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HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



/ 



BREVIORA 



MUSEUM OF COMPARATIVE ZOOLOGY 
Harvard University 



Numbers 179-230 
1963-1965 



CAMBRIDGE, MASS., U.S.A. 
1965 



Edited 
By 

Nelda Wright 



CONTENTS 

BREVIORA 
Museum of Comparative Zoology 

Numbers 179-230 

1963 

No. 179, The holothurians of Clipperton Island in the eastern 
tropical Pacific. By Elisabeth Deichmann. 5 pp. 
January 16. 

No. 180. A new fresh-water amphipod crustacean from Oregon. 
By E. L. Bousfield. 6 pp. January 17. 

No. 181. Systematic notes on the land snails of the genus Tomo- 
cyclus (Cyclophoridae). By Fred G. Thompson. 
11 pp., 1 pi. February 1. 

No. 182. Birds from Flores, Lesser Sunda Islands. By Raymond 
A. Paynter, Jr. 5 pp. February 1. 

No. 183. Australian carabid beetles XIII. Further notes on 
Agonini, and a genus of Licinini new to Australia. By 
P. J. Darlington, Jr. 10 pp. March 12. 

No. 184. Behavior as a taxonomic clue: Relationships of Lisso- 
nyderis (Chiroptera). By Barbara Lawrence and 
Alvin Novick. 16 pp. April 18. 

No. 185. Eleutherodactylus hedricki, a new species of frog from 
Puerto Rico (Salientia, Leptodactylidae). By Juan 
A. Rivero. 7 pp., 1 pi. April 18. 

No. 186. Notes on Hispaniolan herpetology. 8. The forms re- 
lated to Anolis hendersoni Cochran. By Ernest E. 
Williams. 13 pp. April 18. 

No. 187. The labyrinthodont dentition. By John Newland Chase. 
13 pp. July 10. 

No. 188. Redescription of some cavernicolous pseudoscorpions 
(Arachnida, Chelonethida) in the collection of the 
Museum of Comparative Zoology. By William B. 
Muchmore. 16 pp. July 29. 



No. 189. Notes on amphisbaenids (Amphisbaenia; Reptilia). 
10. Redescription and redefinition of Amphisbaena 
pericensis Noble from the mountains of northwestern 
Peru. By Carl Cans. 15 pp. August 29. 

No. 190. Characters and synonymies among the genera of ants. 
Part III. Some members of the tribe Ponerini 
(Ponerinae, Formicidae). By William L. Brown, Jr. 

10 pp. September 30. 

No. 191. Three new species of M angora (Araneae, Argiopidae) 
from Central America. By Arthur M. Chickering. 

11 pp. December 5. 

No. 192. A description of Dinopis longipes F. P.-Cambridge, 
1902 (Araneae, Dinopidae). By Arthur M. Chicker- 
ing. 6 pp. December 5. 

No. 193. A Miocene toad from Colombia, South America. By 
Richard Estes and Richard Wassersug. 13 pp. 
December 5. 

No. 194. A new subspecies of Tropidophis greenwayi from the 
Caicos Bank. By Albert Schwartz. 6 pp. December 
31. 

No. 195. Cayman Islands Tropidophis (Reptilia, Serpentes). By 
Richard Thomas. 8 pp. December 31. 

No. 196. A new genus and species of bathypelagic ophidioid fish 
from the western North Atlantic. By Daniel M. 
Cohen. 8 pp. December 31. 

No. 197. Anolis whitemani, new species from Hispaniola (Sauria, 
Iguanidae). By Ernest E. Williams. 8 pp. December 
31. 

1964 

No. 198. Amphisbaena schmidti, a third species of the genus from 
Puerto Rico (Amphisbaenia: Reptilia). By Carl 
Cans. 11 pp. March 10. 

No. 199. A new subspecies of Varanus exanthematicus (Sauria, 
Varanidae). By R. F. Laurent. 9 pp. March 10. 

No. 200. An anguid lizard from the Leeward Islands. By Garth 
Underwood. 10 pp. April 10. 

No. 201. Food habits and young stages of North Atlantic Alepi- 
saiirus (Pisces, Iniomi). By Richard L. Hacdrich. 
15 pp. April 10. 

No, 202. The blind snakes (Typhlops) of Haiti with descriptions 
of three new species. By Neil D. Richmond. 12 pp. 
April 10. 



A new capromyid rodent from the Quaternary of His- 
paniola. By Clayton E. Ray. 4 pp. April 10. 

The status of Pseudogekko shebae and observations on 
the geckos of the Solomon Islands. By Walter C. 
Brown. 8 pp. May 15. 

Redescription of Amyhishaena duhia Miiller (Amphis- 
baenia: Reptiha). By Carl Gans. 11pp. July 15. 

The aistopod amphibians surveyed. By Donald Baird. 
17 pp. July 15. 

Notes on the horseshoe bats Hipposideros caffer, ruber 
and beatus. By Barbara Lawrence. 5 pp. July 28. 

Three new species of frogs (Leptodactylidae, Eleuthero- 
dactylus) from Hispaniola. By Albert Schwartz. 
15 pp. November 12. 

A new skate, Raja cervigoni, from Venezuela and the 
Guianas. By Henry B. Bigelow and William C. 
Schroeder. 5 pp. November 12. 

The ants of the Florida Keys. By Edward 0. Wilson. 
14 pp. November 12. 

A new species of the snake Leptotyphlops from Colombia- 
By Benjamin Shreve. 4 pp. December 21. 

A new species of freshwater gastropod mollusc of the 
genus Saulea from the Miocene of Kenya. By 
Thomas Pain and Dawn Beatty. 5 pp., 2 pis. 
December 30. 

1965 

No. 213. A hitherto overlooked Anodonta (Mollusca: Unionidae) 
from the Gulf drainage of Florida. By Richard I. 
Johnson. 7 pp., 2 pis. January 11. 

No. 214. A revised classification of the dendrochirote holo- 
thurians. By David L. Pawson and H. Barraclough 
Fell. 7 pp. February 15. 

No. 215. Two new subspecies of Amphisbaena (Amphisbaenia, 
Reptilia) from the Barahona Peninsula of Hispaniola. 
By Richard Thomas. 14 pp. February 15. 

No. 216. The geographical variation of the frog Hyperolius 
marmoratus (Family Hyperoliidae) in Rhodesia, 
Nyasaland and Tanganyika. By R. F. Laurent. 
9 pp. February 15. 



No. 


203. 


No. 


204. 


No. 


205. 


No. 


206. 


No. 


207. 


No. 


208. 


No. 


209. 


No. 


210. 


No. 


211. 


No. 


212. 



No. 217. The auditory region of the borhyaenid marsupial 
Cladosictis. By Bryan Patterson. 9 pp. March 1. 

No. 218. New frogs of the genus Cornufer (Ranidae) from the 
Solomon Islands. By Walter C. Brown. 16 pp., 2 
pis. May 7. 

No. 219. The early evolution of the Echinozoa. By H. Barra- 
clough Fell. 17 pp. May 7. 

No. 220. A new species of Eleutherodactylus from Guadeloupe, 
West Indies. By John D. Lynch. 7 pp. May 7. 

No. 221. New Melanesian ants of the genera Simopone and 
Amblyopone (Hymenoptera — Formicidae) of zoo- 
geographic significance. By Robert W. Taylor. 11 pp. 
May 7. 

No. 222. The genus Leptotyphlops in the West Indies with des- 
cription of a new species from Hispaniola (Serpentes, 
Leptotyphlopidae). By Richard Thomas. 12 pp. 
May 28. 

No. 223. A new subspecies of Clelia clelia (Serpentes: Colubridae) 
from the island of Grenada. By Allen E. Greer. 6 pp. 
May 28. 

No. 224. New species of land mollusks with notes on other 
species from the Solomon Islands. By William J. 
Clench. 8 pp., 2 pis. July 15. 

No. 225. The Asian species of Galeritula Strand (Coleoptera, 
Carabidae). By Hans Reichardt. 16 pp. July 15. 

No. 226. The larval form of the Heteromi (Pisces). By Giles W. 
Mead. 5 pp. July 15. 

No. 227. The species of Hispaniolan green anoles (Sauria, 
Iguanidae). By Ernest E. Williams. 16 pp. Septem- 
ber 10. 

No. 228. Relationships among Indo-Australian Zosteropidae 
(Aves). By Ernst Mayr. 6 pp. September 15. 

No. 229. The genus DarUngionia (Serpentes) in Hispaniola, in- 
cluding a new subspecies from the Dominican Re- 
public. By Albert Schwartz and Richard Thomas. 
10 pp. September 15. 

No. 230. Notes on some non-passerine birds from eastern Ecua- 
dor. By David W. Norton. 11 pp. September 15. 



INDEX OF AUTHORS 

BREVIORA 

MUSEUM OF COMPARATIVE ZOOLOGY 
Numbers 179-230 

1963-1965 

No. 

Baird, Donald 206 

Beatty, Dawn 212 

BiGELOW, Henry B 209 

BOUSFIELD, E. L 180 

Brown, Walter C 204, 218 

Brown, William L., Jr 190 

Chase, John Newland 187 

Chickering, Arthur M 191, 192 

Clench, William J 224 

Cohen, Daniel M 196 

Darlington, P. J., Jr 183 

Deichmann, Elisabeth 179 

EsTES, Richard 193 

Fell, H. Barraclough 214, 219 

Gans, Carl . 189, 198, 205 

Greer, Allen E 223 

Haedrich, Richard L 201 

Johnson, Richard 1 213 

Laurent, R. F 199, 216 



Lawrence, Barbara 184, 207 

Lynch, John D 220 

Mayr, Ernst 228 

Mead, Giles W 226 

MucHMORE, William B 188 

Norton, David W 230 

NovicK, Alvin 184 

Pain, Thomas 212 

Patterson, Bryan 217 

Pawson, David L 214 

Paynter, Raymond A., Jr 182 

Ray, Clayton E 203 

Reichardt, Hans 225 

Richmond, Neil D 202 

Rivero, Juan A 185 

schroeder, william c 209 

Schwartz, Albert 194, 208, 229 

Shreve, Benjamin 211 

Taylor, Robert W 221 

Thomas, Richard 195, 215, 222, 229 

Thompson, Fred G 181 

Underwood, Garth 200 

Wassersug, Richard 193 

Williams, Ernest E 186, 197, 227 

Wilson, Edward 210 



BREVIORA 

Mmseiiioi of Comparative Zoology 



Cambridge, Mass. Jaxiary 16, 1963 Xtmber 179 

THE HOLOTHURIANS OF CLIPPERTON ISLAND IN 
THE EASTERN TROPICAL PACIFIC 

By Elisabeth Deichmann 

Museum of Comparative Zoology 
Cambridge, Massachusetts 

In 1902 H. L. Clark listed two typical Iiido-West-Pacific 
aspidochirote holothurians from Clipperton Island, though at 
that time he considered one of them a new species. Later expe- 
ditions brought no holothurian material from this island. It 
was not until 1958 that Deichmann was able to include in her 
Hancock report some material of a third species, collected by 
the Scripps Institution's expedition a few years before. Finally, 
in 1958, the University of California Clipperton Expedition, 
which used more refined methods, with diving, etc., brought 
back four species, two of which had never been reported from 
any locality in the eastern tropical Pacific. 

Of the five species now known from Clipperton Island, only 
one, Microtliele difficilis (Semper) appears to be well established 
on the mainland of the Pacific coast of America. Two other spe- 
cies had previously been listed from some of the outlying islands, 
Semperothuria atra (Jaeger) and Mertensiothuria leucospilota 
(Brandt). Of the remaining two forms, Stichopus korrens 
Selenka represents a typical Hawaiian form, while Sempero- 
thuria flavomaculata (Semper) is considered rare; it was orig- 
inally described from Samoa and later listed from Tahiti, and 
(?)Batavia in the East Indies. 

The conclusions one can draw from this short list are that 
more Indo-West-Pacific forms are able to cross, at least inter- 
mittently, the ' ' Ekman Barrier ' ' than hitherto assumed, and that 
some of these forms may have escaped notice because they are 
living at a greater depth here than that in which they normally 
live in the more favorable localities of the Indo-West-Pacific. 



2 BBEVIORA No. 179 

STICHOPODIDAE 

1. Stichopus horrens (Selenka) 1867 

Stichopus horrens Selenka, 1867, p 316, pi. 18, figs. 27-29. H. L. Clark, 

1922, p. 64, pi. 2, figs. 19-23. 
Stichopus godeffroyi Semper, 1868, p. 7.5, pi. 130, fig. 4; var. pygmaeus, 

p. 75, var. b, p. 246. 
Stichopus tropicalis Fisher, 1907, p. 676, pi. 70, figs. 1-11. 

The five specimens secured measure 15-20 cm. in length and 
agree well with Fisher's description of his material from Hawaii. 
All the specimens have numerous C-shaped bodies of all sizes in 
the skin ; this is the character which is supposed to separate 
tropicalis (= Semper 's var. b.) from the typical godeffroyi, but 
I fully agree with H. L. Clark in regarding the presence or 
absence of these structures as most unimportant. The typical 
tables — with a pointed spire — are present, although not in 
large numbers in some individuals, and likewise the peculiar 
rosettes could be located in all five individuals. 

The color of four of the specimens is indicated to be "mottled 
with orange and jiale cream," while the fifth is "mottled dull 
greenish," a color range which agrees well with H. L. Clark's 
observations on horrens. Fisher Avrites : ''dark olive green, mot- 
tled with deep brownish green." His material, 16 cm. long, in 
preserved condition, came from tide pools in Haw^aii. 

While most earlier records (from Samoa, Fiji and Hawaii) 
are from shallow water, the present material (Clipperton A- 
588-6) came from 10-20 meters depth, off the edge of "the 10 
fathoms terrace" on the northern side of the atoll: it was noted 
that the species did not occur at greater depth (40 meters) off the 
edge of that terrace. It was also observed, at similar depth, on 
the other side of the island, and the species appears therefore to 
be well established in this locality. 

HOLOTHURIIDAE 

2. MiCROTHELE DiFFiciLis (Semper) 

Holothuria diffidlis Semper, 1868, p. 92, pi. 30, fig. 21. 
Microthele difficilis:—'Deichmann, 1958, p. 288, pi. 1, figs. 6-9 (list of ref- 
erences). 

Four dark brown individuals, 6-7 cm. long, were collected 
(Clipperton A-r)88-8) on the reef flats "from the sbore out, at 
least to the weak algal ridge." 



1963 llOLOTIirRIANS OF CLII'PEKTON ISLAND 3 

The pr('seiu'(> of this sjx'cies in Clipperton is not surprising, 
since this atoll was the type locality for IT. L. Clark's '"Holo- 
iliuria frcquentiamcnsis," described in 1902 on the basis of 17 
individuals, about 4 cm. long-, of which one had immature geni- 
tal organs. 

The species ranges from the east coast of Africa to the Panamic 
region, where it is known from the Galapagos Islands, north- 
wards to Espiritu Santos Island in the Gulf of California. 

.■). Se:\iperotiiuria flavomaculata (Semper) 

nolotlturia flavomaculata Semper, 1868, pp. 87, 277, pi. 30, fig. 26. Pan- 
ning, 1934, pt. II, p. 42, text-fig. 35 (list of references). 

Scmperotliiiria flavomncuhiia : — Deichmann 1958, p. 303 (treated in the key 
for the genus). 

lentil the present study, there were only a few^ species belong- 
ing to the old genus llolothiiria of which the Museum of Com- 
parative Zoology did not possess representatives, and flavomacu- 
lata was one of these. When the key for the few^ members of the 
genus 8cmperothuria was worked out in 1958, it was necessary 
to rely entirely on the literature. It was consequently a satis- 
faction to discover two specimens of this comparatively rare 
species in the Clipperton material and to find that they agreed 
with the key. 

Both specimens came from Clipperton A-588-6, from the 10 
fatlioms terrace, in w-ater varying from 1-20 meters, off the edge 
of the tidal flat. The species was not noted by divers off the 
edge of that terrace, in water up to 40 meters depth. 

The two individuals (M.C.Z. no. 3008) measure 8 and 12 cm. 
in length; they are dark, with few feet in scattered rows on the 
ventrum and more sparingly developed as papillae on the dor- 
sum, here with a yellow area around their base. The spicules 
agree completely with the earlier descriptions: a crowded layer 
of tables with no disk, but a base which tapers to a point, four 
pillars and the spire ending in a double Maltese cross. In the 
deeper layer are scattered short rods, covered by coarse, rough 
spines, or clusters of spines. 

The type, 11 cm. long, came from Samoa, and according to 
Panning the species has also been taken in Tahiti and (?)Bata- 
via. It may possil)ly. when contracted, have been mistaken for 
Ludwi(jothuria atra, or some of the other dark-skinned forms 
in the Indo-Pacific. It was at first assumed that the material 



4 BREVIORA No. 179 

represented atra, previously known from Clipperton Island, but 
the roughness of the skin eliminated that possibility, even before 
the spicules had been examined. 

4. Mertensiothuria leucospilota (Brandt) 

Stichopus leucospilota Brandt, 1835, p. 51. 

Mertensiothuria leucospilota: — Deiclimann, 1958, p. 297, pi. 3, figs. 1-9 
(list of references). 

Three large, reddish-brown individuals (Clipperton A-588-8) 
were collected on reef flats, identical in size and spiculation 
with those taken some years ago by the Scripps Clipperton Ex- 
pedition, and included in Deichmann's 1958 report. 

The species has the same wide range as Microthele difficilis, 
from the east coast of Africa to the Panamic region, but so far 
it has never been found established on the mainland. The Han- 
cock expeditions reported it from Galapagos, Clarion and Socorro 
islands. The largest individuals, 20 cm. long, in preserved con- 
dition, have all come from Clipperton Island. 

5. LUDWIGOTHURIA ATRA (JaCgCr) 

Holothuriu atra Jaeger, 1833, p. 22. Panning, 1934, pt. II, p. 30, text-fig. 

22 (list of literature). 
Ludwigothuria atra: — Deiclimann, 1958, p. 312, pi. 2, figs. 18-23. 

Of this species H. L. Clark reported nine specimens from 
Clipperton Island in 1902. The Hancock expeditions collected 
thirteen in the Galapagos Islands and one in Cocos Island. The 
species is known to range from Mozambique to Hawaii ; in the 
latter locality it is stated by Fisher to be "one of the common- 
est holothurians inhabiting Hawaiian shores." 

The fact that the recent expeditions to Clipperton Island did 
not collect this large and conspicuous species, in spite of the 
intensive collecting undertaken, may indicate that this is one 
of the species which only intermittently crosses the "Ekman 
Barrier" but is unable to become permanently established in 
the less favorable localities which it finds in the eastern Pacific. 

REFEEENCES 

Brandt, J. F. 

1835. Ecliinoderniata ordo Holothurina. In Prodroaius descriptionis 
aninialium. . . . Petropoli, Fasc. 1, pp. 242-262. 



1963 HOLOTIIURIANS OF CLIPPERTON ISLAND 5 

Clabk, II. L. 

1902. Papers from the Hopkins Stanford Galapagos Expedition. 
Vol.4, art. 12, Echinoderniata. Proc. Washington Acad. Science, 
vol. 4, pp. 521-531. 
1922. The holothurians of the genus Stichopus. Bull. Mus. Comp. 
Zool., vol. 65, no. 3, pp. 39-74, pis. 1-2. 
Deighmann, E. 

1958. The Holothurioidea collected by the Velero III and IV during 
the years 1932 to 1954. Part II. Aspidochirota. Allan Hancock 
Pacific Expeditions, vol. 11, no. 2, pp. 251-346, pis. 1-9. (Con- 
tains a number of new generic names for various members of 
the old genus Holothuria.) 
Fisher, W. K. 

1907. The holothurians of the Hawaiian Islands. Proc. U. S. Na- 
tional Museum, vol. 32, pp. 637-744, pis. 66-82. 
Jaeger, G. F. 

1833. De Holothuriis. Diss. Inaug. Turici (Torino, Italia), 40 pp., 
3 pis. 
Panning, A. 

1928- Die Gattung Eoloihuria. Parts I-V. Mitt. Staats Inst, und 

1935. Zool. Museum Hamburg, vol. 44, pp. 91-138, 21 text-figs., 1 

chart; vol. 45, pp. 24-50, 26 text-figs., 3 charts; ibid., pp. 65-84, 

27 text-figs.; ibid., pp. 85-107, 32 text-figs.; vol. 46, pp. 1-18, 

19 text-figs., and index to all five parts. 

Selenka, E. 

1867. Beitrage zur Anatomic und Systematik der Holothurien. Zeit- 
schrift Wiss. Zool., vol. 17, pp. 291-374, pis. 17-20. 

Semper, C. 

1868. Reisen im Archipel der Philippinen, II. Wissenschaftliche Re- 
sultate, vol. 1, Holothurien, 299 pp., 10 pis. 



^^z 



^U 



BREVIORA 

MmsemitTn of Comparative Zoology 



Cambuidge, Mass. January 17, 196;> Number 180 

A NEW FRESH-WATER AMPHIPOD CRUSTACEAN 

FROM OREGON 

By E. L. BousFiELD 
National Museum of Canada 



In a recent account of the fresh-water amphipod fauna of 
Oregon (Bousfield, 1961), the writer listed ten known species 
including two species of Crangonyx, C. richmondensis occiden- 
talis Hubricht and Harrison and C. pseudogracilis Bousfield. Dur- 
ing a further examination of Crangonyx material from small 
alpine lakes in southwestern Oregon, a third species was identi- 
fied. It proved to be distinct from other North American species 
studied by the writer (Bousfield, 1958) and from other known 
species of the genus and is herewith described as Crangonyx 
alpinus sp. nov. 

The present study is part of a general survey of gammaridean 
amphipods in the collection of the Museum of Comparative 
Zoology, Harvard University, conducted in May, 1962, with the 
aid of a grant from the National Science Foundation. The writer 
wishes to thank Dr. Elisabeth Deichmann for instigating the 
study and for her many kindnesses and co-operation during the 
undertaking. 

In the list of specimens, MCZ refers to the Museum of Com- 
parative Zoology, and NMC to the National Museum of Canada. 

Family GAMMARIDAE 

Crangonyx alpinus n. sp. 

(Figs. 1, 2)  

Material examined: A total of 52 specimens, collected in alpine 
lakes of Lane and Douglas counties, Oregon, by F. Ziesenheim 
during the summer of 1937. as follows : Corner L. (alt. 4800 ft.), 
Lane Co., July 29, MCZ No. 10027 — 3 fem., 12 juv. ; Ledge L. 



2 BREVIORA No. 180 

(5350 ft.). Lane Co., Aug. 5, MCZ No. 10028 — 1 ovig. fern. 
(TYPE), 1 jiiv.; Pork L. (4820 ft.). Lane Co.. Aug. 9, MCZ 
No. 10029 — 3 fern. imm. ; Plumb L.. Lane Co., Aug. 21, MCZ 
No. 10030 — 6 juv. : Opal L. (5480 ft.). Douglas Co., Sept. 7, 
MCZ No. 10031 — 8 imm.: Whig. L. (5270 ft.), Douglas Co. 
Sept. 24, MCZ No. 10032 — 2 imm.; Emma L. (5190 ft.). Lane 
Co., Sept. 25, MCZ No. 10033 — 6 fem. imm.. 4 male imm.; 
Easter Brook L. (5050 ft.), Lane Co.. Sept. 27, MCZ No. 10034 
— 1 fem. (Br. 1), 3 juv.; Mud L. (4950 ft.). Lane Co., Sept. 29, 
NMC No. 10035 — 1 fem. imm. 

Diagnosis: A small species of the richtnonderisis group hav- 
ing elongate antennae, peraeopods and uropods, reduced mouth- 
parts, shallowly cleft telson, and powerful gnathopods. but 
distinguished by the acuminate abdominal side plates, sharply 
serrated ba.sal segments of peraeopods 3-5, singly inserted pos- 
terior marginal setae of segment 6 of gnathopod 1, long marginal 
spines of uropod 3 and telson, and by the small eye. 

Female (10.5 mm.). Eye small, black, irregularly oval, re- 
moved from anterior head margin. Antenna 1. flagellum of 25 
segments; accessory flagellum shorter than 1st flagellar segment. 
Antenna 2. flagellum of 9 segments. 

Lower lip, inner lobes, distinct, rather broad. Mandil)ular 
palp, terminal segment slender, with 3 outer marginal setae. 
Maxilla 1, inner plate with only two plumose marginal setae. 
Maxilla 2, plates relatively narrow; inner plate with only one 
elongate plumose facial seta. Maxilliped, inner plate short, 
truncate apex with 5 slender spine-teeth ; outer plate small, 
outer margin convex ; dactyl of palp stout. 

Lower corners of coxal plates 1 and 2 rounded, each with 3-5 
short marginal setae. Gnathopod 1, posterior margin of segment 
2 with numerous slender setae, anterior margin nearly bare ; 
posterior margin of segment 5 with 3-4 clusters of long, slender, 
distally flexed spines, some minutely pectinate ; segment 6 
(propodus) subquadrate, widest distally; palmar margin evenly 
convex, slightly oblique, armed with about 12 medium-small 
spine-teeth on each side, strongest and closely crowded near 
posterior angle ; posterior margin with 6-7 slender setae appear- 
ing singly inserted ; dactyl fairly heavy, closely fitting palm. 
Gnathopod 2, segment 2 distally broadening, margins with sev- 
eral long setae ; posterior margin of segment 5 with 5 posterior 
groups of setae ; segment 6 subrectangular, distally broadening ; 



1963 NEW FRESH -WATER AMPIIIPOD 3 

palm smoothly convex, oblique, lined with widely-spaced medium- 
strong spine-teeth ; posterior angle with one prominent spine and 
another smaller spine ; posterior margin nearly two-thirds the 
anterior ; inner face of propodus with 4 groups of superior lateral 
setae. 1-3 setae per cluster ; dactyl rather slender. 

Peraeopods 1 and 2 slender, subequal ; posterior margins of 
segments 4-6 moderately spinose. Coxal plate of peraeopod 2 
nearly as broad as deep, proximally emarginate behind. Peraeo- 
pods 3-5 long and slender, 4th longest. Basal segments of 
peraeopods are similar in size and shape ; posterior margins 
gently convex, with about 7-9 rather deep serrations, distally 
sharpest ; length of dactyls y^ to ^ the propods. 

Brood plates moderately large ; setae numerous, elongate, 
minutely cleft at tip. Coxal gills present on segments 2-7, paired 
sternal gills on segment 6, two pairs on segment 7. 

Pleopods powerful, 1st strongest ; rami about twice the length 
of the peduncle, inner ramus somewhat longer than outer. 
Peduncles of pleopods each with a few simple marginal setae 
and 3-4 coupling spines of at least two types. Proximal setae 
of inner ramus are non-plumose and bifid at the tip. 

Abdominal side plates, posterior margin shallowly incurved 
distally, corners sharply acuminate and produced posteriorly, 
most strongly in side plate 2. 

Uropods 1 and 2 rather long and slender, lateral margins of 
both rami armed with short spines. Uropod 3, outer ramus 
slender, about twice the peduncle, lateral margins armed with 
about 4 groups of longish spines ; inner ramus with 1 sub-apical 
spine ; peduncle with 1 or 2 lateral marginal spines. Telson about 
as broad as long, shallowly emarginate, each lobe terminated by 
2 longish spines. 

Remarks: Mature males were not present in material at hand. 
The principal breeding period is probably June and early July 
following which the adults die off. 



References 

BOUSFIELD, E. L. 

1958. Fresh-water amphipod crustaceans of glaciated North America. 

Can. Field-Nat. 72(2) : 55-113. 
1961. New Eecords of fresh-water amphipod crustaceans from Oregon. 

Nat. Mus. Can. Nat. Hist. Paper No. 12, 7 pp. 



BREVIORA 



No. 180 



Abbreviations for the Figures 



A 1 — Antenna 1 
A2 — Antenna 2 
LL — lower lip 
Lit Md — Left mandible 
Rt Md —Right mandible 
Mxl — Maxilla 1 
Mx2 — Maxilla 2 
Mxpd — Maxilliped 
Gnl — Gnathopod 1 
(jn2 — (Juathopod 2 
I'l — peraeopod 1 
P2 — peraeopod 2 



P3- 
P4- 
P5- 
PLl 



peraeopod 3 
peraeopod 4 
peraeopod 5 
— pleopod 1 



PL3 — pleopod 3 

Ul — uropod 1 

U2 — uropod 2 

U3 — uropod 3 

T — telson 

EPl — abdominal side plate 1 

EP2 — abdominal side plate 2 

EPS — abdominal side plate 3 



1963 



NEW FKESII-WATER AMIMIIPOD 




Figure 1. Crangonyx alpinus n. sp. Ledge L., Lane Co., Oregon, August 5, 
1937, female (TYPE) 10.5 mm. 



6 



BltEVIORA 



No. 180 




Figure 2. Crangonyx alpinnfi ii. sp. Ledge L., Lane Co., Oi'egon, August 5, 
U);37, fomnlo (TYPE) 10.5 mm. 



BREVIORA 



MiasemtM of Coimpsirative Zoology 



(\\MBRinGi-:, Mass. Kkhkiaky 1, 1963 Numbkr ISl 

SYSTEMATIC NOTES ()X THE LAND SXAILS OK THE 

(JEXCS TOMOC'YCLrs 
(CVCLOIMIORIDAE) 

By Fkki) (i. Tiio.Mi'sox 

Depiiitiiieiit of Zoology 
University of Mi;mii 

The <i•ellll^s TomociicUis includes iieotroi)ifal t'y('l()i)lu)i-id siuuIn 
with a ehondroid oiiorciihini, bearino; a spiral lamella, an eloii- 
yate-turrite shell and <i double peristome. The outer peristome 
flares to pi-oduee a bi-oad eollar around the aperture. The parietal 
region of the outer jxTistome is bent anteriorly to pass under 
the i)reeeding whoii. At this point a notch. th(» iiseudosiphon. 
oeeurs in the reflected outei- peristome. 

Present knowled<ie of the anatomy of To)nocuchis is based on 
the account of Fischer and Crosse (1886). The ver<re is lono. 
simple, tapering- and unbranclied. Tt originates on the right 
side of the neck, behind the tentacle, and ])ossesses an open 
seminal groove. The jaw is typically cyclophorid, with a median 
I)rojection. The radular formula is 3-3-3-3. Morrison (1955: 
152) classified the American cyclophorid snails, and i)laced 
Tonwcyclus in the subfamily Neopupinae. an allocation comply- 
ing with most ])revious systems of classification. Torre and 
Bartsch (1942: 3) proposed the name Megalomastominae for 
this same subfamily, because the name Neopupinae was proposed 
five years prior to its assumed type genus, Ncopupina (Kobelt. 
1902: 261), and because they recognized Xcopupina as a sub- 
genus of Mcgalomasfoma. Whether Xcopupina is or is not dis- 
tinct from McgaIomasto))io, the name ]\Iegalomastominae shoidd 
be retained for this subfamily. 

The genus Tomocyclus is known from British Honduras, east- 
ci-n (ruatemala, and in ]Mexico from Chiapas and the isolated 
mountain region of San Andi-cs Tuxthi. Vera Cvw/.. 



2 BKEVIORA No. 181 

MEASUREMENTS 

For purposes of This study, ineasnrcinonts of height were 
made to iiicliule only the last four I'diiainiiiu' whorls. Mature 
specimeus of TotnociicJus arc generally decollate, and shells in 
a single lot may have 4-8 whorls remaining. Therefore, measure- 
ments must be made from a ]ioint that is consistently ])resent. 
and represents the same relative growth. 

