed ventral scales. 



1968 NEW SPECIES OF TRIBOLONOTUS 15 

REPRODUCTION IN TRIBOLONOTUS 

The mode of reproduction is known for all the species of Tribo- 
lonotus except ponceleti which is known from only 3 male speci- 
mens. From an examination of gravid cmncctens (also see Zweifel. 
1966:2), blanchardi , gracilis and novaeguineae, as well as pseudo- 
ponceleti, it would appear that these species are oviparous and pro- 
duce but a single oblong leather shelled egg in a clutch in which 
little or no embryonic development takes place prior to deposition. 

Schniidti is atypical of the genus in producing living young but 
typical in producing but one young at a time. Roux (1930:133) 
says of a single female he received from Guadalcanal, Solomon 
Islands, "Elle renferme un embryon bien developpe." The adap- 
tive significance of live-bearing habits in schniidti are difficult to 
imagine and will probably remain obscure until more is known of 
the species' behavior and ecology. 

It is interesting to note that blanchardi and schniidti are similar 
to pseudoponceleti in having only one functional oviduct, the 
right. In blanchardi and schniidti as in pseudoponceleti the left 
oviduct is reduced to a small vestigial appendage of the cloaca. In 
blanchardi the left ovary is still functional, and, as in pseudopon- 
celeti, is usually displaced to the right side prior to ovulation. In 
schniidti, on the other hand, the left ovary is either absent or greatly 
reduced in size; in the latter case it is situated well posteriorly in the 
body cavity and is probably not functional. 

Unfortunately the state of preservation of the four gravid gracilis 
and single gravid novaeguineae available for study was such that 
the condition of the left ovary and oviduct could not be interpreted. 

These data are of further interest on two counts. First, it may 
well be that a single egg or young is characteristic of the taxon 
Tribolonotus. Such specificity in the number of eggs produced is 
not unusual in some groups, e.g. geckos, but it is unreported in any 
supraspecific taxa of skinks.^ 

Second, a single functional oviduct has been previously unknown 
in reptiles except in some leptotyphlopids and typhlopids (Guibe, 
1948; Robb, 1960; Fox and Dessauer, 1962). In these "worm 
snakes," as in Tribolonotus, it is the left oviduct that is non-func- 
tional. The left ovary in the leptotyphlopids and typhlopids is 
slighdy smaller in size than the right ovary, although it is still 



'Many Emoia, however, especially the smaller species, have but two 
eggs in a clutch (Greer, 1968). 



16 BREVIORA No. 291 

functional. The left ovary seems to have lost all function in Tribo- 
lonotus sclunidti, but the large ova discovered in the left ovary of 
a few individuals of both blanchardi and pseudoponceleti would 
indicate that this organ is in some cases still functional in these 
species. 

ACKNOWLEDGMENTS 

We would like to thank Mr. Harold Cogger of the Australian 
Museum for his many contributions to the development of this man- 
uscript. Not only did Mr. Cogger supply us with information on 
the type of ponceleti and his field experiences with gracilis, he also 
made many helpful criticisms and corrected several errors in an 
earlier draft of the paper. 

Dr. E. E. Williams of the Museum of Comparative Zoology has 
also read the manuscript in several versions and has smoothed 
over many "rough spots" with each reading. 

Part of the research (done by Greer) for this paper was sup- 
ported by the Evolutionary Biology Fund which is administered 
for the National Science Foundation by the Evolutionary Biology 
Committee of the Biological Laboratories, Harvard University. 

Dr. Richard G. Zweifel of the American Museum of Natural His- 
tory very kindly read the final draft of the manuscript, and Miss 
Nancy Uhlar typed the final version of the paper. 

Publication has been financed by NSF Grant GB 6944 to Dr. 
E. E. Williams. 

A part of Parker's collecting expenses during his 1966 field work 
m the Solomon Islands was covered by a grant from the Science 
and Industry Endowment Fund of the Commonwealth Scientific 
and Industrial Research Organization of Australia. 

LITERATURE CITED 

Brongersma, L. D. 

1942. On the arrangement of the scales on the dorsal surface of the 
digits in L\gosonui and allied genera. Zoologische Mededeelin- 
gen. 24 ( 1-2) : 152-158. 

Fox. W. and H. C. Dessauer 

1962. The single right oviduct and other urogenital structures of 
female Tvplilops and Leptotyphlops. Copeia (1962) No. 3: 
590-597. 

Greer, A. E. 

1968. Clutch size in the genus Einoia. Copeia (1968), in press. 



1968 NEW SPECIES OF TRIBOLONOTUS 17 

GUIBE, J. 

1948. Contribution a Fetude de Fappareil genital des typhlopides 
(Ophidiens). Bull. Soc. Zool. France, 73: 224-228. 

KiNGHORN, J. R. 

1937. A new species of scink from the Solomon Islands. Rec. 
Australian Mus., 20(1 ) : 1-2. 

Mayr, E., E. G. Linsley, and R. L. Usinger 

1953. Methods and Principles of Systematic Zoology. McGraw Hill 
Book Co., Inc., New York, 336 pp. 

Parker. H. W. 

1940. Undescribed anatomical structures and new species of rep- 
tiles and amphibians. Ann. Mag. Nat. Hist., (11) 5: 257-274. 

ROBB. J. 

1960. The internal anatomy of Typhlops Schneider (Reptilia). 
Australian J. Zool.. 8(2): 181-216. 

Roux, J. 

1930. Note sur un reptile scincide des lies Salomon presentant des 
pres pediaux. Verhandl. Naturforsch. Ges. Basel, 41: 129-135. 

1934. Contribution a la connaissance de la faune erpetologique des 
lies Salomon. Verhandl. Naturforsch. Ges. Basel, 45: 77-81. 

ZWEIFEL, R. G. 

1966. A new lizard of the genus Triholonotiis (Scincidae) from New 
Britain. Amer. Mus. Novitates, No. 2264: 1-12. 

(Received 21 November 1967.) 



18 BREVIORA No. 291 

TABLE 1 

Frequency distribution for several meristic characters in the two 
island populations of Tribolnotus pseiidoponceleti 

BOUGAINVILLE 

Enlarged, paired middorsal scales 
X 21 22 23 24 25 

f(x) 1 3 32 64 13 

Gular + ventral scales 
X 45 46 47 48 49 50 51 52 53 54 

f(x) 8 8 17 26 22 10 9 1 3 2 

Subdigital lamellae 4th toe 

X 17 20 21 22 23 24 25 26 27 28 29 N = 211 

x = 23.9 
f(x) 1 2 18 24 44 41 44 18 15 3 1 cr = 1.83 

BUKA ISLAND 

Enlarged, paired and middorsal scales 



f(x) 



f(x) 



6 
1 


N - 114 
x = 23.8 
a - 0.75 




N= 106 
x = 48.4 
<7= 1.97 



f(x) 



22 




23 


24 25 






N=:22 

x = 23.8 


1 




6 


12 3 

Gular + ventral scales 






<T = 0.77 


42 


43 


44 


45 46 47 48 49 






N = 22 
x = 45.0 


1 


6 


3 


3 3 4 11 






<r= 1.95 








Subdigital lamellae 4th toe 






20 




21 


22 23 


24 


25 


N = 42 
xi=21.8 


3 




14 


15 8 


1 


1 


a- 1.06 



1968 NEW SPECIES OF TRIBOLONOTUS 19 



TABLE 2 



Distribution of volar pores and abdominal glands in the sexes of 
the seven known species of Tribolonotus 





Palmar 

c5 


pores 


Plantar Pores 

$ 2 


Abdominal 


glands 
5 


annectens 


? 


— 


? 


— 


7 


+ 


pseitdoponceleti 


+ 


— 


+ 


+ 


+ 


+ 


ponceleti 


— 


? 


+ 


7 


+ 


7 


gracilis 


+ 


— 


+ 


— 


+ 


— 


novcieguineae 


7 


— 


+ 


— 


7 


— 


hhuichanli 


+ 


— 


+ 


— 


+ 


— 


scliDiidti 


+ 


— 


+ 


— 


+ 


+ 



TABLE 3 

Species, number of specimens, snout-vent length, number of eggs/ 
female, and comments on reproductive state of gravid Tribolonotus 

examined 

Snout- 
Number vent Number 
of gravid length of 
Species 9 9 examined ( mm ) eggs/ 9 Comments 

annectens 2 49-50 1 "One large egg evidently nearly 

fully developed" in the type 
(Zweifel, 1966). Second egg in 
Copenhagen specimen with 
thick shell. 

pseudoponceleti 101 44-58 1 Yolky ovarian eggs to large, 

leathery-shelled oviducal eggs. 

gracilis 4 87-100 1 1 yolky ovarian egg; 3 large. 

leathery-shelled oviducal eggs. 

novaegidneae 1 75 11 leathery-shelled oviducal egg. 

blanchardi 9 31-36 1 Yolky ovarian eggs to large, 

leathery-shelled oviducal eggs. 

schmidti 12 34-38 1 Yolky ovarian eggs to terminal 

young. 



20 BREVIORA No. 291 



Figure 2. Paratypes of Triholonotiis pseudoponceleti from Bougainville 
(MCZ 72914) and Buka ( MCZ 67713), exemplifying the differences in 
dorsal coloration between the two island populations. 



1968 



NEW SPECIES OF TRIBOLONOTUS 



21 




22 BREVIORA No. 291 



Figure 3. Tribolonotiis poncclcti from Lofung, Shortland Island. Notice 
the pair of ventral scales overl\ing the abdominal glands, and the plantar 
pores in the ventral view of the specimen. 



1968 



NEW SPECIES OF TRIBOLONOTUS 



23 




"■^rT' '  



B R E V I O R A 

Miiseiuim of Comparative Zoology 

Cambridge. Mass. 29 May. 196 8 Number 292 

HERPETOGEOGRAPHY OF PUERTO RICO 

V. DESCRIPTION OF A NEW SPECIES 

OF SPHAERODACTYLUS 

FROM DESECHEO ISLAND 

Harold Heatwole ' 



INTRODUCTION 

The Puerto Rican island sheif (Greater Puerto Rico) has re- 
ceived considerable attention from herpetologists, and the herpeto- 
fauna of the larger islands is now reasonably well known. However, 
the small islands and keys, with some exceptions, have not been 
studied in a thorough way. A faunal survey of these islands is now 
in progress (Levins and Heatwole, 1963). and some of the results 
have been published (Heatwole et ciL, 1963, 1965; RoUe ct al., 
1964; Heatwole and Torres. 1967). The present paper represents 
a further contribution in that direction. 

ACKNOWLEDGMENTS 

I am indebted to many students and staff members of the Uni- 
versity of Puerto Rico for their assistance in the field, to Audry 
Heatwole for aid in preparation of the manuscript, and to the Na- 
tional Science Foundation for financial support of the project 
through Grant GB-2906. 

DESECHEO ISLAND 

Desecheo is a hot. dry. relatively steep-sided island located in the 
Mona Channel 12 miles west of Punta Higuero. P. R. It consists 
largely of volcanic rock, although some limestone deposits are pres- 
ent. It has a diameter of 1 mile and a maximum elevation of 715 
ft. (U. S. Dept. Comm. Coast and Geodetic Survey. 1962). The 
vegetation is cactus scrub and xeric woodland. Large numbers of 

1 Dept. Zoology, Univ. New England, Armidale, N. S. W., Australia. 



2 BREVIORA No. 292 

Boobies nest on the island; terrestrial hermit crabs form a conspic- 
uous faunal element. Though uninhabited by humans, a small herd 
of feral goats is present. 

Its known herpetofauna consists of 4 species: Alsophis portori- 
censis (Grant. 1932), Aineiva dcsechensis (Heatwole and Torres, 
1967), an Anolis reported by Wetmore (1918) to be intermediate 
between A. cristatcUus and A. moneusis (now being studied by the 
author), and the new species of Sphaerodactylus described below. 

No specimens of Sphaerodactylus from Desecheo were avail- 
able to Thomas and Schwartz (1966), and they singled out this 
island as "one of the few West Indian islands from which Sphaero- 
dactylus has not been reported." The present study fills in this gap. 
All scale counts were made after the method of King ( 1962). 

Sphaerodactylus levinsi sp. nov.' 

Holotype: UPRRP 4566 (donated to the Museum of Compara- 
tive Zoology, Harvard University, MCZ No. 100274), a male taken 
28 May 1965 on Desecheo Island, Puerto Rico, by the author. 

Paratypes: UPRRP 4565, 4567, 4570 (males), and UPRRP 
4571-72 (females), collected 27-29 May 1965 on Desecheo Island, 
P. R., by the author. R. Levins, and F. MacKenzic. 

Distribution: Known only from the type locality, Desecheo 
Island, where it is not abundant; found chiefly under stones and 
dead wood at the edges of dry drainage gullies. 

