/ •
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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1
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
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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 * *
• • • /
3
/
^31
- A .
*' y
«.
•
V • •
/
ca.
1-
-
-• /
•
>a
—
^27
/ ••
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~
O
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
BREVIORA
No. 275
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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
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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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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
No. 280
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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 ....
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'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
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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
13
11
Vi
^
o
1—
>x
^
ae
UJ
8
8
bO
K
o
7
LLm
o
6
6
OS
LU
ea
^s
13
4
3
6-9 1013 14-17 18-21 22-25 26-29 30-33
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
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a
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u
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<u
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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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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,
22: 101-133.
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
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