Measurements of length and diameter were made with vernier 
calipers, and are standard except for the special condition men- 
tioned above. Measurements of tlie ai)erture and outer peri- 
.stome were made with a calibrated Whipple disc. The diameter 
of the aperture Avas measured from the outside of the extended 
inner peristome. The width of the outer peristome (the collar) 
was measured at four places, and the readings were averaged. 

ACKNOWLEDGEMENTS 

I wish to express my gratitude to the following people for 
lending specimens in their charge and foi' other courtesies that 
have facilitated this study: Dr. Tucker xVbbott, Academy of 
Natural Sciences of Philadelphia (ANSP): Dr. AVilliam J. 
Clench, Museum of Comparative Zoology, Harvard T'niversity 
(MCZ) : Dr. Harold A. Kehder, United States National Museum 
(T'SNM) ; and Dr. Henry van der Schalie, Museum of Zoology. 
T^nivcrsity of Michigan (UMMZ). The photographs for Plate 
I were taken by William L. Bruden, staff artist of the ]\Iuseum 
of Zoology, University of Michigan. 

Key to the Species of Tomocyclus 

1. Basal caiiiia present 2 

la. Basal rariiia aljsent 4 

2. Axial sculpture consisting of well developed riblets gealci 

2a. Axial sculpture consisting of fine striatious 3 

3. Basal carina passing in front of reflected outer peristome; diameter of 

aperture 7.45-9.50 mm. fistulosus 
3a. Basal carina pas.sing beneath or behind redected outer peristome; diam- 
eter of aperture 5.73-7.13 mm. simulacrum 

4. Width-height ratio aliout 0.4 anatemalcyisU 
4a. Width-height ratio nbout 0.5 hnxac 

Tu-MucYcLUS GEALEi Crossc aud Fischer 

Tomocyclus gcalel Trosse and Fischer, T<72, .lour. Couchyl., 20: 70; Fisclier 
aiul Crosse, 1880, Miss. Sei. Mex. Aiiicr. Cent., 2 (7): 118-120, pi. 40, 



l!»Go XOTKS ox 'I'O.MOCYCLIS 3 

ti<rs. 1-;;: Ilintscli ami Mdirisdii. ^9A•2. I'.ull. V. S. Xnt. Mus., 181: 143-144, 

1)1. lit, fitr. 1. 
Mrcialomastoma {Tomncyclns ) (/lalci (Crosse ;iii(I I'^ischer ). vim Martens, 

18!t(l, Biol. (Vnt. Auu'v., 9: 10. 

Tiip( hicdlitji: State of Chiapas. Moxico. 

lUcurds: (;rATE:\IALA: AHa Vrra I'a:: woods hctwocii Tac- 
tic and Tainahu; Polocliic \'all('y above Pan/.os and Seiialni 
(von :\Iartens. 18!)(): 10). 

To.Mocvci.rs I'lSTlLOsrs. new species 

(PI. I. tio's. 1-8) 

Deso-ipflmi: Sliell dull yellow, with early whorls and last 
quarter of last whorl beeoniino^ ciiinanion. or rosy: translucent, 
lar^e, clono-ate-turrite ; decollate. 5I4 whorls reniaininti' in the 
liolotype : whorls gradually increasinp- in size, fourth from last 
whorl ahont ().() times diameter of last whorl, rate of increase 
in diametei- of whorls nearly constant; spire slio-htly convex, 
nearly straight sided; whoi-ls eonv(>x, suture^ moderately im- 
pressed; whorls crossed by fine, ])oorly developed, ])osteriorly 
arched jirowth wriidvles. which lie about 0.5 nun. ajiart ; wrinkles 
faintly discernible, di. linctness and s])acin.o' of wrinkles stead- 
ily increasinp' on last whorl ; wrinkles continuin<i- onto reflected 
outer peristome ; whorls also with numerous fine spiral stria- 
tions. which are frequently bi'oken alono- their courses; base of 
last whoi'I with a stronji' carina, which passes in front of the 
aperture (PI. 1, fi<;'. 3) ; carina orijiinating- on last half of penulti- 
mate whorl, and continuing' nearly to reflected peristome; last 
half of last whorl lying- just inside of crest of carina; peristome 
double; outer peristome dull white, forming a slight irregular 
collar, 0.97-"2.()(j mm. wide, 0.1-0.24 times the diameter of the 
aperture; face of collar witli several annulations, which are 
crossed l)y numerous close, fine, irregular, granular ridges that 
occasionally anastomose; parietal region of collar indented by a 
nearly rectangular pseudosiphon notch, about 1 mm. deep and 
2 mm. wide ; aperture dull white, circular ; columella tubular, 
continuing spirally through length of shell, open at first remain- 
ing whorl and umbilicus as a narrow slit : umbilicus nearly ob- 
scured by collar. Operculum dark brown, typically tomocycloid. 
consisting of a thin inner chondroid plate, and an external spiral 
lamella ; inner surface of chondroid plate bearing a central knob ; 
spiral lamella originating in center of operculum, and reflected 
to lie i)arallel to chondroid plate; edge of lamella frequently 
broken to produce a fringed appearance. 



4 BREVIORA Xo. 181 

McasiirojK )ifs of holotypc: total leiijith, 32.5 mm.: leno'th of 
last four whorls 30.0 mm.: diameter. 11.2 mm.; major heio'ht of 
aperture, 10.9 mm.: major width of aperture. 11.02 mm.: height 
of inner peristome, 7.56 mm. ; width of inner peristome, 7.78 mm. 

Mensii}r))irjifs of pnrnfiiprs: 



Length 


DiamettM- 


W 


idtli (1 


<! ( 


:'nll; 


ir 


Tuner Peristome 


:28.5 mm. 


in.9 mm. 




0.97 


mm. 




9.50 mm. 


29.7 


10.8 




1.78 








7.83 


28.6 


10.7 




1.48 








7.4.3 


33.1 


11.8 




2.0(5 








8.48 



Major variations involve only size of the shell and width of the 
collar, and even these variations are relatively slight. 

Holutypc TMMZ 194095. high rainforest at Vallentine (^amp. 
50 miles southwest of t'ayo, British Honduras. Collected by 
C. L. Lundell and E. B. Mains in 1936 while conducting botani- 
cal investigations. 

/'(Ddfupis: T'MMZ 66226 (4 specimens) : same data as the 
holotype. 

T. fisfiilosus appears to be most closely related to T. sim}(lac- 
nou. Characters which separate these two species from the 
other species of the genus are the presence of a basal carina and 
fine axial sculpture. 

T. fistulosus is separated from T. .nmulacru))i on the basis of 
five characters: (1) the diameter of the fourth from the last 
whorl is about 0.60 times the diameter of the last whorl; (2) the 
aperture is 7.45-9.50 mm. in diameter; (3) the collar is relatively 
narrow, 0.10-0.24 times the diameter of the aperture; (4) the 
pseudosiphon is a rectangular indentation; (5) the basal carina 
is larger in circumference, passing in front of the reflected 
outer peristome, and continuing to the last quarter of the last 
whorl (PL I, tig. 3). 

T. simnlacruni may be recognized by the following characters: 
(1) the diameter of the fourth from the last whorl is about 0.75 
times the diameter of the last whorl; (2) the aperture is 5.73- 
7.13 mm. in diameter; (3) the relative width of the collar is 
0.24-0.38 times the diameter of the aperture; (4) the pseudo- 
siphon is circular, and is open or closed; (5) the basal carina is 
smaller in circumference, passes beneath or behind the reflected 
outer peristome, and extends only a little beyond the last half 
of the last wli()i-l. 



1<){):^ XUTKS ON TOMOCYCH'S O 

T().M()( ^Cl.rs SIMTLACKI'M ( Mofclcl ) 

Ctlcl<>!<Unii<i .siiiiiildfni.iii Mdrclct, 1S49, Test. Nov. Iiisul. Ciih. Anicr. Criit.: 

•2-2 (Tyiic Idc.-ility: PcttMi, ( iiKitciii.-ila ). 
CydontoDia copanensc Sowerliy, is.-)(i, Tlies. Coiichyl., 1 (Rupiil.) : Uvl, pi. 

i:?B, fiss. ;U0, 311 (Typf liic-ility: Colinii, (Tuntciiinla ). 
Mcfldloiud.stoma simuhicriini v;u-. iiiimi.s von .Martens, ISiMi, IViol. ('out. 

AiiUT., 9: 10. 
M, ijaUniiastonui .siiiiulacnnii v;ir. c/raciUs von Martens, 1S!)0, Biol. Tont. 

AniiM-., 9: 10 (Type locality: lietAveen Tactic and Ttunahu, Guatemala i. 
Tomocijchis xiphonis Bartscli and Morrison, 1942, Bull. U. S. Nat. Mus., 

181: 145, pi. 19, tig. .3 (Tyjie locality: Alta Vera Paz, Guatemala). 
Tomocychts coiistricius Bartsch and Morrison, 1942, Bull. XT. S. Nat. Mus., 

181: 145-146, pi. 19, fis. 2 (Type locality: Coban, (Juatemala). 

Shell einnaniou or rosy, early whorls and last half of last 
whorl dull yellow brown; opaque or only sli«ihtly translucent; 
of moderate or large size (measurements of height and width, 
and their relationship, are expressed in Text-figure 1) ; pupiform 
to elongate-turrite ; decollate. 4-8 whorls remaining: spire nearly 
straight sided or moderately convex (Bartsch and ]\Iorrison. 
1942: pi. 19, figs. 2, 3, 5, 6) ; diameter of fourth whorl about 
0.75 times the diameter of last whorl; whorls convex, depth of 
suture variable, usually moderately impressed; whorls crossed 
by numerous fine posteriorly arched axial striations, which con- 
tinue onto reflected peristome ; raised spiral scul])ture only 
faintly evident at irregular intervals; base of last whorl with 
a distinct carina which passes below or behind the reflected outer 
peristome, carina usually evident only on earlier half of last 
whorl ; peristome double ; outer peristome usually forming a 
broad, irregular collar 1.55-2.89 mm. wide, 0.24-0.38 times the 
diameter of the aperture; face of collar dull white, with a few 
annulations which are usually distinguishable only near the 
inner peristome : parietal region of collar with a psendosii)hon 
which may vary from an open crescent to a rouiul hole con- 
nected to the outside by a narrow slit; aperture dull white, cir- 
cular, 5.73-7.13 mm. wide; columella tubular, spiral, open at 
first remaining whorl, and at umbilicus as a circular chink. 

T. siiuidacrum is highly variable, as is indicated by its lengthy 
synonymy. The name ])r(>])()sed by Morelet has l)een properly 
applied by subse((uent authors. Cyclostonia copanense Sowerby 
was described as being smaller than simulacrum. Later authors 
found that the two forms were not as distinct as Sowerby be- 
lieved, and Fischer and Crosse (1886: 121), von Martens (1890: 
10) and Kolx'lt (lf)()2: 271) considered copaxcusc as a variet\" 



BREVIORA 



No. 181 



of simulacruin. Recently, Bartseh and Morrison (1942: I-IT-MS) 
reallocated copancnse to specific status. Specimens under 27 
mm. in total length were assigned to copancnse, and specimens 
over 35 nnn. in total length were assigned to siniuJacnon. 

As is demonstrated in the diagram (Text-figure 1) no signifi- 
cant diflPerence in size will distinguish the two forms. Different 
specimens in various lots are represented near both extremes 



35 



30 



H 
X 

UJ 

X 



25 . 



20 



• • • 

• 

• • • 

• • 



10 

WIDTH 



15 



Text-figure 1 

Tliis (liagr.'iiii ilenionstrntes the relationsship between height nnd widtli 
of the shell of Tinnuciiclus .siiiiiihicrioii (Morelet). Meiisui-enieuts of heiglit 
were iiiiide to iiiclufle tlie l;ist t'oui- reinaiiiing whoi-ls. .Mc'isnri'iiieuts of 
widtli are slaiidaid. .Ml iiicasiiit'iiu'iils are in millimeters. 



1968 NOTES 0.\ TU.MOCVCLUS 7 

of the same <;rai)h, and the rcinaiiiinji' spcciincns in these same 
lots are re])resented at many intermediate loei. Since siicli vari- 
ability of size eommonly oeeiirs within a sin«il(> lot, copnnensc 
cannot be recognized even as a variety of siniulaci-Kiii. 

Migaloinasioma siiniilacniiii var. iniittts was [)roi)osed by von 
Martens as a synonym of copatirsi . This is clear by his nse of the 
two names. Mcgalo))Kist()ma siiniildcfion var. (jfdcilis is also in- 
distinguishable from simulacnDii, for tlie slij>ht ditlterence of 
shape used to separate the two forms is hiohly variable and lots 
containing' only a few sj^ecimens cannot be satisfactorily sorted 
into two groups by use of this character. 

Tomocychis siplto)iis Bartseh and Morrison was distinguished 
from sitnuIacrKiit by the presence of a closed pseudosiphon as op 
posed to an open one. Comparison of tiie type o\' sipJuniis (TSXIM 
162511) with many specimens of sinuilacnon shows that this 
character varies from a completely enclosed hole connected to 
the outside by a narrow slit, to a broad open crescent. The dis- 
tinction between a closed and an open pseudosiphon is obscured 
by many intermediate stages, thus preventing the differentiation 
l)etween sipJionis and simulacrun}. 

Toniocyclus co7ist7-ictits Bartseh and Morrison was recognized 
on the basis of a closed pseudosiphon and a relatively deeply 
impressed suture. As shown above, the natui-e of the ]isendo- 
siphon is an unreliable charactei-. The degree of impression of 
the suture is also variable. Several lots examined contain speci- 
mens that completely bridge the diflFerence from a deep suture 
with strongly rounded whorls, to a shallow suture with slightly 
convex whorls. Occasional sp(\-imens have a deep suture between 
the early whorls, and a shallow sutui-e between the lat<M' whorls. 
Thus, constricius cannot be separated fi'om siniulacruiii. 

Specimen.^ (xautiittd. MEXICO: lu, iuldiiional ddia. TMMZ 
87073 (2). GUATEMALA: im addiiimud data, ANSP 45648 
(3). T'MMZ 87075 d). r:\IMZ 8707(i (^). Alfa Vna Paz: 
Chama. AXSP 13105 (1). AXSP 13107 (3). I'SXM 484861 (1 ) : 
Col)an. MCZ 10061 (6), MCZ ex I^.land (4). MCZ ex Kolison (2). 
MCZ ex Bequaert (6), IJSXM 250603 (4). rSX:\r 516028 (1) : 
Samac, TSXM 515763 (2) ; P^inca dc Provideiicia. T^SX:\1 32070 
(2), TTSXM 203656 (1), USXM 316385 (3). PSXM 821005 
(1), T^SXM 321080 (1). TTSXM 515764 (1) ; nr. Arroya Yalcha- 
tila. 4 mi. sw of Seiba. UMMZ 64717 (2) : 1 km. n of TTacienda 
Fiiica Samac. UMMZ 132317 (2): 55 mi. ne Co) an. VMSIZ 
105169 (1); nr. Chinaja. C.AIMZ 208418 (1). EI Quiche 4-6 



8 I'.KI.VIOKA No. 181 

km. w of Hacienda Fiiica Paeala. V^n\7. 182318 (5). NO 
DATA: M(;Z 4r)!)2 (li. MCZ (will (4). :\r('Z 1-!88:?() (:5). 

rsxM 86:);}()5 d i. 

To.MOcYciAs I.I WAK Bartscli 

Tomocydus giiat.emalensis (in part) Fischer and Crosse, 1886, Miss. Sei. 

Mex. Amer. Cent., 2: 124, pi. 40, fig. Ha. 
Tomocydus lunae Bartsch, 1945, Proc. Biol. Soc. Wash., 58: 63. 

Tyi^e locality: 8anteeoniapaii. \'ciacru/.. INIexico. 

Specimens examined. ]\IEXI('(): \'(i'acn(:: soutli slojic X'ol- 
ean San Martin. 1040 ft. alt., UM.MZ IJir^lTO (1 i ; \'()l<-an San 
Martin. MCZ (1). 

ToiMOCYCLUS GUATEM.' I.r.XSIS lIM'cift'ciO 

Cydoxioiini guatemdlensis Pfeiffer, 1851, Proc. Zool. Soc. Loud.: 24."). 
MegaJomusloma fiuatcmaJensis Pfeiffer, IS.l'J. .MdiKi;^. Pncniiion. A'ivent., 

1 : 132. 
Tomocydus i/iKiti iiiah nsis (Pfeiffer). Crosse and Fis(dier, 1872, .Tom-. 

Conchy]., 2ll: 7(i ; Fischer and Crosse, ISSii, .Miss. Sci. Mex. Amer. Cent., 

2: 124, pi. 4(1, fifT. ]l. 

Tj/pt local ill/: W'VH Paz. (rnatemala. 

Known only from the type locality. 

T. (jnalemalensis is of donbtful slalus. 1 liaxc seen only a 
single s]iecimen identified as this species (T^^IMZ S7()7'2. e.\ 
Bryant Walker;. In most respects tins I'oiin rcscmlilcs 7'. siniiil- 
acnini, hiit it lacks the basal carina that is ])resent in that si)e- 
cies. The distinction between T. (/uaUntah iisis aiul 7'. lunai is 
also uncertain, for the characters used by liartsch (l!)4r)) to 
separate these two species are subject to variation. 

LlTFKATUliE ( irKD 

Bartscit, I'. 

194"). ,\ new Toiuocydns from Mexico. I'loc. I!iol. Soc. W'asldiij^lon, 
58: 63. 
Bartsch, I', and .1. P. E. Morrison 

1942. 'I'lie cycdophorid mollnsks of the iiiainl;i iid of .\inerica. I'.ull. 
V. S. Xat. Mus., 181: 142-291 pis. 19-42. 
FisciiKK. 1'. .and II. Crosse 

188(5. .Mission Scientifirpie an .Mcxicinc ct d.ans 1 '.Ainciiinie Centialc. 
.Moilns.-.a. (7 I 2: 1 i:;-12C.. 

KoKKl.T, \V. 

1902. CS-cIoplioiidae. Das Tici reicli, 16: i-xxxi.\. l-()()2, ti<is. Mid. 



liKi:; NO IKS ox TOMOCVCKIS 9 

MORRISD.V. J. P. E. 

liK").". Xdtrs (111 Anicric.-in cvcloijliorid l.-iiul sii.-iils, with two now iKiiiics, 
tlirec new jicner.'i, iind tlic fjiniily Ainpliicydotiiliu' sci);ir;it('(l on 
nninnil cliMrncteis. Jour. Wasliin^i'ton Ai:nl. Sci., 45 (o ) : 149 
102, figs. 1-31. 
Torre, C. de la and T*. Bartscm 

19-f2. The cytdophorid niollusks of Cuba. Bull. I'. S. Xat. Mus.. 181: 
3-42, pis. 1-8. 
vox .Martexs, E. 

ISOO- Land and ficshwator Arollusca. r.i(d. Ci'ut. AnuT.. 9: ixxvii, 
1S91. l-70(>, pis. 1-44. 



10 BREVIORA No. 181 



Plate I 

II()loty])e of Tonmeiichis tistulo.sii.s', new ^spe^'ies (UM^rZ 194095) ; liigli 
lainfoiest nt Valleiitine Camp, 50 miles Southwest of Cayo, British ITon- 
iluias. 

Fig. 1. Frontal view. 

Fig. 2. Lateral view. 

Fig. 3. Basal view. 



lOO.'S 



NOTES OX TOMOCYCLrS 



11 






C)'0<-^ 



BREVIORA 

Miiseiim of Comparative Zoology 

Cambridge, Mass. I'EBRrARY 1, 1963 Number 182 

BIRDS FRO.M FLORES. LESSER Sl'NDA ISLANDS 
Bv RAY.MONTn A. Paynter, Jr. 



During recent years tlie Musenni of Comparative Zoolooy has 
received from the Rev. J. A. J. \'erheijen, S.V.D., several collec- 
tions of birds from the western part of Flores Island. Without 
using a gun, Father \'erheijen and his local assistants have se- 
cured representatives of 70 species. Eleven of these have not 
been previously recorded from the island and several others are 
of taxonomic importance. It is the purpose of this paper to note 
briefly these more Intercast ing species. 

Nycticorax caledonicus subsp. 

A bird in partial adult plumage was snared at Tjantjar, Ra- 
hong, on 19 May 1957. LTntil 1940, when an example of N. cale- 
donicus was found paired with one of N. nycticorax in western 
Java (Hoogerwerf, 1952), this night-heron was not known west 
of Timor. This is the only record from Flores. 

Dendrocygna javanica 

A wing has been preserved of a bird collected in 1956. Sum- 
bawa is the farthest east it had been recorded previously. 

Elanus caeeuleus subsp. 

The easternmost localities for this hawk were Sumba and 
Celebes. A nestling, just about to fledge, was taken near Ruteng, 
at about 1000 meters, in July, 1957. 

Rallus pectoralis exsul 

Three birds, the first known since the type specimen was se- 
cured, were collected in 1958 and 1959. One is an adult; un- 
fortunately it was not sexed. The other two are juvenals which 



2 BREVIORA No. 182 

have little barring on the ventrum, lack the rufescent head, and 
have no bright olive margins to the feathers of the back. 

T have examined two specimens of Rallus mirificus Parkes and 
Amadon (1959) from Luzon, Philippine Islands, and believe 
that this form is a race of R. pecioralis. From Australia through 
New Guinea to Flores, R. pcctoralis exhibits a fairly orderly 
cline of decreasing dorsal streaking, darkening head color, and 
shortening of the dorsal feathers. The characters of mi7-ificus 
are the culmination of these trends and are those one might have 
predicted at the end of a cline extending from Australia to the 
Philippines. The differences between the Philippine bird and 
the Australian races are marked, but the subspecies from Flores 
is almost perfectly intermediate. Additional races may well be 
discovered between Luzon and Flores, showing that the cline is 
less disjunctive than it now appears. Support for this prediction 
may be found in the knowledge that R. p. mirificus remained un- 
known until only four years ago, in spite of its presence within 
50 miles of Manila, long a center of ornithological research. 

PORZANA PUSILLA PUSILLA 

Three specimens were collected : two at Tjara in late April and 
mid-May, 1957, and one at Wangjung in late April, 1956. One 
is a male, and two were not sexed, but on plumage characters 
these seem to be a male and a female. The species apparently 
breeds on Flores, although there are no prior records of its 
presence. 

It may be that these specimens represent an undescribed race. 
However, our birds are too poorly preserved and few in number 
to be certain that the apparent racial characters are real. The 
specimens are close to nominate pusilla but are somewhat darker 
dorsally, with more extensive black centers to the feathers, par- 
ticularly on the tail, and have heavier, but not longer, bills. The 
pale area on the chin and upper throat of the males seems whiter 
and more sharply demarcated than in the nominate form. The 
bills of two birds are black with very narrow yellowish markings 
on the anterior edges of both the maxilla and mandible. The 
third specimen is similar but also has a small pale area near the 
tip of the mandible. In a series of 26 s])ecimens of P. p. pusillo 
one has a bill similarly colored to the third Flores bird, and none 
of the series resembles the other two birds. 

The dried legs of the P'lores specimens are considerably dai-ker 
tiian any of P. p. pusilla. 



1963 BIRDS FROM FLORES 3 

The three birds from Flores are easily distinguished from 
mayri of New (iuinea and from palustris of Australia by their 
larger size (wing 84-86 mm.). T have not seen an exami)le of 
inird, from East Borneo, but the original description (Riley, 
1938) notes that the bill is olive-yellow and bronze, which seems 
vastly different from the Flores birds. 

POLIOLIMNAS CINEREUS CINEREUS 

This widespread rail is new for Flores. 

Gallicrex cinerea 

A specimen was collected at Tjantjar on 14 May 1957. The 
species apparently has not been found before on the Malay 
Archipelago east of Java, where it seems to be a winter visitor 
(Kuroda. 1936). Its presence on Flores in mid-May suggests 
that it may breed there but, unfortunately, the gonads of our 
specimen were not examined. 

Gallinula chloropus orientalis 

Nine specimens were collected, one of which contained an egg 
in mid-May. No prior records from Flores exist. 

KOSTRATULA BEN'GIIALENSIS BENGHALENSIS 

This is another bird not reported from the island before and 
which represents an eastward range extension. A specimen col- 
lected in July was breeding. 

Gallinago stenura 

Three specimens establish this snipe as a winter visitor on 
Flores. 

Gallinago megala 

A long series, collected over several winters, represents a 
new record for the island. 

Otus (bakkamoena) silvicola 

This endemic owl is probably a giant geographical representa- 
tive of 0. hakkarnodia. It is similar to 0. b. semitorquts of 
Japan, but it is considerably larger and lacks the buff crescent 
on the upper back. This was noted by Hartert (1897), who also 



4 BKEVIORA No. 182 

remarked on the similarity of silvicola to 0. b. whiteheadi of 
Luzon. The Philippine form has a white hindneck and is darker 
below than silvicola, but in size more nearly approaches the 
Plores bird than any other race of 0. bakkamoena. Compared to 
the o'eographieally closer Javan and Bornean populations, silvi- 
cola is markedly larger. This is suggestive of the sitviation in the 
Philippines, where whiteheadi is considerably larger than any 
nearby races. 

Tyto capensis subsp. 

A specimen obtained in March, 1956. is the tirst record of the 
species from the Suncla Islands, thus confirming Hartert's sug- 
gestion (1929) that it might some day be found there. 

I am following Amadon and Jewett (1946) in treating longi- 
memhris and capensis as conspecific. 

In view of the variability of the species and the paucity of 
material, particularly from the southwest Pacific, I am reluctant 
to assign this specimen to any race. Amadon (1959) tentatively 
placed two individuals from Celebes and Kalidupa with T. c. 
ivallcri (type locality Queensland); the Flores bird may also 
belong here. 

CORACINA DOHERTYI 

An immature bird was collected in early March. Rensch 
(1931) recorded several specimens from the island, but Flores 
was inadvertently omitted from the range of the species as given 
in Volume 9 of the "Check-list of Birds of the World" (Mayr 
and Greenway, 1960) . 

ACKNOWLEDGEMENTS 

1 am grateful to James C. Greenway, Jr. and Ernst Mayr for 
assistance' with several problems, and to Father Verheijen for 
depositing his collection in this museum. Dean Amadon gener- 
ously lent specimens from the collections of the American Mu- 
seum of Natural llistor}-. 

LITERATURE CITED 

Amadon, 1). 

IStf)'). Remarks on the subspecies of the Grass Owl, I'l/lo eapensLs. 
.Jouni. liombav Nat. Hist. Sue. 56: ;U4-:54(i. 



1963 BIRDS FROM FLORES 5 

Amadon, D. and S. G. Jewktt, Jr. 

1946. Notes on Philippine birds. Auk, 63 : 541-559. 
Habtert, E. 

1897. On the birds collected by Mr. Everett in Soiih Flores. 
Novit. Zool., 4: 513-528. 

1929. On various forms of the genus Tyto. Novit. Zool., 35: 93-104. 

HOOQERWERF, A. 

1952. Voorkomeu van Nycticorax caledonicus subsp. in W.- Java. 
Limosa, 25: 29-32. 
KURODA, N. 

1936. Birds of the Island of Java. Vol. 2. Tokyo, vi + pp. 371-794. 
Mayr, E. and J. 0. Grbenvtay, Jr., editors 

1960. Check-list of birds of the world. Vol. 9. Cambridge, Mass. 
Museum of Comparative Zoology, xii + 506 pp. 
Parkes, K. C. and D. Amadon 

1959. A new species of rail from the Philippine Islands. Wilson Bull., 
71: 303-306. 
Peters, J. L. 

1934. Check-list of birds of the world. Vol. 2. Cambridge, Mass. 
Harvard Univ. Press, xvii + *01 PP- 
Rensch, B. 

1931. Die Vogelwelt von Lombok, Sumbawa und Flores. Mitt. Zool. 
Mus. Berlin, 17: 451-637. 
Riley, J. H. 

1938. Three new birds from Banka and Borneo. Proc. Biol. Sot. 
Washington, 51 : 95-96. 



vu€<y 



BREVIORA 



Museiioi of Coampsirative Zoology 

Cambridge, Mass. March 12, 1963 Number 183 

AUSTRALIAX CARABID BEETLES XTIT. 

FURTHER NOTES ON AGONINI, AND A 

GENUS OF LICININI NEW TO AUSTRALIA 

By p. J. Darlington, Jr. 
Museum of Comparative Zoology, Cambridge, Mass. 

Since publication of my "Notes on the [Australian] Agonini" 
(1956) I have spent nineteen months in Australia (Dec. 1956- 
June 1958), mostly collecting in the eastern forested areas. My 
itinerary, with list of localities, is summarized in a recent paper 
(1961). The following additional notes on Agonini are based 
on material secured during this trip and borrowed from the 
Queensland Museum, and on examination of some of Sloane's 
types. The most exciting new discovery is an Australian species 
of the Indian genus Dilonchus, which is a licinine rather than 
an agonine but which is in some ways suitable to be ancestral 
to the supposedly agonine genus Tlomothcs. The finding of 
dimorphism of "fixed" setae in Notagoniim macleayi (Sloane) is 
noteworthy too. 

Tribe AGONINI 

Before considering the Agonini proper, I have to say that one 
supposed Australian agonine does not belong in this tribe. It is : 

CoPTOGLOSsus carteri (Sloaiie) (new combination) 

Platynus carteri Sloane 191.'i, p. 460. 

The type of this species, from Dorrigo, New South Wales, is 
in the Sloane Collection at Canberra. I have compared one of 
my specimens with it. It is a broad, depressed, dull black carab 
that looks superficially as if it might be either an agonine or a 
lebiine. The elytra are broadly rounded apically, not obliquely 



2 BREVIORA No. 183 

truncate as usual in Lebiini, but this is not an invariable tribal 
character. Two other characters show that the insect is a lebiine 
and that Sloane was wrong in assigning it to the Agonini 
("Sphodrini"). One character is that the middle tibiae are 
sparsely pubescent rather than spinose externally. The other is 
that the basal bulb of the aedeagus is small and not much bent 
down, not large and strongly curved as in the Agonini. "Platy- 
mis" carteri Sloane should therefore be transferred to the tribe 
Lebiini, where it can be placed tentatively in the genus Cop- 
toglossus. 

NOTAGONUM Darlington 

Darlington 1952, p. 127. 

In my recent collecting I found 4 species assignable to Nota- 
gonum in tropical North Queensland. However, this is a genus 
of convenience, and I am not sure that these species are all 
related to each other. They may represent 4 independent stocks, 
so far as the Australian fauna is concerned. The 4 species are 
distinguished in the key and briefly discussed thereafter. In 
addition, the following agonines were found in North Queensland : 
Lorostemma {"Platynus") cooki (Sloane), distinguished from 
Notagonum by sole of hind tarsus with a single, regular row of 
bristles on each side; Dicranoncus queenslandicns (Sloane), a 
slender brown agonine with toothed tarsal claws; Viola gonum 
{"Colpodes") violaceum (Chaudoir), a blue or purple species 
with spined elytra; Colpodcs hahilis Sloane, larger, with bright 
green elytra; the well known HomofJirs ; and Odontagonum, a 
convex, shining black, flightless agonine with dentate humeri. 