Diagnosis: A species of Sphaerodactyhis characterized by: (1) 
a prominent light-colored frontal bar, bordered by brown com- 
pletely separating it from other light-colored areas (Fig. I), (2) 
shoulder patch not bordered by a lighter color but enclosing two 
white, round ocelli, (3) lack of sexual dichromism. (4) scale rows 
around mid-body 46-48, and (5) lamellae under 4th toe 7-8. 

Description of holotxpe: An adult male with snout-vent length 
of 28.2 mm; dorsal scales axilla to groin 28, ventral scales axilla 
to groin 31; mid-body rows of scales 48; 4th toe lamellae 8; inter- 
nasals 1; supralabials to mid-eye 3; escutcheon 5 by 17 scales; dor- 
sal scales keeled, imbricate, no microscopic hairs or knobs detected; 
no granular dorsal scales; ventral scales smooth, nearly round. 



' This species is named in honor of Dr. Richard Levins, who was a stimu- 
lating and pleasant companion on expeditions to more than 100 islands and 
keys of the Puerto Rican shelf, and who contributed to this and other studies 
by collecting many herpetological specimens. The abbreviation UPRRP 
refers to the collections of the University of Puerto Rico (Rio Piedras). 



1968 A NEW SPHAERODACTYLUS 3 

venter dirty white; dorsal ground-color brown with darker brown 
spots tending to be aligned in longitudinal rows; a U-shaped, dark 
brown mark over sacrum; head pattern with a dark brown medial 
preorbital stripe diverging anterior to the eyes to enclose a longi- 
tudinal, conspicuous, light-colored bar (frontal bar), extending 
nearly to the occipital region, where the brown stripes re-converge, 
isolating bar from other light-colored areas; lateral, dark brown 
preorbital stripes diverge around eye and extend separately across 
the temporal region to break up into spots near the shoulder; promi- 
nent, round dark brown occipital spot followed posteriorly by two 
pairs of faint spots; no well-defined nuchal spot; oval, dark brown 
shoulder patch enclosing a pair of light-colored ocelli. Ocelli and 
frontal bar are near-white and are conspicously lighter than any 
other features; occipital spot and shoulder patch (near-black) con- 
spicuously darker than any other feature. 

Variation: Coloration was extremely uniform, and the descrip- 
tion of the holotype could serve almost equally well for any of the 
other specimens examined, including both sexes, except for ( 1 ) 
varying tendencies of the lateral head-stripes to coalesce with the 
faint spots behind the occipital spot, and (2) a narrow line run- 
ning anteriorly from the occipital spot to connect with the brown 
color enclosing the frontal bar in two specimens (males). Escutch- 
eons were 4-6 by 12-17 in scale number; 4th toe lamellae were 7 
or 8; upper labials were 3 or 4; internasals did not vary (always 
1); scale rows around mid-body were 48 with one exception (46); 
dorsal rows axilla to groin varied from 27 to 32 (mean 29.0; stand- 
ard error 0.82) and ventral ones axilla to groin from 28 to 32 
(mean 29.8; standard error 0.66). 

Relationships: S. levinsi is most closely related to S. monensis 
and S. macrolepis. It is distinguished from S. tnacrolepis by its 
higher number of dorsal scales, its lower number of toe lamellae, 
and by lacking sexual dichromism. It differs from S. monensis by 
lacking a light border around the scapular patch and by having 
round rather than transverse ocelli. The conspicuous frontal bar 
( Fig. 1 ) is not shared with either species. 

Thomas and Schwartz (1966) have suggested that with the pos- 
sible exception of S. beattxi of St. Croix all the Sphaerodactylus on 
the Puerto Rican shelf are the result of radiation from a single 
ancestral invading stock. According to their scheme, a proto- 
niacrolepis form separately gave rise to S. roosevelti and S. monen- 
sis, before continuing on to modern macrolepis, which in turn gave 
rise to two other species groups as well as a variety of subspecies. 



4 BREVIORA No. 292 

S. monensis is now found only on Mona Island (about 40 miles 
west of Puerto Rico), and S. leviusi only on Desecheo Island (12 
miles west of Puerto Rico). I suggest that the proto-niacrolepis 
ancestor succeeded in reaching Mona and Desecheo. where it sep- 
arately gave rise to the two present-day forms. Heatwole and 
MacKenzie (1967) have shown that during the Pleistocene, con- 
ditions were much more favorable for flotsam transport from 
Puerto Rico to both Mona and Desecheo than they are now. The 
southwestern coast of Puerto Rico extended considerably farther 
into the Mona Channel, thereby reducing inter-island distances and 
also dellectinii currents in more favorable directions. At maximum 
extension, the current passing westward along the southern coast 
of Puerto Rico would have proceeded almost directly toward Mona, 
and it was probably at this time that prolo-/nacrolepis arrived there. 
As the southwestern coast of Puerto Rico subsequently receded, 
it would have permitted the current to swing increasingly toward 
its present northward direction through the Mona Channel. Then 
Desecheo, rather than Mona, would have been in the more direct 
path of the current. Land configuration was also conducive to 
eddy currents between Desecheo and Punta Higliero, Puerto Rico. 
On the one hand, this probably caused increased isolation of the 
Mona population, permitting its differentiation into S. monensis, 
and on the other, made possible flotsam transport of proto-macro- 
lepis to Desecheo. Gene flow via flotsam transport of animals was 
probably maximal between Puerto Rico and Desecheo at that time, 
as further recession of the southwestern coast increased the effective 
inter-island distance and resulted in currents less favorable for 
flotsam transport between them. Today, any flotsam reaching Des- 
echeo would have its most likely origin on the extreme southwest- 
ern tip of Puerto Rico, rather than geographically nearer points. 
This increased isolation undoubtedly facilitated divergence of the 
Desecheo population to form S. levinsi. 

Additional evidence for the isolation of the Mona population be- 
fore that of Desecheo is that the former has retained a light border 
around the scapular patch, a character no longer present in the 
western populations of S. macrolepis or in 5. levinsi. 



1968 A NEW SPHAERODACTYLUS 5 

REFERENCES CITED 

Grant, C. 

1932. The genus Alsnphis in the Puerto Rico area. J. Dept. Agric. 
Puerto Rico, 16: 149-151. 

Heatwole, H., and F. Mackenzie 

1967. Herpetogeography of Puerto Rico. IV. Paieogeography, fauna! 
similarity and endemism. Evolution, 21: 429-438. 

Heatwole. H., D. S. Sade, and R. Hildreth 

1963. Herpetogeography of Puerto Rico. I. Herpetofauna of Cayo 
Santiago and Cayo Batata. Carib. J. Sci., 3: 1-5. 

Heatwole, H., and F. Torres 

1967. Distribution and geographic variation of the ameivas of Puerto 
Rico and the Virgin Islands. Herpetogeography of Puerto Rico 
III. Studies Fauna Curasao, 24: 63-1 1 1. 

Heatwole, H., F. Torres, and A. Heatwole 

1965. Herpetogeography of Puerto Rico. II. Distributional records for 
some of the outlying islands. Stahlia, No. 4: 1-4. 

King, W. 

1962. Systematics of Lesser Antillean lizards of the genus Sphaero- 
ilactyliis. Bull. Florida State Mus., 7: 1-52. 

Levins, R., and H. Heatwole 

1963. On the distribution of organisms on islands. Carib. J. Sci., 3: 

173-177. 

RoLLE, F. J., H. Heatwole, R. Levins, and F. Torres 

1964. Faunal notes on Monito Island, Puerto Rico. Carib. I. Sci., 4: 
321-322. 

Thomas, R., and A. Schwartz 

1966. Sphaerodactyliis (Gekkonidae) in the Greater Puerto Rico 
region. Bull. Florida State Mus., 10: 193-260. 

U. S. Dept. Comm. Coast and Geod. Survey 

1962. United States Coast Pilot 5. Atlantic Coast, Gulf of Mexico, 
Puerto Rico and Virgin Islands. 5th ed. U. S. Gov't Printing Off., 
Washington, D. C, 264 pp. 

Wetmore, a. 

1918. The birds of Desecheo Island, Puerto Rico. Auk, 35: 333-340. 

(Received 21 November 1967.) 



BREVIORA 



No. 292 




Figure 1. Head of holotype of Sphaerodactylus levinsi. 



B R E V I O R A 

Meseiuiim of Comparative Zoology 



Cambridge, Mass. 29 May. 1968 Number 293 

INTRA- AND INTERSPECIFIC CHROMOSOME VARIATION 

IN THE LIZARD ANOUS CRISTATELLUS AND ITS 

CLOSEST RELATIVES 

George C. Gorman,' Richard Thomas,'- and Leonard Atkins"' 



INTRODUCTION 

Widespread on the Puerto Rican bank (Puerto Rico, its offshore 
keys, and the Virgin Islands, excluding St. Croix) is the common 
lizard Anolis cristatellus. Data on its ecological distribution have 
been summarized by Rand (1964). The last taxonomic revision 
(Grant, 1931 ) recognized three subspecies: A. c. cristatellus, found 
virtually throughout Puerto Rico, A. c. wileyae^ from the islands 
east of Puerto Rico, including both the adjacent keys and the 
Virgin Islands, and A. c. cooki from the extremely dry, desert-like 
southwestern part of Puerto Rico. Thomas (1966) used the com- 
bination "Anolis cooki" in a discussion of the endemicity of the 
herpetofauna of southwestern Puerto Rico but did not justify his 
usa^e. Anolis monensis on the island of Mona to the west of Puerto 
Rico has been considered a separate species, but it is very close to 
cristatellus and its taxonomic status needs reassessment. Unfor- 
tunately, specimens of this species were unobtainable for chro- 
mosomal analysis. 

1 Present address. Museum of Vertebrate Zoology. University of Califor- 
nia, Berkeley. California 94720. 

■-' Department of Zoology, University of South Florida. Tampa. Florida 
33620. 

;^ Department of Pathology and the Joseph P. Kennedy, Jr., Laboratories 
of the Department of Neurology, Massachusetts General Hospital. Boston. 
Massachusetts 02114. 

^ Ahhough described by Grant as Anolis cristatellus wileyi, the name 
must be changed to take the feminine ending, as the subspecies was named 
after Grace Olive Wiley (Grant. 1931). (See articles 31 and 32, Interna- 
tional code of Zoological Nomenclature.) 



2 BREVIORA No. 293 

Evidence that cooki is a separate species was obtained by Albert 
Sciiwartz and David Leber in 1961 when they collected two kinds 
of cristatellus-Wke anoles on the islet of Caja de Muertos off the 
south coast of Puerto Rico. Later, Thomas observed that where 
cooki is sympatric with crisuitellus, along coastal areas of southwest 
Puerto Rico, it appears to split the structural niche (terminology 
from Rand, 1964) by perching lower than cristatellus, often on 
rocks or on the ground rather than in trees. Although morphologic- 
ally very similar to cristatellus, cooki has larger dorsal scales (Fig. 
1). It also has lower 4th toe lamellar counts and keeled ventral 
scales, and in coloration it is typically paler than cristatellus and 
has a redder dewlap. The morphological differences are most valid 
when contrasted with sympatric cristatellus. For these reasons and 
for reasons presented below, we consider cooki a separate species. 
Independently, E. E. Williams has come to similar conclusions by 






Anegada 

Virgin Gorda   ■l»l -.l  

Tortola -   I !■_ 

St. John   



St.Thonnas 

Culebra 

Vieques 



■■iiili..!,  



PR Cays ,-iliill.  



ljilUjiuIui^^ 



A.c.cristatellus 

A. monensis ■- ■■■■!■ u 



A. cooki 



lmJ»Ij_ 



A.scriptus 




30 40 50 60 

Figure 1. Dorsal scale counts (taken dorsolaterally in the standard dis- 
tance) for Aiiolis cristatellus and its closest relatives. The samples for 
Tortola and St. Thomas include specimens from satellite islets: specimens 
examined are in the Albert Schwartz Field Series and the Museum of Com- 
parative Zoology. The smallest vertical units represent single individuals. 



1968 ANOLIS CRIST ATELLUS 3 

comparison of paratypes of cooki with cristatellus from nearby 
localities. 

The final species to be considered is A . scriptus, from the south- 
ern Bahamas. This species had been synonymized with cristatellus 
by Barbour (1914), but Rand (1962) showed that there were 
minor scale diflferences. He considered scriptus a valid species 
although close to cristatellus. Of the four described subspecies, all 
were recognized by Rand, and two were available for chromosomal 
analysis. 

The forms discussed here are distinguishable by differences in 
coloration, especially dewlap color; their external morphology, 
although not always diagnostic, shows interesting regional variation. 
The variation in dorsal scale counts of cristatellus ( Fig. 1 ) may be 
taken as a paradigm of trends shown by other morphological 
characters in this species. Tail crest height, degree of ventral scale 
keeling, and counts of 4th toe lamellae and loreal scales show the 
same pattern of variation, although the trends are generally less 
pronounced than are those shown by the dorsal scale counts. A. 
cooki and A. nionensis also follow this trend in having low loreal 
and lamellar counts (cooki is extreme in its low number of lamel- 
lae). It is of interest that the cooki and eastern populations of 
wileyae agree with one another in having large dorsal scales, few 
4th toe lamellae, few loreals. and relatively pronounced ventral 
keeling. This may be part of a similar adaptive constellation of 
characters; it should be noted that all of the small island forms 
(sensu lato: including cooki) show tendencies towards enlarged 
dorsal scales in comparison with nominate cristatellus. 

All of these lizards belong to the cristatellus series, following the 
classification of Anolis into species series by Etheridge (1960). 
Also included in this series are four other Puerto Rican species: 
krugi. pulchellus, poncensis (grass anoles), and gundlachi, a species 
which resembles cristatellus and appears to replace it ecologically 
at high altitudes (Rand, 1964); and three species of Hispaniolan 
anoles, cvbotes, whitemani, and shrevei, which appear very closely 
related inter se and seem to resemble most closely A. cristatellus. 

Considerable data have already been presented on the chromo- 
somes of Anolis. A karyotype consisting of six pairs of metacentric 
macrochromosomes and twelve pairs of microchromosomes 
(2n=36) is found in numerous species of Anolis and other iguanid 
lizards. This formula is considered primitive for the family Iguani- 
dae (Gorman. Atkins, and Holzinger. 1967). Lizards with this 
karyotype lack obvious sex chromosomal heteromorphism, as il- 
lustrated by A. cybotes, a Hispaniolan member of the cristatellus 



4 BREVIORA No. 293 

series (Gorman and Atkins, 1966, fig. 1 d-f). Other anoles are 
characterized by a reduced diploid number, and a complex sex 
chromosome system in which males are XiXoY and females 
XiXiX-Xj (Gorman and Atkins, 1966). The karyotypes of the 
four Puerto Rican cristatellus group anoles that have already been 
reported on (pulchelliis, poncensis, krugi, and gundlachi) are of this 
sort. The karyotypic difference between the Puerto Rican and His- 
paniolan members of the cristatellus series and its phylogenetic 
implications are discussed elsewhere (Gorman and Atkins, in 
press) and will not be repeated here. 

AAATERIALS AND METHODS 

Chromosome spreads were obtained by direct preparation of 
gonads or by whole blood tissue culture using a modification of the 
technique of Moorhead. et cil. (1960). Techniques were outlined 
in Gorman and Atkins (1966). Chromosome data were obtained 
from 19 A. c. cristatellus from Puerto Rico, 14 A. c. wileyae from 
the islands east of Puerto Rico, 6 A. cooki from southwestern 
Puerto Rico. 3 A. scriptus leucophaeus from Great Inagua. and 
one A. scriptus mariguanae from Mayaguana. Figures 2-4 show 
the localities sampled in this study. For comparative purposes, data 
are presented on two additional Puerto Rican members of the 
cristatellus series. 



/' '""■"■^-— ^ 


^^ ~_ 


-^ 


"^®©^ 


">S 


1 


\D 


PUERTO 


RICO 






^ 


ijL.^-vSS^ 




Vx-x. 


^>-/-^— ^^ 






m^^\\) 













Figure 2. Localities on Puerto Rico sampled for A. c. cristatellus (circles) 
and A. cooki (squares). The number within each symbol is the number 
of specimens sampled for that locality. 



968 



ANOLIS CRISTATELLUS 



CAYO ICACOS ( 2 1 

CAYO LOBOS ( 4 ' 



JSLETA MARINA 

'( I ) 



( I ) 
HANS LOLLIK 





. / J>^^ VIRGIN 'GORDA 



V ^-""^ BT' 



ST. THOMAS 
( I ) 




VIEQUES 



Figure 3. Map of the eastern portion of the Puerto Rico bank; islands 
where specimens were sampled are named. Number in parentheses is the 
number of specimens from each locahty. 




Figure 4. Map of the southern Bahamas and eastern Greater Antilles. 
The area enclosed in solid black represents the distribution of A. scriptiis. 
Inagua and Mayaguana are the localities sampled. Dotted areas represent 
submerged island banks. Arrows show general trend of the currents. 



BREVIORA No. 293 



RESULTS 



Male meiosis. The four species in the cristatellus group previ- 
ously studied (gundlachi, pulchellus, poncensis, krugi, see Gorman 
and Atkins, in press) are all very similar to one another. In diaki- 
nesis there are six large bivalents, two intermediate-sized bivalents, 
five small bivalents (13 autosomal bivalents in all), and a sex 
trivalent. Figure 5a shows this in A. gundlachi. Among the forms 
reported on here, only A. cooki has 13 autosomal bivalents and a 
sex trivalent. A. cooki differs from gundlachi and pulchellus only 
in the detail that the smallest bivalents do not break sharply in size 
when compared with the intermediates (see Figure 5b). 

All A . c. cristatellus and both the scriptus subspecies differ from 
cooki and the other members of the cristatellus series by having one 
fewer autosomal bivalent. Again there are six macrobivalents; 
however, there are only six smaller ones, which apparently can 
be divided into two classes of three intermediate-sized and three 
small, but certainly with gradation (Figs. 5c, d). 

It is among the offshore populations of A. cristatellus wileyae 
that we have made the most unusual finding. Twelve of the animals 
sampled appeared identical to the Puerto Rican cristatellus and 
Bahaman scriptus, but two others differ in having a moderate-sized 
body that might be a univalent. One of these animals is from the 
small key Cayo Lobos, the second is from Vieques. Three of the 
four males sampled from each of the two islands had normal 
meiosis (Fig. 6). 

Mitosis. The mitotic metaphase of males of cooki is characterized 
by 29 chromosomes. There are six pairs of metacentric macro- 
chromosomes and 1 7 smaller elements. Resolution of these ele- 
ments is difficult, but there appear to be three pairs of small meta- 
centric chromosomes, four dot-like pairs (presumably acrocentric), 
and three unpaired chromosomes, the largest of which is clearly 
metacentric and is presumably the Y (Fig. 7b). Females have 30 
chromosomes and lack the unpaired metacentric. The male karyo- 
type is similar to pulchellus (Fig. 7a), but again, as in meiosis, 
the break between size classes of chromosomes is greater in pul- 
chellus. There are six pairs of metacentric macrochromosomes, two 
intermediate pairs of metacentric chromosomes, five pairs of micro- 
chromosomes, and, in the male, three unpaired chromosomes 
(2n=29). 

A. scriptus and A. c. cristatellus have male diploid numbers of 
27; female scriptus have 2nr=28 (female cristatellus were not 
studied). The male diploid number of 27 is also found m A. c. 
wileyae (one exception will be discussed below). 



1968 



ANOLIS CRISTATELLUS 




Figure 5. Meiosis in cristatellus group anoles. Giemsa stain. 

A. A. ^iiiidUichi. There are six large hivalents. two of intermediate size, 
and five distinctly smaller bivalents; a sex trivalent is on the far right. 

B. A. cooki. There are six large bivalents. seven ranging from interme- 
diate to small size with no sharp break between them; and a sex trivalent. 

C. A. c. cristatellus. There are six large bivalents, and only six ranging 
from intermediate to small size with no sharp break between them: and a 
sex trivalent. 

D. A. scripliis Icucophaeiis. Quite similar to cristatellus directly above. 



In details of karyotype, cristatellus and scriptus are very similar. 
There are six pairs of metacentric macrochromosomes and six 
autosomal pairs of small chromosomes, of which the largest three 
pairs are also metacentric. Males have three unpaired sex chromo- 
somes (Figs. 7c, d). 

The one exceptional animal was the A. c. wileyae from Vieques 
which had the extra body (presumed univalent) in meiosis (no 
mitotic divisions were seen in the specimen from Cayo Lobos that 



8 BREVIORA No. 293 



Figure 6. Meiosis in A. cnstatelliis wileyae. Giemsa stain. 

A and B. Two different males from the island of Vieques. Male A is 
identical to typical crisicitellus, male B has a dark staining supernumerary 
body (arrow). 

C. A male from Cayo Lobos that also shows a dark staining supernu- 
merary body (arrow). 

D. A male from St. Thomas, Virgin Islands, which has the typical 
cristate'lliis complement. 



had the same meiosis). Only 10 mitotic spreads could be counted 
from the testis of this animal, of which 4 had 27 chromosomes 
(the same as other cristatellus) and 6 had 28, or one extra chromo- 
some. The extra chromosome appears to be a microchromosome. 
Possibly, the animal is a 27/28 mosaic. However, interpretation 
is difficult. Because of the small size of the microchromosomes, it 
is easy to have one masked by a large arm, and it is indeed possible 
that the real diploid number is 28. Ten counts are far too few 
to establish the karyotype with certainty. It should be emphasized 
that the two chromosomally aberrant wileyae did not differ mor- 
phologically from the other lizards sampled. 



[968 



ANOLIS CRISTATELLUS 




imi"'"' 



9Sli# •!»• 



• •■ 



• fl 



B 




m^ Jlii ^tS 



m ii ♦,* • « • * * 



c 




ll K H « « <«►«» .^ « • • 



m^ ^ 



D 




Figure 7. Mitosis in males of ciistatelliis group anoles. Giemsa stain. 

A. Aiiolis pulclieUits (2n=29). There are six pairs of metacentric mac- 
rochromosomes (top row), two pairs of small metacentric chrcmoscmes 
(far left of second row), a sharp break between these and the five pairs 
of microchromosomes, and three unpaired sex chromosomes. 

B. A. cooki (2n=:29). Macrochromosomes as above (top row). Pairs 
seven and eight, small metacentrics, are as in piilchelliis, but there is no 
sharp break between pair eight and pair nine. There are again three un- 
paired sex chromosomes. 

C. A. ciistatelliis wileyae (2n=27). Macrochromosomes as in the 
above species. Microchromosomes most similar to cooki. but one pair 
fewer. Sex chromosomes similar. 

D. A. SLiiptiis leiicophacns (2n=27). Resembles c;/.s7{(/(7////s. 



10 BREVIORA No. 293 

DISCUSSION 

The chromosomal data support the hypothesis that cooki is a 
species distinct from cristatellus. This is in accord with observations 
made by Thomas in the field. There are two alternative explanations 
which may account for the observed chromosomal complements of 
these two species and their present-day distribution: 

\. A. cristatellus has the lowest diploid number in its species 
group. Reduction in chromosome number is usually the derived 
condition and. if this be so in this case, cristatellus is a derived form. 
Chromosomal heterozygotes experience meiotic difficulties, lead- 
ing to reduced fitness; the resultant structural mutants would be 
swamped in large populations. Chromosome structural changes, 
therefore, are probably established in small isolated populations. 
Thus it is logical to assume that the stock leading to cristatellus 
evolved in the limited population of a small island off Puerto Rico. 
Once chromosome loss had been established, cristatellus reinvaded 
the main island, successfully filling the lowland arboreal niche 
throughout Puerto Rico. 

A. cooki has a very restricted range (southwestern Puerto Rico) 
and is specialized for a very dry rocky area. One hypothesis ac- 
counting for its origin is that it may have been isolated from its 
ancestral stock on an insular region corresponding to the present 
coastal hills of southwestern Puerto Rico. This stock, quite pos- 
sibly the same one that gave rise to cristatellus (for the two species 
are remarkably similar in appearance), might have had the prim- 
itive karyotype for the species group. 

The diploid number of cooki (29) is the same as that of gund- 
lachi and the grass anoles of the cristatellus series, but there are 
some differences in the details of karyotype. The six pairs of macro- 
chromosomes are comparable in all the forms. In cooki there is 
then a gradation from pairs 7 to 13. of which the largest three are 
metacentric (see Fig. 7), while in the grass anoles and gundlachi 
pairs 7 and 8 are medium-sized metacentrics which break sharply 
in size with the smaller pairs (9-13). 

With respect to the microchromosomes, cristatellus resembles 
cooki in that pairs 7, 8, and 9 are all metacentric and grade in size 
into the smaller elements. The major difference, of course, is that 
cristatellus has one fewer pair of microchromosomes. 

Hence we might postulate that cooki, although morphologically 
specialized, retains the primitive karyotype. A. cristatellus, then, 
is similar but has suffered chromosome loss, while A. gundlachi 
and the three grass anole species are the products of a single 



1968 ANOLIS CRISTATELLUS 11 

radiation in which a new karyotype was established by reciprocal 
translocations from a species with the primitive (cooki-Uke) karyo- 
type. 

2. An alternative hypothesis would consider the gundlachi-type 
of karyotype primitive for the species group. A. cristatellus would 
again have been derived from a 2n=29 ancestor by chromosome 
loss, followed by minor rearrangements. The apparently closely 
related A. cooki might then have been directly derived from cris- 
tatellus by centric fission, which secondarily raised the male diploid 
number back to 29. 

Karyotypic data, in any case, show that giindlachi is closer to 
the grass anoles than it is to cristatellus. This is not obvious from 
external morphology but is in accord with serum protein data 
(Maldonado and Ortiz, 1966) and osteological data (Etheridge, 
1960). In karyotype, gundlachi is virtually indistinguishable from 
the grass anoles. 