Key to Species of Notagonum of North Queensland 

1. Slender; head more than .75 width prothorax; 4th segment hind tarsus 
broadly emarginate (Fig. 5) ; (length c. 9.5 mm.) submetalUciim 

— Less slender; head less than .75 width prothorax; 4th segment hind 
tarsus narrower, usually more loljed 2 

2. Larger (9-10 mm) ; aeneous, elytral striae shallow, intervals flat 

lafertei 

— Smaller (6-8 mm.) ; brown, sometimes slightly iridescent, but not 
aeneous; elytral striae deeper, intervals slightly convex 3 

3. Apex of elytron not subtruncate, usually not distinctly denticulate 
(Fig. 9) ; 4tli segment hind tarsus wider apically, with angle between 
lobes c. right (Fig. 4). macleayi 



1963 ATTSTRALTAN CARABID BEETLES 3 

- — Apex of elytron subtrunciite, usually denticulate (Fig. 10) ; 4tli seg- 
ment hind tarsus oval, strongly narrowed apieally, with angle between 
lobes acute (Fig. 3) dentellum 

NoTAGONUM suBMETALLiciTM (White) (new combination) 

Colpodes submetallicus White 1846, p. 2. 

Tliis is a very common species in south temperate Australia 
and Tasmania, and it occurs also in New Zealand. It lives in a 
variety of situations on the ground by water. It is apparently 
rare in tropical Australia but does occur there. I found 2 speci- 
mens by tramping down tall grass and thick herbage growing in 
sluggishly flowing water in open country near Atherton, North 
Queensland, in February, 1958. This is an example of a phe- 
nomenon which is probably zoogeographically significant : pene- 
tration of the tropics by a temperate carabid associated with 
water. Some other, north temperate C'arabidae seem to have 
crossed the whole width of the tropics in waterside habitats. I 
have recently given examples in Bcmbidion and the Trechini 
(1959, esp. pp. 332, 342). 

NoTAGONUM LAFERTEi (Montrouzier) (new combination) 

Anchomenus lafertcl Montrouzier 1860, p. 238. 

Although lafertci has been placed in Colpodes, the 4th hind 
tarsal segment is only weakly lobed (Fig. 6). I tentatively assign 
the species to Nofagommi because of absence of any obvious dis- 
tinguishing characters. N. lafertei is common from the vicinity 
of Cooktown south through Queensland and part of New South 
Wales. I did not find it on the Cape York Peninsula north of 
Cooktown and it is unknown in New Guinea, but it occurs on 
New Caledonia. It is frequently found with the two following 
species in debris and among dead leaves on the ground by water, 
but it often occurs also away from water and in more arboreal 
habitats, especially in clumps of wilted leaves on fallen trees in 
rain forest. 

NoTAGONUM MACLEAYi (Sloauc) (ucw Combination) 
Platynus macleayi Sloane 1910, p. 454. 

Sloane described macleayi from Kuranda, North Queensland. 
The type in the Sloane Collection is now reduced to one elytron 



4 BREVIORA No. 183 

which, however, is identifiable. I have topotypes, one of which 
matches the type elytron almost exactly. (The form of the elytral 
apex is somewhat variable in both this and the following species 
but does serve to identify most individuals satisfactorily.) 

This species is unknown in New Guinea and I did not find it 
near the tip of Cape York, but it occurs from the mid-peninsular 
forests (Iron Range, Rocky Scrub, etc.) south to the Atherton 
Tableland and the Kirrama Range, which is west of Cardwell. 
It lives beside brooks and rivers, often in masses of drift caught 
on obstructions above the water or in accumulations of dead 
leaves deposited on stream banks at bends or beside pools. 

N. macleayi proves to be dimorphic in presence or absence of 
anterior lateral prothoracic and anterior discal elytral setae 
(Figs. 1, lA). These particular setae are apparently inherited as 
a group : all 4 of them are present or all absent in most indi- 
viduals. Tliis dimorphism is presumably due to a single mutation 
inherited in Mendelian fashion. The only possible exception to 
strict dimorphism is a female from Shipton's Flat which lacks 
the setae in question except for an apparent trace of the anterior 
discal elytral puncture on the right elytron only. With this ex- 
ception, my 48 specimens of the species divide as follows. 





With designated 


Without designated 


Locality 


setae 




setae 


Tozer Gap 


3 5 9 




1 9 


(W. of Iron Range) 








Iron Eange 


16 S $,12 


9 9 




Rocky River 


S $ s 






Shipton's Flat 


1 $ 




3 9 9 


(S. of Cooktown) 








Kuranda 


2 S $ 




2 ,J <?,39 9 


Kirrama Range 


1 i 







These figures suggest that the population of macleayi in the 
base-of-peninsular forests (from the Atherton Tableland to near 
Cooktown, tabulated below the broken line) includes a consider- 
able proportion of individuals without the setae in question, but 
that the population in the mid-peninsular forests of the Cape 
York peninsula (tabulated above the broken line) includes only 
an occasional individual that has lost setae. 

In my paper on New Guinean Agoniiii (1952, pp. 89-252) T 
used presence or absence of the anterior lateral prothoracic setae 



1963 AUSTRALIAN CARABID BEETLES 5 

as a character to help distinguish genera, especially to distinguish 
the two "genera of convenience" Notagonum and Altayonum. 
However, I made it clear in that paper (bottom of p. 97) that 
the character was only a useful "tag" and not a character of 
generic value in itself. The setae in question are not known to 
vary in any species of agonine in New Guinea, and they do pro- 
vide useful tags for dividing the New Guinean species into con- 
venient groups. 

Notagonum dentellum Darlington 

Darlington 1952, p. 147. 

In New Guinea, this species is common, widely distributed, 
and somewhat variable. In Australia, it occurs from the tip of 
the Cape York Peninsula (Lockerbie and Bamaga) south to the 
Atherton Tableland and the Kirrama Range. It lives in about 
the same situations as the preceding species. 

CoLPODES HABiLis Sloane 

Sloane 1907, pp. 178, 179. 
Darlington 1952, p. 164. 

Colpocles hahilis is black with green elytra and is 13-17 mm. 
long (in New Guinea). It has a wide range in the eastern Malay 
Archipelago, including Buru, New Guinea, New Britain, the 
Solomons, and the Santa Cruz Islands. It has not previously 
been reported from Australia, but I have seen specimens from 
Cairns District, Coen, and Port Douglas (near Mossman) (all in 
the Queensland Museum), and an individual flew into the lighted 
window of a house where I was staying in Cairns in February, 
1958. The species' usual habitat (in New Guinea) is in foliage, 
including clumps of wilted leaves on fallen trees in rain forest. 

Odontagonum nigrum Darlington 
Darlington 1956, p. 9. 

This flightless species is the type of a very distinct monotypic 
genus of unknown relationships. It was described from three 
individuals from Millaa Millaa and Lake Barrine, on the Ather- 
ton Tableland, North Queensland. It proves to be fairly widely 
distributed on the Tableland but apparently does not reach either 
the Dividing Range west of Atherton or the vicinity of Kuranda, 
although apparently suitable rain forests exist in both these 
places. It lives on the ground in rain forest. 



6 BREVIORA No. 183 

Tribe LICININI 

DiLONCHUS Andrewes 
Andrewes 1936, p. 179. 

This genus was proposed for one medium sized Agonum-like 
species from India. The Australian species described below seems 
congeneric. It combines characters of the tribe Licinini with 
color pattern suggesting Homothes. Generic characters are in- 
cluded in the following specific description, and the place of 
the genus among other Australian Licinini is discussed after the 
description. 

DiLONCHUS PICTUS U. Sp. 

Form (Fig. 2) like Agonum but with elytra more ample; color 
black ; reflexed margins of prothorax testaceous ; elytra with hu- 
meri, epipleuri, and outer margins testaceous, the pale color 
extending to the 9th intervals anteriorly and forming separated 
spots on these intervals posteriorly; a small testaceous spot at 
each dorsal seta ; lower surface testaceous with episterna more 
or less darker ; femora pale, tibiae, tarsi, antennae, and palpi 
browner; upper surface rather dull, Avith close reticulate micro- 
sculpture isodiametrie on head and pronotum and slightly trans- 
verse on elytra. 

Head short, .66 & .64 width prothorax (from measurement of 
$ type and 9 paratype) ; mandibles slightly sinuate externally, 
then abruptly curved and bent down toward apex, each with very 
large triangular inner tooth and short terebra ; eyes large and 
prominent ; 2 supra-ocular setae each side ; antennae slender, 
pubescent from near base 4th segment, with segment 1 f. 5 X 
long as wide, segments 3 and 4 slightly shorter and subequal, 
segment 2 much shorter ; maxillary palpi with last segment 
somewhat thickened, labial palpi with last segment subsecuriform 
in both sexes (wider than in hkle^is) ; clypeus truncate or nearly 
so ; labrum moderately emarginate, 4-setose ; front nearly evenly 
convex, with neck impression and frontal impressions weak, 
clypeal suture finely impressed; mentum joining gula without 
distinct intervening sclerites, broadly emarginate, without tooth. 

Prothorax: width/length 1.29 & 1.33; base/apex 1.39 & 1.41; 
base/head 1.31 & 1.33; apex broadly emarginate; anterior angles 
not otherwise advanced, moderately rounded ; base very broadly 



1963 AUSTRALIAN CARABID BEETLES 7 

arcuate, almost truncate at middle; sides broadly rounded an- 
teriorly, slightly so posteriorly, slightly sinuate before posterior 
angles; latter slightly obtuse, slightly blunted; lateral margins 
rather wide, moderately reflexed, each with usual 2 setae about 
1/3 from apex and at basal angle; disc moderately convex, de- 
pressed at sides, margined at base and apex ; median line distinct, 
apical and basal transverse impressions weak ; surface faintly 
sparsely transversely wrinkled at middle, more strongly longi- 
tudinally so at apex and base; basal foveae broad, not well de- 
fined, impunctate or nearly so. Elytra long-oval, about 2/3 
wider than prothorax (E/P — & 1.64) ; anterior margin entire, 
liumeral margins vaguely angulate, lateral margins slightly sinu- 
ate before apex but not interrupted, apices simple ; scutellar 
striae well developed ; striae entire, impunctate ; intervals nearly 
flat, not specially modified; no 10th interval; 3rd interval 2- 
punctate near 2nd stria behind basal 1/3 and near apical 1/3. 
Inner wings fully developed. Legs slender ; tibiae spinescent ; 
hind tibiae not s}:)ecially grooved externally ; hind tarsi with first 
3 segments finely grooved each side above, 4th segment shallowly 
emarginate but not lobed, 5tli segment with conspicuous acces- 
sory setae ; claws simple ; sole of hind tarsus with a single regular 
row of setae each side. Secondary sexual characters: S with 
first 3 segments each front tarsus dilated, squamulose below, 
the squamae small and forming 4 longitudinal rows; S with 
apparently only 1, $ v/ith 4 or 5 (asymmetrical) setae near apex 
each side last ventral segment. Length c. 7.5-8.0; width c. 3.1- 
3.3 mm. 

Holotype $ (M. C. Z. Type No. 30,394) from Longlands Gap, 
Atherton Tableland, North Queensland, about 3000 ft. altitude, 
Feb. 1958 ; and 1 5 paratype fi'om Kirrama Range, W. of Card- 
well, North Queensland, about 2000 ft. altitude, Dec. 1957. Both 
specimens taken by myself in accumulations of dead leaves on 
the ground under the heads of small fallen trees in partly felled 
rain forest. The S type is teneral and warped, so that width of 
elytra cannot be measured and the genitalia cannot be dissected. 

This species runs to Microfcronia in Sloane's (1898, p. 488) 
key to Australian genera of Licinini but has a more Agonum-\\ke 
form, simpler clypeus (almost truncate, with only a narrow 
transverse membrane anteriorly), longer and less emarginate lab- 
rum, more distinct subapical sinuations of elytra, longer scutellar 
striae (these are characters of Dilonchus), and different color 
pattern. As compared with the geiiotype of Dilovchns; (hidcns 



8 BREVIORA No. 183 

Andrewes, of which I have a cotype), the new Australian species 
has a relatively narrower prothorax with better defined posterior 
angles, more distinct elytral microsculpture, wider last segment 
of labial palpi, and more extensive elytral markings. However 
hidcns has indications of the same pattern of markings, espe- 
cially laterally, that pictus has. (Two apparent errors in An- 
drewes' description of hidens should be noted. The insect is not 
apterous. My "cotype" has fully developed inner wings. And 
the clypeus is probably not emarginate. It seems squarely trun- 
cate in front but semicircularly impressed, and the impressed 
area is pale and easily mistaken for an emargination — but dis- 
section would be necessary to make sure of this detail.) 

The rather Agonum-\\kQ form, untoothed mentum, and 
Homot lies -like color pattern (and habitat) together suggest that 
the present new species may represent the ancestral stock of 
Homotlies. If so, Homothcs has lost some of the specializations 
(of mouth parts) that characterize most Licinini, while develop- 
ing other specializations of its own, some of which suggest 
agonine rather than licinine affinites. To determine the inter- 
relationship (if any) and probable evolution of Dilonchus and 
Homothcs would require both more material and more time than 
I now have. 

REFERENCES 

Andrewes, H. E. 

1936. Some new Carabidae from India. Indian Forest Records (new 
series), Ent., 2: 177-180. 
Darlington, P. J., Jr. 

1952. Tlie earabid beetles of New Guinea. Part 2. The Agonini. 

Bull. Mus. Comp. Zool., 107: 89-252, 4 pis. 
1956. Australian earabid beetles III. Notes on the Agonini. Psyche, 

63: 1-10. 
1959. The Bemhidion and Trechus of the Malay Archipelago. Pacific 

Insects, 1: 331-345. 
1961. Australian earabid beetles IV. List of localities, 1956-1958. 
Psyche, 67: 111-126. 
MoNTRorziER, Rev. Pere 

1860. Essai sur la faune entomologiquc de la Nouvelle-Caledonie ... 
Ann. Sue. Ent. France, (3) 8: 229-308. 
Sloane, T. G. 

1898. [Australian Licinini.] Proc. Linn. Soc. New South Wales, 
23: 487-492. 



1963 



AUSTRALIAN CARABID BEETLES 



1907. New Carabiflnc from German New Guinea and its dependencies. 

Deutsche Ent. Zeitschrift, 1907: 177-185. 
1910. [Platynus macleayi.] Proc. Linn. Soc. New South Wales, 35: 

454-455. 
1915. [Platynus oarteri.] Proc. Linn. Soc. New South Wales, 40: 

460-461. 
White, A. 

1846. [Colpodes suh metallic us.] Voyage Erebus & Terror, Ent., p. 2. 






Fig. 1. Notagonum macleayi (SI.), with full complement of setae. 
Fig. lA. Same, outline with anterior lateral pronotal and anterior discal 
elytral setae absent. 

Fig. 2. Dilonohus pictus n. sp. 



10 



BREVIORA 



No. 183 








Fig. 3. Fourth segment right liiiid tarsus, outline from above with setae 
omitted, of N otagonum dentelhim Darl. 
Fig. 4. Same of Noiagomim maclcayi (SI.). 
Fig. 5. Same of Notagonmn suhmetalUoum (White). 
Fig. 6. Same of N otagonum lafertei (Montr.). 
Fig. 7. Apex right elytron of Notagonum submetallicum (White). 
Fig. 8. Same of Noiagomim lafertei (Montr.). 
Fig. 9. Same of N otagonum, macleayi (SI.). 
Fig. 10. Same of Notagonum dentellum Darl. 



BREVIORA 



Miiseiiiim of Comparative Zoology 

Cambridgk. Mass. Ai'Kil 18, IWA Numher 184 

BEHAVIOR AS A TAXONOMIC CLl'E : KELATIONSIILPS 
OF LIS.SONYCTERIS (CHIROPTEKA) 

By 

Barbara Lawrence 

MiLseuni uf Cunipiiijitive Zoology, Harvard University 

and 

Alvin Novick^ 

Osborn Memorial Lalioratories and Peabody Museum of 
Natural Historv, Yale University 



Re-examination of the ^enerie status and i-elationships of 
Ro}(scffus, sensu stricfo, and Lissonyctrris, a supposed su])g'enus 
of Rouscttus, has been prompted by the great ditferenees observed 
between live specimens of the two genera. In 1955 and 1956, 
Novick was able to make extensive observations of living bats 
not only of these two genera but of six other pterojuds as well 
(Xovick, 1958). Of these, according to Andersen (1912), Eidolon 
helrioii and Ptcropus g. (jigantcus as well as Rouscttus belong 
in the Rouscttus section of the Pteropinae, Cynoptcrus brachyotis 
luzonicnsis, Cynoptcrus sphinx ccylonensis. and Ptcnochirus 
joyorii belong in the Cynoptcrus section of the Pteropinae, 
while Eonyctcris spclaca glandifcra, Eonyctcris rob}istn, and 
Macroglossus J. lagochilus are in tlie subfamily Macroglossinae. 

BEHAVIORAL COMPARISONS 

A single wild colony of about twenty Lissonyctcris angolcnsis 
was observed in the Belgian Congo and various wild colonies of 
Rouscttus amplcxicaudatus, R. scnii)iudus, and R. acgyptiacus 
were observed in the Philippines, Ceylon, and the Belgian 
Congo, respectively. Two Lissonyctcris angolcnsis were captured 

1 Fornicrly ;it tin' lliolojiical lialxtrMturics, Ilarvaril I'nivtTsit.v. 



2 BREVIORA No. 184 

and one, a matnre female, lived in eaptivity for fifteen months. 
Many R. acgyptian(.s were also taken alive and observed for a 
period of some weeks in the Belgian Congo. One, a mature male, 
was kept for over nine months concurrently with the Lissonycteris 
mentioned above. In addition, some six K. se)nini(di(s and a dozen 
K. amplcricaudafus were observed in captivity for periods of six 
weeks to more than three months. 

From the very first, the differences in behavior of Lissonycteris 
and RoHsettus were grossly apparent. These can best be described 
as differences in limb use and in orientation. 

Limb Use 

Rousettiis ordinarily occupy diml}- lighted caves with large 
entrances and sheltered retreats. Here they commonly hang in 
large clusters along the walls or ceiling. On the wall, they hang 
by their hind feet with their legs so turned that their backs are 
to the wall, their wings folded at their sides. They take flight by 
swinging their body up and rotating around their feet Avhich 
remain temporarily in position here though the legs are now 
crossed. This position is well illustrated by Kulzer (1958), in 
an excellent paper on the biology of Roust ft us. Roxsctfus also 
roost in crevices. Here one cannot see their undisturbed resting 
position but they must use all four limbs in entering and leaving 
such crevices. Rousettiis can fold their wings considerably and 
walk awkwardly on their wrists and hind feet or they can climb 
vertically, head first, or move along the irregular surface of a 
cave or crevice ceiling using the claws of their thumbs and hind 
feet. When Rouseftus roost in trees, as they do when disturbed in 
their caves or at night when feeding, they almost always hang 
by both hind feet plus one or both thumbs in a sloth-like posi- 
tion, one foot and one thumb on each side of the branch and 
their wings folded. They frecpiently move along the undersid(> 
of the branch in this position using all four limbs. Thus, wild 
Rousettus use their wrists for walking and their thumbs for 
climbing, roosting, and moving along the underside of branches 
or fruits. 

Captive Rouseftus would roost in all of the positions desr-ribed 
above. When they were exposed to light, they would fre(ni(Mitly 
crawl into a corner of the cage floor or into their feeding disli 
where they would ])r('ss tliciuselves as closely as possible against 
theii- sui-roundings, avoiding th(> light on Iheir eyes. Crawling 
is a not uncommon mode of locomotion for Rousettus. whcthei' 



1[)68 RELATIOXSIIIPS OK LISSON YCTERIS 3 

rcstrictod to a cajic oi' allowed to move freely about a vooiii. 
While feedinji', foi- example, tliey would crawl between the liands 
of a stem of bananas. In addition, they oeeasionally used their 
wrists and tliumbs as pushers to reailjust morsels of fruit in 
their mouths and R. avgyptiacus once was seen using the claws 
of one hind foot to manipulate the food in its mouth. 

Lissonyctcris never use their win^s for locomotion other than 
flight. Tn the wild, and in captivity in a large flight room, 
IAss<t)nfrf< ris always roosted hanging free from the ceiling or 
a branch or, occasionally, hanging from an irregularity of the 
wall, their ventral surface out as with Rousettns, but never 
using their wrists or thumbs for support and never entering a 
crevice. Liss(mycteris would hang from a branch or the ceiling 
by the claws of one or both hind feet. If they used both feet, 
characteristically both would be on the same side of the branch 
or irregularity. Lissouycicris seemed incapable of folding their 
wings tightly as Rouseffiis do; when roosting they hold their 
wings onlv slightlv folded enveloping their ventral surface (see 
Fig. 5). "  

Lissonifcfen's were incajjable (»f walking or climbing since both 
activities recjuire that the wrists or thumbs be used. They moved 
along their roost only by releasing their hold with one foot, 
moving this foot along to a new hold and then following with 
the other foot. Such movements, always slow and inefficient, 
were suitable only foi' slightly changing their roosting position. 
Lissonyctcris never landed on a horizontal surface liy choice 
and, when forced to do so to take its fruit from a dish, its move- 
ments were awkward and incompetent. After picking up a 
piece of fruit, it would flap otf without crawling. 

Lissonyctcris regularly use tlieii- hind feet, but rarely their 
wrists or thumbs, for handling their food. When one Avas handed 
a piece of banana, it would grasj) this with its teeth. Then, having 
readjusted its roosting foothold, it would bring one foot dowai 
(either one with apparently e(|ual facility) to its mouth and 
hold the piece of fruit with its widely spread toes while biting 
or breaking off a manageable fragment (see Fig. 1). Then, 
while chewing this fragment, it would hold the bulk of the piece 
of fruit, with its toes, against its chest or abdomen, frequently 
partly or completely covered by its wings (see Fig. 4). IJarely, 
when through carelessness or fragmentation it was about to drop 
the piece of fruit Avhich it was chewing, the bat would regain 
its tooth-hold by using its wrists and thumbs as awkward pushers. 
Thus Rousettus, sensu stricto, differs from Lissonyctcris in its 



4 BREVIORA No. 184 

use of wrists, tlnimbs, and hind feet in roostinp', (•limbin<r, moving 
along branches, walking, and handling its food. 

Among the other pteropids, Eidulon and Pfcropiis, though 
they never enter caves, roost in trees in much the same posture as 
Ronseitiis, holding on with both hind feet as Avell as with one or 
both thumbs. Eoiiycfcris, in caves and in captivity, roost and 
move in all ways like Roiisdfus. In all of these genera the wings 
can be considerably folded ; some also commonly roost Avith their 
wings enveloping their ventral surface. Neither Eidolon nor 
Pier opus can walk well on horizontal surfaces and both would 
probably be incapable of entering cave crevices as do Poiisdfus 
and Eonjfcfrris. Pteropiis, at least, use their Avrists and thumbs 
skillfully to handle fruit and to orient the morsels in their mouth. 

Cynopfcrns, Ptcnochirus and, to a certain extent, Macroglossus 
also use their limbs very much as does Lissonyctcris. All three 
hang from above, by their hind feet only, though, unlike Lis- 
sor}yctcris, they appear to use their thumbs for climbing up walls 
or along branches. The wings of Cyuopirrus and Macro(/lostsus 
do not seem to fold in life any more tightly than those of Lis- 
sonycteris. Judging from pictures of Epomophorus (Allen, 
Lang, and Chapin, 1917, pi. 46) and the position of Mifonyctcris' 
wings in alcoholic specimens, their posture in life is also like that 
of Lissonyctcris. 

Lissonyctcris (as also ('ynopt< rus and Ptcnochirus) tended 
to hold large amounts of food in their cheeks and to carry food 
to their roost, there to chew very slowly. AVhen recpiired to fly 
while eating, they would sometimes drop a really large piece. 
If Lissonyctcris Avas holding a piece of fruit with its hind foot, 
it would either transfer this to its mouth for flight or drop it. 
Rouscttus never seemed to store fruit in its cheeks or to fly with 
any in its mouth but, like Eonyctcris, Eidolon, and Ptcropus, 
generally stayed at the food source, biting and woi-ryinu off 
fragments and chewing these at once. 

Lissonyctcris would chew and crush fruit, such as melon and 
pineapple, until all of the juice had lieen expressed and swallowed 
and then push the remaining small bolus of fiber out of its mouth 
with its tongue. Banana, being non-fibrous, was swallowed com- 
pletely after chewing. Rousettus usually ate banana and. like 
Lissonyctcris, swallowed the whole fruit hut when Rousettus ate 
mango, ])apaya, or melon, uidike LisstDiycto'is. they seemed to 
swallow the whole substance. 



^\)C)^^ RELATIOXSIIIPS OF LISSOX Vf "PKKIS 5 

( )Kli:.\TA'l'I()N 

jAssonf/ctfris and Noiisrffus differ strikiiij>ly in their orienta- 
tion. Lisson!irt( ris oi-ient entirely by vision (Novielv, 1958). The 
captive hat was helph'ss in the dark and refused to fly in the 
dark oi- when hlindfoUh'd. When it was forced to fly witliout 
vision. I)y Ijein""' tlirown into the air. it always crashed into the 
first ol)staele it met. 

Roiisdfus anip1( .ricanddl IIS, R. sciniiiiidiis, and R. argyptiacus 
all orient visually and acoustically (Mohres and Kulzer, 1956; 
Kulzer, 1956, 1958; Novick, 1958; Griffin, Noviek, and Korn- 
tield, 1958). The sin^ile R. acgyptiacus ol)served for over nine 
months in captivity oriented largely visually in strong light but 
oriented acoustically in the dark, in dim light, when avoiding 
intricate obstacles, and when landing or taking off. Such acoustic 
orientation obviously demands specializations for sound ])roduc- 
tion, emission, reception, and interpretation. The brain, in 
particular, must be highly specialized for handling acoustic 
information. 

In the pteropids, acoustic orientation has so far been found 
oidy in Rousettus, scusii stricto, and not in Eidolon, Pteropus, 
Cynoptcrus, PtcHocJiiriis, Eonycteris, Macroglossus, or Lissonyc- 
teris (Novick, 1958). The orientation of Ste^wnycteris, a sub- 
genus of Roiiscttiis, has not been observed. The distinctness of 
Rouscitus, not only from Lissonycteris but from a fair sample of 
other pteropids as well, is clearly establisiied by its acoustic 
orientation and the behavioral, physiological, and anatomical 
features associated with it. 

Summary 

To summarize, Lissoiiycfi ris differ from Rouscitus, scnsu 
stricto, in their roosting posture, in their non-flight locomotion, 
and in their iiuU)ility to orient acoustically. Comparing the 
roosting and locomotory behavior of those bats observed alive, 
we find that Rouscttus resembles Eidolon, Pteropus, and Eonyc- 
teris, while Lissonycteris resembles Cynopterus most closely. 
Brief ol)servati()n of live epomophorine bats and examination of 
pictures and preserved specimens of epomophorine bats and of 
Myonycteris suggest that in behavioi- Lisso7iycteris resembles 
these bats as well. 



6 BREVIORA No. 184 

MORlMIOLOdiCAL C0.MPAK180NS 

The behavioral differenees between tlie live bats are reinforced 
by less speetaeular, l)iit equally definite, characters of the more 
conventional taxononiic sort. These make it clear that Lis- 
soin/rteris is far closer to Myonycteris, a supposed intermediate 
genus between the rousettine and cynopterine "roups, than to 
Rousettus. They also show that Lissonycteris and Myonycteris 
form a natural gi-oup probably intermediate between the rouset- 
tine and epomophorine groups, and one whose resemblances to 
the cynopterines are more apparent than real. 

The rousettine and cynopterine sections of the Pteropinae are 
chiefly distinguished by the following characters possessed by the 
latter and not by the former: The rostrum is shortened and the 
facial axis is not deflected, Ms and usually M- are lost, the eyes 
are larger, and there is a tendency to form tubular nostrils. The 
differences in the ridges of the soft palate described by Andersen 
(1912, pp. Ix, 485, 591, figs. 29A, 50) are also characteristic. 

On the basis, largely, of these characters, Andersen points out 
that Myonycteris is somewhat intermediate between the cynop- 
terine and rousettine sections. He says (1912, pp. Ivi-lvii) : 
"Myonycteris . . . has in many respects remained on the Rouset- 
tine level of development, while in others it exhibits modifications 
approaching those of Cynopterus. The general appearance, the 
dental formula, and the palate ridges are qviite or nearly as in 
Botisettus, but the rostrum is conspicuously shortened, the facial 
axis less deflected, mn and m- (last lower and upper molar) 
reduced almost to rudiments, the orbits larger, the nostrils more 
prominent and the calcar weaker," and later (p. Ixi) : "The 
fact is that this genus has retained many characters of Rousettus, 
while in jn-actically all the features in which it differs from 
Rousdtus it more or less closely approaches to Cynopterus. 
Whether a genus exhibiting characters of this description ought, 
in a linear arrangement, to be classed at or near the end of the 
Rousettine section oi' as the ■opening' genus of the Cynopterine 
section, must necessarily remain a matter of opinion." 

The resemblance of Lisso)iycieris to Myonycteris in all the 
above traits, except the shortening of the rosti'uiii and i-eduction 
of the dentition, was not noted by Andersen when he classified 
tile foniici-asan ahcrrjint form related to but {)ossibly generically 
distinct from /t<nis(ltHs {(ip. r//., 1!I12. p. 814). At tlie same 
time he does note ( 1912, p. xlix ) : "the In-ain case [Lissonycteris] 
is |)eculiarly flattened postei'ioi-ly and tlie facial axis even less 



1!)6."{ RELATIONSHIPS OF LISSONYCTERIS 7 

deflected tlian in Rouscttus, both eliaraeters giving the skull, 
vi(»we(l in pi-otih', a i-itliei- striking i-esemblanee to Ejjoinops." 

External Characters 

Aetna lly, in addition to the ratiier generalized niyonyeterine 
characters given above, both Lissonijcteris and Mijuinjcteris have 
a number of traits in common which the small size of Myonycteris 
and the superficial i-esemblance of Lissonycteris to Rousettus have 
tended to mask. Chief among these, externally, is the develop- 
ment of the wing in these two genera Avhich greatly exceeds that 
of Rousettus. Though it is more similar in size to that in Cynop- 
tcnis, it is ditt'erent in proportions. The size ditference shows 
most conspicuously in the greater length of the combined meta- 
carpal and first phalanx of digits three to five as compared with 
length of forearm. The following comparison of finger lengths 
refers only to these two joints. In Lissonycteris, the fifth finger 
is conspicuously longer than the forearm; in Myonycteris, the 
slightly smaller wing has digit five subequal in length to the 
forearm. In Rousettus, the fingers are much shorter; usually 
only digit three has the upper joints as long as the forearm ; 
occasionally three is somewhat longer and four is subequal to 
the forearm. In all three genera, these two joints of digit five 
are shortest though the relative lengths vary specifically. Cynop- 
terus, while longer fingered than Rousettus, is less extreme than 
the other two genera and differently proportioned. The meta- 
carpal and first phalanx of digits four and five are subequal in 
length, with four being slightly shorter than five which is about 
the same length as the forearm, while digit three is conspicuously 
longer. 

Other differences in ratios of wing bones can be worked out, 
but the most significant fact is that in development of the wing 
Lissonycteris and Myonycteris are not intermediate between 
Rousettus and Cynopterus but differ equally from both. The 
same is true of the attachment of the wing wiiich in the two 
former genera is near the middle of the first phalanx of the 
second toe, in Cynopterus is near the distal end of the first 
phalanx of the first toe, and in Rousettus is usually between 
metatarsals one and two, sometimes near the bases of the first 
phalanges, often well proximal to this. 