It is a further point of interest that the chromosomal evidence 
confirms the close relationship of scriptus and cristatellus. No 
other anoles of the 70 species and subspecies that we have studied 
have a male diploid number of 27, and even details of karyotype 
are identical. 

From a glance at a map, one misht expect that the southern 
Bahamas would be colonized from Hispaniola rather than Puerto 
Rico. However, as E. E. Williams pointed out to us. the general 
current flow is northwesterly, and there are several barely sub- 
merged banks between Puerto Rico and the southern Bahamas, 
banks which may have served as stepping stones for colonization. 
It is therefore not surprising to find a Puerto Rican derivative on 
these islands (see Fig. 4). The widespread Hispaniolan member 
of the cristatellus group, A. cy botes, has a diploid number of 36 
in both males and females with no heteromorphism and could not 
possibly be the direct ancestor of scriptus. 

The finding of apparent supernumerary chromosomes in two 
island populations of A. c. wileyae is of cytological interest, but 
lack of data at the present time on the extent of this variation 
within the populations and on the behavior of the supernumerary 
precludes discussion. 

ACKNOWLEDGMENTS 

We thank Dr. E. E. Williams for his constant interest and advice 
during the course of this work. Mr. W. P. Hall, III, has carefully 
read the manuscript and provided valuable criticism. Dr. Albert 



12 BREVIORA No. 293 

Schwartz supported field work in Puerto Rico done by himself 
and Thomas jointly and by Thomas alone. Specimens from the 
small keys ofT Puerto Rico were obtained through the aid of Drs. 
H. Heatwole and R. Levins, and Mr. F. MacKenzie, in conjunction 
with their work supported by NSF Grant GB-2906. Specimens 
were collected by the senior author on field trips sponsored by 
NSF Grant GB-2444 to Dr. E. E. Williams and the Evolutionary 
Biology Committee of Harvard University. Additional specimens 
were sent by Mr. A. Laska. Mr. R. McAndless, Mr. D. Norton, 
Mr. C. R. Warren, and Miss A. Swidler. Laboratory work was 
sponsored by Children's Bureau Project No. 906. We thank Mrs. 
C. Kayavas and Miss L Leone for technical assistance, and Mrs. 
P. Kerfoot for the illustrations. Additional support was provided 
by NSF Grant GB-6944 to Dr. E. E. Williams. 

LITERATURE CITED 

Barbour, T. 

1914. A contribution to the zoogeography of the West Indies, with 
especial reference to amphibians and reptiles. Mem. Mus. 
Comp. ZooL, 44: 209-359. 

Etheridge, R. 

1960. The relationships of the anoles (Reptilia, Sauria, Iguanidae): 
an interpretation based on skeletal morphology. Ph.D. thesis. 
University of Michigan. University Microfilms. Inc., Ann Ar- 
bor, Michigan. 

Gorman, G. C. and L. Atkins 

1966. Chromosomal heteromorphism in some male lizards of the genus 
Anolis. Amer. Nat., 100: 579-583. 

In press. The zoogeography of Lesser Antillean AnoUs. An inter- 
pretation based upon chromosomes and lactic dehydrogenases. 
Bull. Mus. Comp. Zool. 

Gorman, G. C, L. Atkins, and T. Holzinger 

1967. New karyotypic data on 15 genera of lizards in the family 
Iguanidae, with a discussion of taxonomic and cytological 
implications. Cytogenetics. 6: 286-299. 

Grant, C. 

1931. A new species and two new subspecies of the genus Anolis. 
J. Dept. Agriculture, Puerto Rico, 15: 219-222. 

Maldonado, a. a., and E. Ortiz 

1966. Electrophoretic patterns of serum proteins of some West Indian 
Anolis. Copeia, 1966: 179-182. 



1968 ANOLIS CRISTATELLUS 13 

MooRHEAD, p. S., P. C. NowELL, W. J. Mellman, D. M. Battips, and 

D. A. HUNGERFORD 

1960. Chromosome preparations of leukocytes cultured from human 
peripheral blood. Exp. Cell Res., 20: 613-616. 

Rand, A. S. 

1962. Anolis scriptiis Garman 1887, an earlier name for Aiiolis leiico- 
phaeiis Garman 1888. Breviora, Mus. Comp. Zool., No. 153: 
1-5. 
1964. Ecological distribution in anoline lizards of Puerto Rico. Ecol- 
ogy, 45: 745-752. 

Thomas, R. 

1966. Additional notes on the amphisbaenids of greater Puerto Rico. 
Breviora, Mus. Comp. Zool., No. 249: 1-23. 

(Received 21 November, 1967.) 



BREVIORA 

Museiuiinn of Comnparative Zoology 



Cambridge, Mass. 29 May, 1968 Number 294 

DISTRIBUTION AND BIOLOGY OF THE 

OPISTHOPROCTID FISH 
WINJERIA TELESCOPA BRAUER 190P 

Richard L. Haedrich and James E. Craddock^ 



INTRODUCTION 

First taken by the German VALDIVIA Expedition in 1898, 
Winteria telescopa Brauer 1901 remained a unique oddity until 
Marshall (1960) reported a second specimen, Blache (1963) a 
third and fourth, and Bertelsen and Munk (1964) a fifth. Recent 
midwater trawl cruises, in particular those of the Woods Hole 
Oceanographic Institution in the Atlantic and of the ANTON 
BRUUN in the Indian and Eastern Pacific Oceans, have added sis- 
nificantly to this number. Twenty-three specimens are known, of 
which 18 have not been reported previously. This note records 
the specimens, compares them taxonomically, and offers some ob- 
servations on the distribution and natural history of Winteria. 

ACKNOWLEDGMENTS 

For access to specimens we extend our thanks to E. Bertel- 
sen, Carlsberg Foundation's DANA Expedition, Charlottenlund 
(DANA); Robert J. Lavenberg, Los Angeles County Museum, 
Los Angeles (LACM); Giles W. Mead, Museum of Comparative 
Zoology, Harvard (MCZ); Richard H. Rosenblatt and Robert L. 
Wisner, Scripps Institution of Oceanography, La Jolla (SIO); 
Frederick H. Berry, Tropical Atlantic Biological Laboratory, Bur- 
eau of Commercial Fisheries, Miami (TABL); C. Richard Robins, 
Institute of Marine Science, University of Miami, Miami (UMML); 



1 Contribution No. 2031 from the Woods Hole Oceanographic Institution, 
and Papers from the "Dana" Oceanographical Collections No. 66. 

- Woods Hole Oceanographic Institution, Woods Hole, Mass., and 
Museum of Comparative Zoology, Harvard University. 



BREVIORA 



No. 294 









5 mm 
I 1 



Figure 1. A young Wintcria telescopa. 25.5 mm in standard length, from 
the Eastern Pacific (MCZ — 33°3rS, 77°29'W). 




5 mm 



Figure 2. Head of an adult male Winteria telescopa, 91.5 mm in standard 
length, from the Eastern Pacific (USNM — 32"02'S, 73°48'W). Note the 
retinal diverticulum on the eye. The pinniform structure below the eye is 
a muscle, probably the adductor mandibulae, seen through the thin cover- 
ing bone. Drawn by Martha M. Howbert. 



1968 NOTES ON WINTERIA 3 

and Robert H. Gibbs, Jr., United States National Museum, Wash- 
ington, D. C. (USNM). Don Dockires, SIO, provided informa- 
tion on the sex of two Scripps specimens. The manuscript was 
read by Richard H. Backus and Giles W. Mead. Financial support 
has been provided in part by a United States Government Grant 
under the Fulbright-Hays Act to the senior author, and by the 
National Science Foundation through its support of the U. S. Pro- 
gram in Biology of the International Indian Ocean Expedition 
(IIOE) and the Southeastern Pacific Biological Oceanographic 
Program, and its grants GB-4424 (principal investigator Giles W. 
Mead) and GB-4431 (principal investigator Richard H. Backus) 
to the Woods Hole Oceanographic Institution. 

MATERIAL EXAMINED 

The following list of material, by ocean, indicates source institu- 
tion (many specimens are as yet uncatalogued and hence have no 
catalog numbers), number of specimens, sex (if known), standard 
length in mm, and (in parentheses) vessel, cruise, station number 
or numbers, position, date, primary depths fished (no closing gear 
was used), and gear (IKMT = Isaacs-Kidd midwater trawl): 

Atlantic — MCZ, 1 spec, $ , 55 mm (CHAIN, Cruise 35, RHB- 
962, 5°24'N, 39°55'W, 13 Feb. 1963, 510-860 m, 10' IKMT); 
MCZ, 1 spec, 9 , 83 mm (CHAIN, Cruise 35, RHB-976, 0°03'N, 
27°3rW, 26 Feb. 1963, 565-675 m, 10' IKMT); MCZ. 1 spec, 
9, 98.5 mm (ATLANTIS II, Cruise 20, RHB-1206, 11°01'N, 
34°18'W, 23 Feb. 1966, 430-490 m, 10' IKMT); TABL 14, 1 
spec, 86 mm (GERONIMO. Cruise 2, sta. 82, 3°28'S, 01°14'W, 
6 Aug. 1963); UMML 19998, 1 spec, 50 mm (PILLSBURY, sta. 
295, 0°25'N, 5°09'E, 23 May 1965, 850 m, 10' IKMT). 

Indian — DANA, 1 spec, 100 mm (DANA. sta. 3847(3), 
12°02'S, 96°43'E, 11 Oct. 1929, 2500 mwo, 300 cm Ring-trawl); 
MCZ, 1 spec, 9 , 1 15 mm (ANTON BRUUN, Cruise 3, Trawl 10 
(AE-14), 2°06'S, 60^02'E, 21 Aug. 1963, 1600 m, 10' IKMT); 
MCZ, 1 spec, $, 130 mm (ANTON BRUUN, Cruise 3, Trawl 
11 (AE-15), 5°03'S, 63°10'E, 23 Aug. 1963, 685 m, 10' IKMT). 

Pacific — DANA, 1 spec, 82.5 mm (DANA, sta. 3716(3), 
19°18.5'S, 120°13'E, 22 May 1929, 2000 mwo, 300 cm Ring- 
trawl); LACM, 1 spec, $ , 124 mm (ELTANIN, Cruise 14, biol. 
sta. 1186, 52°10'S, 159°21'W, 4 Aug. 1964, 895-1030 m, 10' 
IKMT); LACM, 1 spec, 9 , 147 mm (ELTANIN, Cruise 27, biol. 
sta. 1986, 45°33'S, 147°18'E, 26 Feb. 1967, 2840-3001 m, 5' 



4 BREVIORA No. 294 

Blake Trawl); MCZ, 1 spec, 25.5 mm (ANTON BRUUN, Cruise 
13, collection 40, 33°3rS, 77°29'W, 28 Jan. 1966, 425-820 m, 
lO'IKMT); MCZ, 1 spec, 27 mm (ANTON BRUUN, Cruise 13, 
collection 43, 33°25'S, 77°38'W, 28-29 Jan. 1966, 100-380 m, 10' 
IKMT); MCZ, 1 spec, 69 mm (ANTON BRUUN, Cruise 13, col- 
lection 58, 33°42'S, 72°17'W, 3 Feb. 1966, 400 m, 10' IKMT); 
SIO, 1 spec, 33 mm (ANTON BRUUN, Cruise 12, SIO 65-664, 
33°31'S, 75°18'W, 17-18 Dec 1965, 530 m, 10' IKMT); USNM, 
1 spec, $, 91.5 mm (ANTON BRUUN, Cruise 14, sta. 550-A 
(RHG-66-9), 32°02'S, 73°48'W, 15 Feb. 1966, 200-500 m, 10' 
IKMT). 

Dr. Richard H. Rosenblatt has very kindly sent us counts and 
X-rays of three specimens in the collections at SIO. Pertinent data 
are: SIO 61-33, 1 spec, 70.5 mm (Monsoon Exped., sta. III-5, 
10°39'S, 98°51'E, 22/23 Nov. 1960, 0-1500 m, 10' IKMT. 
Counts: D 7, A 7, Pect. 12-12, Pelv. 8-8, vert. 34); SIO 61-37, 2 
spec, 121.3 and 144.7 mm (Monsoon Exped., sta. IV-19, 33°19'S, 
72°34'E, 19 Dec. 1960, 0-2000 m, 10' IKMT. Counts: D 9 and 7, 
A 7 and 7, Pect. 12-13 and 12-13, Pelv. 9-9 and 8-9, Lat. line 
scales 35 and ca. 35, vert. 35 and 35, respectively). 