Lissonycteris and Myonycteris are also equally distinct from 
tlie other two genera in shape and arrangement of the odontoid 
papillae. These papillae, which border the lips, inside, from the 



8 BREVIORA No. 184 

angle of the mouth forward, are rather liigh and pointed and, 
in Lissontjctcris, form a single row extending about to the eanines. 
In Myonycferis, the shape and arrangement of the papillae are 
about the same, but in some instances on the upper lip toward 
the angle of the mouth there is a poorly defined second row. In 
RoHscttKS, the single row of small papillae is much reduced in 
extent and size. In Cynoptcnis, the more numerous, larger 
I)apillae are irregularly arranged in a broad band two or three 
rows wide and somewhat better developed on the upper than the 
lower lip. 

The arrangement of the palatal ridges in Lissonijcten's is very 
different from that in Cynopicnis and, while more like that in 
RoiiscttKs, ditt'ers in certain details which again point up Lis- 
sonycfcris' close resemblance to Myonycieris. In the two latter, 
the three anteriormost ridges are not divided and scarcely, if at 
all, bowed forward; four and five are divided in the middle, 
slojic forward (as in Rousetfits), and have the inner ends re- 
curved ; six and seven, which lie behind the tooth row, are 
similarly sliaped and these last four converge somewhat at their 
medial ends. Behind this, there are two or three other poorly 
defined ridges. Rousettus differs in general in having the anterior 
ridges more bowed forward, number four usually not divided, 
and the posterior ridges more nearly parallel, with each other. 

Other external characters which set Lissonycteris and Myonyc- 
feris apart from Ro)isettns are the shorter, less robust tail, the 
longer, denser fur on the notopatagium and on the proximal, 
dorsal surface of the til)ia, the smaller foot with the basal quarter 
to third of the first j)lialanges wel)bed (see Pig. 1), the more 
slendei- claws, the moi'e delicate calcar, and the extensive patch 
of ulandular fur on the throat of adult males. 



o' 



Hair Stri'cture 

Additional evidence of tlie close relationship of Lissonyctcris 
and M yoiiffcicris comes from Benedict (1})57) who says that they 
have essentially the same liair structure: the pigment distribu- 
tion, absence of a medulla, and form, an-angement and dimen- 
sions of the scales being alike. The slight difference she describes 
in the more distal maximum diameter of the hair in M yonyctcris 
seems scarcely imj)or1ant. The hairs of Cynopicvus are so dif- 
ferent as not to need comparison here. Those of Ii(/its( ft its are 
less compellingly so, being similar in pigment dist i-itmt ion and 



in68 RELATIONSI Ill's oi' I ,ISSONYCTERIS 9 

at ISC I lee of ;i iiicdiil l;i. Tlicy appjirciit ly ditVci- liowcvfT in arraiige- 
iiu'iil aiul size of s(';;|cs ;iii(l. 1(» a (•ci'tiiiii cxtciil in llicii- form. 
In ucnt'i'al, Ih.rsc (lirfci-cnccs seem more of dcHi-cc ilmn kind. 

Cranial Characters 

C'r;inially, as cxtcrnjilly, Lissoiijich iis ;ind Ihioinich )is i)Ossess 
a innnl)('i- of fcatui-cs in coininon wliicli dist inuuisli tlicni ('({iially 
from R<jus(llus and (' iinopivnis. In addition to the jjropor- 
tions of the skull as a whole, llic shape and spaeiii<i' of the 
teeth, the occlusion i)a1tcrn, and the structure of the inter- 
orbital retiion are particularly iinpoi-tant. In <i'cneral jiropor- 
tions. tlie doiniuant featui-e of the small M i/oinich ris skull, and 
one ill which it aui-ees with the much lar<i'(M' Lissonyctcris. is the 
relatively ^reat leni>th of tlie anterior part of the skull (measured 
Irom behind the postorbital processes to th(^ tips of the premaxil- 
laries) as compared with both leniith of the brain ease (measured 
from behind the postorlatal jiroeesses) and its bulk. Typically, 
also, the bi-aiii case in M i/oiii/ch rIs is elon<iatcd behind the 
temi)oral root of the zyo'omatie arch, ami the rostrum is slender 
with the posterior ends of the nasals depressed giving the skull 
in profile a dish-faced ajipeai'ance, an appearance that is ac- 
centuati'd by tlie great develoi)ment of the lateral of the two 
pairs of frontal sinnses. 

Compared with Iti)iis( ff us. Llss<)inict( ris and M ijoinjctcris have 
a more slender rostrum. In the two latter, a projection of a line 
along the top of the nasals ])asses ventral to the top of the orbit 
and the brain case. Tn Roiisctfus. sciis}/ <iiricio, snch a line passes 
dorsal to both of tlK^se points. In the myonycterines, the brain 
case is elongated and flattened so that the distance from the 
bottom of the occipital coiulyles to the top of the occipital crest 
is less than the distance fi-om the postglenoid jirocess to the back 
of the comlyle. whereas in Ixousciius the first distance is ecjual to 
(u- greater than the s(M'ond. The largei- orbit, though difficult to 
measure, is easy to sec. In Lissoiiijcfi fis and Mnonijctcris the 
diameter of the orbit taken approximately parallel to its antero- 
rostral margin is e(pial to oi- greater than the laci-ymal width; 
in Nousdtiis it is less. Further, the antero-venti-al border of 
the oi'bit foi-med by the i-oot of the zygomatic arch is a better 
develojx'd, sharper edged rim; it is thin and i)late-like Avhere 
the infraorbital canal pierces it. The thicker root in Roiisctfus 
results in a noticeablv longer canal. 



10 BREVIORA No. 184 

When Lissoinjcferis and Myo)tyctcris are compared with Cij- 
nopterns, the ang:le of the nasals, tlie elongation of the back of the 
skull, and the rather large orhit with a short infraorbital canal 
show no striking differences. That these similarities are indicative 
more of parallelism than i-elationship is suggested by the pro- 
nounced ditf'erences in overall ])roportions of the skull. The 
short, heavy rostrum and relatively- large brain case of Cynopterns 
contrast sharj^ly with the more slender nosed, less chunky skulls 
of Lissonyctois and Myonycivris. Such features are hard to 
measure, but in Cynopterns the distance from the postorbital 
foramen to the back of the occiput is much longer than the dis- 
tance from this foramen to the premaxillary suture, while in 
Lissonyctcris and M yonyctcris these distanees are almost equal; 
further, the heiglit of the brain case vertically above the in- 
terpterygoid region is usually a third or more of the length of 
the skull in the former, while in the two latter it is usually con- 
sideral)ly less than one-tliii"d of this length. 

Certain features of the frontal sinuses are also very distinctive. 
Characteristically, those of Lissonyctcris and Myonyctcris have 
the lateral pair nnu-h more inflated than the medial. This, to- 
gether with the tilting U]) of the orbital margin ])Osteriorly, 
makes the interorbital region concave. In Ronscttns, the reverse 
is true; the smaller, lateral pair of sinuses are relatively less 
inflated, the medial more, and the orbital margins are not tilted 
up, so the interorbital region is flattened or domed, sometimes 
with a median sulcus anteriorl>-. In all three of these genera, the 
lateral i)air of sinuses end in front of the postorbital foramina. 
In Cynopterns, on the other hand, the well inflated lateral ]iair 
extend at least to the le\('l of the postorbital foramina which are 
thus crowded outwai'ds into the postorbital ])rocesses, and the 
much smallci'. medial pair show scarcely any inflation. 

Dentition 

In past classifications, the i-eduction of the last molai's has 
been one of the most im])oi-tant reasons for grouping Myonyctcris 
with Cyni)pt( riis. while the numl)er of teeth has been used to 
relate the former to R()iis(ftiis. Actually, the shape of the an- 
terior teeth is fai' more important than eitlu'i- of these in showing 
generic relationships and is one of the most clearcut characters 
relating Lissoiiyrtt ris and M yonych ris. \'ai-iation at the end of 
the molar row is, on the other hand, an impoi'tant differential 
charactei- separatiinj these latter two. Ty|)ically. the cyno])terine 



1963 RELATIONSHIPS OF LISSONYCTERIS 11 

bats have lost tlic last u|)|)('r and lower molars, while these teeth 
are retained and well di^veloped in the rousettine j^roup. Both 
Lissonycteris and M i/oinirtf ris have the rousettine formula hut, 
in the latter, the last molars are tiny and elearly obsolescent, 
while in tiie former they are tlie oi)posite. Reduetion of the 
molars in M i/nuifctcris is no moi-e r-eason than their stronger 
development in Lissoiitfcfcri.s for jj^rouping- the one with Cy- 
tiopterus and the other with Houscttus. 

The structure of the teeth, their spacing, and the occlusion 
pattern are remarlval)ly similar in Mijonycteris and Lissonycteris 
and differ equally from both Cynopierus and Rousettus. The 
most striking feature of the tooth rows of Lissonycteris and 
Myonycteris is the wide si)aeing of the short, broad teeth; in 
addition, the anterior teeth have the main external cusps placed 
well back on the outer margin so that in i)rotile the teeth look 
more equilaterally triangular. The teeth in Cynopterus differ 
sharply in being larger and more crowded, and in the arrange- 
ment of the cusps, with the main external cusp more anterior 
and a deeper sulcus separating the inner and outer halves of the 
teeth. In the upper jaw of Cynopterus, F'-' has a well marked 
ridge or internal cusp, whereas in Lissonycteris and Myonycteris 
this tooth, though thick, is sim])le. In the lower jaw of Cynop- 
terus, Pj is relatively large and P- compared to P4 is conspicu- 
ously higher, almost as broad, but shorter with a less well de- 
veloped heel. In Lissonycteris and Myonycteris P^ is tiny and F^ 
reduced, much narrower and only slightly higher than P4, wdth 
no trace of a heel ; P4 is relatively large with a high, almost 
central, main cusp, bilobed in Lissonycteris and in Myonycteris 
with sometimes a suggestion of a heel. In genei-al, these pre- 
molars are more similar to those of Rousettus ; in the latter. P-''. 
Po, and P4 are simple, high, and differ from Lissonycteris and 
Myonycteris chiefly in having the single cusp set farther forward 
with a trace of a heel posteriorly. P"* has the more usual pteropid 
elongated shape with the main cusps forward and the inner 
margin sometimes bilobed. In the lower jaw, neither of the first 
two premolars of Rousdtus is reduced, as in the former genera, 
while the molars are longei' and narrower. 

The wdde spacing and simjile shape of the teeth in Ljissonycteris 
and Myonycteris relate to a very characteristic occlusion pattern 
in which the P;{ and P-' alternate with each other, P4 barely 
occludes with the posterior edge of P"\ and there is scarcely any 
more contact between P"^ and the posterior edge of Pj. In Rouset- 
tus, the overlapping of the teeth is more pronounced, correlated 



12 BREVIORA No. 184 

with the ))('<i'iiiiiiiio- of a hoel. In Cynoptcrus, it is oven more so 
with the crowded teeth slanting forwards and Py reaching the 
back of the np])er canine. 

On tile liasis of the matei'ial at liand, Myonycteris and Lis- 
S07iyct('ris differ sufficiently from each other to warrant generic 
distinction. However, it may be significant that M. (Phygctis) 
is distinguislied from M. (^Jyonyctcris) by its relatively larger 
teeth, especially ^I- and M.{, and the hilobed main cusp of P4. 
characters which also helj) distinguish Lissonycteris from Myo- 
nycteris, sensu stricto. Further, two small species of Lissonycteris, 
SDiitJtH and cfypticohi, which are described as having reduced 
dentition, may prove to be souK^what intermediate between the 
two genera. Study of these possibly annectant forms may make 
it necessary to reconsider the generic status of Lissonycteris. 
For the present it seems best to retain both as full genera. 

DTSCrSSTON 

While the results of this study point up the distinctness of 
the myonycterine from both the cynopterine and the rousettine 
sections of the Pteropinae, they also suggest that the former may 
be more closely related to the epomoi)horine section than has 
previously been supposed. A casual inspection of the epomo- 
phorines and myonycterines shows such (\\treme external dif- 
ferences, such differences in structure of the palate, posterior 
bony nares, shai)e of the molars and tooth formula, and such a 
greatly elongated rostnnn in many cjiomojihorines that any close 
relationship niiglit seem most unlikely. However, in the rather 
unspecialized genus Epomops, the little or not deflected skull, 
the relative size as well as shape of the brain case and rostrum, 
the details of the interorbital region, and the size of the orbits all 
show a strong resemblance to Lissonycteris. In similar ways. 
Epinnophorns Jahiafiis. exce|)t for its elongated rosti-um. re- 
sembles Myonyeii lis. Thei-e are othei' resemblances. In the 
epomophorines, the wide si)acing of the teeth and the conse(pieni 
alternating more than ovei'lapping occlusion |)attcrn of the 
anterior ones as well as the iiioi-e posterior position of the mail. 
cusj)s of both up|)er and lower thii'd and four-th premolars and 
the small size of the fii-st lowei- premolar- all suggest the myo- 
nycterine gi'dup. The charactei-istic tooth formula of the 
eponiophoi'ines is foi-eshadowed by the great reduction in M y<>- 
nycttns of those teeth which ai'e lacking in the foi'nier. Ex- 
ternally, the long-fingered wiiiii, in which the metacarpal and 



V)G'] RELATIONS! I IPS Ol' MSSON •^(•TI•:HIS 13 

first phalanx of (li<>'it five is a little loiij-cr fliaii the forearm in 
Kpomops, and the attaclmient of the \vin<i' nieitibrane in this 
and most of the epomophorine hats to the tirst phalanx of the 
seeoiid toe are also chai'aeti'rist ically myonycteriiic. h'inally. 
thon<j:h the ai'i'an^u'eiuent of the palatal ridj^cs is lii<>hly Aariahle 
from species to sjx'eies. in one form at least of Epomops the di- 
vided fourth and undivided, straight first three rid<>'es are not 
unlike those in Lissotijich vis. 

The epomophorine affinities of Liss()ii!icf< ris and M i/oin/ctt ris 
ai'e further indicated by Benedict (IDf)?. p. L'!)2) : "Strikingly, 
the scale form of //. diif/oh )isis [Lissonifcicris aiKjolcnsis] is 
more similar to that of the Epomopliorus section than to the 
Roiisfffiis section to which AndersiMi assigns it." and "The scales 
of Mfjoiiycfcris are virtnally indistinguishable from those of the 
Epomophorns section. 

Analyzing the ditl^'erences l)et^\■een the ei)omo])horines and 
myonycterines. we fiml that the most conspicnoiis ones iisnally 
are found in the regions of the skull where, Avithin the epomo- 
l)horine group, there is nnich generic variation, or that these 
differences are more extreme developments of traits which 
Lisaonyctcris and M]ion}ict< ris already show. This suggests that 
the epomophorine is a specialized branch of the myonycterine 
group which in turn is fairly close to an early Eousitins-M^e 
stock. These annectant forms, the myonycterines, help to confirm 
Andersen's (1912, liii-lvi) supposition that the origin of the 
epomo})horine group was from a primitive and Bonscttus-Y\k(' 
stock. 

The cynopterine bats are clearly (juite different. Resem- 
blances between these and Myonycfcris such as tooth formula, 
shortening of the rostrum, slight elongation of the back of the 
skull, reduction of the tail, and d(»vel()])ment of the Aving as well 
as roosting and locomotory behavior are probably parallel 
developments in groups Avhich diverged early. Andersen also 
supposes that the cynopterine bats arose from early RoKsrftus- 
like ancestors, but. with M yonyctcris eliminated as an inter- 
mediate form, the two groups are more sharply distinct than was 
previously su]iposed. 

Geograi)hically, the j)icture pi-esented by the morphological 
evidence is a very logical one. The cynopterine bats become 
exclusively an Oriental and Austro-Malayan group, the epomo- 
phorines are restricted to Africa where they center in the 
westei-n forc^stcnl I'egions, and in this same g(Mieral area we find 
the rather more primitive but fairly closely related myonycterine 



14 BREVIORA No. 184 

group. Common to l)oth regions is the rousettine group con- 
sidered, on the basis of morphological characters, to be the most 
primitive and therefore })rol)ably the most direct descendant of 
the ancestral stock from which all arose. The occurrence of a 
highly developed s(niar system in the otherwise not particularly 
specialized RoKSfftus suggests, however, that they are farther 
removed from the ancestral stock than was previously supposed. 

ACKNOWLEDGMENTS 

This work was supported in jiart by the Office of Naval 
Research^, the United States Public Health Service, the Sigma 
Xi-RESA Fund, the Belgian American Educational Foundation, 
and Harvard T^niversity. During jiart of this period, Novick 
was a Fellow of the National Institute^ of Neurological Diseases 
and Blindness. For help in capturing and keeping these bats, 
we are grateful to the personnel of the Sangley Point Naval Air 
Station and Clark Air Base in the Philippines, to the 
Naval Attaches and other personnel of the American Embassies 
in the Philijipines and Ceylon, to Mr. Pablo Tanciojo and the 
Negritos of the Zambales Mountains of Luzon, to Capt. J. A. 
Simon, T^SAF, to Major A. Weinman of the Dehiwela Zoo, 
Ceylon, and to ]\Ir. and Mrs. W. W. A. Phillips formerly of 
Namunukula, Ceylon, to Dr. Louis van den Berghe. Messrs. J. 
Moureau, P. Pirlot, and (i. Marlier of the histitiff pour la 
Recherche 8cietififi(/U( ( n Afri(j}i( ('eufrnh . T^wiro, Belgian 
Congo, and to countless others who made these studies possible 
by giving of their time or experience. We are most deeply in- 
debted to Dr. D. R. Griffin of Harvard University for help of 
every description. 

LITERATl^BE CITED 

Allen, J. A., H. T.ans, and J. P. riiapin 

1917. The American Museum Congo Exjieilition colleition of bats. 
Bull. Amer. Mus. Xat. Hist., 2^^Am-:^^^?,, pis. XLIV-LV, figs. 1-26. 
Andersen, K. 

1912. Catalogue of tlic Cliiro])tera in the collection of the British 
Museum, ^'ol. 1. Megachiroi)tera. London, British Museum: 
i-ei, 1-854, figs. 1-79. 

Benedict, F. A. 

1957. Flair structure as a generic (diaracter in Iiats. Uuiv. ('alif.. 
Paid. Zool., 59 (8):285-548, pis. -lA-?,'!, Mgs. 1-4. 



1 Rpitroflnctiiui of this jinpcr in wlioli' or in pnrt is iifrniittfil for .my i)nrposc 
of the ITnitt'd Stati-s (Jovcrnnicnt. 



1963 



RELATIOXSIITI'S OF USSONYCTKIUS 



15 



Grittiii, I). 1{., A. Xovi.k, .-iiid M. Kornlii'M 

1958. Till' sensitivitv of ccliohM-jitidii in the I'niit li;it, RousetUis. 
Hiol. I'.iill., 115:1<I7-1 i;!. 
Kulzer, K. 

ly.jCi. Fliiglumdc (■r/iirK«'ii ( )ri('nt itTuiiKslnuti' (luicli Zungenschlag. 

Xaturwissi'iiscli.-irtcn, 43:117 IIS, 1 fi^. 
1958. Untersuclmngon iil)i'r die Biologie von Flutrliiiiidcn der Gattimg 
Ronscttiis (ir.-iy. Zcitsclir. Moi'i'li. Okol. Ticrc, 47:;574-402, figs. 
1-23. 
Miiliros, F. P. ;iiid K. Kiilzcr 

1956. UIkt (lie Orient iciuiig der Flughundc (Cliiruptera-Pteropodidae). 
Zeitschr. vergl. I'livsiol., 38:1-29, figs. 1-7. 
Noviik, A. 

1958. Orientation in paleotropical hats. II. Megaeiiiroiitera. J. Exp. 
Zool., 137:44:!-4(;l\ iil. 1. 




Figure 1. Lissonyetei-is angolensis hanging from slat by the left foot (not 
visible), holding a piece of banana with its right foot, while biting off a 

manageable portion. 



16 



BREVIORA 



No. 184 




J^^ 





Figures '2, '.'>, niid 4. Lismui i/cl cris a inioh iisis li.-iinlliii^' iiicccs (if li;iii;iii,i. 
The use (if tlic fddt ill Ji.-iiiijiiiii;' fddii .■iml tlic piistnri' of tlic wiii^is wiiiic 
doing so is t-lciirly siiown. 

Figure ~). Li.s.son!jcteri.'< (tiifiolnisis resting, Imiigiiig on ;i (■;iii\;is \v;ill .-it 
its junction witli tlie ceiling. Xote tiie position of the feet wiiicli c;in hold 
onto ;i I'oost equally well in this or the oiijiosite orientation. The wings are 
being held well o\'erla|iiieii, sulistnnt ially enwiapping tlie body. If the bat 
had not liecn disturbed by the photographer, its chiii would luive heen 
tucked against its chest and its eyes closed. 



BREVIORA 

Meseemi of Coimiparsitive Zoology 



Cambridge, Mass. 



April IN. l<)(;;i 



NlIMHER 185 



ELEUTHERODACTVLUS IIEDinCKh A NEW SPP]CIES 

OF FROG FKO.M PFFHTO KK^O 

(SALIENTIA, LEPTODACTYLIDAE) 

By Juan A. Rivero 

Department of Biology 

University of Puerto Rieo 

Mayngiiez, Puerto Rico 



The finding of a new spet'ies of frog in Puerto Rieo has seemed 
very iniprobalile. The island has been thoroughly searched by a 
ininiber of herpetologists, and since the calls of the described 
forms are well known, the recognition of a new voice is not diffi- 
cult. Yet the author has traveled miles and miles in the island, at 
night, and no unfamiliar voice was ever heard in recent years. 
He was almost convinced that a new species was out of the ques- 
tion, but it appears that this conclusion may be ])remature in 
almost any country, no matter how well the fauna of the ])lace 
is known. The reason is clear: there is a tendency to look for 
frogs in places where the known species occur, and heiu'c many 
potential habitats remain uninvestigated. 

Eleutherodactylus hedricki was heard wliih' collecting E. karl- 
schmidti in a mountain stream at El Yuncjue (El Verde). It 
was immediately evident that the voice heard was that of a new" 
species, and the animal was desperately hunted, until at last col- 
lected. It is interesting to note that this frog was recognized as 
new before being seen — a proof of the usefulness of voice, 
especially Avhere the fauna is well known, in the collecting of 
frogs new to science. 

The name hedricki has been given in honor of Hedrick J. 
Rivero. aged 9, who has declared himself assistant to his father 
and who follows him through creeks, caves and mountains dur- 
ing any time of the day or night. The author also wishes to 
express his appreciation to his wife, Eneida, and to his son, 



i BREVIORA No. 185 

Jiiaii Jr., 16, who always join him and Hedriek in th(> search 
for frogs. Iloraeio Mayorga, research assistant to tlie author, 
has also been (wtreniely useful, not only in field work, but also 
in the tedious curatorial and laboratory work that usually fol- 
lows. Dr. John Randall kindly took tlie photouraplis of E. 
hedricki, and the Galiiiaues family of San Juan made their El 
Verde house available as a (MMiter of activities for the colleetiuff 
party. To all these jx'opic, the autlioi- feels (iee])ly indebted. 

Elet'tiierodactylus iiKnRrcKi s]). n. 

Type. Museum of Coniparative Zoology Xo. ;}()!)()8, c^ from 
El Verde, west flank of El Yuiuinc Puerto Hico. I.IOO ft. 
Coll. J. A. Kivero, 11 Aug-. I!)(i2. 

Paratypes. University of Puerto Rico (]\Iaya<iiiez) Nos. 1132- 
1135, same data as type. UPRM Xos. n3(i-113S, El Yunque, 
10.9 km on road l)etween Palmer and La ]\Iina, elevation 1765 
ft., September 1962. Coll. Rivero and Mayorga. 

Difif/uosis. A medium-sized Elcutlirroddcti/JKs with short 
snout, the eye diann^ter more or less equal to the distance be- 
tween eye and nostril ; an indistinct tympanum, webless toes, 
and two, externally concave, lijjrbt-colorcd marking's on eadi side 
of the back, from behind the nape to the sacral i-egion. 

Description of type. Head much broader than long': snout 
short, sub-triangular, with a small vertical ridge at the tip ; 
tongue large, free ; vomerine odontoids distinct, almost trans- 
verse, separated, and well in back of the choanae, their exterior 
margin not quite reaching the inner l)orders of the latter; eye 
diameter almost as long as the snout ; interorbital space uuich 
broader than an upper eyelid; tympaniun indistinct, about % 
the eye diameter; canthus indistinct; loreal region almost ver- 
tical; a slight supratympanic fold to the shoulder; first finger 
shorter than second, which is shorter than the fourth ; disks 
large, fan shaped, the one on tlic first fiiigei' smaller than the 
others; two elongated metacarpal tubercles, aiul a third, smaller 
outside one; subarticular tul)ercles moderate; metatarsal tu- 
bercles small, the inner oval, the outer round and inconspicuous; 
heel of the adpressed hind limb extends to the posterior border 
of the eye; heels overlaj) when tibia and femur are placed |)er- 
pendicular to the body. Skin above and on the sides of the 
head and upper flanks studded with small warts and granules. 
Below, th(> belly, lower flanks and buttocks granular; male with 
a large subgular vocal sac. 



196."] i:i,i:i"nii:i{()i)A("rYi,i's iiiiuKicKr 3 

(\tl())-. Above, liray, willi viiric^at ions and vcrniicnlaTions of 
a li<ilit(M- ii'ray oi- wliitisli; snout dark <ira\\ except foi- some 
li<i-litei-, indistinct inai'kin<i's belo\v the eye; a dark supratyni- 
panic streak; a wliitisli interoi'hital bar followed j)osteriorly by 
a broader dark bar; two distinct, whitish, externally concave 
markings on the back from beliind the na|)e to tiie sacrum; be- 
tween these, on the anterior rei>ion, two shorter and more diffuse 
marking's of the same color; a wliitish transverse blotch with 
irreo'ular margins posterior to the sacral region; seat blackish 
with a whitish blotch on each side; thighs cinnamon brown, with 
a longitudinal series of whitish blotches from base to knee; 
two whitish, irregularly margined, transverse bars on the tibiae, 
and two on the tarsii; throat of male intensely infuscated; chest 
and belly slightly infuscated. 

Measuremenis. ( c^ , in nnn ) Snout-vent 34.8 ; head length 
]1.8; head breadth 13; femur l-t.5 ; tibia 16; foot 21. 

Description of paratypcs. UPRM No. 1182 is a 6 specimen 
with the same data as the type and with the following measure- 
ments : Snout-vent 34.1; head length 12; head breadth 13.1; 
femur 14.19; tibia 15; foot 20.8. 

The snout has a more rounded appearance than in the type, 
because the ridge at the tip is more indistinct ; the vomerine 
odontoids are rounded, set well apart and considerably behind 
the ehoanae ; the tongue is indented behind, and the tympanum 
is fairly distinct. 

The color above is blackish all over, with iiulistinct darker 
areas behind the nape and on the sacrum; the interorbital bars 
and externally concave dorsolateral lines are not evident, but 
there are three, small, light-colored areas that appear as if the 
skin had been ero(h'd fi'om those areas. The whitish thigh 
blotches are not loo distinct and there is only one broad, dark 
band on the tibia and another on the tarsus. The venter is nuu*h 
darker than in the type, but still lighter than its own throat, 
which is (hM'ply infuscated. This specimen (no. 1132) and the 
following were ])i'eserved during the daytime, while the type 
and paratype No. 1136 were preserved at night. This may 
account for the dai'k(>r coloration of 1132 and 1133. 

rPRM No. 1133, is also a 6 with the same data and the fol- 
lowing measurements: Snout-vent 3.3; head length 11; head 
breadth 12.3; femur 13.3; tibia 14; foot 20. 

This animal is almost as dark above as UPKM 1132, but the 
dorsolateral lines are still evident and tlie liglit blotches on the 



4 BREVIORA No. 185 

thighs are present, altlioujili not as distinctly as in the type. 
The axillae have eolorless areas which appear like spots; there 
ai'c two fine, liyht-colored lines marg'inino- a dark band on the 
tibia and the same ^itnation is repeated on the tarsns. The belly 
is ninch liiihtei- than in the specirnen just described. 

CPRM Xos. n;}4 and 11.35 are juvenile specimens just coming- 
out of the eggs. The dorsolateral markings are present in both 
examples, but one is almost smooth abov(% while the other has 
a number of small warts and tubercles. The liml) l)lotches are 
also more distinct in the warty example (UPK^r 11:54). They 
are 7 and 8 nnn in snout-vent length. 

UPRM Xo. 1186 is another i. from El Yunque, 10.9 km. be- 
tween Palmer and La Mina, elevation 1765 ft., 2 Sept. 1962. 
Coll. Rivero and Mayorga. Its measurements are as follows : 
Snout-vent 35.3; head length 12.1; head breadth 14.8: femur 
15.5; tibia 16; foot 22.2. 

The snout in this specimen looks rounder and higher than in 
the type and the snout ridge is only shown by a small ])ro- 
tuberance. In coloration it looks more like the type than any 
of the other examples, but it is more jirofusely reticulated and 
there is a ferruginous tinge on the dorsum, especially on the 
area of the dorsolat(M'al lines. The colorless area of the axillae 
is very distinct, and the whitish thigh spots are transverse and 
well indicated, but the seat is not as dark as in the type. The 
venter is of a light color whih^ the throat is as infuscated as in 
the other examples. 

T^PRM No. 1137 is a 7 mm juvenile from the egg clutch of 
the above paratype. It siiows the dorsolateral nuirkings (tlu)Ugh 
much broader than in the adults) but the thiaiis are moiv or 
less marbled and llie ventei- is pin-])ointed with black. 

UPRM No. 1138 is also from the egg clutch of 1136. It is 6.2 
mm in length and has a small tail, 'AS) nnn in length. Appai'entjy 
it had just come out of the ('g<x. 

Disfrihitfidit. Although E. hcdrichi has not been collected out- 
side of El Yuncfde, its voice was recently heard at the Toro 
Negro Reserve Forest and it thus seems tliat the s|)eeies i-anges 
throughout the high eh'vatioiis of the ('ordillera <'eiiti'al. The 
female is thus far unknown. 

H(lati(inshi})s. FjUhHu rodachjlus Indrichi is not too closely 
related to any Puei-to Piean frog and its habits ai-e distinct fi'om 
those of any of the known species. A coinpai'at ive skeletal study 
of the Puerto Kicaii I'Ui ulht itnlacl i/l us is under \va\- and this 



1963 ELEITTTTKHOnACTYr-rs IIKDRICKI 5 

may reveal its true affinities. In the nieatit iiiie, it is interesting 
to note that the two externally concave lines that are eharac- 
teristie of this si)eeies are shared with diminutive E. cochranae, 
E. locustus and E. gri/llus. However, these seem to be basic in 
the jjenus and ai-e shown here and th(M'e in various of the An- 
tillean species. 

Habits. EhuthcnxhichilKs Indiicki is a dense forest frog 
which breeds and apparently stays during the daytime in tree 
holes and cracks. The male guards the ego^ clutch, which in 
the case of the type consisted of 32, 4 to 4.5 mm eggs, while one 
of the paratypes had 14 eggs and 2 juveniles, and another 
had 15 eggs and 2 juveniles. In all cases where eggs were 
found, these were attached by gelatinous threads to the wood 
encircling the i)ith cavity. At the extreme ends of the cavity 
where the eggs were deposited, the i)itli was rotten and cracked 
and it does not appear improbable that juveniles wander through 
the pith for some time, eating the termites and small ants that 
abound in those areas. 

The voice of the male E. hcdricki is a sonorous "ping, ping, 
ping," quite distinct from anything known in Puerto Rico, but 
perhaps more similar to the call of E. locustus. The first speci- 
men heard was calling from high up in a tree and could not 
be collected. One of the adult ])aratyi)es called from a dead 
tree trunk, three to four inches in diameter, about six feet from 
the ground and one inch from a hole that communicated with 
a small cavity inside. The third adult specimen was in a cavity 
inside the branch of a living tree about four feet from the 
o-round. It was calling from the outside and retreated into the 
hole when collection was attempted. 

The capture of the \y\w and of paratype UPRM 1136 was 
interesting enough so that in both cases it will be related in 
detail. The call of the tyj^e was heard coming from a bushy 
area that included three, 10 to 12 ft. high young trees and a 
4 to 5 in. diameter, rotten tree trunk with a hollowed upper 
end and one or two holes a few feet from the ground. Bending 
of the trees did not stop the animal from calling, so that it 
was suspected that it was hiding somewhere in the dead trunk. 
When this was broken at the base, the frog stopped calling, but 
a thorough search did not reveal its presence. However, one 
of the members of the party thought that something that ap- 
peared like eggs was deposited at the bottom of one of the 
cavities in the trunk. To the author, these appeared to be 



6 BREVIORA No. 185 

funo'ous o'l'OAvtlis, aiul the seart-li was abandoned and preference 
ojiven to another call, eoniino- from about 50 ft. away. This 
animal conld not be found, l)nt it was during this interval, Avhieh 
may have lastetl one houi-, tliat two of the adnlt paratypes were 
obtained. On I'etnrning to the original site, the frog was im- 
mediately located, calling from the entrance of the hole. A 
thorongh search revealed the eggs that have already been men- 
tioned. The collection of this specimen is described because, if 
the animal was not hiding deep inside the tree cavity when its 
search was abandoned, then it escaped to the outside and re- 
turned to the ('^^i^ clutch in a trunk that was now in a completely 
ditferent position, that is, lying down on the ground instead of 
standing erect. 

In obtaining paratype No. 1186, a Cecropia tree had to be 
felled, since the animal was calling from a hole at about 25 ft. 
from the ground. TIk^ frog was seen to retreat further back 
into the hole when the tree hit the ground, but after that it 
managed to escajx' to the outside, perhaps because vigilance 
under the light of two Hasldights whose battei-ies were almost 
dead, was not adecpiate. The cavity contained an egg clutch 
and two juveniles (paraty]ies 1187 and 1188). It was round 
and smooth inside and perhaj)s not larger than two inches in 
diameter. No jiossibility existed for the animal to escape into 
the jntli cavity of the tree which, in Cecropia, is divided into 
compartments. 

After the eggs and juveniles were obtained, the whole section 
of the tree containing the egg clutch was removed with a machete. 
Since a thorough search of the surroumling area did not reveal 
the presence of the adult animal, the ])lace was abandoned and 
other voices traced. After aliout half an hour of unfruitfvil work, 
tlie call of h<(h-irhi led the pai-ty to the fallen Cecropia, fi'om 
which the male was again calling, only two or tliree inches away 
fi'om the place where the v^xg clutch had been i-emoved. It thus 
a])pears that this fi-og has some way of guiding itself to the 
breeding site and that this ability may not depend exclusively, 
perhaps not at all, on visual cues. 

Paratype UPRM 118(5 had 5 yellowish eggs in an advanced 
state of develo]iment. 10 white egus that wei-e iu)t as well de- 
veloped, 5 empty egg cases and 2 juveniles, it a])j)eared as if 
the two groups of eggs were laid at different times. It took 22 
days i'or all the little frogs from tlie egg clutch of tlu' ty])e to 
hatch. 



l!)(i:} 



KLKir 1 1 KRODACTYLUS IIEDRK'KI 




Leff: Eleutherodactiiliis hedricl-i s^p. iiov. type. MCZ 36903. Ilipht : Egg 
clutch of Eleutherodactylus hedricki inside a tree trunk. Some wood re- 
moved to expose the eggs. 



1 

I 



BREVIORA 

Mmseimm of Coimparative Zoology 

Cambridge, Mass. April 18, 1963 Number 186 



NOTES ON HISPANIOLAN HERPETOLOGY 

8. THE FORMS RELATED TO 
ANOLIS HENDERSONT COC!IIRAN. 

By Ernest E. Williams 

In 1923 Doris Cochran described Anolis heyidersoni from 
Petionville, Haiti. This long-headed species with a reduced dew- 
lap was represented by a single adult male collected by J. B. 
Henderson and Dr. Paul Bartsch in 1917. 

Ten years later, without making any reference to Anolis herider- 
soni, Noble and Hassler (1933) described the very similar Anolis 
baharucoensis from the mountains of Barahona peninsula to the 
east of the type locality of A. hcndersoni. This new species was 
represented by 76 specimens. The series demonstrated marked 
sexual dichromatism, and there was for the first time information 
on the habits and habitat of the species. Noble and Hassler (1933, 
p. 12) reported collecting it between the altitudes of 1500 and 
3700 feet and stated : " It is believed that the lower mountain 
sides are not humid enough for the species. It is much more 
terrestrial in its habits than the other species of Anolis in the 
same locality, being found on the ground and on low brush in 
coffee groves and forested areas. Its favorite habitat, however, 
was on low plants, leaves and trash along mountain streams and 
in humid ravines. ' ' 

Since these two descriptions the only additional published 
information is in Cochran's (1941) "Herpetology of Hispan- 
iola." She reported six additional specimens of hendersoni: 
one (USNM 82566) is a misidentified young coelestinus: the 
others are a Fond des Negres specimen (USNM 72629) collected 
by A. Wetmore and four specimens (MCZ 13792, 13794. 13795, 
13797) collected at Port-au-Prince by G. M. Allen. Cochran 
recognized the strong resemblance between hendersoni and 
haharucoensis ]nit retained the latter as a full species. She said: 
"A. hendersoni has a close ally in A. haharucoensis, as they are 



2 BREVIORA No. 186 

both distinguished by the same elongated head and body. Al- 
though at first glance the immaculate dorsum of hendersoni does 
not suggest alliance with the heavily banded baharucoensis, other 
details of coloration are more suggestive of the relationship. The 
lips are spotted in a nearly identical manner. The white lateral 
line, so striking a feature of hendersoni, is developed in a slighter 
degree in haharucoensis. The peculiarly crowded appearance of 
the scales just behind the mentals and their extreme convexity 
which makes them appear to be ridged without actually being 
keeled, is duplicated exactly in both species. Their dif- 
ferences are equally distinct and set them off in the same way 
that chlorocyanus and coelestinus are differentiated. The very 
fine bod}^ granules of haharucoensis and its rather square snout 
are not to be confused with the coarser scales and more rounded 
profile of hendersoni'^." No additional material of baharucoensis 
was recorded. 

Recent collections (1959-61) in Haiti have greatly increased 
our knowledge of Anolis hendersoni and have permitted recogni- 
tion of a third member of the hendersoni complex from the west- 
ern tip of the southwestern peninsula. This new form is recogniz- 
able on the color pattern and probably on head length of the 
adult male, but the wealth of new material appears to show that 
there are no consistent squamation differences between any of 
the members of the complex, and, therefore, despite the absence 
of any clear and positive instance of intergradation, I regard 
these three as subspecies of a single species. This problem is 
further discussed below. 

We may now redefine Anolis hendersoni as a polytypic species 
of Anolis belonging to the Hispaniolan radiation of Etheridge's 
alpha section and carolinensis series- (i.e. an Anolis without cau- 
dal transverse processes, with the lateral processes of the inter- 
clavicle in close contact with the expanded proximal parts of the 
clavicles, and with three parasternal chevrons attached to the 
dorsal ribs followed by a single chevron not so attached) and 
with the following diagnostic external characters. Head and body 
slender (head ca. Yj, or more snout-vent length), dewlap very 
reduced, not extensible. Dorsal scales small, ca. four middorsal 
rows slightly enlarged and distinctly keeled but grading into 



1 Cochran (1941, p. 1S7) was able ti> cxamino only a singlf paratype of 
hfihanicoinKis. 

2 Thcsp terms (for present purposes sufficiently defined liy the data witliin the 
parentheses ahove) are diM'ived from the doctoral thesis of Kicliarcl Kl heiiclLTc at 
the University of Michigan (availahle on microfilm). 



1963 ANOLIS HENDERSONI 3 

granular flank scales. Ventral scales smooth, polygonal, sub- 
imbricate. Digital dilations moderate. About 19-21 lamellae 
under phalanges ii and iii of fourth toe. Tail not compressed nor 
with crest. Verticils obscure. Sexually dimorphic in size and 
color pattern. Anterior head scales smooth. Supraorbital semi- 
circles separated by one scale row. Interparietal scale separated 
from semicircles by 4-6 scales. Loreal rows 6-7, canthals 6-7, 
supralabials to center of eye 6-7, suboculars in contact with 
supralabials. Mentals much longer than wide, throat scales 
medially deeply inserted between them. Three populations con- 
form to this definition but differ strongly in male color pattern 
(and in one instance in the relative head length of the adult 
male ) . 

TAXONOMIC DESCRIPTION 

AnOLIS HENDERSONI HENDERSONI Cochran^ 

Anolis hendersoni Cochran 1923. Jour. Washington Acad. Sci. 
13:25. (Type locality: Petionville, Haiti) — Cochran 1941, 
p. 181. 

Specimen list. Haiti. Departement du Nord : Citadelle MCZ 
25484, 25486, W. J. Eyerdam, 1927. Departement du Quest : Port- 
au-Prince MCZ 13792, 13794, 13795, 13797, G. M. Allen, 1919. 
Petionville USNM 59210 (type), J. B. Henderson and P. Bartsch, 
1917. Boutillier Road MCZ 59951-7, E. Williams and A. S. Rand, 
1959; MCZ 62956-9, A. S. Rand and J. Lazell, 1960. Morne 
Decay ette MCZ 62960-8, A. S. Rand and J. Lazell, 1960; MCZ 
62969-75, 63443, L. Whiteman, I960; MCZ 65635-46, L. White- 
man, 1961. Diqiiini MCZ 64824-40, L. Whiteman, 1961. Below 
Kenskoff MCZ 59950, L. Bonfil, 1959. Penault MCZ 63437-42, L. 
Whiteman, 1960. Furcij MCZ 64823, L. Whiteman, 1961. Mar- 
bial, 21 km NE Jacmel MCZ 65170-8, CM 3812-17, L. Whiteman, 
1961. Croix Joseph, Marhial, 21 km NE Jacmel MCZ 65183-202, 
CM 37818 (16), L. Whiteman, 1961. Source Fleury, Mayerre, 
8 km E Jacmel MCZ 65179-82, CM 37819-22, L. Whiteman, 1961. 
Departement du Sud : Butete near Miragoane MCZ 66029-62, 
CM 37919 (32), L. Whiteman coll. 13-viii-61. Mingrette near 
Miragoane MCZ 66063-79, CM 37920 (15), L. Whiteman coll. 



1 The following abbreviations have been used for the museums or collections 
from which specimens of anoles of this complex have been examined : AMNH. 
American Museum of Natural History ; CM, Carnegie Museum ; MCZ, Museum 
of Comparative Zoologv ; UMMZ. Universitv of Michigan Museum of Zoology : 
USNM, United States National Museum ; YPM, Yale Peabody Museum ; AS-X, 
Albert Schwartz, personal collection, 



BREVIORA 



No. 186 



1961. Risque near Miragoanc MCZ 66080-85, L. Whiteman coll. 
1961. 

Diagnosis. Head length in adult male about 33 per cent of 
snout-vent length. Male coloration distinctive. (Head brown. 
Nape vermiculate, lighter on darker brown. Dorsum brown 
anteriorly, greenish posteriorly, without transverse saddles or 
other markings. Flank stripe extending to groin bordered above 
by intense black and below at sides of belly by black vermicula- 
tion. Belly bluish.) 




Figure 1. Anolis hendersoni hendersoni, MCZ 59949. Lateral and dorsal 
views. N. Strekalovsky del. 



Color in life. Alive, hendersoni is quite spectacularly beautiful 
— especially the male. The differences between the two sexes are 
best shown by a tabular comparison of the descriptions of two 
specimens. I add a few remarks on color variation. 



$ from below Kenskoff 

Head brown and anterior back brown 
merging into green at sacral region. 



9 from Boutillier Eoad 

Head, nape and center of back dark 
brown, head somewhat mottled. The 
central dorsal area bordered by a 
cream line with an irregular bound- 
ary. 



Nape with vermiculation behind 
interparietal which is white. 



Nape without vermiculation. 
parietal white. 



Inter- 



1963 



ANOLIS HENDERSONI 



Blue spots on upper and lower la- 
bials. Limbs red brown. Tail with 
greenish tinge, becoming black 
posteriorly. 



Upper and lower labials obscurely 
marked with darker. Jjinibs and tail 
brown, obscurely mottled. 



A broad blue-black band from be- 
hind eye over shoulder, becoming less 
well defined posteriorly. 

Below this a light line from upper 
labials to groin, white below eye, 
yellow from shoulder % of way to 
hind leg, beyond this with greenish 
tinge. 

Flanks I)elow light line lioldly mot- 
tled with black. 

Throat yellow green. 



A broad brown band from snout 
through eye along flanks, lighter in 
center, dark edged above and below. 

A light line from upijer labials above 
shoulder to groin, nearly white be- 
low eye, yellow green in front of 
slioulder, purplish and indistinct on 
Hanks. 

Flanks below light line darker, not 
mottled. 

Throat yellow green with two narrow 
dark longitudinal lines. 



Belly blue green, some orange under jBelly and underside of tail pale 
base of tail. Jgreenish. 

Comments. The female, as the table indicates, is basically 
similar to the male in pattern but with duller colors and with 
frequently a light dorsal longitudinal band. The light flank 
stripe of the male is very brightly colored and brought into bold 
relief by the black mottling below it and the blue-black of the 
dark band above it. The same stripe is always obscure or absent 
posteriorly in females and may be very little evident anteriorly. 
The green of the hind quarters of the male is absent in the female 
and so also is the vermiculation on the nape. On the other hand, 
the two narrow black lines on the sides of the throat may be 
present or absent regardless of sex. 

There is a striking consistency in pattern in the animals from 
well-separated localities (i.e. Diquini and Mayerre). Such varia- 
tion as exists seems to be individual only. In the males this 
appears to be a matter of clarity of expression of various elements 
of the pattern ; this has undoubtedly been influenced by vagaries 
of preservation. In the females, along with this same variability 
in the boldness or obscurity of pattern, there seems to be also real 
pattern variation in the dorsal zone from interparietal to sacral 
region. In the live specimen described above the impression is of 
a wide dark middorsal stripe bordered by a narrow light area on 



6 BREVIORA No. 186 

each side. In numerous other females, however, the middorsal 
dark stripe is narrow, broken or irregular with usually a narrow 
light center and bordered on each side by a wide light area. This 
appears to be the more frequent of the two conditions. 

This long-headed and slender form has until now been con- 
sidered very rare. While nowhere as abundant as the ubiquitous 
species cy botes and distichus or even the two common green 
anoles, it is, as is often true of "rare" species, not really un- 
common in certain restricted situations. It seems to be a bush 
anole of middle elevations and associated especially with certain 
bushy thickets. Its habits are thus similar to those which Noble 
and Hassler (1933, p. 14) reported for haharucoensis. Hender- 
soni is, however, apparently less restricted to humid situations 
than Noble and Hassler believed haharucoensis to be. 

A. S. Rand made field notes (July 30, 1961) for this species at 
Morne de Cayette. "A. hendersoni were on stems and branches 
close to the ground, three or four feet up at most. They were 
crawling about in the bushes, jumping from branch to branch, 
seldom coming to the ground, though one did do so to catch an 
insect. They are very shy and escape by dodging away through 
the stems, neither climbing nor hiding. Relatively slow moving 
normally. ' ' 

James Lazell, Jr. provides in Figure 4 sketches from life of 
female hendersoni from Boutillier Road (August 9, 1961). These 
show in excellent fashion the characteristic postures and attitudes 
of the species. 

Anolis hendersoni haharucoensis Noble and Hassler 

Anolis haharucoensis Noble and Hassler 1933. Amer. Mus. Novi- 
tates No. 652: 12. (Type locality: "Valley of Polo, Barahona 
Province, D. R.")— Cochran 1941: 184. 

Specimen list. Haiti. Departement de Quest : Road to Sal Trou, 
on south side of range of Mt. La Selle AMNH 50096, W. G. 
Hassler, 1935. Caroije near Sal Trou MCZ 68693-714, TJSNM 
146616-8, YPM 3704-13. UMMZ 123372-81, CM 38497 (11), G. 
Whiteman, 1962. ^ Doimnican Republic. Barahona Province: 
Vicinity of Polo AMNH 49516. VaJle de Polo AMNH 51081-106, 
51108-19, 51123-27, 51128 (type), 51129-52, 51154-56, MCZ 
43822, 45952-53, 56138. Polo AMNH 50317, 50322-23. Palomino 



1 The recorrts listeil above for the vifinity of Sal Trou are the first for Haiti. 
At Caroy^, in addition to .4. bahanicotuKis, Georjre Whiteman obtained also the 
rare form, Chinndcliiioropis icvtmorti. 



1963 



ANOLIS HENDERSONI 



Springs near Barahona AMNH 49840-42, 49884-85, MCZ 43827, 
56137. Barahona AMNH 50263.1 

Diagnosis. Head not more than 33 per cent of snout-vent 
length. Coloration of male distinctive. (Head green. Nape 
green spotted with brown. Dorsum with broad brown saddles 
on a green ground. Flank stripe extending to groin, not sharply 
defined, sometimes broken, bordered irregularly above and be- 
low. No black vermiculations on sides of belly. Belly cream 
tinged with brown and green.) 

Color in life. Noble and Hassler (1933, p. 13) : "In life the 
male is an extremely beautiful lizard. In its usual and brightest 
phase the dorsal surface of the head is an olive green ; the neck 
a lighter green, spotted with brown. The back is a bluer green, 




Figure 2. A^wll.s hendersoni halianicoeiisis, MCZ 68917. Lateral and dorsal 
views. N. Strekalovsky del. 



1 Sonip confusion exists regardintf the labelling of American Museum of Natural 
History specimens. Seventy-three paratypes of bahdnicoeitsis — AMNH 51081-127, 
.">1129-r)0. all apparently from Valle de Polo, are listed above. (The specimens 
missiuf,' from the series are all presunuibly exchanged.) However, Noble and 
Hassler's text mentions specimens from five additional localities, only two of 
which are now accounted for by specimens so labelled in the American Museum 
collections. Thus, unrepresented in the extant collections, though mentioned by 
Noble and Hassler. is material from Maniel Viejo, from the coffee finca of Senor 
Luis E. Del Monte near Barahona, and from the property of Mr. G. HeiTnann 
near Paradis. In addition, some of the specimens now catalogued in the American 
Museum as baharucoensis from Polo are pulchcllus, presumably from Puerto Rico 
or the Virgin Islands. Doubtless, as is known to have occurred with Hassler's 
and other material at this period, some of the lizards were kept alive for a period 
and only later and rather randomly catalogued. 

Certain of the MCZ material (MCZ 43822, 43827) at present catalogued as 
paratypes, is listeu as received in exchange from J. C. Armstrong and presumably 
was never at the American Museum and thus is not, in fact, paratypic. 



8 BREVIORA No. 186 

while the four broad saddles or cross bars are a tone of burnt 
umber. The dorsal surface of the tail, for the anterior two-thirds, 
is a yellowish green with several dark brown bars. The posterior 
end of the tail is brown. The side of the head is greenish brown 
back to the eye. The upper eyelid may be vivid golden yellow. 
The posterior corner of the lower lid may be blue or purplish. 
Just posterior to the eye is a narrow patch of dark brown, fol- 
lowed by a crescent of light blue or white. Posterior to this the 
side of the head is a brownish green merging into the lighter 
green of the side of the body which is peppered and veined with 
brown. Extending from a i)oint on the upper labials anterior to 
the eye along the sides of tlie body nearly to tiie hind leg is a 
slightly broken white or cream-colored line edged with brown 
and suffused in the region above the front leg with yellowish 
green. The legs are light brown above with slightly darker bars. 
They are nearly white beneath. The ventral surface of the ab- 
domen is cream-colored, faintly tinged with brown and green. The 
throat is the same color with several rows of very faint brown 
spots along the side. 

"This species changes color rapidly and to a marked degree. 
When the lizard is caught or frightened, these colors almost 
instantly become darker, the green changing to gray or dark 
brown with the brown cross bars growing darker and almost 
black-edged. The liead l)ecomes dark l)rown and the labials green- 
ish. The pineal region becomes white and very conspicuous. The 
ventral surfaces turn greenish or yellowish and the spots become 
more distinct." 

AnOLIS IIENDERSONI DOIilCHOCEPIIALUS Sul)Sl). U. 

Type: M('Z 64510, adult male. Place Negre near Jeremie, 
Depart emeiit du Sud, Haiti. Luc and George Whiteman coll. 
12-xii-G(). 

Paratypes: Haiti. Departement du Sud. Place Negre near 
Jeremie MCZ 64507-9, 64511-36, Luc and George Whiteman coll. 
12-xii-60. Les Platans above Carrefour Canon near Ducifi MCZ 
62f)76-82, A. S. Hand and J. Lazell coll. 4-viii-60. Carrefour 
Canon near Dueis MCZ 62!)8;}-f)2, A. S. Hand and J. Lazell coll. 
4-5-viii-60. Tomheau Cheval MCZ 62!);)3-7, A. S. Kaiul and J. 
Lazell coll. 7-viii-60. Mountains on road 1o Jeremie MCZ 56145, 
AMNII 49504, W. G. Ilassler coll. 1!)35. About 8 miles from 
Camp Perrin AMNH 50098, W. G. Hassler. 1935. Five miles 
from Camp Perrin on J (route Road AMNH 50127, W.(}. Ilassler, 



1963 



ANOLIS HENDERSONI 



1935. Camp Perrin AS-X 2664, 2800-2802, 2923-25, A. Schwartz 
coll. 1962. 13 km N Cavaillon AS-X 3646, A. Schwartz coll. 1962. 
Diagnosiii. A subspecies of Anolis liendersoni differing in the 
greater elongation of the head in large males (more than 33 per 
cent snout-vent length) and in coloration. (Head brown, nape 
verniicuhite ligliter brown on darker. On dorsum of male a few 
small middorsal liglit -edged transverse bars at intervals, the 
widest just behind nape. Flaidv stripe extending only to mid- 
body, narrowing and terminating rather abruptly. A narrow 
bhick border above the stripe and black vermiculations below it, 
both disappearing abruptly along with the stripe itself. The 
postei-ior flanks unimtterned. Belly yellowish.) 




Figure 3. Anoli.s lu-ndcrsoni dolirhorrphahis suhap. iiov. Type, MCZ 64510. 
Lateral ami dorsal views. N. Strekalovsk.v del. 



Color in life. Description by W. G. Hassler of AMNH 50098 9 
from mountains on Jeremie road about 8 miles from (,'amp Perrin, 
2000-3000 ft.: "Dark brown, striped with lighter brown or 
yellowish lines. Belly and throat yellowish. Belly spotted. No 



green. 



A. S. Rand for specimens from Les Platons : "5 Uniform 
light brown above, grayer on head and neck. Darker brown on 
side of neck. A yellow stripe, black bordered above and below 
from below eye to midbody. Below pale yellowish. Faint dark 
striping on throat. Dewlap area with greenish tint. Eye brown. 

" ? A middorsal stripe edged with light gray. Sides brown 
with white, black bordei-ed stripe from below eye to mid body. 
Belly yellow. Throat and chin white." 



10 



BBEVIORA 



No. 186 



A. Schwartz for specimens from Camp Perrin : " 9 Dorsal 
ground color yellow-tan with a fine pale hairline with brown suf- 
fusions on each side. Sides dark brown with a light lower line 
on sides. Upper labials cream, occipital creamy. Ventral ground 
color pale greenish yellow." 

Comments. Again the color pattern is remarkably consistent 
in the several populations sampled. Males from the south side of 
the Massif de La Hotte (Les Platons etc.) differ from those from 
the vicinity of Jeremie only in the weaker expression of certain 
features, i.e. the vermiculation of the nape and the small mid- 
dorsal transverse markings. The striking way, however, in which 
the flank stripe, the black line above it, and the vermiculation 
below stop abruptly at the same place is exactly repeated in all 




Figure 4. AnoUs hendersoni hendersoni. A Boutillier Road female at the 
height of one foot above tlie ground. Characteristic poses. J. Lazell, Jr. del. 



1963 ANOLIS HENDERSONI 11 

male specimens. The extreme long-headedness of fully adult 
males is again characteristic of all samples. 

Female coloration is very like that of typical hendersoni except 
that in topotypic specimens from tlie vicinity of Jeremie the 
lateral light stripe fades rather abruptly at midbody, while in 
females from south of the Massif de La Hotte the stripe tends to 
continue very faintly to the groin as is usual in typical hender- 
soni. 

Dolichocephalus at Tombeau Cheval (3000 ft. elevation) was 
captured along with Anolis monticola, A. distichus subsp., A. 
cyhotes subsp. and A. coelestinus in the vicinity of a great heap 
of jagged limestone boulders overgrown with bushes and with 
much leaf litter — cool and shady. A. S. Rand reports that here 
while the monticola were found ' ' around the cliffs on the rocks, 
roots, twigs, small branches and leaves close to the ground," 
dolichocephalus was "... up on small branches, vertical stems 
one to four feet up and to a less extent among the rocks. A. 
distichus was common on the larger trees and less frequently 
on big rocks. A coelestinus was seen and a few A. cyhotes on 
rocks, trees, and bush stems. These last were much more common 
along the open trail and in coffee." 

SPECIES OR SUBSPECIES 

The three members of the hendersoni complex are allopatric 
and in most respects extraordinarily similar. Despite Cochran's 
statement quoted above, I do not find the structural differences 
between these three taxa at all clear. Abundance of material has 
reduced rather than strengthened any suggestion of shape or 
squamation difference. 

The color differences, on the other hand, are very marked. 
What is the biological significance of such differences in anoles 
without a functional dewlap? There is no objective evidence. I 
have reduced haharucoensis to a subspecies and have described 
dolichocephalus at this level for the following reasons: Wliile 
color is very important in Anolis and while the importance of 
pattern and color may be expected to be still more important in 
forms which lack a functional dewlap, anoles which do differ 
strongly in color and pattern (and even in structure) may inter- 
grade (e.g. Lazell, 1962). Further, every sibling Anolis species 
of which I am aware is discovered to show at least average struc- 
tural differences once the sample is adequate. The apparently 
contradictory case of .4. alter newly described by me (Williams, 



12 



BREVIORA 



No. 186 



1962b) I believe to be an instance of inadequate sample size. The 
material of the hendersoni complex is now quite sufficient to 
demonstrate average differences, were they in fact present. It 
is true that the largest males of dolichocephalus are slightly but 
distinctly longer headed than any other members of the complex, 
but this is quite obviously not the sort of evidence that suggests 
species status. 




HISPANIOLA 



SCALE OF KILOMETERS 



hendersoni 9 
bahorucoensis  
dolichocephalus -f- 

Fi^iiie "). M;i]) of tlu' (lisliibution of the races of A. licnilcrsoni. 



ZOOGEOORAPHY 

The two specimens from tlie Citadelle indicate that much is 
still to l)e found out about this gi-oup in Hispaniola. Rare as these 
forms have been in previous collections, they are abundant in 
some of the most recent. It is therefore impossible to reason on 
the basis of negative evidence that A. hendersoni is really absent 
in any part of Hispaniola north of the Cul de Sac — whether in 
Haiti or in the Dominican Republic. 



1963 ANOLIS HENDERSONI 13 

It is thus too early to discuss the origiu or history of the 
hvndcrsoni complex. It is, however, worth calling attention to 
the tripartite division within what I have called (1961, 1962a) 
the "southern island" — Ilispaniola south of the Cul de Sac. 
This singular pattern occurs in several other instances and will 
be discussed further in later papers of this series. 

ACKNOWLEDGMENTS 

I am indebted to A. S. Rand and J. D. Lazell, Jr. who first 
collected the new subspecies of hcndersoni described herein and 
to Luc and George Whiteman of Port-au-Prince, Haiti, who ob- 
tained most of the other material reported here. I am grateful 
also to Dr. Doris M. Cochran and Mr. C. M. Bogert who permitted 
me to examine material in their charge and to Dr. Albert Schwartz 
who has allowed me to study material he has very recently col- 
lected. A grant from the American Philosophical Society and 
National Foundation Grant NSF G5634 have supported part of 
the collecting reported here. Work on these and other Haitian 
anoles has continued under NSF G16066. 

EEFEEENCES CITED 

Cochran, D. 

1923. A new Annlis from Haiti. Jour. Washington Acad. Sci. 13: 225- 

226. 
1941. The herpetology of Hispaniola. Bull. U. S. Nat Mus 177: 1-398. 
Noble, G. K. and W. G. Hassler 

1933. Two species of frogs, five new species and a new race of lizards 
from the Dominican Republic. Amer. Mus. Novit. No. 652: 1-17. 
Lazell, J. D. Jr. 

1962. The anoles of the eastern Caribbean (Sauria, Iguanidae). V. 
Geographic differentiation in A7wlis oculatus on Dominica. Bull. 
Mus. Conip. Zool. 127: 466-475. 
Williams, E. E. 

1961. Notes on Hispaniolan herpetology. 3. The evolution and rela- 
tionships of the Anolis semilineatus group. Breviora, No. 136: 
1-8. 
1962a. Notes on Hispaniolan herpetology. 6. The giant anoles. Breviora, 

No. 155: 1-15. 
1962b. The anoles of the eastern Caribbean (Sauria, Iguanidae). IV. 
The anoles of the northern Leewards, Anguilla to Montserrat: 
New data and a new species. Bull. Mus. Comp. Zool. 127: 453-465. 



BREVIORA 

Museiiwii of Comparative Zoology 



Cambridge, Mass. .July 10, 1963 Number 187 

THE LABYRINTIIODONT DENTITION^ 

By John Newland Chase 

Department of Zoology, Ohio Wesleyan University 

Because of the pliylogenetic importance of the Labyriiitho- 
dontia, numerous studies have been made of the cranial anatomy 
of members of this oreat amphibian group. Little, however, has 
been written regarding their dentition. My attention was called 
to this subject because of the unusual nature of the dentition of 
a newly discovered trimerorhachoid rhachitome from the Texas 
Permian, which is to be described in a forthcoming paper as 
Neldasauriis wrujhiae? In the present paper I shall review our 
current knowledge of the dentition of the Labyrinthodontia as a 
w]u>le. The data on which this review is based, summarized in 
Table 1, were drawn directly from specimens in a few instances 
but most were obtained from the literature ; measurements re- 
corded are tliose quoted by authors or determined from scale 
drawings of original or reconstructed specimens. Fifty-one gen- 
era, including temnospondyls, anthracosaurs and a crossoptery- 
gian Avere reviewed. The arrangement of labyrinthodont genera 
in the table is essentially as in Romer's 1947 classification (pp. 
310-319). 