DESCRIPTION 

The following descriptive notes are provided to complement the 
characterizations of other opisthoproctids by Cohen (1964): 

Body cylindrical, plump, becoming somewhat compressed at 
caudal peduncle, not particularly elongate. Belly lacking a sole. 
A rectal bulb and reflector organ present (Bertelsen and Munk, 
1964). Eyes tubular and directed anteriorly. Interorbital very 
narrow. A dark-colored retinal diverticulum located anterolaterally 
on each eye. Snout elongate and hyaline; nasal capsules easily 
discerned. Branchiostegal rays 3, short and blunt. Gill membranes 
united across the isthmus. Maxillaries very small, thin and scale- 
like. Premaxillaries appear to be absent. No apparent teeth on the 
dentary. Swimbladder present (Marshall, 1960). Dorsal adipose 
fin present. Dorsal, anal, and ventral fins posteriorly placed; rays 
perhaps prolonged. Pectoral fins on peduncles, horizontally ori- 
ented high on the sides. Anus between pelvic fins, well before anal 
fin. Scales very deciduous. 

Color in life deep blue-black with silvery overtones on the head. 
Fresh-caught specimens have a very plump body; preserved speci- 
mens are much shrunken in comparison. 



1968 NOTES ON WINTERIA 



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6 BREVIORA No. 294 

COMPARISON OF SPECIMENS 

Morphometric and meristic data for 17 specimens, including 
Brauer's holotype of Winteria telescopa, are presented in Table 1. 
The small range of variation in both proportions and counts is 
strong evidence that there is but one species represented, and thus 
all our material is referred to Winteria telescopa Brauer 1901. 
There is perhaps some geographical variation in the number of 
vertebrae. The range is 33-35 vertebrae in the Atlantic, 34-35 in 
the Indian, and 35-36 in the Pacific. No pattern of variation is 
evident in the numbers of fin rays. 

Little allometry occurs between 25 and 147 mm SL (Table 1). 
The only regular change is in the length of the head, which becomes 
relatively shorter with growth. This may be also true in the fins, 
although the rays are extremely fragile and are usually damaged. 

As has been observed in the young of other opisthoproctids (Co- 
hen, 1964), young Winteria (Fig. 1) somewhat resemble Rhyncho- 
hyalus, the genus linking the two divergent lines of the 
Opisthoproctidae. Characters found also in Rhynchohyalus are 
the elongate snout, the tubular eyes, the retinal diverticula ("orbi- 
tal light organs" of Cohen, 1964, see Bertelsen et al., 1965), the 
raised horizontal placement of the pectoral fins, and the insertion 
of the pelvic fins above the anus. With growth, the profile of the 
head in Winteria changes. The head becomes somewhat deeper in 
respect to its length, and the angle between the snout and the 
interorbital region is more pronounced (Fig. 2). This is in con- 
tradistinction to Brauer (1906: pi. I), who shows a smooth arc 
from the tip of the snout to the occiput. 

DISTRIBUTION 

Four of the previously known specimens of Winteria are from 
the tropical Atlantic off Africa (Brauer, 1901, holotype; Marshall, 
1960, 1 spec; Blache, 1963, 2 spec), and have been referred by 
their recorders to Winteria telescopa. The fifth specimen was in- 
cidentally reported from the eastern Indian Ocean by Bertelsen and 
Munk (1964). The concern of their paper was anatomical, and 
they made no judgement as to the specfic status of their specimen, 
referring to it simply as Winteria. 

As shown above, all specimens so far known can be referred to 
Winteria telescopa Brauer 1901. The distribution of this species 
is circumglobal (Fig. 3). The shallowest possible depth-of -capture 
(ANTON BRUUN 13-43) was certainly greater than 200 meters; 



1968 



NOTES ON WINTERIA 




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1968 NOTES ON WINTERIA 9 

the deepest tow (ELTANIN 1986) reached 3000 meters. The 
majority of captures probably occurred in the range 500-700 
meters. Wiuteria then, like other opisthoproctids (Cohen, 1964), 
is a relatively deep-living fish. 

Hydrographic casts had been made nearby or were made in con- 
nection with nine of the trawls which took Winteria. The tem- 
perature-salinity relationships at these stations, from 200 meters 
to the maximum possible depth-of-capture, are plotted in Figure 4. 
With the exception of the two western Indian Ocean stations 
(IIOE 150, 151), none falls entirely within a discrete water mass. 
They occur, for the most part, in the transitional regions between 
water masses, areas where the productivity is probably higher than 
in the generally poorer waters of the central gyres. 

On the T-S curves of Figure 4, most probable depths-of-capture 
are indicated. When at surfaces (density isopleths dependent upon 
temperature and salinity) are superimposed on the T-S plot, these 
depths fall within a at range of 27.00 to 27.40. This suggests that 
the distribution of Winteria may be determined at least in part by 
density, a not unreasonable proposition. Pickford (1946) found 
that the distribution of Vampyroteuthis, a bathypelagic cephalopod, 
was restricted to the at range 27.40 to 27.80. 

REPRODUCTION 

The gonads could not be distinguished in fishes smaller than 
50 mm SL. Fishes between 50 and 90 mm appeared immature, 
with the onset of maturity occurring at about 90 mm SL. Both 
males and females could be recognized, and the gonads appear 
to be normal. There is no indication of hermaphroditism or vivi- 
parity. The ovaries of the three large females examined each con- 
tained two size classes of eggs, a 0.5-1.2 mm diameter group and 
a 1.8-2.5 mm diameter group (Table 2). These two size classes 

Table 2. Standard length of female Winteria 
and numbers and sizes of eggs in one ovary. 



Std. length 




98 mm 


115 mm 


147 mm 


N" small eggs 




80 


101 


125 


N*^ large eggs 




59 


102 


171 


Size range small 


eggs 


.8-1.1 mm 


.9-1.2 mm 


.5-1.0 mm 


Size range large 


eggs 


1.8-2.2 mm 


2.2-2.5 mm 


2.0-2.5 mm 



10 BREVIORA No. 294 

are present in about equal numbers. The data suggest a direct 
relationship between size of the fish and the number of eggs. 

With the exception of the two Scripps specimens (SIO 61-37), 
all the Winteria were taken as single captures. The larger of the 
two fish which were taken together is a female containing large 
orange eggs; the smaller specimen appears to be a ripe or near- 
ripe male. Winteria is probably a solitary fish, encountering an- 
other of its kind only occasionally and pairing when ready to 
spawn. 

FEEDING 

The stomachs of the six largest specimens were opened. One 
was empty; the rest contained a finely divided, light yellowish, 
pulpy material. No structure could be discerned in this material, 
and we assume it was not composed of crustacean remains. It 
most closely resembled the finely shredded tissues of siphonophores, 
medusae, and salps found by Haedrich (unpublished) in the 
stomachs of stromateoid fishes. 

LITERATURE CITED 

Bertelsen, E. and Ole Munk 

1964. Rectal light organs in the argentinoid fishes Opisthopr actus and 
Winteria. Dana-Report No. 62: 1-21. 

Bertelsen, E., Birgit Theisen, and Ole Munk 

1965. On a postlarval specimen, anal light organ, and tubular eyes of 
the argentinoid fish Rliyncholiyalus natalensis (Gilchrist and 
von Bonde). Vidensk. Medd. Dansk Naturh. Foren., 128: 
357-371. 

Blache, J. 

1963. Poissons bathypelagiques rares ou peu connus provenant des 
eaux de I'Atlantique oriental tropical. 1"" Note. Famille des 
Opisthoproctidae. Cahiers O.R.S.T.O.M., Oceanographie, No. 
5: 89-95, 4 figs. 

Brauer, a. 

1901. Ueber einige von der Valdivia-Expedition gesammelten Tiefsee- 
fische und ihre Augen. Sitzungsber. Gesell. Beford. Ges. Nat- 
urwiss. Marburg, 1901, No. 8: 115-130. 
1906. Die Tiefsee-Fische. I. Systematischer Teil. Wiss. Ergebn. "Val- 
divia", 15: 1-420, 16 pi. 
Cohen, Daniel M. 

1964. Suborder Argentinoidea in Fishes of the Western North Atlan- 
tic. Sears Found. Mar. Res., Mem. 1, part 4: 1-70, 20 figs. 



1968 NOTES ON WINTERIA H 

Marshall. N. B. 

1960. Swimbladder structure of deep-sea fishes in relation to their 
systematics and biology. D/icovery Reports, 31: 1-122. 
PicKFORD, Grace E. 

1946. Vampyroteiithis infernalis CHUN. An archaic dibranchiate 
cephalopod. I. Natural history and distribution. Dana-Report 
No. 29: 1-40. 
SvERDRUP, H. U., Martin W. Johnson, and Richard H. Fleming 
1942. The oceans. Prentice-Hall, New Jersey, 1087 pp. 



BREVIORA 

Mesemm of Contiparative Zoology 



Cambridge, Mass. 29 May, 1968 Number 295 

THE CHANARES (ARGENTINA) TRIASSIC REPTILE FAUNA 

IV. THE DICYNODONT FAUNA 

C. Barry Cox^ 

The dicynodonts described in this paper were collected by the 
1964-1965 expedition of the Museo de la Plata and the Museum 
of Comparative Zoology to the Permo-Triassic of western Argen- 
tina (Romer, 1966). Nearly all come from the Chanares Forma- 
tion, the lithology and stratigraphic relationships of which have 
been described by Romer and Jensen (1966). They state (p. 12) 
that, "With one exception, all of the numerous vertebrate fossils 
found in the Chaiiares were from the lowest 10 meters or so of 
the formation." Eighteen dicynodont specimens were found in this 
fossiliferous band. One of these belongs to a new genus and species 
of dicynodont, described here as Chanaha platyceps. Eight speci- 
mens belong to a new species closely allied to Dinodontosaurus 
turpior from the Santa Maria Formation of Brazil (Cox, 1965); 
it is here named Dinodontosaurus brevirostris. Another specimen 
belongs to a second new species of this genus, here named D. platy- 
gnathus. Finally, an isolated large scapulocoracoid and clavicle 
show the presence of another genus of dicynodont. The remaining 
seven specimens are incomplete or badly damaged and therefore 
cannot be assigned with certainty to any of the named taxa. 

In the accompanying figures, oblique shading indicates broken 
bone surface, horizontal shading indicates the presence of matrix, 
and broken lines indicate restored outlines of bone. The following 
abbreviations are used: MCZ — the Museum of Comparative Zo- 
ology, Harvard University; BYU — the Earth Sciences Museum, 
Brigham Young University, Provo, Utah; DGM — Divisao de 
Geologia e Mineralogia, Ministerio das Minas e Energia, Rio de 
Janeiro. 



1 Zoology Dept., King's College, Strand, London W.C.2., England. 



2 BREVIORA No. 295 

Chanaria gen. nov. 
Type species: C. platyceps sp. nov. 

Diagnosis. Dicynodont of moderate size: skull length 33 cm. 
Teeth absent except for upper tusks. Greatest width of skull is 
across occipital region. Blunt anterior end to snout. Tusk pro- 
jects anteroventrally. Lacrimal extends forwards on face, prob- 
ably meets septomaxilla. No bony bosses on snout or on pineal 
region. Interorbital bar fairly wide. Frontal extends anteriorly 
into nasal as rectangular midline projection. Quite large pre- 
frontal. Preparietal present, forming anterior border of pineal 
foramen. Postorbital bone extends well behind pineal foramen. 
Intertemporal bar narrow but concave in cross-section, not ridged. 
Squamosal does not extend on to intertemporal bar, posterior end 
of which is formed by interparietal. Short, wide temporal opening. 
Low, wide occiput; supraoccipital does not extend high up in mid- 
line. 

Chanaria platyceps sp. nov. 

Holotype. No. 65-XI-14-3 Museo de la Plata, La Plata, Ar- 
gentina, skull. Collected by the 1964-1965 expedition of the Museo 
de la Plata and the Museum of Comparative Zoology. 

Horizon and locality. From an exposure in the Triassic Chan- 
ares Formation about 6 miles east of the point where the Chanares 
River emerges into the Campo de Talampaya, in western La Rioja 
Province, Argentina. 

Description. The type skull is 33 cm long and 30 cm wide. 
The mid-region of the palate, both quadrate regions, and part of 
the left side of the skull are missing. 

Dorsal view (Fig. 1). The premaxilla extends a considerable 
distance posteriorly in the midline. There is only a short midline 
nasal suture, for the posterior regions of the nasals are separated 
by a rectangular anterior projection of the frontals. There is quite 
a large prefrontal; the posterior end of its suture with the frontal 
is uncertain, owing to damage. The interorbital bar is of moderate 
breadth. 

The postorbital bone extends back to a point halfway between 
the pineal foramen and the posterior end of the intertemporal bar. 
The intertemporal bar is quite narrow, but it is slightly concave 
from side to side, without any midline ridge. The extreme pos- 
terior end of the bar is formed by the interparietal. 



1968 



CHANARES DICYNODONT FAUNA 



The occipital wing of the squamosal extends laterally, not pos- 
terolaterally. As a result, the posterior edge of the skull is almost 
straight and runs transversely, and the temporal opening is almost 
quadrilateral, its width approximately equalling its length. 