Characteristic of labyrinthodonts generally is a dentition 
which includes, in addition to marginal tooth rows, palatal tusks 
and often, at least, a shagreen of denticles on the palate and the 
coronoid region of the lower jaw. The primitive condition, it 
would seem, was one in which a relatively small number of 
marginal teeth were present and the palatal dentition consisted 



1 This paper is part of a tliesis submitteil to the Department of Biology of 
Harvard University as partial fultillnient of the reqnirements for the degree of 
Doctor of Philosophy, August, 1U(52. 

2 This is a nomen nudum here, and will enter scientific nomenclature with the 
publication of my projected descrijition of this new form. 



2 BREVIORA NXD. 187 

of a siiiti'le stout tu.sk-j)air on each of tlie tliroe lateral ]ialatal 
elements. There is, however, wide variation in the number of 
marginal teeth, and in various instances a trend toward reduc- 
tion in size and concomitant increase in numbers of palatal tusks. 

Tn this paper I have confined attention to the dentition of the 
upper jaw and palate, since, apart from the occasional presence 
of symphysial tusks, the lower dentition in great measure mirrors 
the marginal dentition of the upper .iaw. Further, no attempt 
is made to ascertain how widely a shagreen of tiny denticles is 
present on the jialate, since, even if present, such a shagreen is 
often destroyed in preparation and consequently unrecorded. 

The data ])resented in the tal)ulatio]i include : 

(1) A formula representing the palatal dentition, recorded in 
terms of the numbers of tusks (or tusk pits) and small 
teeth on the vomer, the palatine and the ectopterygoid of 
one side. For example, the formula for the palatal denti- 
tion of Lyroccphalus is 2 — 2(3) — (13), which means 
there are two tusks on the vomer, two tusks and three 
smaller teeth on the palatine, and thirteen small teeth on 
the ectopterygoid. 

(2) Information concerning the marginal upper jaw denti- 
tion such as (a) the number of premaxillary and maxil- 
lary teeth, (b) the presence of regionally enlarged teeth 
forming "canine peaks" Avhenever it could be determined, 
indicated on the table by a plus sign. 

(3) Skull length taken as the distance from the tip of the 
snout to the end of the occipital condyle. (This, taken 
together Avith the figure on the number of teeth, will give 
an indication of relative tooth spacing.) 

(4) Sources, placed here to avoid repetition in the text. 

It is generally and reasonably assumed that labyrinthodonts 
are descended from rhipidistian crossopterygians. The dentition 
is known in only a few crosso])terygiaiis, such as E nsthenoptcron, 
MegalicJifJnjs and Eciostcorhachis. In Eusilioioptcron of the 
Upper Devonian we find: (1) a series of very nuiiu'i-ous small 
marginal teeth, (2) a row of numerous small teeth along the 
outer margins of the vomer, palatine and ectopterygoid, (3) 
larger tusks, few in mnnber, ])lace(l more medially on these thive 
elements, with some indication of tusk and ])it pairing; as far as 
known the vomer bears only one pair of tusks. 

In the following discussion we will consider marginal and 
palatal dentitions separately. 



1963 LABYRINTHODONT DENTITION 



MARGINAL TKETII 



T1h> oldest known labyriiitliodoiit, Ichthyostcf/a from East 
Greenland beds near the Devonian-Carboniferons bonndary, is 
somewhat oft' the main line of" labyi-inthodont evolntion bnt shows 
a nio(h^st number of well spaced teeth — 8-9 premaxillary, 16-18 
maxillary — of fairly ji'ood size, most al)ont the same length but 
with th(> posterioi- premaxillary teeth somewhat eidarfjed. Acan- 
thostega, an iehthyosteo-alian of comparable age, appears to have 
had more maxillary teeth — about 30. Romer (1947) provision- 
ally associated the Colosteidae of Linton with the ichthyostegals. 
The marginal dentition of the colosteid Erpciosaurus, however, 
does not agree with that of the ichthyostegals, for here, in con- 
trast, there is a reduced number of rather large premaxillary 
teeth and very numerous small maxillary teeth. 

All further well-known labyrinthodonts can be clearly divided 
into temnospondyls and anthracosaurs. The marginal dentition 
of temnospondyls will be dealt with first. 

The most primitive temnospondyls (apart from their peculiar 
''keyhole orbit") are clearly the loxommids. such as Baphetcs 
and Mcgaloccplialns. In the number of premaxillary teeth both 
appear primitive, but the maxillary series — 2\ in Bophctes, 40 
in Mcgolocrphah(s — are in contrast, the larger number being 
perhaps proportional to skull elongation in Mcgaloccphalus. 

Exemplifying the next higher stage are the edopsoids, Edops 
and Eugyrinus (the last was considered a trimerorhachoid by 
Romer, but recent studies by Carroll indicate that Eugyrin\is is 
more primitive and generalized than the trimerorhachoids) . Here 
the premaxillary teeth are primitive in number (9, ?7) and the 
number of maxillary teeth (24, 29) is fairly low. 

Among trimerorhachoids, some agree fairly well with the prim- 
itive edopsoid pattern (the maxillary count of 19 in Saurerpc- 
ton is lower) but within the group there is a notable tendency 
toward an increase in the number of marginal teeth. The pre- 
maxillary count increases from 9 in Sourerpcton to 12 in Trinic- 
rorlKfchis and Eohrachyops and to 15 in Neldasaurus and Dvino- 
mnrns. The number of maxillary teeth is somewhat increased in 
Eohrachyops, Dvinosaurns and Trimerorhachis and increased to 
the spectacularly high number of approximately 93 in Nelda- 
sdunis. 

Certain, forms which Romer grouped as the Micropholoidc^a 
show characters more or less intermediate between primitive 
edopsoids and the "typical" rhachitomes of the Lower Permian. 



4 BREVIORA No. 187 

Three members of the group — Archegosaurus, Chenoprosopus 
and Lysiptcrygium — show a "normal" rhachitomous tooth 
count with 8-11 premaxillary and 27-29 maxillary teeth; Micro- 
pholis, however, has a reduced dentition with 5 premaxillary and 
16 maxillary teeth, Avhile PJatyops, in correlation with an ex- 
tremely elongate skull, has a total of about 65 marginal teeth in 
the upper jaw. 

The "typical" rhachitomes, the Eryopsoidea, include a wide 
variety of Permian labyrinthodonts and a few Carboniferous 
predecessors. Dental formulas are given for nine members of this 
group (nos. 20-28, Table 1) and show some variation within the 
group. Premaxillary teeth range from a maximum of 15 in the 
broad-snouted Zatrachys and its Carboniferous relative Acan- 
tJtostoma, through 13 in Eryops to a low figure of 5-8 for Acti- 
nndon, Cacops and trematopsids. Some eryopsoids have a 
reduced maxillary count with a low of 12 in Cacops; Eryops, 
on the other hand, has an increased count of 38. 

The trematosaurs, a persistently rhachitomous, early Triassic, 
fish-eating group, reflect accompanying snout elongation in tooth 
numbers. Although the premaxillary teeth retain a count of 
10-12, in the long-snouted forms the number of maxillary teeth 
increases, reaching a figure of 50 in Trcmatosuchns. 

The neorhachitomes of the late Permian are presumed to have 
been derived from eryopsoids. Such a typical form as Rhine- 
suchns lias a marginal dentition almost exactly like that of its 
morphological ancestor Eryops; Lydekkcrina, however, has a 
reduced formula, with only 20 maxillary teeth. Still more ad- 
vanced neorhachitomes of the early Triassic, those apparently 
leading to the capitosaurs, are such forms as WcUngasaurus, 
VoUjasaurus and Benthosuchus. Here, facial elongation is accom- 
panied by an increase in the number of maxillary teeth to 52, 56 
and 61 in these genera, respectively. A high number of maxil- 
lary teeth occurs in capitosaurs, reaching a peak in Mastodon- 
saurus, which has 23 premaxillary and about 75 maxillary teeth. 
Parallel in development to the capitosaurs are those forms 
grouped as the metoposaurs. Metoposaurus itself has a "general- 
ized" count of 10 premaxillary and 34 maxillary teeth, but in 
the American species Eupdor browni and the European Ey prior 
frnasi the count increases to about 60 maxillary te(>th in the 
former and to 1)0 in the latter. Certain short-faced Triassic forms 
tend to have reduced counts; the brachyopid Batrachosnchus 
having but 17 maxillary teeth and the plagiosaur Gerrothorax 
having somewhat over 26. 



1963 LABYRINTIIODONT DENTITION 5 

A group by group account of variations of marginal tootli 
count in tenniospondyls has been presented above. As can be 
seen, no consistent ]iattern emerges. One gains the general im- 
pression of a probable early temnospomlyl condition of 9 or so 
premaxillary teeth and a maxillary count in the 20 's, with a 
modest increase, on the average, in typical rhachitomes. Occa- 
sionally there are reductions to lower figures (as, for example, 
in Sclerocephalus, Actinodon, or the dissorophid Cacops). On 
the whole, however, variations are toward higher figures in later 
forms. In some cases increases in tooth count are definitely 
associated either with notable snout elongation, as in Platyops 
and the trematosaurs, or, as in capitosaurs and metoposaurs, with 
a combination of moderate skull elongation and absolute increase 
in size, the teeth failing to increase in proportion to the total size 
of the animal. Most exceptional of all are the trimerorhachoids, 
particularly Neldasaurns, where an exceedingly high tooth count 
may be found in a skull of modest proportions. No consistent 
correlation between tooth number and skull size can be demon- 
strated. 

Anthracosaurian dentition is known in only a few, mainly 
Carboniferous forms. Here the premaxillary teeth are low in 
number, related to the usually narrow snout and the fairly large 
size of the individual teeth. Marginal teeth are essentially of 
two types : either large and few in number, as in Anthracosaurus 
and Pteroplax, or small and more numerous, as in Pholiderpeton, 
Neoptcroplax and, especially, Archeria, where the exact count is 
indeterminate although the teeth are certainly very numerous. 
Marginal tooth counts in the premaxilla and maxilla of ^'('^/- 
mouria are low; Kotlassia shows a slight increase over the primi- 
tive number. 

Regionally enlarged teeth forming "canine peaks" occur 
sporadically in different labyrinthodont groups but appear to 
have been fairly common in primitive forms. 

PALATAL TEETH 

Known rhipidistian crossopterygians have, as mentioned above, 
(1) a very large number of small teeth in a row along the mar- 
gins of the vomer, the palatine and the ectopterygoid, (2) larger 
tusks more medially placed on these three bones. As is well 
known, in all typical labyrinthodonts the larger teeth are re- 
tained while the lateral row of smaller teeth is lost. But, surpris- 
ingly, in IchtJiyostcga there is no trace at all of the "tusk" row; 



6 BREVIORA No. 187 

instead it appears that the lateral row of numerous, essentially 
even-sized teeth has been retained. In this regard Ichthyostcga 
is very far removed from the condition expected in the ancestor 
of later labyrinthodonts. 

Primitive temnospondyls proper almost always have one pair 
of tusks on each of the three bones, the vomer, palatine and ec- 
topterygoid. This is true almost without exception in loxommids, 
edopsoids and most eryopsoids. Occasionally, a replacement pit 
may be absent ( ? or indeterminate) but exceptions are rare. In 
palatal dentition, colosteids (e.g. Erpetosanrus) are typically 
temnospondyl in character, thus arguing strongly against 
Romer's suggestion of ichthyostegal relationships for these 
animals. 

In numerous more specialized or advanced labyrinthodonts 
there is an increase in the number of smaller palatal teeth and 
a tendency toward reduction of the more prominent tusks, 
usually those on the ectopterygoid. The presence of this pattern 
in different groups indicates that this trend has occurred sev- 
eral times in parallel fashion. It is, for example, seen in tri- 
merorhachoids (except Eohrachyops), mieropholoids and trema- 
tosaurs. In Rhinesuchus, which appears closely related to typical 
eryopsoids, this trend is seen again, leading to conditions in 
capitosaurs where, as in metoposaurs, the increase in the number 
of small teeth is very prominent. In some short-faced Triassic 
forms the increase in smaller palatal teeth is not so pronounced 
— a condition no doubt correlated with the short palate in these 
animals. 

Distinctive of tlie anthracosaurs is the absence of vomerine 
teeth — apparently related to the narrow palatal exposure of 
the vomers in these forms. In the few palates known, there are 
generally a pair of large palatine tusks and a modest number 
of ectopterygoid teeth, the front ones tusk-like. In Pliolidogaster, 
with six tusk-like teeth on the ectopterygoid, and in Anthraco- 
saurus, where a tusk pair is accompanied by a pair of fairly large 
teeth, we encounter conditions reminiscent of the crossoptery- 
gians, in which the palatine had a row of large teeth. 

Seymouriamorphs are unusual among anthracosaurs. Scy- 
mouria, in contrast to other anthracosaurs, has a pair of vomerine 
tusks and — in parallel fashion to some temnospondyl groups — 
has lost the ectopterygoid teeth. Kotlassia resembles other anthra- 
cosaurs in lacking vomerine teeth but has a close-set row of small 
teeth on the palatine and ectopterygoid, decreasing gradually in 



]963 LABYRINTIIODONT DENTITION 7 

size from front to back. These differences between known anthra- 
eosanrians sngwest much variation within the group, but the 
extent of this variation is at pi-esent incompletf^ly known. 

I wish to express my appreciation to Dr. Alfred S. Konier of 
the Museum of Comi)arative ZooU)gy for his generous aid and 
many helpful suggestions during the preparation of this paper. 
1 am also indebted to Professor Bryan Patterson and Dr. Ernest 
E. Williams of the Museum of Comparative Zoology for eon- 
.stinu'tive criticism of the manuserii)t. 

EErERENCES 

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BrOILI, F. and J. RCHROEDER 

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Bystrow, a. p. 

1935. Morphologische Untersuchungen der Deckknoclien des Schiidels 
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8 BREVIORA No. 187 

Efremov, J. A. 

1933. Xeuljesi-hreil)U]ig fles Labyrinthodonten PJaiyops stuchcnbergi 
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1940. Pieliminary description of the new Permian and Triassic Tetra- 
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Fraas, E. 

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1952. On the fish-like tail in the ichthyostegid stegocephalians. Medd. 
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Langston, W., Jr. 

1953. Permian amphibians from New Mexico. Bull. Dept. Geol. Univ. 
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NiLSSON, T. 

1934. Vorliiufige Mitteihing iiber einen Stegocephalenfund aus dem 
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1937. Ein Plagiosauride aus dem Rhiit Schonens. Beitrage zur Kenntnis 
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Olson, E. C. 

1941. The family Trematopsidae. Jour. Geol., 49:149-176. 
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1930. The Pennsylvanian tetrnpods of Linton, Ohio. Bull. Anier. Mus. 
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1942. Edops, a primitive rhacliitomous auipliiliinn from the Texas red- 
beds. Jour. Geol., 50:925-960. 



1963 LABYBINTHODONT DENTITION 9 

Save-Soderbergh, G. 

193(). On the moiplioloKy of Triassic stegocephalians from Si)itzberKen, 

and the interpiotation of the endocraniuni in the Labyiintho- 

dontia. K. Sveuska Vetenskapsakad. Handl., (3)16(1) :1-181. 
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1938. On the fossil Amphibia from the Gas Coal of Nyfany and other 
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1936. Notes on the pre-Jurassic tetrapods from USSR. III. Dvino- 
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10 BREVIORA No. 187 

Whittard, W. F. 

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1911. Restoration of Seymouria baylvrensis Broili, an American 
cotylosaur. Jour. Geol., 19:232-237. 



1963 LA15YK1NTJ10DONT DENTITION 11 



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BREVIORA 

Meseium of Coimparatlve Zoology 



Cambridge, Mass. Jily 2<). 196:5 Number 1S8 

REDESCKIPTION OF SOME CAVERNICOLOUS 

PSEUDOSCORPIONS (ARACHNIDA, CHELONETHIDA) 

IN THE COLLECTION OF THE 

MUSEUM OF COMPARATIVE ZOOLOGY 

By 

William B. Muchmore 

University of Eochester 

Throu<i'li the courtesy of Dr. H. W. Levi, the author was able 
to examine the collections of pseucloscorpions in the Museum of 
Comparative Zoology at Harvard College. The material consid- 
ered in this paper was discovered among- undetermined collec- 
tions and recognized as type material by the characteristics 
mentioned in the treatment of the individual species. Included 
here are the types of three little-known, cavernicolous forms, 
Blothi'us packardi Hagen, 1879, Chthonius coccus Packard, 1884, 
and Ohisinm cavicola Packard, 1884. Detailed redescriptions of 
these species based on mounted specimens are presented. In 
addition, the specimens from Mammoth Cave, Kentucky, con- 
sidered conspecific with B. packarcli (from Wyandotte Cave, 
Indiana) hy Hagen, Packard and others, are separated from that 
species and made the basis of a new species. 

Suborder HETEROSPIIYRONIDA Chamberlin 

Family CHTHONIIDAE Hansen 

Subfamily CHTHONIINAE Daday 

Tribe CHTHONIINI Chamberlin 

Genus KlEPTOCHTHONIUS Chamberlin 
Subgenus Chamberlinochthonius Vachon 



2 BREVIORA No. 188 

Kleptochthonius (Chamberlinochthonius) packardi 

(Hageii), new eombination 

Blothrus paclcardi Hagen, 1879, Zool. Aiiz., vol. 2, p. 399. 

ChtJionius paclcardi [pro parte] : Hubbard, 1880, Amer. Eiit., vol. 3, p. 83. 

Chthonius packai'dii [pro parte] : Packard, 1888, Mem. Natl. Acad. Sei. 

Washington, vol. 4, p. 43. 
Chthonius pacl'ardi [pro parte] : Banks, 1895, Jour. New York Eut. Soc, 

vol. 3, p. 13. 
Chthonius? paclcardi: Beier, 1932, Das Tierreich, vol. 57, p. 61. 
Genus? paclcardi: Hoff, 1949, Bull. Illinois Nat. Hist. Surv., vol. 24, p. 443. 
Chthonius? paclcardi: Hoff, 1958, Amer. Mus. Novitates, no. 1875, p. 4. 

Material: The type collection, consisting- of 5 males and 1 
female, was identified by a label reading "Wyandotte Cave, 
Ind." in the presumed handwriting of H. Hagen. Another 
female speoimon with the label "Wyandotte New Cave, Paelvard" 
is taken to be that specimen mentioned by Packard (1888, p. 44). 
All of these specimens, witli the exception of one male from the 
type collection, were mounted on slides for study. One of the 
mounted males has been designated the lectotype. Although 
Packard and Hul)bard have given fairly accurate general de- 
scriptions (cf. Packard, 1888, p. 43-45), a redescription based 
on tiie mounted s})ecimens is desirable. 

Diagnosis: Male: (Measurements are given first for the lecto- 
type, while in parentheses are given the ranges for the three 
paratypes.) Body and appendages generally of chthoniid facies 
but more slender than in epigean species (see Packard, 1888, fig. 
12, p. 45) ; lightly sclerotized and xery light brown in color, the 
chelicerae and palps a little darlvcr than other parts. Carapace 
nearly square in dorsal outline, the lateral margins usually a 
little convex ; surface lightly sculptured ; no epistome ; no eyes 
or eye spots; total setae 16 (15-18), of which 3 (2-4) are on the 
anterior margin and 2 (2-3) near the posterior margin. Abdomen 
elongate ; tergal and sternal scuta entire and smooth ; pleural 
membranes very finely granulate ; tergites of lectotype with 

2 :2 :2 :2 :4 :4 :6 :6 :6 :2 long, acuminate setae, paratypes with 2-3 :2- 

3 :2-3 :2-4 :4 :4 :5-6 :6 :6 :2-5 setae ; sternites 4-8 with 7-10 acuminate 
setae, shorter and more slender than those of the tergites. 

Chelicerae generally of chthoniid size and proportions; hand 
with 7 setae ; fixed finger with a row of 6-9 teeth, the movable 
finger with a row of 4-7 teeth, the teeth of each finger being 
heavy and triangular and becoming progressively lower and 



1963 CAVERNICOLOMS PSEUDOSCORPIONS 3 

sinHlIcr lowai'd \he proximal eiul of tlic fiiijjfer; no isolated sul)- 
distal tootli on (ntlicr finy-er; tlio p'aloa represented by a small, 
but distinct, elevation ; g-aleal seta distal to the midpoint of the 
finofer; serrula exterior with about 18 teeth; flapellnm of 8 uni- 
laterally branehed setae. 

Palps lon<i- and slender ; surfaces smooth ; setae lonp: and acumi- 
nate ; general proportions of ]iodomeres shown in Figure 1 ; 
chelal hand rather long and narrow, fingers long and gently 
curved; tactile setae as shown in Figure 2; fixed finger with a 
marginal row of alternating large and small teeth — 28 (28-29) 
widely-spaced large, acutely triangular teeth and 20 (20-22) 
small, low, triangular teeth between ; movable finger distally with 
teeth similar to those on fixed finger, 17 (16-21) large and 16 
(14-18) small, while proximally the teeth abruptly become low, 
rounded and contiguous, 11 (9-11) in number; proximal end of 
the movable finger with a long cylindrical process from its dorso- 
median border, parallel with the long axis of the finger and ex- 
tending well beyond the proximo-ventral margin (this projection 
apparently serves for broader attachment of the muscles which 
move the finger and provides increased leverage). Trochanter 
1.9 (1.7-2.0)! femur 7.2 (7.1-7.2), tibia 2.5 (2.4-2.7), chela 7.6 
(7.6-8.0) and hand 3.0 (2.8-3.1) times as long as broad; movable 
finger 1.69 (1.61-1.69) times as long as hand. 

Legs generally of chthoniid form but elongate and slender. 
First leg with coxa bearing spines as characteristic of the genus, 
5 (7) on the right coxa and 6 (6-8) on the left; fourth leg with 
tactile setae on tibia 0.40 (0.39-0.41), on metatarsus 0.24 (0.23- 
0.28), and on telotarsus 0.16 (0.21-0.26) the length of the seg- 
ment from the proximal end. 

Genitalia essentially as figured by Chamberlin (1931, fig. 50E) 
for K. croshyi, and Vachon (1952, fig. 2) for K. henroti; about 15 
setae on the anterior operculum ; 10-12 setae on each side of the 
aperture; 7-8 setae on the posterior operculum between the 
spiracles. 

Female: (The first measurements given are those for the fe- 
male from Hagen's type series, which is designated the allotype, 
Avhile in parentheses are given those for the specimen from 
"Wyandotte New Cave.") The female is essentially similar to 
the male except for slightly larger size. Carapace with a total 
of 16 (17) setae, of which 4 are on the anterior margin and 
2 (3) at the posterior margin; no eyes or eye spots. Abdominal 



4 BREVIORA No. 188 

tergites with 2 :2 :2 .-S A A :5 :6 :6 :2 (2 :2 :3 :4 :4 :6 :4 :6 :6 :4) setae ; 
sternites with 6-10 setae. 

Chelicera slightly larger than in the male ; galea somewhat 
larger and more distinct than in the male, thongh not so sharply 
set off from the finger as indicated by Packard (1888, fig. 12d). 

Palps like those of the male but slightly larger ; fixed finger 
A\dth 29 (29) large and 21 (20) small teeth; movable finger with 
20 (18) large and 16 (14) small teeth distally and 10 (11) low, 
rounded teeth proximally; trochanter 1.8 (1.9), femur 7.0 (6.7), 
tibia 2.6 (2.5), chela 7.6 (7.1) and hand 2.9 (2.7) times as long 
as broad; movable finger 1.67 (1.64) times as long as hand. 

Legs as in the male but slightly larger. Coxal spines 7 (6) 
on the right coxa and 7 (6) on the left. Leg IV with tactile setae 
on tibia 0.42 (0.40), on metatarsus 0.22 (0.24) and on telotarsus 
0.20 the length of the segment from the proximal margin. 

Genitalia essentially as figured by Chamberlin (1931, fig. 52 A) 
for Chthoiiiiis ischnochcles but Avith setae of anterior operculum 
grouped more toward the midline and the row of setae on the 
posterior operculum extending farther lateral on each side ; an- 
terior operculum with 10 irregularly grouped setae ; posterior 
operculum with a row of 6 (8) setae between the spiracles. 

Measurements (in mm.) : Male lectotype and paratypes (range 
of latter in parentheses) : Body length 1.97 (1.79-1.89) ; carapace 
0.60 (0.55-0.58) long, greatest width 0.57 (0.55-0.57); alidomen 
0.73 (0.80-0.88) broad. Chelicera 0.45 (0.45-0.47) by 0.22 (0.20- 
0.23). Palpal trochanter 0.26 (0.23-0.27) ])y 0.14 (0.13-0.14); 
femur 0.90 (0.90-0.93) by 0.12 (0.12-0.13) ; tibia 0.34 (0.33-0.34) 
by 0.14 (0.12-0.14) ; chela 1.32 (1.29-1.33) by 0.18 (0.16-0.18) ; 
hand 0.50 (0.50-0.51) by 0.17 (0.16-0.18) ; movable finger 0.84 
(0.83-0.84) long; proximal process of movable finger 0.033 (0.039- 
0.043) long. Leg I: basifemur 0.56 (0.55-0.57) bv 0.08 (0.08); 
telofemur 0.23 (0.23-0.25) by 0.07 (0.07) ; tibia 0.28 (0.29-0.31) 
by 0.06 (0.06) ; tarsus 0.59 (0.58-0.60) by 0.05 (0.05-0.06). Leg 
IV: entire femur 0.77 (0.76-0.80) long; basifemur 0.28 (0.30- 
0.31) by 0.21 (0.18-0.23) ; telofemur 0.55 (0.53-0.57) by 0.19 
(0.16-0.20) ; tibia 0.51 (0.51-0.54) by 0.10 (0.09-0.10) ; metatar- 
sus 0.25 (0.25-0.26) by 0.07 (0.07) ; telotarsus 0.66 (0.64-0.66) by 
0.06 (0.05-0.06). 

Female allotype and paratype (latter in parentheses) : Body 
length 1.94 (1.89) ; carapace 0.59 (0.61) long, greatest width 
0.61 (0.60); abdomen 0.88 (0.84) broad; chelicera 0.50 (0.48) 
by 0.23 (0.22). Palpal femur 0.96 (0.98) by 0.14 (0.15) ; tibia 



1963 CAVERNICOLOUS PSEUDOSCORPIONS 5 

0.3G (0.87) by O.U (0.15); chela 1.39 (1.40) by 0.18 (0.20); 
luuulO.53 (0.54) by 0.18 (0.20). Movable finger 0.88 (0.88) long; 
])roxiinal process of movable finger 0.045 (0.043) long. Leg I: 
basifenuir 0.57 (0.57) by 0.08 (0.09) ; telofemur 0.24 (0.24) by 
0.07 (0.08); tibia 0.31 (0.31) by 0.07 (0.07) and tarsus 0.58 
(0.61). Leg IV: entire femur 0.81 (0.82) long; basifemur 0.30 
(0.32) by 0.22 (0.23) ; telofemur 0.58 (0.58) by 0.20 (0.21) ; 
tibia 0.54 (0.56) by 0.10 (0.10) ; metatarsus 0.26 (0.25) by 0.07 
(0.08) ; telotarsus 6.66 by 0.06. 

Remarks: This species is restricted to the type locality as far 
as is known at present. The specimens from Mammoth Cave, Ky., 
and vicinity recorded as Chthonius packardi by Hagen, Packard, 
and others, actually belong to Klcptochthonuis ccrherus Malcolm 
and Chamberlin or to K. hagcni n. sp. (see below). This species 
and K. gertschi Malcolm and Chamberlin are unique in the genus 
in the complete absence of eyes or eye spots. These two species 
may be differentiated readily by the number of setae of the 
cheliceral hand (7 in packardi, 9 in gertschi) and the number 
of setae at the posterior border of the carapace (2 in packardi, 
6 in gertschi). 

Kleptochthonius (Chamberlinochthonius) hageni 

new species 

Material: Holotype male and allotype "with a dead bat" at 
bottom of Mammoth Dome, Edmonson Co., Ky., collected Sept. 
11, 1874 by F. W. Putnam. ^lale paratype from Mammoth Cave, 
Edmonson Co., Ky. (no other data). Female paratypes from 
Mannnoth Cave collected by A. S. Packard (?), Apr! 30, 1874, 
and by L. llubricht. Dee. 15, 1956, and from Long Cave, Glasgow 
Junction, Ky., collected by F. G. Sanborn, Nov. 5, 1874. 

Diagnosis: Male: (Measurements are given first for the holo- 
type, while in parentheses are given those for the male paratype.) 
Body and appendages generally of chthoniid facies but more 
slender than in epigean species; lightly sclerotized and pale 
brown in color, the chelicerae and palps somewhat darker than 
the body and legs. Carapace nearly square in dorsal outline, the 
lateral margins a little convex ; no epistome ; 2 eyes present in the 
anterior position with moderately well-developed corneas; no 
trace of posterior eyes or eye spots; total setae 20, of which 6 
are on the anterior margin and 4 are situated near the posterior 
margin; surface generally smooth, but lightly sculptured on the 
sides. Abdomen elongate ; tergal and sternal scuta entire and 



6 BREVIORA No. 188 

smooth ; pleural membranes weakly marked with fine granula- 
tions. Tergites of holotype with 2 :2 :2 :3 :5 :4 :6 :6 :7 :7 long acumi- 
nate setae, paratj^pe with 2 :3 A A :5 :5 :6 :6 :8 :6 setae ; sternites 4-8 
with 8-11 acuminate setae, shorter and more slender than those 
of the dorsum. 

Chelicera of chthoiuid fades ; hand with 7 setae ; fixed finger 
with a row of 8-10 teeth, the movable finger with a row of 5-6 
teeth, the teeth of each finger being generally heavy and triangu- 
lar but becoming progressively smaller toward the proximal end 
of the row ; no isolated sub-distal tooth on either finger ; the galea 
represented by a low elevation on the movable finger ; galeal seta 
distal to the midpoint of the finger ; serrula exterior with about 
18 teeth ; flagellum consisting of 8 unilaterally branched setae. 

Palps long and slender ; surfaces smooth ; setae long and acumi- 
nate ; general proportions of podom.eres shown in Figure 3 ; tac- 
tile setae of chela as shown in Figure 4; fixed finger of chela 
with a marginal row of alternating large and small teeth : 44 (39) 
large, acutely-pointed, widely-spaced teeth with 20 (18) tiny, 
rounded teeth between. Movable finger distally with teeth simi- 
lar to those on the fixed finger, 30 (29) large teeth and 20 (10) 
tiny teeth between, while proximally the teeth abruptly become 
low, rounded and contiguous, 7 (7) in number; proximal end of 
the movable finger with onh' a short, broad projection from the 
dorso-median border, not extending beyond the proximo-ventral 
margin. Trochanter 2.0 (2.2), femur 6.9 (6.4), tibia 2.3 (2.1), 
chela 7.1 (6.8), and hand 2.8 (2.6) times as long as broad; mov- 
able finger 1.62 (1.66) times as long as the hand. 

Legs generally of chthoniid facies but elongate and slender. 
First leg with coxa bearing spines as characteristic of the genus, 

7 (8) on the right and 8 (8) on the left coxa; fourth leg with 
tactile setae on tibia 0.38 (0.38), on metatarsus 0.24 (0.23) and 
on telotarsus 0.26 (0.20) the length of the segment from the 
proximal end. 