Occipital view (Fig. 2). The occiput is rather low, its midline 
region being little higher than its more lateral parts. Most of the 
midline surface above the foramen magnum is formed by the in- 
terparietal rather than by the supraoccipital. The bones of the 
central occipital plate are fused together. The quadrate region and 
outer edge of the lateral wing of the squamosal are missing, but 
the dotted outline shown is the minimum extent of squamosal 
necessary to accommodate a quadrate of normal size. 




Figure 1. Chanaria platyceps, type specimen. Dorsal view of skull, x V3 
(for abbreviations, see p. 26). 



BREVIORA 



No. 295 




Figure 2. Chanaria platyceps, type specimen. Occipital view of skull. 
X V3 (for abbreviations, see p. 26). 



Lateral view (Fig. 3). The powerful tusks project antero- 
ventrally. The squamosal runs forward to meet the maxilla under 
the front half of the orbit. 

A groove runs anteroventrally from the anterior corner of the 
nostril to the edge of the snout. A ridge which forms the lower 
edge of this groove also extends posteriorly along the maxilla be- 
hind the nostril. Above this ridge both the maxilla and the lacri- 
mal curve slightly inwards, so that there is no sharply defined pos- 
terior border to the nostril. The lacrimal extends a considerable 
distance anteriorly and, though its anterior end and most of the 
septomaxilla are missing, it seems very probable that these two 
bones met, separating the nasal from the maxilla. 

The palatal surface of the premaxilla bears the pair of anterior 
ridges and the median posterior ridge that are normal in the dicy- 
nodonts. The remainder of the ventral region of the skull is too 
incomplete to show any features of value. 

Taxonomic position. In an earlier paper (Cox, 1965) I sug- 
gested that two main families of Triassic dicynodonts could be 
distinguished on the basis of the shape of the snout, the presence 
or absence of a median crest along the intertemporal bar, and the 
proportions of the occiput and of the temporal opening. In Chan- 
aria the snout is wide and blunt, there is no median crest along 



1968 



CHANARES DICYNODONT FAUNA 




Figure 3. Clianaria platyceps, type specimen. Lateral view of skull, x V3 
(for abbreviations, see p. 26). 



the intertemporal bar, the occiput is wide but low, and the tem- 
poral opening is short. In all these characteristics, Chanaria 
clearly belongs to the family Stahleckeriidae. 

Bonaparte (1966a) has since disagreed with my proposed classi- 
fication of the Triassic dicynodonts into separate families. He be- 
lieves that these forms are very uniform in structure, and that they 
are monophyletic in origin. Though I do not agree with either of 
these beliefs, Bonaparte's views and my own are, in the absence 
of knowledge of the Permian ancestry of the Triassic forms, sub- 
jective views of equal validity. The truth will emerge only from 
further work on the dicynodonts of the late Permian and early 
Triassic, and further discussion on these alternative classifications 
may be postponed until relevant material has been found and 
studied. 



DlNODONTOSAURUS BREVIROSTRIS Sp. HOV. 

Holotype. No. 65-XI-14-4 Museo de la Plata, La Plata, Argen- 
tina, skull. Collected by the 1964-1965 expedition of the Museo 
de la Plata and the Museum of Comparative Zoology. 

Horizon and locality. From an exposure in the Triassic Chan- 
ares Formation about 6 miles east of the point where the Chanares 
River emerges into the Campo de Talampaya, in western La Rioja 
Province, Argentina. 



BREVIORA 



No. 295 



Referred specimens. Specimen Nos. 3452-3457, Museum of 
Comparative Zoology, Harvard; specimen No. MR- 120, Earth 
Sciences Museum, Brigham Young University, Provo, Utah. 




Figure 4. Dinodontosaiirus brevirostris, type specimen. Dorsal view of 
skull, X V3 (for abbreviations, see p. 26). 



Description. The skull of the type specimen, and those of speci- 
mens MCZ Nos. 3453 and 3457 and of specimen BYU No. MR- 
120, are all about 30 cm long. Specimen MCZ No. 3454 is 23 cm 
long and appears to be a juvenile. Other dicynodont remains from 
the Chanares Formation which cannot be certainly identified but 
which probably belong to this, the commonest dicynodont in the 
fauna, show that specimens nearly twice as large as the type were 
present. 



1968 



CHANARES DICYNODONT FAUNA 









Figure 5. Dorsal views of pineal areas of specimens of Dinodontosaunis, 
reduced to equal size. A, D. turpior, MCZ No. 1628; B, D. turpior, MCZ 
No. 1687; C, D. brevirostris, MCZ No. 3454; D, D. brevirostris, type speci- 
men; E, D. turpior, DGM No. 530R; F, D. turpior, DGM No. 213R. 



8 



BREVIORA 



No. 295 



Dorsal view (Fig. 4). D. brevirostris has the short temporal 
region and narrow, ridged intertemporal bar which are character- 
istic of the genus. However, there are some differences between 
D. brevirostris and the type species, D. lurpior, in the relation- 
ships of the postorbital to the pineal opening. The postorbital of 
D. brevirostris approaches quite closely to the side of the pineal 
opening, and also extends posteriorly well beyond it. Though 
there is considerable range of variation in these characteristics in 
D. turpior (Fig. 5), the postorbital in that species is not normally 
as extensive as that of D. brevirostris. 

Ventral view. The palate of the type specimen of D. brevirostris 
is too poorly preserved to show anything of value, but this area is 
well shown in the smaller specimen, MCZ No. 3454 (Fig. 6). The 
secondary palate extends posteriorly to a point level with the root 
of the canine tusks. The vomer and palatines do not form a bony 




Figure 6. Dinodontosaunis brevirostris, MCZ No. 3454. Ventral view, 
X 4/9 (for abbreviations, see p. 26). 



1968 



CHANARES DICYNODONT FAUNA 



roof to the anterior part of the primary palate, between the anterior 
rami of the pterygoids. On either side of the median blade of the 
vomer there is therefore a clear passage dorsally (in the bony skull) 
to the ventral side of the bones of the skull roof (see also Fig. 9B). 
Behind the vomer, only a relic of the interpterygoid vacuity re- 
mains; through it is visible the ventral surface of the processus 
cultriformis of the parasphenoid. 

The only specimen of D. turpior in which the median region of 
the palate is well preserved is specimen DGM No. 530R. In this^ 
the vomer and palatines form a bony roof over a greater extent of 
the posterior part of the primary palate, and the median fusion of 
the pterygoids has extended further forward within the interptery- 
goid vacuity, so that the processus cultriformis is not visible from 




Figure 7. Dinodontosaums brevirostris, type specimen. 
1/3 (for abbreviations, see p. 26). 



Lateral view, x 



iThis specimen is of particular interest because it was described in 1936 
by Tupi Caldas under the name Diodontosaurus pedroanum, a name which 
predates Romer's description of Dinodontosaums (1943). However, Tupi 
Caldas's description is unfortunately so brief and so poorly illustrated that 
it cannot be regarded as a satisfactory basis for the identification of a new 
taxon, and Diodontosaurus pedroanum must therefore be regarded as a 
nomen nudum. The specimen has recently been more fully figured by 
Beltrao (1966). 



10 BREVIORA No. 295 

below. However, it would be unjustifiable to assume that the two 
species difi'er in these features, as the smaller extent of these bones 
in specimen MCZ No. 3454 could merely be a result of its relative 
immaturity. 

Side view (Fig. 7). The external naris is rather damaged but 
appears to be shallow, and there are no large foramina in the re- 
gion of the septomaxilla. Though the edge of the maxilla where 
it forms the margin of the mouth is somewhat damaged, it is thin 
and was probably sharp-edged. This edge descends ventroposteri- 
orly so that it reaches a considerable distance down the anterior 
border of the tusk. 

Comparison of D. brevirostris with the specimens of D. turpior 
in the Museum of Comparative Zoology suggested that the snout 
of the Argentinian species was very much shorter, and the tusk 
much more ventrally directed, than in the Brazilian species. How- 
ever, I was fortunately able to visit Rio de Janeiro and study the 
excellent material of D. turpior in the Divisao de Geologia e Min- 
eralogia, Ministerio das Minas e Energia. This material has been 
prepared under the direction of Mr. L. I. Price, who has also made 
extensive studies of it. Examination of this material soon showed 
that the skull of specimen MCZ No. 1670 is very unusual in its long, 
low shape, and that there is a considerable range of variation in this 
region in D. turpior (Fig. 8). Nevertheless, there do appear to 
be two constant difi"erences between the snouts of the two species. 
Firstly, the external naris of D. brevirostris is much closer to the 
anterior edge of the premaxilla than is that of D. turpior. Secondly, 
the tusk of D. brevirostris always projects directly ventrally, so 
that there is an abrupt angle between the body of the maxilla and 
the anterior end of the zygomatic arch, whereas there is a smoother 
transition between these regions in D. turpior. It is worth noting 
also that the lacrimal extends further forward on the face and 
meets the septomaxilla in all specimens of D. brevirostris in which 
the sutures of these regions are visible. This condition is known in 
D. turpior (Fig. 8F), but in that species these two bones are often 
separated by the maxilla. 

There is a prominent boss on the dorsal surface of the palate, 
anterior to the base of the epipterygoid. This boss lies above a 
very distinct curving suture which separates it from the pterygoid, 
and it is therefore formed by the basisphenoid. Restudy of this 
area in D. turpior shows an identical situation, despite my state- 
ment that the boss in that species "is clearly part of the pterygoid" 
(Cox, 1965:482). Camp and Welles (1956:fig. 49) also show 
this boss as part of the basisphenoid in Daptocephalus leoniceps. 



1968 



CHANARES DICYNODONT FAUNA 



11 








Figure 8. Lateral views of snouts of specimens of Dinodontosaiinis, re- 
duced to equal size. A, D. turpior, MCZ No. 1670; B, D. turpior, MCZ 
No. 1687; C, D. turpior, MCZ No. 1628; D, D. brevirostris, MCZ No. 3454; 
E, D. turpior, DGM No. 530R; F, D. turpior, DGM No. 309. 



12 



BREVIORA 



No. 295 




Vi-;: 





pmx 



B 



Figure 9. Dinodontosaurus brevirostris, MCZ No. 3454. Lateral views, 
X 4/9, A, of complete skull; B, after removal of suborbital bar, postorbital 
bar, anterior end of left half of palate, and part of left maxilla. Dotted 
lines in figure B show ventral outlines of midline regions of premaxilla. 
vomer and pterygoid (for abbreviations, see p. 26). 



1968 CHANARES DICYNODONT FAUNA 



13 



Further information on the structure of the dorsal roof of the 
palate and of the median interorbital-internasal septum is provided 
by specimen MCZ No. 3454 (Fig. 9A, B). This shows clearly 
that the ectopterygoid is present, confirming my eadier tentative 
identification of this bone in D. turpior (Cox, 1965). The palatine 
has an extensive exposure on the dorsal surface of the palate. It 
forms the lower border of the large foramen which runs forwards 
into the dental cavity of the maxDIa; it extends dorsally to contact 
the parasphenoid, and runs back to meet the basisphenoid. Much 
of the palatine overlies the lateral wing of the vomer, which is 
visible in front of the palatine and also above its posterior exten- 
sion, where the vomer forms the lower border of the dorsal end 
of the canal which runs forwards and upwards from the interptery- 
goid vacuity. As Klaauw and Roon (1942) have pointed out, this 
opening is not the simple equivalent of the old interpterygoid 
vacuity, and they have suggested that it be called the fenestra 
medio-palatinalis. The canal was probably traversed by a blood 
vessel. Since no such blood vessel is known in either Sphenodon 
(O'Donoghue, 1920) or Ctenosaura (Oelrich, 1956), it is im- 
possible to identify it, though it seems most likely to have been 
a branch of the palatine artery. 

Above the fenestra medio-palatinalis lies the medial region of 
the parasphenoid-basisphenoid complex. The processus cultri- 
formis of the parasphenoid, as is normal in synapsids, continues 
anteriorly beyond the basisphenoid. and receives the lower edge of 
the sphenethmoid. The processus cultriformis is underlain by the 
vomer. This medial part of the vomer forms the lower part of the 
interorbital septum and also the posterior part of the internasal 
septum. It extends ventrally to meet the posterior end of the me- 
dian palatal ridge of the premaxilla and has a total depth of 5.5 
cm. The more dorsal portion of the interorbital septum is formed 
by the sphenethmoid (Fig. 9B). 

More posteriorly, much of the rod-like epipterygoid is preserved 
in the young specimen, but its lower edge is damaged and the posi- 
tion of its suture with the parietal dorsally is uncertain. There are 
clear sutures between the prootic and the parietal, and between 
the prootic and the parasphenoid-basisphenoid complex. 

All the specimens of D. brevirostris are tusked. Most of the 
tusks are broken off short, but the whole of the right tusk of speci- 
men MCZ No. 3453 is preserved and shows clear signs of wear. 
The outer surface of the distal end of the tusk is worn, so that its 
tip lies near its posteromedial edge. Traces of other wear facets 



14 BREVIORA No. 295 

are also visible on the posteromedial surface of this specimen, but 
are better shown in an isolated tusk, specimen MCZ No. 3452 (Fig. 
lOA-C). 