Genitalia essentially as in other species of the genus ; with 
12-15 setae on the anterior operculum; 7-12 setae on each side 
of the aperture ; and 6-9 setae on the posterior margin of the 4th 
sternite between the spiracles. 

Female: (The first measurements given are those for the allo- 
type, while in parentheses are given the ranges for the three 
paratype females.) The female is essentially similar to the male. 
Carapace with a total of 20 (19-20) setae, of which 6 (5-6) are 
at the anterior margin and 4 (3-4) near the posterior margin; 



]9G3 CAVERNICOLOUS PSEUDOSCOKPIONS 7 

2 eyes in the anterior position with moderately well developed 
eorneas; no trace of eyes or eye spots at the posterior position. 
Abdominal terg-ites Avith 2:2 :3 :4:4:6 :6 :6 :? : ? :(2 :2 :2-4 :3-4:4:5- 
() :6 :(j-7 :6-7 :4-6) setae ; sternites with 8-12 setae. 

Chelieera similar to that of the male; 7 (7) setae on the hand; 
galeal tnberele somewhat larger and more distinct than that of 
the male. Palps as in the male but somewhat larger and heavier. 
Fixed finger with 40 (37-39) large and 19 (15-17) small teeth; 
movable finger with 28 (25-27) large and 15 (15-17) small teeth 
distally and 7 (7-9) low, rounded teeth proximally. Trochanter 
2.0 (1.9-2.0), femur 6.7 (6.3-6.6), tibia 2.3 (2.3), chela 6.7 (6.2- 
6.7) and hand 2.7 (2.4-2.7) times as long as broad; movable 
finger 1.63 (1.59-1.64) times as long as hand. 

Legs as in the male ; coxal spines 6 (7-9) on the right coxa and 
7 (6-9) on the left. Leg IV with tactile setae on tibia 0.37 (0.29- 
0.36), on metatarsus 0.27 (0.26-0.29) and on telotarsus 0.28 
(0.23-0.27) the length of the segment from the proximal margin. 

Genitalia essentially as in K. packardi; anterior operculum 
with 9 (8-9) grouped setae; posterior operculum with a mar- 
ginal row of 8 (7-8) setae between the spiracles. 

Measurements (in mm.) : Male holotype and paratype (latter 
in parentheses) : Body length 2.14 (2.08) ; carapace 0.65 (0.62) 
long, greatest width 0.64; abdomen 0.84 (0.95) broad. Chelieera 
0.56 (0.53) by 0.26 (0.25). Palpal trochanter 0.31 (0.31) by 
0.16 (0.15) ; femur 1.06 (0.99) by 0.15 (0.15) ; tibia 0.41 (0.37) 
by 0.18 (0.18) ; chela 1.50 (1.45) by 0.21 (0.21) ; movable finger 
0.96 (0.92) long; proximal process of movable finger 0.020 
(0.016) long. Leg I: basifemur 0.63 (0.60) by 0.09 (0.09); 
telofemur 0.25 (0.25) by 0.08 (0.08) ; tibia 0.32 (0.33) by 0.07 
(0.07) ; tarsus 0.64 (0.62) by 0.06 (0.06). Leg IV: entire femur 
0.93 (0.86) long ; basifemur 0.36 (0.34) by 0.28 (0.25) ; telofemur 
0.66 (0.64) by 0.25 (0.23) ; tibia 0.60 (0.58) by 0.11 (0.11) ; 
metatarsus 0.31 (0.29) by 0.09 (0.09) ; telotarsus 0.74 (0.69) by 
0.06 (0.06). 

Female allotype and three paratypes (ranges of latter in paren- 
theses) : Body length 2.20 (2.30-2.55) ; carapace 0.60 (0.62-0.67) 
long, greatest width 0.60 (0.58-0.71) ; abdomen 0.95 (0.80-1.09) 
broad. Chelieera 0.56 (0.58-0.64) by 0.26 (0.27-0.29). Palpal 
femur 1.03 (1.05-1.18) by 0.15 (0.17-0.18) ; tibia 0.40 (0.42- 
0.46) by 0.18 (0.18-0.20) ; chela 1.48 (1.53-1.72) by 0.22 (0.23- 
0.28) ; hand 0.58 (0.58-0.67) by 0.21 (0.22-0.28) ; movable finger 
0.94 (0.94-1.07) long; proximal process of movable finger 0.019 



8 BREVIORA No. 188 

(0.017-0.025). Leg I: basifemur 0.60 (0.64-0.72) by 0.09 (0.09- 
0.10) ; telofemur 0.26 (0.26-0.31) by 0.08 (0.08-0.09) ; tibia 0.31 
(0.31-0.36) by 0.07 (0.07) ; tarsus 0.62 (0.64-0.72) by 0.06 (0.06- 
0.07). Leg iV: entire femur 0.87 (0.94-1.00) long; basifemur 
0.34 (0.36-0.37) by 0.25 (0.26-0.34); telofemur 0.62 (0.66-0.75) 
by 0.23 (0.23-0.30) ; tibia 0.57 (0.62-0.69) by 0.11 (0.11-0.13) ; 
metatarsus 0.30 (0.31-0.37) by 0.08 (0.09-0.10) ; telotarsus 0.72 
(0.71-0.83) by 0.06 (0.06-0.07). 

A deutonymph is at hand wliieli probably pertains to this 
species. It was collected in Audubon Avenue, Mammoth Cave, 
Kentucky, by L. Hubricht on Dec. 15, 1956. It is of interest to 
note some points of difference evidently associated with imma- 
turity. The appendages are all considerably stouter than those 
of the adults. Only 18 setae are present on the carapace, with 6 
at the anterior edge and 2 at the posterior. Two coxal spines 
are present on each coxa I and the discal seta is wanting. The 
galeal tubercle is small but distinct (as in the adult). There are 
only 6 blades in the flagellum and only 5 setae on the cheliceral 
hand. The movable finger of the chela has only two setae (prob- 
ably st and sh), while the fixed finger is lacking ist and one 
seta from the dorsum of the hand. 

Remarks: This material from Mammoth Cave was considered 
by Ilagen, Packard, and other early workers to be conspecific 
with K. packardi from Wyandotte Cave, Indiana, although they 
remarked the presence of eyes in the former and their absence 
in the latter. Later workers have been sceptical of this rela- 
tionship (cf. Hoff, 1958, p. 4; Chamberlin and Malcolm, 1960, p. 
Ill), but only recently has material become available to clarify 
the situation. Restudy of the type material has made obvious 
the differences between the Indiana specimens and those de- 
scribed here as K. hageyii. K. liagcni can be distinguished from 
K. Cerberus Malcolm and Chamberlin, which it closely resembles, 
by the pcssession of small, but distinct, microdenticles between 
the macrodenticles of both fixed and movable chelal fingers, by 
the possession of a small but distinct galeal tubercle, and by the 
smooth inner edge of the movable cheliceral finger distal to the 
row of teeth. 

In addition there are a number of other specimens from Mam- 
moth Cave and vicinity, collected by Packard and others, and 
moi'c recently by T. C. Barr, Jr., L. Ilubricht, and C. Krekeler, 
which ditt'er from the described forms in certain details, particu- 
larly number of eyes, chaetotaxy, and proportions of the proximal 



1963 CAVERNICOLOUS PSEUDOSCORPIONS 9 

l)roeess of the movable chelal finger. The relations of these forms 
to K. hageni and to the cave species recently described by Mal- 
colm and Chamberlin (1961) will be treated in a later paper. 

This species is named in honor of II. A. Hagen who initiated 
the study of cave pseudoscorpions in the United States. 

Genus APOCHTHONIUS Chamberlin 

Apochthonius coecus (Packard), new combination 

Chthonius coecus Packard, 1884, Amer. Nat., vol. IS, p. 203; 1888, Mem. 

Natl. Acad. Sci. Washington, vol. 4, p. 46. 
Chthoniufi? coecus: Beier, 1932, Das Tierreieh, vol. 57, p. 61. 
Chiltonius? coccus: Hoff, 1958, Amer. Mas. Novitates, no. 1875, p. 4. 

Material: The two type specimens in the collection of the 
M.C.Z. have been mounted on slides for study. They are a male, 
which has been designated the lectotype, and a female, the allo- 
type. Both were collected by Packard in Weyer's Cave (Grand 
Caverns), Augusta Co., Virginia, in 1874. The descriptions of 
the species by Packard (1884, 1888) are generally accurate, but 
a detailed description based on the mounted specimens is de- 
sirable. It is obvious that the species belongs in Apochthonius. 

Diagnosis: Male lectotype: Body and appendages typical of 
the genus; very lightly sclerotized and pale brownish in color, 
the chelicerae and palps being slightly darker than the body and 
legs. Carapace slightly broader than long, being slightly nar- 
rower tow^ard the posterior end ; no epistome ; no eyes or eye 
spots; total setae 22, of which 8 are at the anterior margin and 
4 near the posterior margin ; surface smooth. Abdomen longer 
than broad and smoothly rounded in outline; tergal and sternal 
scuta entire and smooth ; pleural membranes very faintly, longi- 
tudinally striate. Tergal chaetotaxy 4 :4 :5 -A^ :6 :7 :7 :8 :8 :6 ; ster- 
nites with 8-10 setae. 

Chelicera large and heavy, typical of the genus; hand with 7 
setae ; fixed finger with a row of 7 triangular teeth which become 
progressively smaller toward the base ; movable finger with a row" 
of 7-8 similar teeth and an additional tooth situated half way 
between the distal end of the row and the tip of the finger; galea 
represented by a low but distinct elevation. 

Left palp of typical facies (right palp missing) ; general pro- 
portions of podomeres shown in Figure 5 ; tactile setae of chela 
as shown in Figure 6 ; fixed finger with a marginal row of 72 
nearly contiguous teeth, which are tall and rectangular at the 



10 BREVIORA No. 188 

distal end of the row, becoming low and broadly rounded toward 
the proximal end ; movable finger with 71 similar teeth. Trochan- 
ter 1.8, femur 4.5, tibia 1.9, chela 5.5, and hand 1.7 times as 
long as broad ; movable finger 2.27 times as long as the hand. 

Legs of typical facies. First leg with coxa bearing spines as 
characteristic of the genus, with 2 spines on the right coxa and 
3 on the left. Fourth leg with tactile setae on tibia 0.46, on 
metatarsus 0.25, and on telotarsus 0.20 the length of the segment 
from the proximal margin. 

Genitalia as in other members of the genus ; anterior operculum 
with an irregular group of 6 setae anteriorly and a row of 9 
setae along the posterior margin; posterior operculum with 7 
setae on each side of the aperture and a marginal row of 8 setae 
between the spiracles. 

Female allotype: Essentially similar to the male. Carapace 
wath a total of 22 setae of which 8 are at the anterior margin 
and 4 near the posterior border ; no eyes or eye spots. Abdominal 
tergites with 4 A :6 :6 :6 :7 :7 :8 : ? : ? : setae. 

Right chelicera missing; left apparently similar to that of 
male. 

Right palp missing ; left palp similar to that of male. Fixed 
chelal finger with 72 teeth and movable finger with 70 teeth. 
Trochanter 1.9; femur 4.8; tibia 1.8; chela. 4. 9 and hand 1.6 
times as long as broad ; movable finger 2.12 times as long as 
hand. Legs as in tlie male. Leg I with 3 spines on each coxa. 
Leg IV with tactile setae on tibia 0.46, on metatarsus 0.31, and 
on telotarsus 0.24 the length of the segment from tlie proxi- 
mal margin. 

Genitalia essentially the same as for A', jjackardi; anterior 
operculum with 8 grouped setae ; posterior operculum with 7 
setae in a row between the spiracles. 

Measurements (in mm.) : Male lectotype and female allotype 
(latter in parentheses) : Body length 1.13 (1.07) ; carapace 0.40 
(0.41) long, greatest width 0.44 (0.44); abdomen 0.51 (0.51) 
broad. Chelicera 0.36 (0.38) by 0.18 (0.18). Palpal trochanter 
0.18 (0.20) by 0.10 (0.10) ; femur 0.43 (0.46) by 0.10 (0.10) ; 
tibia 0.22 (0.23) by 0.12 (0.12); chela 0.68 (0.72) by 0.12 
(0.15) ; hand 0.21 (0.23) by 0.12 (0.15) ; movable finger 0.48 
(0.50) long. Leg I (missing in female) : basifemur 0.24 by 0.05; 
telofemur 0.12 by 0.05; tibia 0.14 by 0.04; tarsus 0.26 by 0.04. 
Leg IV: entire femur 0.35 (0.36) long; basifemur 0.17 (0.17) 
by 0.15 (0.15) ; telofemur 0.23 (0.23) by 0.12 (0.13) ; tibia 0.26 



1963 CAVERNICOLOUS PSEUDOSCORPIONS 11 

(0.27) by 0.07 (0.07); metatarsus 0.12 (0.12) by 0.05 (0.05); 
telotarsus 0.26 (0.25) by 0.04 (0.04). 

Remarks: This is the first species of Apochthoymis known to 
be cavernicolous. Unlike the trog'lobious species of Klcptochtho- 
nius it is not markedly modified for life in caves. It is generally 
similar to the epigean species, A. moestus. From this it differs 
in the following respects : no trace of eyes ; derm less heavily 
sclerotized and lighter in color ; slightly more slender palpal 
podomeres; slightly reduced chaetotaxy of the carapace and the 
abdominal tergites. 

It is pertinent at this point to mention that we have at hand 
a tritonymph collected in Madison Cave, Augusta Co., Virginia, 
Aug. 28, 1958 by T. C. Barr, Jr. This specimen is perhaps re- 
ferable to A. coecus, since Madison Cave is close to Weyer's cave. 
It is similar in most details to the type adults, but with numbers 
and measurements somewhat reduced as would be expected of a 
tritonymph. Unexpectedly, however, this individual clearly pos- 
sesses two eyes in the anterior position, weakly but definitely 
corneate. The explanation of this situation is not certain. It 
may be that this nymph belongs to a separate, new species. Or 
it may be that the species, A. coecus, may contain individuals 
both with and without eyes. A third possibility is that nymphs 
of A. coecus possess eyes which are then lost at the adult molt 
(as is suggested for Chitrclla archeri below). Only study of 
larger, as yet uncollected, series of specimens can throw further 
light on this problem. 

Suborder DIPLOSPHYRONIDA Chamberlin 
Family SYARINIDAE Chamberlin 
Subfamily CHITRELLINAE Beier 

Oenus ChITRELLA Beier 

Chitrella cavicola (Packard), new combination. 
Obisium cavicola Packard, 1884, Amer. Nat., vol. 18, p. 202. 
Obisium cavicola: Packard, 1888, Mem. Nutl. Acad. Sci. Washington, vol. 

4, p. 42. 
Microcreagris? cavicola: Beier, 1932, Das Tierreicli, vol. 57, p. 157. 
Microcreagris? cavicola: Iloff, 1958, Amer. Mus. Novitates, No. 1875, p. 12. 

The specimen upon which Packard based the species Obisium 
cavicola was preserved in the M.C.Z. with a label bearing only 
the notation "New Market Cave, Packard." There can be no 



12 BREVIORA No. 188 

doubt that it is the specimen in question, inasmuch as it retained 
the peculiar pose of the appendages and outline of body as de- 
picted in Packard's illustrations (1884, fig. 1, and 1888, fig. 11). 
The figure and description are, however, at variance with the 
specimen in two major respects: 1) there is no cleft in the 
carapace, which is, indeed, perfectly normal ; and 2) there are 
actually four eyes present with distinct, albeit weak, corneas. 
In the alcoholic specimen the illusion of a cleft carapace was 
given at first glance, due to the transparency of the carapace and 
the converging medial borders of the rather opaque cheliceral 
muscles. It is difficult to understand how the eyes could have 
been overlooked since they were readily visible in the alcoholic 
specimen. The specimen was mounted on a slide for detailed 
study. It was revealed to be a tritonymph of a species belonging 
in the genus CJiitrclla Beier. It is regrettable but necessary to 
base the description of the species upon this single, immature 
specimen. 

Diagnosis: Tritonymph holotype : Body and appendages of 
typical chitrelline facies (cf. Malcolm and Chamberlin, 1960, fig. 
4) ; very lightly sclerotized and pale in color ; carapace slightly 
longer than broad, with its greatest breadth at about the middle ; 
no epistome ; four eyes weakly but definitely corneate ; total setae 
28, of which 6 are on the anterior margin and 6 at the posterior 
margin. Abdomen elongate; tergal and sternal scuta entire; 
pleui-al membranes longitudinally striate. Tergal chaetotaxy: 
9:11:11:11:12:1:3:13:11:8:?. Sternal chaetotaxy 3 :(3)8(3) :(3) 

2m 
8(3) :12:—— : 15:14:12:12:?, the two microsetae of sternite 6 

lying side by side at the center of the scutum. 

Chelicera typical, about twice as long as broad ; chaetotaxy 
normal, with 5 setae on hand and a galeal seta; flagellum ap- 
parently consisting of 6 or 7 blades; fixed finger with 16-18 
triangular teeth, largest in the middle of the row ; movable finger 
with 8-10 irregular teeth, very heavy distally but smaller toward 
the base of the finger; galea absent. 

Palps typical, showing no special adaptation to subterranean 
life ; general proportions of podomeres as shown in Figure 7. 
Tactile setae of chela as shown in Figure 8 ; seta ish absent from 
the fixed finger and sh from the movable finger ; fixed finger with 
a marginal row of 42 contiguous teeth, quadrangular in the dis- 
tal third but low and rounded in the basal two-thirds of the row; 



1963 CAVERNirOLOMS PSEUDOSCOKI'IONS 13 

moTable finger with 49 eontigiioiis teeth, all low, broad and some- 
what rounded. Trochanter 2.1 ; femur 3.8 ; tibia 2.4 ; chela 3.fi, 
and hand 1.5 times as long as broad; movable finger L52 times 
as long as hand. 

Legs generally tyi^ieal of the germs but with the telotarsus of 
each leg swollen subbasally so that its depth is equal to or greater 
than that of the metatarsus. Leg IV with a tactile seta on the 
tibia 0.67 the length of the segment from the proximal margin ; 
apparently no tactile seta present on the metatarsus. 

Measurements (in mm.) : Tritonymph holotype : Body length 
1.90; carapace 0.48 long, greatest width 0.45; abdomen 0.81 
broad. Chelicera 0.29 by 0.15. Palps: trochanter 0.26 by 0.12; 
femur 0.44 by 0.12; tibia 0.37 by 0.15; chela 0.72 (0.77 with 
pedicel) by 0.20; hand 0.30 by 0.19; movable finger 0.45 long. 
Leg I : basifemur 0.22 by 0.07 ; telofemur 0.15 by 0.07 ; tibia 0.20 
by 0.05; metatarsus 0.11 by 0.044; telotarsus 0.15 by 0.047. Leg 
IV: entire femur 0.38 long; basifemur 0.15 by 0.13; telofemur 
0.23 by 0.14; tibia 0.31 by 0.07; metatarsus 0.13 by 0.058; telo- 
tarsus" 0.19 by 0.058. 

Remarks: Though this species is presently known only from 
a single tritonymph some idea of the adult form may be surmised 
from a comparison with C. archeri Malcolm and Chamberlin, of 
which 3 tritonymphs have been studied (unpublished). In most 
respects the tritonymph of C. cavicola is similar to those of C. 
archeri, but it is a little smaller and slightly less slender both 
in body and in the appendages. It has fewer teeth on the chelal 
fingers (fixed finger 42 and movable finger 49 as compared with 
49-54 and 56-58, respectively) ; it has 9 setae ou the first tergal 
scutum as compared with 4-6 ; and tactile seta t of the movable 
chelal finger is slightly closer to the tip of the finger (index 
0.328 as compared with 0.361-0.370). On the other hand, it 
shares with C. archeri tritonymphs the possession of two micro- 
setae on the sixth sternite ; the subbasal swelling of the telotarsi 
(unpublished observation; cf. also Vachon, 1954, p. 219); and 
(with one specimen) the lack of a tactile seta on metatarsus IV. 
Furthermore, it should be mentioned here that, while the adults 
of C. archeri are without demonstrable eyes, the tritonymphs 
possess definite, though weakly developed, corneas corresponding 
to, but weaker than, the anterior eyes of C. cavicola. 

From the foregoing it seems warranted to assume that the 
adult of C. cavicola is similar to that of C. archeri, but slightly 
smaller and more robust and differing in details of chaetotaxy, 



14 BREVIORA No. 188 

etc., and perhaps very close to the adult of C. muesebecki Mal- 
colm and Chamberlin. 

EEFEEENCES 

Chamberlin, J. C. 

1931. The ;ir,R-hni(l ordfr Clielonethida. Stanford Univ. Pubis. Biol. 
Sci., 7: 284 pp. 
Chamberlin, J. C. and D. E. Malcolm 

1960. The occurrence of false scorpions in caves witli special reference 
to cavernicolous adaptation and to cave species in the North 
American fauna (Araclmida — Chelonethida). Amer. Midi. Nat., 
64: 105-115. 
HoFP, C. C. 

1958. List of the pseudoscorpions of North America north of Mexico. 
Anier. Mus. Novitates, no. 1875, 1-50. 
Malcolm, D. E. and J. C. Chamberlin 

1960. The pseudoscorpion genus ChiircUa (Chelonethida, Syarinidae). 
Amer. Mus. Novitates, no. 1989, 1-19. 

1961. Tlie pseudoscorpion genus KleiHochtJwnius Chamberlin (Clielone- 
thida, Chthoniidae). Amer. Mus. Novitates, no. 2063, 1-35. 

Packard, A. S. 

1884. New cave arachnids. Amer. Naturalist, 18 : 2012-204. 

1888. The cave fauna of North America, with remarks on the nnatoniy 
of the brain and origin of the blind species. Mem. Natl. Acad. 
Sci. Washington, 4 : 3-1.16. 
Vachon, M. 

1952. A propos d 'on pseudoscorpion cavernicole. Notes Biospeo- 
logiques, 7: 105-112. 

1954. Eemarques morphologiques et anatomiques sur les pseudoscor- 
pions (Arachnides) appartenant au genre Psciidoblothnas 
(Beier) (Earn. Syarinidae J.C.C). Bull. Mus. Nat. Hist. Nat- 
urelle, Paris, (2° ser.) 26: 212-219. 



1963 



CAVERNICOLOUS PSEUDOSCORPIONS 



15 




Figures 1 and 2. KleptochtJionius {Chaviberlinochthonms) pacTcardi (Ha- 
gen), allotype female. 1. Dorsal view of right palp. 2. Lateral view of left 
chela (setae missing from specimen). 

Figures 3 and 4. Eleptochthonius (Chamherlinochthonius) hageni, new 
species, holotype male. 3. Dorsal view of left palp. 4. Lateral view of right 
chela. 



16 



BREVIORA 



No. 188 




Figures 5 and (5. ApochtJioniiis coccus (Packard), lectotype male. 5. 
Dorsal view of left palp. 6. Lateral view of left chela (setae missing from 
specimen and teeth omitted). 

Figures 7 and 8. ChitrcUa cavicola (Packard), holotype tritonymph. 7. 
Dorsal view of right palp. 8. Lateral view of left chela (setae missing from 
specimen and teeth omitted). 



BREVIORA 

Miaseiiam of Comparaitive Zoology 

Cambridge. Mass. Arorsx 2!). 1963 Number 1S9 

NOTES ON AMPHISBAENIDS ( AMPIIISBAENTA ; REP- 

TILIA). 10. REDESCKTPTION AND REDEFINITION 

OF AMPHISBAENA PERiri^JXSlS NOBLE FRO]\I THE 

MOUNTAINS OF NORTHWESTERN PERU. 

By Carl Cans 

Department of Biology 

State University of New York 

at Buffalo, Buffalo 14, New York 

In 1921 Noble described the new species Amphishaena peri- 
censis, basing his description upon a single specimen (M.C.Z. 
No. 14631/ collected by him, wliile a member of the "Harvard 
Peruvian Expedition of 1916." The holotype came from "Perico, 
Pern," l)ut Noble added that the species was represented in the 
collection by a "large series of specimens." He neither listed, 
nor otherwise commented upon these, except to state the range 
of the form as the ''arid valleys of the Chinchipe and Marafion, 
from Perico on the north to Bellavista on the south." 

The series of specimens referred to has never been discussed 
in the subsequent literature. The only reference to additional 
specimens was made in error. S.U. 8314-8315 (Burt and Myers, 
1942, p. 48) actually were A. occidcntalis townsendi from the 
Pariiias Valley. Our conce])t of A. pfvicensis thus still rests 
u])()n the description of the holotype. Half of the remaining 
specimens, referred to as paratypes with but faint justification, 
have been traded to at least eight other museums. Tlie form has 
had repeated mention in catalogs and specimen lists (Barliour 
and Loveridge, 1929 ; Burt and Burt, 1930, 1931 ; Cochran, 1961 ; 
Marx, 1958). 

The present notes are based upon the re-assembly of a signifi- 
cant fraction of the original series. They include a redefinition 
and standardized redescription ( cf . Gans and Alexander, 1962), 
coupled with the first illustration of the form. 



BREVIORA 



No. 189 



I am indebted to the following curators of institutions (identi- 
fied throughout by the abbreviations given in parentheses) for 
permission to borrow or examine material in their care : Mr. 
Charles M. Bogert of the American Museum of Natural Historv 




Fig. 1. Amphishnrrui pcricciisis. Mjip showiii}^ lot-alitics iiu'iitioiu'd in tlip 
text. Tlie two small dots in the small scale map indicate the ty])(> localities 
of AmpMnhaena o. occiflentalis Cope [bottom or south], and ./. <>. toirnfiendi 
Stejneger Ilo]' or iioitli|. 'Die dottrd line on llic lar^c scale |dctail| map 
indicates the I'rovince of Jacn of the Department of Cajamaii-a, I'eni. 



1963 AMPHISBAENA PERICENSIS 3 

(A.M. X. II.), :\Iiss Alice (i. (\ (iraiulisoii of the I'.ritish :\Ins(Mnn 
(Natural History) (B.M.), Dr. Alan Lcviton of the California 
Academy of Sciences at San Francisco (C.A.S.), Mr. Hymen 
Marx and Dr. Kohert F. Injicr of the Chicago Natural Ilisloiy 
Museum (C.N.II.M.), Dr. Ernest E. Williams of the Museum 
of Comparative Zoology (IM.C.Z.). Dr. Koni-ad Klemmer of the 
Senckenbergische Naturforschende Gesellschaft of Frankfurt a. 
M., Germany (S.M.F.), Dr. George S. Myers of Stanford T'ni- 
versity. Museum of Natural History (S.U.), Dr. Hobart M. 
Smith of the University of Hlinois Museum of Natural Histor^^ 
(U.I.M.N.H.), Dr. Doris M. Cochran of the United States Na- 
tional Museum (U.S.N.M.), and Dr. Heinz Wennuth [formerly] 
of the Zoologisches Institut der Universitat in Berlin (Z.M.U.). 
Dr. Virginia Cummings figured the specimen and ^liss Charlyn 
Rhodes furnished technical assistance. The overall project has 
been supported by grants NSF G-9054 and G-21819 from the 
National Science Foundation. 

Amphisbaena pekicensis Nol)Ie, 1921 

Ampliishaena pericensis Noble, 1921, p. 141. Terra typic-a : "Perieo, Peru." 
HOLOTYPE: M.C.Z. 14(i31. PAEATYPES: A.M.N.H. 28501-03; 
B.M. 1929.6.1. 85-RR 1946.8.31.83; C.A.S. .■:46]4; C.N.H.M. 16106-07, 
73371; B.Z. [ ?] ; M.C.Z. 14764-6.5, 14767-68, 14770, 14772-75; S.M.F. 
11826, 11887-88; U.I.M.N.H. 41494; U.S.N.:M. 75970; Z.M.U. 29659 
[Perieo]. M.C.Z. 14789-90 [Bellavista]. U.R.N. :\t. 59926 [Chiiic-liipe 
River]. U.S.N.:\1. 60057-58 [Marafion River]. 

Status of the types: The status of the various s})ecimeiis col- 
lected by G. K. Noble is thoroughly confused, possibly because 
the collection does not appear to have been catalogued at the 
M.C.Z. in its entirety, and certainly because Noble neither gave 
the total number of specimens examined, nor otherwise identified 
the "paratypes." (See table on p. 13.) 

There does not seem to be any difficulty about the liolutyi)e, 
M.C.Z. 14631. It Avas listed both by Noble and by Barbour and 
Loveridge (1929, p. 215). 

The latter authors also listed M.C.Z. 1476-1-68, 14770, 14772-80 
and 14789-90 as paratypes then remaining in the ^luseum of 
Comparative Zoology. A cheek of the catalog disclosed that tlie 
series originally ran from 14764 to 14790. Of these I have been 
able to identify 14764-65, 14767-80 and 14786-90 in various 
collections, relying always on the presence of one of the original 
parchment labels. Since 14766 is marked "Amaral" in the 



4 BREVIORA No. 189 

M.C.Z. eataloo' (hence presumably in the Departamento cle Zo- 
ohjgia, Sao Paulo collection = D. Z.), only 14781-85 remain in 
question. 

All of the questionable specimens have their catalog entry 
marked " Anomalepis aspinosus Taylor" in A. Loveridge's 
handwi'iting. The astonishing thing is that Dunn's first report 
(1923, p. 185) on the collection mentions only three specimens 
of Anomalepis, while Taylor's original description (1939, p. 92) 
refers to six including 14781-85. Taylor apparently omitted 
mention of two other M.C.Z. specimens of this form, and the 
total of eight once in the M.C.Z. then suggests (1) that Dunn 
never saw M.C.Z. 14781-85, and (2) by implication that Noble 
did not check his ''paratype" series very carefully. Neverthe- 
less, the remaining 17 specimens may well be considered to be 
paratypes. 

An even more complex problem is posed by seven specimens 
also supposedly from the Noble collections; four of these are 
labelled paratype, stem from Perico, and are respectively in 
the American Museum (A.M.N.II. 28501-03) and California 
Academy (C.A.S. 54614) collections. Since Noble was curator 
of the American ^luseum collection at the time they were cata- 
logued, it remains possible that these are indeed part of his mate- 
rial. Less probability attaches to three specimens from the 
''Chinchipe River" (U.S.N.M. 59926) and the "Rio Maranon" 
(U.S.N.M. 60057-58). Stejneger marked the catalog ''topopara- 
type" and these specimens Avere received from the M.C.Z., but 
were never catalogued there. Yet even they may be considered 
to have been included as paratypes by Noble's statement of 
species range. ^ 

Diagnosis: A form of Amphishacna without major fusions of 
head shields ; with pairs of very large first and slightly reduced 
second parietals ; with the head relatively blunt and not par- 
ticularly set off from the neck; with a cylindrical blunt-tipped 
tail ; with a faint autotomy constriction at the sixth to eighth 
caudal annulus where autotomy takes place. The form has 198 
to 218 body annuli ; generally 18 or 19, occasionally 16 or 17 
caudal annuli: 12 to 16 (generally 14) dorsal and 16 to 20 (gen- 
erally 18) ventral segments to a midbody annulus; and 4 small 



1 Two spcoinicns (T'.M.AF.Z. ."."ITCiT A it B) just (liscdvorccl in the (•ipllcctioii of tlip 
Univorsity of Mirlii;,'aii Mnsciiin of Zoology contirm tli;it Xoblc iiippiii'cntly ilis- 
trilmfcd "pariitypcs" witlioiit first {riving tliciii M.C.Z. niinilitTs. 'I'licir jar contains 
an M.C.Z. lalirl reading (aiiimrcntly in Nolilo's liand) ■'Aniidiisliacna sp. nov. in 
nis. — I'prico, N. W. Pern. — Ilarv. IN'rn. Kxp." Ttirlr couiils and nicasurcnionts 
fall within the range of the present des('rii>tion. 