In longitudinal section of a tusk, the dentine appears to consist 
of a series of V-shaped bands; in a tusk about 15 cm long and 2.5 
cm in diameter at its base, these bands are about 2.5 mm thick 
(Fig. lOE). They are caused by alternating light and dark zones 
of dentine, which are apparently due to slight variations in the con- 
centration or thickness of the dentinal tubules. Further bands, 
which are similar but only a few tenths of a millimetre in diameter, 
are also visible in thin sections examined under a microscope. 

A system of cracks, which in general parallel this system of V- 
shaped bands in the dentine, can also be seen in longitudinal sec- 
tion (Fig. lOD), and appear as concentric circles in transverse 
section of the tusk. A similar system has been described in Placerias 
and Kannemeyeria by Camp and Welles (1956). However, these 
cracks are not related to the above-mentioned variations in the 
structure of the dentine, and appear to be post-mortem. The 
cracks merely reflect the main plane of structural weakness in the 
dentine, which in turn is related to its mode of deposition. 

The base of the tusk is open and it probably grew continuously. 
A series of annular grooves can also be seen around that portion of 
the tusk which lies within the maxilla; this has also been noted by 
Camp and Welles (1956). Similar annular grooves and variations 
in dentinal structure (alternating between columnar and marbled 
dentine) have been described in the elephant seal Mirounga 
leonina by Laws ( 1953 ) . He has shown that the pattern of banding 
in that animal is complex, but that there is a regular annual repe- 
tition of this pattern. The details of the annual pattern also differ 
between the sexes, due to their different cycles of activity during 
the breeding season. It is unfortunately impossible to verify 
whether these features in Dinodontosaiirus are similarly related to 
age or sex. This would be expected only if the climate were suffi- 
ciently seasonal to cause variations in the rate of growth of both 
sexes, or if variations resulted from such seasonal activities as egg- 
laying. 

Postcranial material. Specimens MCZ Nos. 3454, 3455 and 
3456 all included postcranial material; that belonging to specimen 
MCZ No. 3455 is particularly well preserved. However, none of 
this material shows significant differences from the corresponding 
bones of D. turpior, or adds to our knowledge of the postcranial 
skeleton of the genus. 



1968 



CHANARES DICYNODONT FAUNA 



15 





B 






Figure 10. Dinodontosaiiriis brevirostris. A-C, right tusk of specimen 
MCZ No. 3452 showing wear facets, x %. A, lateral view; B, anterior view; 
C, medial view. D, E, longitudinal sections of part of tusk of specimen 
MCZ No. 3456. D, showing system of cracks, x V^; E, showing pattern of 
bands in dentine, x 1. 



16 



BREVIORA 



No. 295 



DiNODONTOSAURUS PLATYGNATHUS sp. nOV. 

Holotype. No. 65-XI-14-5 Museo de la Plata, La Plata, Argen- 
tina, fragmentary skull and lower jaw. Collected by the 1964-1965 
expedition of the Museo de la Plata and the Museum of Com- 
parative Zoology. 

Horizon and locality. From an exposure in the Triassic Chah- 
ares Formation, about P/i miles north-north-west of the point 
where the Gualo River emerges from the Piano del Gualo, in west- 
ern La Rioja Province, Argentina. 

Referred material. Specimen No. 149R, Divisao de Geologia 
e Mineralogia, Ministerio das Minas e Energia, Rio de Janeiro. 

Description. The type specimen consists only of the palatal and 
occipital regions of a large skull, and of an almost complete lower 
jaw (Fig. 11). The bluntly-ending snout and the presence of 
downwardly directed canine tusks show that the specimen prob- 
ably belongs to the genus Dinodontosaurus. The anterior end of 
the lower jaw, however, is elongated and tapers to a relatively thin 




Figure II. Dinodontosaurus platygnathus, type specimen. Lateral view 
of lower jaw, x Vi (for abbreviations, see p. 26). 



point, unlike that of Dinodontosaurus turpior. It at first seemed 
likely that this was merely an aberrant type of distortion, but the 
existence of an almost identical lower jaw (specimen No. DGM 
149R) in the Rio de Janeiro collection from the Santa Maria 
Formation of Brazil makes this explanation less plausible, and sug- 
gests instead that a different species of Dinodontosaurus may be 
represented. The fragments of the palate and occiput associated 
with the type specimen unfortunately do not show any other fea- 
tures by which the species could be distinguished from D. turpior, 
but it is felt nevertheless that the characters of the lower jaw merit 
specific distinction as Dinodontosaurus platygnathus. 



1968 CHANARES DICYNODONT FAUNA 17 

KANNEMEYERIID REMAINS FROM THE CHANARES FORMATION 

Specimen MCZ No. 3459 comprises a large left scapulocoracoid 
and clavicle; it was found about two miles east of the Mogote de' 
Gualo (see Romer and Jensen, 1966, fig. 2). 

The scapulocoracoid is badly flattened. As preserved, it has a 
total length of 63.5 cm, the scapula itself being about 48 cm long 
(Fig. 12). The scapula is very markedly constricted, being only 
8 cm across at its narrowest point but expanding to 25 cm wide 
at its upper end. Most of the spine down the outer surface of the 
scapula has been eroded away but, from the width of its base and 
from the contours of the surrounding bone, the spine was clearly 
well developed and its dorsal end rose rapidly upwards from the 
blade. A groove runs down the inner surface of the lower end of 
the scapula. This groove leads to the coracoid foramen, which 
lies within the precoracoid bone. 

The incomplete clavicle measures 33.5 cm along its outer sur- 
face, which is slightly convex longitudinally. 

As discussed in an earlier paper (Cox, 1965), the shape of the 
scapula seems to be one of the diagnostic features of the two main 
families of Triassic dicynodonts. Short, wide scapulae are found 
in the Stahleckeriidae, which includes the other Chaiiares dicyno- 
donts {Chanaria and Dinodontosaurus) . Tall, narrow-waisted 
scapulae are, on the other hand, characteristic of the Kannemey- 
eriidae, which includes the genera Kannemeyeria, Parakanne- 
meyeria, Sinokannemeyeria, Barysoma and Ischigualastia. The 
scapula of MCZ No. 3459 is very similar to that of Barysoma, 
which is known from some postcranial material and an occipital 
plate from the Santa Maria Formation of Brazil (Romer and Price, 
1944; Cox, 1965). The scapula of MCZ No. 3459 and that of 
Barysoma resemble one another closely in shape and in the posi- 
tion and strength of the spine. The scapula of Barysoma is, how- 
ever, considerably larger; though incomplete, it is 55.5 cm long. 

The single scapulocoracoid and clavicle of MCZ No. 3459 do 
not, of course, provide sufficient evidence to claim that Barysoma 
was present in the Chaiiares Formation or, alternatively, to erect 
a new genus or species of dicynodont. The specimen is merely 
evidence that the kannemeyeriid dicynodonts existed in Argentina 
during the time that the Chaiiares Formation was deposited. This 
is not surprising, since kannemeyeriids are known in Argentina 
both from the earlier Puesto Viejo Formation (Bonaparte, 1966a) 
and from the later Ischigualasto Formation (Cox, 1965). 



18 



BREVIORA 



No. 295 




Figure 12. Scapulocoracoid of specimen MCZ No. 3459, x Vs (for ab- 
breviations, see p. 26). 



DICYNODONT REMAINS FROM THE TARJADOS FORMATION 

A few fragmentary dicynodont remains were also collected in the 
transitional beds underlying the thick white sandstones which form 
the upper part of the Tarjados Formation. Specimen MCZ No. 
3468 was collected just north of the Piano del Gualo, about one 
mile west of the Mogote del Gualo (see Romer and Jensen, 1966, 
fig. 2) ; it includes fragments of a skull and lower jaw. The maxilla 
encloses the proximal portion of a powerful tusk about 3.0 cm 
in diameter. This tusk shows traces of the alternating bands de- 
scribed above in Dinodontosaurus brevirostris, but also bears sev- 
eral longitudinal furrows. The remaining two specimens from the 
Tarjados Formation may also belong to the Dicynodontia: two 
fragments of large limb bones (MCZ No. 3469) and fragments 
of ribs (MCZ No. 3467). 



1968 CHANARES DICYNODONT FAUNA 



19 



THE AGE OF THE CHANARES FAUNA 

The commonest Chanares dicynodont is Dinodontosaurus hrev- 
irostris. This species is extremely closely related to Dinodontosaurus 
turpior, which is the commonest dicynodont of the Santa Maria 
Formation of Brazil. The two species differ only in that the nostril 
of D. brevirostris is slightly closer to the front edge of the pre- 
maxilla. and its postorbital bone extends closer to, and further 
behind, the pineal foramen. This degree of similarity between their 
most abundant types of dicynodont is very strong evidence that 
the two faunas are very close in age. This is supported by the pres- 
ence in both faunas of the peculiar elongated type of dicynodont 
jaw, which has here been named Dinodontosaurus platygnathus. 
The Chaiiares fauna, however, lacks Stahleckeria, a dicynodont 
which occurs with Dinodontosaurus in the Santa Maria fauna; this 
suggests that the two faunas are not identical in age. In the slightly 
greater extent of its postorbital, D. brevirostris is more primitive 
than D. turpior, and it may well be ancestral to the Brazilian spe- 
cies; this suggests that the Chanares fauna is slightly earlier than 
the Santa Maria fauna. 

Another fauna which appears to be earlier than that of the Santa 
Maria Formation is that of the Manda Formation of East Africa 
(Cox, 1965). This includes the dicynodont genus Kannemeyeria, 
found also in the early Triassic Cynognathus Zone fauna of South 
Africa, and differs from the Santa Maria fauna in containing a 
more primitive type of rhynchosaur (Colbert, 1958) and in lacking 
dinosaurs. Since both the Manda fauna and the Chanares fauna 
thus appear to be earlier than the Santa Maria fauna, it is necessary 
next to discuss the relative ages of these two faunas. 

As just noted, the strong similarity between the dicynodonts of 
the Chafiares fauna and those of the Santa Maria fauna strongly 
suggests that these two faunas are very close in age. In view of the 
considerable differences between the Santa Maria fauna and the 
Manda fauna, this further implies that the Chafiares fauna is prob- 
ably later than the Manda fauna. However, Romer (1966) has, 
on the contrary, suggested that the Chafiares fauna is the earlier 
of the two. The evidence provided by the other elements of these 
faunas (cynodonts, pseudosuchians and rhynchosaurs) must there- 
fore be examined. 

The traversodontid Massetognathus is the only Chanares cyno- 
dont so far described (Romer, 1967). Each of the upper cheek 
teeth of this genus "shoulders" slightly into the one ahead; their 
crowns each bear two external cusps and a cross-ridge which lies 



20 BREVIORA No. 295 

at the extreme posterior edge of the tooth so that the crown con- 
sists of a single basin. The traversodontid of the Manda fauna is 
Scolenodon, in which the upper cheek teeth do not "shoulder" 
into one another, the crown of the tooth bears only a single external 
cusp, and the cross-ridge lies more anteriorly, so that it divides 
the crown into anterior and posterior basins. The upper cheek 
teeth of later Triassic traversodontids such as Proexaeretodon and 
Exaeretodon of the Ischigualasto Formation, on the other hand, are 
similar to those of Massetognathus, but the "shouldering" of one 
tooth into the next is more pronounced. As Romer (1967: 20) 
states, "These differences suggest that Scolenodon is rather more 
primitive than the described South American forms." Further- 
more, trirachodontid cynodonts are present in both the Manda 
Formation and in the earlier Cynognathus Zone fauna of South 
Africa, but are absent in the Chafiares fauna. The relationships 
among all these cynodont faunas are thus more easily explained if 
the Chafiares fauna is later than that of the Manda. 

Romer (1966) has stated that the Chafiares fauna contains small 
thecodonts similar to Eiiparkeria of the Cynognathus Zone. Other, 
more advanced pseudosuchians are also present, however, and our 
knowledge of the Chafiares pseudosuchians is as yet too incomplete 
to provide any basis for a dating of the fauna. 

Rhynchosaurs are absent from the Chafiares fauna, but are found 
in both the Manda fauna and the Santa Maria fauna. As noted by 
Romer (1967), this fact is more easily explained if the Chafiares 
fauna is older than that of the Manda and represents a time at which 
the rhynchosaurs were not yet developed as a major faunal ele- 
ment. However, rhynchosaurs are also unknown in two of the three 
localities in the Santa Maria Formation, and the composition of 
the fauna of these two localities is exactly the same as that of the 
Chafiares fauna: dicynodonts (mainly Dinodontosaurus) , cyno- 
donts and pseudosuchians. Rhynchosaurs are known in the Santa 
Maria Formation only from a third locality, in which dicynodonts 
are absent (von Huene, 1935-42; Bortoluzzi and Barberena, 
1967). These facts suggest that, though rhynchosaurs and dicyno- 
donts are found together in the earlier Manda fauna, the ecological 
preferences of the South American type of rhynchosaur were dis- 
tinct from those of the dicynodonts of these faunas, so that these 
groups are rarely preserved together — and it is relevant to note 
here the extremely specialised dental apparatus of the rhyncho- 
saurs. If this is true, it is neither surprising nor significant that 
rhynchosaurs have not been found in the Chafiares fauna. 



1968 CHANARES DICYNODONT FAUNA 21 

To summarize, none of the evidence from tiie other vertebrate 
groups provides a convincing reason for rejecting the conclusion 
(based on the dicynodonts) that the Chanares fauna is only slightly 
earlier than that of the Santa Maria Formation of Brazil; it further 
appears to be later than that of the Manda Formation of East 
Africa. 

One may next attempt to assess the relative ages of these South 
American Triassic faunas. The Chafiares Formation of Argentina 
is somewhat older than the Los Rastros Formation, by which it is 
conformably overlain. On the other hand, it is also, on the evidence 
of the species of Dinodontosaurus, somewhat older than the Santa 
Maria Formation of Brazil, which may therefore be regarded as 
equivalent in age to the Los Rastros Formation of Argentina. Un- 
fortunately, vertebrate fossils have not been found in the Los 
Rastros Formation, so it is impossible to confirm the equivalence 
by direct faunal comparison. The footprints of Rigalites from the 
Los Rastros Formation could, however, well have been made by 
one of the large pseudosuchians of the Santa Maria Formation 
(Bonaparte, 1966b). 

There is also no unconformity between the Los Rastros Forma- 
tion and the Ischigualasto Formation in the Chanares area, but 
merely a decrease in the amount of sandstone and an increase in 
the amount of clay and shale. Despite this lack of any break be- 
tween the two formations, and the similarity between their cynodonts 
and rhynchosaurs, the archosaurs of the Ischigualasto fauna are 
considerably more advanced than those of Los Rastros/Santa 
Maria age. 

Finally, it is impossible to equate these Gondwanaland verte- 
brate faunas to the standard German and Alpine divisions of the 
Northern Hemisphere Triassic with any degree of accuracy 
(Romer, 1966; Cox, 1967). One can at present only attempt to 
define the extreme limits between which these faunas must lie. As 
argued earlier, even the Chafiares fauna is younger than the Manda 
fauna of East Africa. The latter contains the pseudosuchian reptile 
Mandasuchus, which is very similar to the genus Ticinosuchus, 
found in a definitely Anisian level of Monte San Giorgio, Switzer- 
land (Krebs, 1965; Charig, MS in preparation). All these South 
American faunas are therefore probably post-Anisian. At the 
other extreme, even the Ischigualasto fauna is earlier than the 
Upper Norian, since it contains armoured pseudosuchians but 
lacks coelurosaurs and large dinosaurs (cf. Chowdhury, 1965). 
In fact, this fauna is probably somewhat earlier than Norian: Bona- 
parte (1966b), after reviewing the whole Ischigualasto fauna, has 
concluded that it is probably of Carnian age. 



22 BREVIORA No. 295 

These conclusions are similar to those of Stipanicic, which are 
based upon evidence from fossil floras and ammonites, and which 
may be briefly considered here. The palaeobotanical evidence 
consists of the presence in the Los Rastros Formation (among 
others) of a flora which contains both Gondwanic Dicroidium ele- 
ments and also a considerable number of Northern Hemisphere 
species (Stipanicic, 1957). Stipanicic points out (In press) that 
the northern species are known in the Northern Hemisphere only in 
deposits of post-Ladinian age. However, the range of these species 
into the earlier Triassic is still unknown, and this evidence therefore 
cannot yet be regarded as conclusive. 

The ammonite evidence consists of the presence of specimens, 
identified by Barthel (1958) as Ciiccoceras n.sp.aff. cuccense and 
Beyrichites sp., in beds lying under the igneous Pastos Grandes 
Group in Chile. According to Stipanicic (1967), this group prac- 
tically interfingers with the Choiyoilitense Group of Argentina. 
Stipanicic (In press) believes that the Choiyoilitense Group is part 
of an igneous complex which is to be found below the various fos- 
sil vertebrate faunas and below the Dicroidium flora. Since the 
ammonites mentioned above indicate an Upper Anisian age, the 
overlying igneous complex and the fossil faunas and floras must all 
be post-Anisian. However, Kummel (personal communication) 
feels that the ammonite specimens are too incomplete for a definite 
taxonomic identification, and this line of argument must therefore 
be regarded as unproved. 

SUMAAARY 

Three new stahleckeriid dicynodonts from the Chaiiares Forma- 
tion of Argentina are described: Chanaria platyceps gen.et sp.nov., 
Dinodontosaurus brevirostris sp.nov., and Dinodontosaurus platy- 
gnathus sp. nov. A few post-cranial remains suggest that a kanne- 
meyeriid dicynodont was also present. 

Dinodontosaurus brevirostris is very closely related to Dinodon- 
tosaurus turpior of the Santa Maria Formation of Brazil, and is 
probably directly ancestral to it. The Santa Maria Formation there- 
fore appears to be only slightly younger than the Chanares Forma- 
tion, and equivalent to the Los Rastros Formation of Argentina. 

The Chanares fauna is therefore younger than the Manda fauna 
of East Africa; this conclusion is supported by the cynodonts of 
these faunas. Though rhynchosaurs are absent from the Chaiiares 
fauna, this is also true of two of the three localities in the Santa 
Maria Formation, and their absence is almost certainly merely 
ecological. 



1968 CHANARES DICYNODONT FAUNA 23 

The Chanares, Santa Maria and Ischigualasto faunas are, col- 
lectively, probably of Ladinian to Carnian age. 



ACKNOWLEDGMENTS 

I am, firstly, very grateful to Dr. A. S. Romer for generously in- 
viting me to describe these dicynodonts, which he collected during 
an expedition which was largely financed by the National Science 
Foundation, under grant No. GB2454. Most of the material was 
prepared and illustrated by my research assistant, Mr. P. Hutch- 
inson; I am greatly indebted to the Natural Environment Research 
Council for the grant which has allowed him to work with me, and 
for a special grant which enabled both of us to carry out this work 
at Harvard. Most of the preparation was done with the aid of NSF 
grant No. GB4615 to Dr. Romer. My grateful thanks are also due 
to the Royal Society, whose travel grant made it possible for me to 
visit Rio de Janeiro. Finally, I should like to thank Mr. L. I. Price 
of the Divisao de Geologia e Mineralogia, Ministerio das Minas e 
Energia, Rio de Janeiro, for allowing me to make use of his draw- 
ings for Figures 5E, F, and 8E, F. 

REFERENCES CITED 

Barthel, K. W. 

1958. Eine marine Faunula aus der mittleren Trias von Chile. Neues 
Jahrb. Geol. Palaont. Abh., 106: 352-382. 

Beltrao, R. 

1966. Paleontologia de Santa Maria e Sao Pedro do Sul, Rio Grande 
do Sul, Brasil. Bol. Inst. Cien. Nat. Univ. Fed. Santa Maria, 
No. 2: 3-114. 

Bonaparte, J. F. 

1966a. Una nueva "fauna" Triasica de Argentina (Therapsida: Cyno- 
dontia, Dicynodontia). Consideraciones filogeneticas y paleo- 
biogeograficas. Rev. Asoc. Pal. Argentina, 4: 243-296. 
1966b. Cronologia de algunas formaciones Triasicas Argentinas ba- 
sadas en restos de tetrapodos. Rev. Asoc. Geol. Argentina, 21: 
20-38. 

BORTOLUZZI, C. A., and M. C. Barberena 

1967. The Santa Maria beds in Rio Grande do Sul (Brazil). In: 
Problems in Brazilian Gondwana Geology, eds. J. J. Bigarella, 
R. D. Becker, and I. D. Pinto, Curitiba, Brazil, pp. 169-195. 



24 BREVIORA No. 295 

Camp, C. L., and S. P. Welles 

1956. Triassic dicynodont reptiles. Part I. The North American 
genus Placerias. Mem. Univ. California, 13: 255-304. 

Chowdhury, T. R. 

1965. A new metoposaurid amphibian from the Upper Triassic Maleri 
Formation of Central India. Phil. Trans. Roy. Soc. London, 
(B) 250: 1-52. 

Colbert, E. H. 

1958. Relationships of the Triassic Maleri fauna. J. Palaecnt. Soc. 
India, 3: 67-81. 

Cox, C. B. 

1965. New Triassic dicynodonts from South America, their origins 
and relationships. Phil. Trans. Roy. Soc. London, (B) 248: 
457-516. 

1967. Changes in terrestrial vertebrate faunas during the Mesozoic. 
In: The Fossil Record, ed. W. B. Harland, et al. London (Geo- 
logical Society), pp. 77-89. 

HUENE, F. VON 

1935-1942. Die fossilen Reptilien des slidamerikanischen Gondwana- 
landes. Munich, 332 pp. 

Klaauw, C. J. VAN DER, and J. M. van Roon 

1942. Fenestra medio-palatinalis und Interpterygoidalspalt. Arch. 
Neerland. Zool., 6: 327-340. 

Krebs, B. 

1965. Ticinosuchus ferox nov. gen. nov. sp. Ein neuer Pseudosuchier 
aus der Trias des Monte San Giorgio. Schweiz. Palaeont. 
Abhandl., 81: 1-140. 

Laws, R. M. 

1953. A new method of age determination in mammals with special 
reference to the elephant seal [Miroiinga leonina, Linn.). Falk- 
land Is. Dep. Surv. Sci. Rep., 2: 1-11. 

O'Donoghue, C. H. 

1920. The blood vascular system of the tuatara, Sphenodon punctotus. 
Phil. Trans. Roy. Soc. London, (B) 210: 175-252. 

Oelrich, T. M. 

1956. The anatomy of the head of Ctenosaura pectinata (Iguanidae). 
Misc. Publ. Mus. Zool. Univ. Michigan, 94: 1-122. 

Romer, A. S. 

1943. Recent mounts of fossil reptiles and amphibians in the Museum 
of Comparative Zoology. Bull. Mus. Comp. Zool., 42: 331-338. 



1968 CHAN ARES DICYNODONT FAUNA 25 

1966. The Chaiiares (Argentina) Triassic reptile fauna. I. Intro- 
duction. Breviora, Mus. Comp. Zool., No. 247: 1-14. 

1967. The Chaiiares (Argentina) Triassic reptile fauna. III. Two new 
gomphodonts, Massetognathiis pascuali and M. teruggii. Bre- 
viora, Mus. Comp. Zool., No. 264: 1-25. 

RoMER, A. S., and J. A. Jensen 

1966. The Chaiiares (Argentina) Triassic reptile fauna. II. Sketch of 
the geology of the Rio Chafiares — Rio Gualo region. Breviora, 
Mus. Comp. Zool., No. 252: 1-20. 

RoMER, A. S., and L. I. Price 

1944. Stahleckeria lenzii, a giant Triassic Brazilian dicynodont. Bull. 
Mus. Comp. Zool., 43: 465-490. 

Stipanicic, p. N. 

1957. El Sistema Triasico en la Argentina. XX Cong. Geol. Int. 
(Mexico, 1956), Section II: 73-112. 

1967. Consideraciones sobre las edades de algunas fases magmaticas 
del Neopaleozoico y Mesozoico. Rev. Asoc. Geol. Argentina, 
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In press. Triasico. Review, I Int. Symp. Gondwana Strat. Paleont. 

TuPi Caldas, J. 

1936. Paleontologia do Rio-Grande-Do-Sul. O fossil de Sao-Pedro. 
Rev. Inst. Hist. Geogr. Rio Grande do Sul., 1936: 243-249. 

(Received 5 January 1968.) 



26 BREVIORA No. 295 

LIST OF ABBREVIATIONS USED IN THE FIGURES 



a 


angular 


art 


articular 


bo 


basioccipital 


bsp 


basisphenoid-parasphenoid complex 


CO 


coracoid 


corf 


coracoid foramen 


d 


dentary 


ect 


ectopterygoid 


eo 


exoccipital 


ept 


epipterygoid 


f 


frontal 


fmp 


fenestra medio-palatinalis 


ip 


interparietal 


J 


jugal 


1 


lacrimal 


mx 


maxilla 


n 


nasal 


op 


opisthotic 


P 


parietal 


pal 


palatine 


pc 


processus cultriformis 


pco 


precoracoid 


pmx 


premaxilla 


po 


postorbital 


PP 


preparietal 


prf 


prefrontal 


pro 


prootic 


psp 


parasphenoid 



1968 CHANARES DICYNODONT FAUNA 27 



pt 


pterygoid 


q 


quadrate 


qj 


quadratojugal 


s 


sphenethmoid 


sa 


surangular 


sc 


scapula 


smx 


septomaxilla 


so 


supraoccipital 


sp 


splenial 


sq 


squamosal 


V 


vomer 




Harvard MCZ Libra 





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