1963 



AMPHISBAENA PERICENSIS 



round preeluacal pores. Tlu' color ol' pi-eser\'i'(l spreimcns is a 
luiiforni dark brown, much darker dorsally than ventrally, light- 
ened on the tip of tail and snout and produced by a dense pig- 
mentation of the segments (contrasted by lighter intersegmental 
sutures). 



PERICO 



" ' 



RIVERS 



BELLAVI STA 



200 208 216 

Body Annuli 



12 14 16 

Max. Dorsals 



Fig. 2. Ampliishaena pericensls. Diagram showing distribution of counts 
of body annuli and maximum dorsal counts of the three groupings collected 
respectively at Perico, along the "rivers" of Chinchipe and Maraiion, and 
at Bella vista. 



Oeographic variation: The entire sample comes from a rela- 
tively restricted area and 24 out of the 29 available specimens 
from the single locality, Perico. Two other specimens bear the 
definite indication Bellavista, while the other three are marked 
only with the names of the river courses traversing the area. 

Number of segments to the dorsal portion of the midbody 
annulus differs clearly between the Perico sample (12 to 14) 
and the Bellavista sample (15 to 16). Two of the river course 
specimens also range to 15. It may also be significant that the 
two Bellavista specimens and one of the river course individuals 
show a slightly higher number of body annuli than does the 
Perico sample (Fig. 2). Here, as in so many other instances, 
we are handicapped by the markedly inadequate samples. 



6 BREVIORA No. 189 

Description: Figure 3 shows views of the head, Figure 4 the 
ventral surface of the cloaca and tail, and 6, 7 and 8 are photo- 
graphs showing coloration and other aspects of the specimens. 
P'igure 5 gives a scatter diagram of tail versus snout-vent lengths. 
Meristie data are listed in the table. 

This is a small species of Amphishacna, of a dark brownish 
color and without pattern, but with marked dorsoventral counter 
shading. The lightened ventral color continues anteriorly onto 
tlie rostral, and caudally to the cloaca and occasionally to the 
distal tip of the tail. The color is clearly restricted to the seg- 
ments, as the intersegmental sutures and the lateral grooves are 
significantly lighter. The specimens thus present the impression 
of a checkerboard under magnification. 

The coh)ration appears to i-esult from two levels of pigmenta- 
tion and two degrees of expression. There is a general darkening 
of the entire segmental surface, plus an additional intensifica- 
tion of pigmentation on tlic i-ectangular segmental centers. The 
dorsal segments of some well jn-eserved specimens liarely show 
tliis midsegmental ])igment dichotomy ; other, possibly more 
faded, s])ecimens show it more clearly though never as markedly 
as in Amphisbatna siU'cstrii (Gans, 1962) in which the margins 
are quite distinct from the centers. The dorsoventral fading 
occurs both by a general weakening of the coloration and by a 
complete dro])ping out of the main jiigmentation by segments. 
(The absence of living siieciinens does not permit decision as to 
whetlKn- these ventral segments are tndy unpigmented, or 
whether the phenomenon liere discussed applies only to those 
pigments least affected by methods of preservation.) The gen- 
eral weakening of the pigmentation may result in disappearance 
or reduction to flecking of the darkening of the entire surface, 
as well as a reduction of pigment density and size of tlie rec- 
tangular segmental centers. The complete depigmentation varies 
in extent from a blanching of most midventral rows to that of 
scattered groups of ventral segments. Such areas as the tail 
are ventrally countershaded by the first phenomenon alone and 
all of their segments retain (more or less faint) dark centers. 

The head segmentation is characterized by lack of major fu- 
sions, and by pairs of very large first and slightly reduced sec- 
ond parietals. The head is slightly flattened dorsoventrally ])ar- 
ticularly on the posterior half. The rostral region projects 
slightly across the tip of the recessed lower jaw. The entire body 
shows some doi-soventral flattening. The tempornl muscles are 



1963 



A.MPllISBAENA PERICENSIS 



scarcely noticeable in juveniles, l)ut are definitely swollen in 
adults in which they change the shape of the head. 

The rostral is slightly smaller than ai'c the first siipralabials 
and only its dorsal tip is visible I'roni above. Pairs of niediuni- 






FiG. 3. AmpJiisbaena pericensis. Dorsal, lateral and voiitial views of the 
head of the holotype, M.C.Z. 14631, from Perico. The line equals 1 mm to 
scale. (V. Cummings del.) 



8 



BREVIORA 



No. 189 



sized nasals, large prefrontals, smaller frontals and wide first 
and narrower second parietals follow in sequence along the dor- 
sal midline of the head. The middle of the parietal lies on the 
level of the angulus oris. There are three supra labials of which 
the second is generally' the largest. The position of the angulus 
oris lies at the posterior edge of this segment and is generally 
quite easy to determine as the intersegmental sutures of the first 
body annulus do not tend to align with the slit of the mouth. 
The suture between the first and second supralabials runs at 
an angle of 45° to the edge of the mouth, while that between the 
second and third runs directly rostracl or only very slightly an- 
teriorly. The ocular is quadrangular, contacting the prefrontals 
and frontals anterodorsally, and the second and third supralabi- 
als anteroventrally. Its po.sterior margins adjoin the segments 
of the first i)odv ainiulus. 




Fit,. 4. A tni>hisb(ii iKi [i< iic( iisis. Yoiitial view uf cloat-a and tail 
li()liitYi)f. Tlie line equals 1 iiini to sc-alo. (V. Ciinimings, del.) 



(if the 



The mental is medium sized and slightly larger than the first 
infralabials which are followed by the very large second infra- 
labials (the largest segments of tlie lower jaw), and these in 
turn by the narrow third infralabials. The oval to pentagonal 
postmental is somewhat larger than the mental and is in contact 
with it, with the first and second pairs of infralabials, and with 
a row of postgenials that clearly exclude it from contact with 
the iiialai-s. The malars are small and fill the spaces between 
the wide second and nari'ow third infralabials. The antei'ior 
row of ])()stgenials generally consists of two tear-droj) shajjcd 
segments, occasionally se]iarated by a small intermediate seg- 
ment (indicated in the table undei- chin segments by a figure 
one in parenthesis). This segment occasioiudly contacts the post- 
mental so that a first row of three postgenials is counted. The 
second postgenial I'ow contains tlwee to foui- segments. Tliei'c 
are no postmalai's. 



1963 



AMl'lllSBAENA PERICENSIS 



Dorsally, the two paii's of very inucli ciilarjjed segments of 
the first body annuliis curve anteriorly and the to|)most pair 
contacts the lateral edges of the frontals. A dorsal half-annuhis 
of one to two seo-nients on each side intercalates behind this, 
including the first parietals. The second parietals are the mid- 



o- 6 
J 3 




16 



E 

^ 14 



C 
0) 



12 



- 10 






8 



• om 



• •• 



7 17 

Snout - Vent 



15 

Length - cm 



Fig. 5. Amphisbaena pericensis. Combined frequency histogram to sliow 
distribution of size classes of snout-vent lengtli (summed in units of 1 cm, 
from 7.0 to 7.9, 8.0 to 8.9, etc.), and scatter diagram of tail versus snout- 
vent length (the hollow dots indicate two coincident points each). 



10 BREVIORA No. 189 

dorsal segments of the second body annulus. The posterior edge 
of this annulus shows no forward curvature, though several of 
its segments are anteriorly elongate, particularly in the tem- 
poral region. 

There are 198 to 218 body annuli from the back of the third 
infralabial, up to and including the pore-bearing precloacals. 
The fourth through sixth or eighth anterior annuli are modified. 
The anterior ones are shorter, and the posterior ones longer 
than the succeeding annuli on the trunk. There is generally no 
complexing of the segments in the "pectoral" region, nor are 
there intercalated dorsal half -annuli. There are 12 to 16 (gen- 
erally 14) dorsal and 16 to 20 (generally 18) ventral segments 
to a midbody annulus, while the normal pattern appears to be 
14/18 (see comments on geographical variation). 

The cloacal region is characterized by 4 round precloacal 
pores, wliich are much less noticeable in juveniles and females. 
There are 6 to 8 precloacal and 11 to 14 postcloacal segments and 
three to five (generally four) lateral rows. The not particularly 
apparent autotomy annulus falls on the sixth to eighth postclo- 
acal annulus and autotomy takes jilace here. Specimens have 
16 to 19 (generally 18 to 19) caudal annuli. The cross section 
of the tail changes from a dorsoventrally flattened ellipse in the 
immediate postcloacal region, to a circle at the autotomy an- 
nulus, to an enlarged vertical ellipse behind this. The enlarged 
distal ti]^ terminates in a blunt vei'tical keel. 

The lateral sulci are clearly marked, starting between the 
twentieth and fortieth body annuli and continuing up to the 
level of the cloaca. At midbody, each of them is represented by 
little more than an elaboration of the normal intersegmental 
suture, which is made more comj^lex by having the extreme cor- 
ners of the adjacent segments diagonally cut off. The dorsal and 
ventral intersegmental sutures are aligned along the middorsal 
and midventral region, but there is no other indication of dorsal 
and ventral sulci. 

The dorsal segments of a midlxidy annulus are approximately 
one and one-half times as long as wide, while the ventral ones 
are one and three-quarter times as wide as long. 

Range: Pern, Depai'tment of C'ajamarca, "arid valleys of the 
Chinchipe and Marafion, from Pci-ico on the north to Bellavista 
on the south." 



1963 AMPHISBAENA PERICENSIS 11 

LTTERATUEE CITED 

Barbotr, Thomas jukI Aktiii r Loveridge 

19-9. Typit'.-il reptiles and anipliiliiaiis. Bull, .\liis. Coiiip. Zool., vol. 
69, no. lU, i)p. 205-360. 
Burt, Charles E. and May DANHEnt Burt 

1930. The Soutli American lizal•d^s in the coUectidn of tlie United 
States National .Miiseiiin. Free. United States Nat. .Mus., vol. 
78, art. G, pp. 1-52. 

1931. South American lizards in the collection of the American Mu- 
seum of Natural History. Bull. Amer. Mus. Nat. Hist., vol. 61, 
art. 7, pp. 227-395. 

Burt, Charles E. and Ceorge S. ^NTyers 

1942. Neotropical lizards in the collection of the natural history mu- 
seum of Stanford University. Stanford Univ. Publ., Univ. ser., 
Biol. Sci., vol. 8, no. 2, pp. 1-52. 
Cochran, Doris Mable 

19(51. Type specimens of reptiles and amphibians in the United States 
National Museum. Bull. United States Nat. Mus., no. 220, pp. 
i-xv-f 1-291. 
Dunn, Em mett Eeid 

1923. Some snakes from northwesti'rn Peru. Proc. Biol. Soc. Wash- 
ington, vol. 36, pp. 185-188. 
Gans, Carl 

1962. Eedefinition and description of the Brasilian reptiles Amphis- 
baena silveslrii Boulenger and A. neglecta Dunn and Piatt. 
Copeia, no. 1, pp. 164-170. 
Gans, Carl and A. Allan Alexander 

1962. Studies on amphisl)aenid8 (Amphisbaenia, Reptilia). 2. On the 
amphisbaenids of the Antilles. Bull. Mus. Comp. Zool., vol. 128, 
no. 3, pp. 65-158. 
Marx, Hymen 

1958. Catalogue of type specim.ens of reptiles and amphibians in the 
Chicago Natural History Museum. Fieldiana : Zool., vol. 36, 
no. 4, pp. 406-496. 
Noble, Gladwvn Kingsley 

1921. Two new lizards from northwestern Peru. Ann. New York 
Acad. Sci., vol. 29, pp. 141-143. 
Taylor, Edward H. 

1939. Two new species of tlie genus Anomalepis Jan, with a proposal 
of a new family of snakes. Proc. New England Zool. Club, vol. 
17, pp. 87-96. 



12 



BREVIORA 



No. 189 



Data for specimens of Amphisbaena pericensis Noble 



Collection 


annul: 


SEGMENTS 


Chin 






and number 


Body/Lat/Tail 


Dors/Vent 


Segments 


Cloaca 


Length 


AMNH 28501 


204+5+ (6) 18 


14/18 


2-4 


4-6-13 


132+13 


AMNH 28502 


200+4+ (6) 16 


13-4/18 


2(l)-4 


4-6-13 


146+13.5 


AMNH 28503 


212+4+ (7) 18 


14/18-20 


2-3 


4-6-12 


128+12 


BM 1929.6.1.85 - 












RR 1946.8.31.83 


202+4+ (7) 18 


14/18 


irreg. 


4-7-13 


140+14 


CAS 54614 


214+4+ (7) 18 


14/18 


2-3 


4-8-12 


150+14 


CNHM 16106 


198+4+ (7) 19 


14/18-19 


2(l)-4 


4-8-12 


134+14 


CNHM 16107 


207+4+ (7) 18 


13-4/18 


2(l)-4 


4-6-11 


101+10 


CNHM 73371 


203+4+ (7) 18 


12-3/18 


2(l)-4 


4-8-12 


110+11 


MCZ 14631 


208+4+ (7) 19 


14/20 


3-4 


4-8-12 


137+14 


MCZ 14764 


212+5+ (6) 18 


12/18 


2-3 


4-8-12 


153+15 


MCZ 14765 


200+4+ (6)x 


14/18 


3-4 


4-7-12 


104+ (3) 


MCZ 14767 


206+4+ (7) 18 


14/18 


2-4 


4-8-14 


77 + 8 


MCZ 14768 


199+4+ (6) 18 


12-14/18 


2-4 


4-7-12 


103+10 


MCZ 14770 


200+4+ (6) 17 


14/16-8 


2-4 


4-8-13 


122+12 


MCZ 14772 


206+4+ (7) 19 
4 


12-3/17-8 


2-3 


4-7-12 


127+12 


MOZ 14773 


207+1+ (7) 19 


13-4/18 


2-3 


4-8-15 


135+14 


MCZ 14774 


212+4+ (7) 18 


12-4/18 


2-4 


4-8-12 


86+8 


MCZ 14775 


215+4+ (6)x 


14/18 


2-4 


4-7-12 


142+6 


SMF 11826 


215+4+ (7) 19 


14/16-8 


2(l)-4 


4-7-13 


122+13 


SMF 11887 


206+i+(7)18 


14/18 


2-3 


4-8-11 


92+10 


SMF 11888 


206+4+f— )- 


13-4/18 


2(2)-5 


4-8-13 


127+11 


UIMNH 41494 


210+4+ (7) 18 


13-4/17-8 


2-4 


4-8-12 


84+8 


USNM 75970 


207 + 4+ (7)18 


12-4/18 


2(l)-4 


4-7-13 


101+10 


ZMU 29659 


199+4+ (6) 18 


13-4/18 


2(l)-3 


4-6-12 


122+12 


MCZ 14789 


217+3+ (6) 17 


16/18 


2(l)-4 


4-8-13 


133+13 


MCZ 14790 


217+3+ (7) 19 


15-6/18-20 


2(l)-4 


4-7-12 


130+13 


USNM 59926 


201+4+ (8) 19 

J 


13-5/18 


2-3 


4-8-12 


143+16 


USNM 60057 


218+4+ (7) X 


14-5/20 


2(l)-4 


4-8-14 


142+ (5) 


USNM 60058 


206+4+ (7) 18 


12-4/18 


2-3 


4-8-11 


126+12 



1963 



AMPHISBAENA PERICENSIS 



13 



Present status of the ^r.C.Z. imiatyix's of A. pcriccn.sifi. 



Orif/inal rnimber 

14764-65 

14766 

14767-68 

14769 

14770 

14771 

14772-75 

14776 

14777-79 

14780 



Prrsoif iiinvhrr 

Same 

"Amaral" = D.Z. ? 

8ame 

Z.M.U. 29659 

Same 

S.M.F. 11826 

Same 

U.I.M.N.H. 41494 

C.N.H.M. 16106-07, 73371 

B.M. 1929.6. 1.85-EE 1946. 

8.31.83 



14781-85 = Anomalepis M.C.Z. and U.S.N. M. 

14786 U.S.N.M. 75970 

14787-88 S.M.F. 11887-88 

14789-90 Same 



u 



BREVIORA 



No. 189 




Fig. 6. AnipliislKK nn pcricriisis. Dors.-il, l.-itcinl and ventral views of the 
head of the holotype, M.C.Z. U(i31. 



1963 



AAll'lllSUAKNA I'ERR'KNSIS 



15 




Fig. 7. AmpJiisbaetia pericensis. Dorsal (left^ and ventral (liglit) views 
of the holotype at midbofly to show size and pigmentation of segments. 




Fig. 8. Amphishaena periccnsis. Ventral view of cdoaca and tail of the 
holotype, to show pigmentation, segment arrangement and the hardly ap- 
l)arent autotomj' constriction. 



BREVIORA 

Musemini of Coimparative Zoology 



Cambridge, IMass. September 30, 1963 Number 190 

CHARACTERS AND SYNONYMIES 
AMONG THE GENERA OP ANTS. 

PART III. SOME MEMBERS OP THE TRIBE 
PONERINI (PONERINAE, PORMICIDAE) 

By William L. Brown, Jr. 

Department of Entomology, Cornell University 

INTRODUCTION 

The present part^ is concerned with some genera and species 
in the tribe Ponerini of subfamily Ponerinae. It consists mainly of 
the justification for some rather radical revisionary changes in the 
tribe at the generic level. It is felt that these findings should 
be presented here in order that they should not distract from 
the "final" reclassification results eventually to be presented 
in synoptic form for the Ponerinae and for all Pormicidae. This 
paper by no means exhausts the necessary changes that must be 
made in the generic classification of the Ponerini, but it deals 
with some of the most fundamental ones that require more than 
sunnnary proposal. Some synonymies proposed or suggested 
at the species level are to be considered a by-product of the re- 
search into generic limits. At various stages during the last 
two years, the work on ponerine reclassification has been sup- 
ported by grants from the Bache Pund of the National Academy 
of Sciences, from the Society of the Sigma Xi, and from the 
National Science Foundation (G-23680). This help is gratefully 
acknowledged. 

SOME OLD AND NEW GENERIC CHARACTERS 

OF PONERINI 

Tribe Ponerini, the largest in subfamily Ponerinae, has under- 
gone no basic classificatory changes since the appearance of 

1 Part I. Breviora, Mus. Comp. Zool., no. 11 : 1-13 (19.3.3). 
Part II. Ibid., no. 18 : 1-8 (1953). 



2 BREVIORA NO. 190 

Emery's fascicle covering the subfamily in the Genera Insectorum 
in 1911. Emery's treatment was really a refinement of his earlier 
reclassification of the ponerines published in 1901. In his well- 
known key to the genera of ants, Wheeler (1922) followed 
Emery's classification of the Ponerini for the most part, but 
Avorked in the additional genera described up to 1922. 

The Emery classification employed a number of characters of 
worker and female in distinguishing his genera of tribe Ponerini, 
but the two most stressed were (1) the number of segments in the 
maxillary and labial palpi, and (2) the number of tibial spurs on 
the middle and hind legs. 

Unfortunately, the palpal character saw extensive use only 
during some of the early years of ant systematics. Emery, Mayr, 
and even Frederick Smith often gave the count of segments in 
their descriptions, but curiously, after Emery used the palpal 
counts as a foundation stone of his 1901 classification, these ap- 
pendages were almost completely ignored by most later describers 
of ponerine species, probably due to the fact that the smaller 
mouthparts are so small and difficult of access. Later authors 
seem to have relied more on habitus than on truly diagnostic 
characters when they placed new (or supposed new) species to 
genus. As a result, some species have been described as new 
over and over again in different genera and subgenera (consult the 
case of "Trachymesopiis" hrunoi and its s^alonyms, discussed 
below). The primitive number of worker-female palpal segments 
appears to be 4, 4 (4 maxillary, 4 labial — the formula always 
stated in that order) ; in the male it is 6, 4, the basic number for 
all ants. In many Ponerini, especially the smaller or crypto- 
biotic species and their derivatives, the number is much reduced, 
and reaches as low as 0, 1 in the worker, while less consistent 
reduction occurs in the male. Palpal segmentation is undoubtedly 
important in generic classification and in the analysis of phyletic 
trends, but due to its neglect in species descriptions, it remains 
unknown for a majority of forms. "Work is now going forward 
to correct this deficiency, and some results are incorporated in 
the present paper. 

The other major character in question is the number and 
state of development of the tibial spurs. The trait is an ambigu- 
ous one, as can be seen from its employment in keys. Wheeler's 
1922 key to the genera of the Ponerini states it this way in the 
very first couplet : 

Middle and hind til)iae with two spurs 2. 

Middle and liind til)iae with a single, well-developed spur, which is always 
pectinate; the lateral spur is rudimentary or absent 19. 



ANT CHARACTERS AND SYNONYMIES 3 

This couplot really expresses the difference thought to separate 
the ''lower" Ponerini, mostly large in size, from the "higher" 
members, which tend to be smaller (e.g., Ponera, Cryptopone) . 
Generally speaking, the genera with smaller-sized species do 
tend to lose the lateral spurs of the middle and hind tibiae, while 
genera having larger-sized species normally retain the extra spur, 
although usually in a more or less reduced condition. The diffi- 
culty, of course, lies with distinguishing between the condition 
"two spurs" and "lateral spur rudimentary," especially when 
one finds that in most species the lateral spur is considerably 
smaller than its mate. Furthermore, the threshold at which a 
"vestigial spur" becomes just another apical seta of the tibia 
is unspecified along a gradual morphocline of species. In short, 
the lateral tibial spur, as an allometric character, cannot be used 
to split the Ponerini into two main groups, and in fact probably 
cannot even be used by itself as a diagnostic character at the 
generic level. 

Other adult characters of value in classification still remain: 
the shape of the clypeus has been and still is an important generic 
character. The tarsal claws may be simple, or may have one or 
more teeth, or may even be pectinate, as in the case of most 
Leptogenys species. I consider the pectination of the claws, 
imperfect in some African species of the genus, to be insufficient 
as a tribal character in view of the several other strong char- 
acters shared by both adult and larval Ponerini and Leptogenys, 
and accordingly I am placing tribe Leptogenyini as an included 
synonym of Ponerini (new synonymy). 

Another character of some importance is the presence of heavy, 
conical, spine-like setae on the extensor surfaces of the middle 
tibiae and tarsi. These structures, which appear to promote 
movement through soil or rotten wood by improving traction, 
are found in certain cryptobiotic genera (e. g., Centromyrmex, 
Wadeura, Cryptopone, Promyopias) as well as in a number of 
fossorial wasps (many pristocerine Bethylidae, most non- 
parasitic Scolioidea, for instance), all of which seek their prey 
underground or in other confined circumstances. Emery cited 
the presence of such spine-like, as well as merely bristle-like, setae 
as characteristic of Euponera subgenus Trackymesopus, which 
he named (in 1911) accordingly. The heterogeneity of 
Trachymesopus in this regard, as well as in other respects, 
made it a perfect catch-all for miscellaneous species of medium- 
to-small Ponerini from the very beginning, and it has continued 
in this role right down to the present. As a matter of fact. 



4 BREVIORA NO. 190 

heavy, conical spine-like setae that arise over half or more of 
the mid-tibial extensor surface are found only in a particular 
group of "Trachymesopus," and this group {ochracea and 
allies) shares this and several other characters with the mem- 
bers of Cri/ptoponc. The relationship between Trachymesopus 
and Cryptopone has long been discussed by E. 0. Wilson and 
myself (see Wilson, 1958: 352), and now the discovery of the 
concordance between the tibial armament and other characters, 
particularly the basal mandibular pit, makes the solution of this 
problem obvious. The "Trachymcsopns" species with these char- 
acters are all really Cryptopone, and are transferred accordingly, 
as discussed under that genus below. 

One of the chief Cryptopone characters just mentioned is a 
particularly interesting one ; this is a prominent oval pit or 
fovea near the base of the mandible in its dorsolateral surface. 
This pit, or its obvious homolog, is also found in all Brachy- 
ponera, in the members of the "Trachymesopus" sharpi group, 
in Hagcnsia, in Euponera sikorae Forel (type species of Eu- 
ponera!), and in a few other African species formerly placed 
in Euponera or other genera, but is lacking in T. stigma, the type 
species of Trachymesopus, and its closest relatives. All known 
species with the basal pit are from the Old World, and chiefly 
from Africa, with the exception of Cryptopone gilva and (per- 
haps) C. guatemalensis, which apparently represent a Crypto- 
pone invasion of the New World that has spread through North 
America and reached Central America. In the other direction, 
Cryptopone has reached southeastern Australia and New Cale- 
donia, but curiously, no true Cryptopone are known to occur in 
Ethiopian Africa. 

The basal mandibular pit in ponerine species was early noted 
as a character by Gustav Mayr and occasionally mentioned by 
later authors in species descriptions, but only Arnold used it as 
a generic character in his 1951 review of Hagensia, a genus in 
which the pits are unusually distinct. In most species that have 
it, the pit has been overlooked completely in descriptions, par- 
ticularly after 1900. Many Ponerini, especially among the larger 
and presumably more primitive species, bear another character 
in the form of an oblique groove across the dorsal face of the 
mandible, curving outward from the inner base and usually 
continuing into the lateral longitudinal mandibular sulcus that 
runs to the apex. This channel, which may be functioning to 



ANT CHARACTERS AND SYNONYMIES 5 

distribute some product from the mandibular glands, is prob- 
ably not homologous with the basal pit. It is oriented differently, 
and at least one species of " Bothroponera" has both the pit and 
the groove. 

In the former Trachymesopus, species without the truly spine- 
like mid -tibial setae (except for 2-3 of these setae at the tibial 
apex) are divided into three groups: the stigma and darwini 
groups, which have no basal mandibular pit and no anal lobe 
on the hind wing of the sexes, and the sharpi group, which has 
the pit, and the lobe in the male only, so far as known. The 
sharpi group is close to Brachyponera, with which it shares the pit 
and the lobe, but further information may dictate a separate 
genus for each of these groups. 

In what remains of T r achy m^eso pits, the stigma group (e. g., 
stigma, cautiis, pachynodiis, rufonigrus) has palpal segmenta- 
tion 3, 3, while darwini has 4, 3. Thus, it may be seen that 
Trachymesopus is a heterogeneous grouping even after the re- 
moval of the Cryptopone species ; its reclassification will have to 
await the study of more of the larvae and adult winged forms. 

Among the most promising of characters to be used in ponerine 
systematics are those of the larvae. The Wheelers (1952) have 
laid the groundwork for a comparative study of the larvae of 
the genera of Ponerini, but for the great majority of species of 
the tribe, the larvae have never been studied. The larvae of 
Ponerini are usuall}^ covered with peculiar medullate projections, 
called by the AVheelers ''tubercles." In consonance with the 
morphological terminology applied to other holometabolous 
larvae, particularly Lepidoptera, I propose that these projections 
be called by the more specifically descriptive term chalaza (sing.), 
chalazae (pL). The chalazae of Ponerini are matched by ap- 
parently homologous structures in tribes Thaumatomyrmicini 
and Odontomachini, which are close to Ponerini on the basis of 
adult characters as well. Some of the smaller ponerine genera 
bear special paired mushroom-shaped chalazae on one or more 
abdominal tergites; these have long been known to function as 
"hangers" by which the larvae are stuck to the ceiling and 
walls of the nest by a glutinous substance covering the head of 
the chalaza. The number and placement of these fungiform 
chalazae is important in generic taxonomy, but they must be 
used with care owing to the fact that they may change in 
number and form, or be lost altogether, as the instars metamor- 
phose one into another. 



6 BREVIORA NO. 190 

REVISIONARY OBSERVATIONS ON SOME 
PARTICULAR GENERA 

Cryptopone Emery 

Worker : Small in size, generally under 4 mm total length, 
depigmented (ferruginous or yellowish) ; eyes absent or ex- 
tremely reduced, body compact ; mandibles downcurved, with a 
few (4-6) coarse teeth set on more or less oblique masticatory 
borders. Dorsolateral mandibular surface with a conspicuous 
basal pit or fovea, circular or elliptical in outline. Middle tibiae 
with stout spinules covering about % or more of their extensor 
surfaces. Palpal segmentation 2, 2 or less. 

Female : Aside from well-developed compound eyes, ocelli, 
thorax and the other obviously female characters, fit the worker 
diagnosis. Hind wings without anal lobe, but with two basal cells. 

Male : Small, slender, dark-colored, with pygidial spine so 
common among Ponerini. Hind wing without anal lobe. 

Species: Those placed here by Wilson (1958), plus the former 
Trachymesopvs species crassicornis (Emery), gilva (Roger), 
probal)ly guatemalensis (Forel), ochracea (Mayr), rotundiceps 
(Emery), sauteri (Wheeler), taivanae (Forel) and its probable 
junior synonym takahashii (Wheeler) . Of species formerly placed 
in Poncra, P. typhla (Karawajew) is clearly a Cryptopone, and 
is probably a synonym of C. testacea (Emery) ; the Australian 
P. mjoehergi Forel is a junior synonym of C. rotundiceps (new 
synonymy). Among species to be deleted from the Cryptopone 
roll, C. rufofcstaceus Donisthorpe belongs in Trachymesopus as 
that genus is presently constituted, and is the same as the 
large variant of T. darivini hitherto known as T. lamarki Santschi 
(new synonymy). True Cryptopone has not been taken in 
Africa south of the Sahara, and C. angustata Santschi (type 
examined) and C. hartwigi Arnold should be transferred to 
P oner a as that genus is currently constituted. I have not checked 
this list of species for exhaustiveness, but it should serve to give 
a general idea of Cryptopone in the sense of the present work. 

Trachymesopus Emery 

As already explained above, Trachymesopus (without the 
Cryptopone species) contains at least three groups of species. 
The type species, T. stigma, is common and widespread in both 
the Indo-Australian region and in the warmer parts of the Ncav 
World. The Old World synonymy is extensive, and has been 



ANT CHARACTERS AND SYNONYMIES 7 

dealt with by Wilson (1958:355). In the New World, the 
synom-my of stigma is not yet clear. T. cognata (Emery) has 
never been differentiated satisfactorily, and T. sticcedanea 
(Roger) could be either stigma or the closely related coAitus 
Mann. T. compressinodis Boro-meier is a synonym of T. cautus 
Mann (types compared ; new synonymy) . Agreeing with stigma 
and cautus in palpal segmentation (3, 3) and in the lack of an 
anal lobe on the hind wings of both sexes, are two Australian 
species, rufonigra (Clark), transferred from Brachyponera, and 
pachynodus Clark (wings unknown in the latter). Two other 
species that apparently are close to this group are lunaris (Emery) 
and ferrugineus (F. Smith), though so little is known about 
these forms now that they cannot be confidently assigned. 

The second group consists of darwini and relatives. These are 
medium-small species known only from the ferruginous-colored 
females, which are taken commonly at light throughout wide 
areas of the Old World tropics. These females lack an anal 
lobe on the hind wing, and their palpal formula is 4, 3. They 
lack a mandibular pit and have no spine-like setae on the outer 
face of the middle tibia. They are found from northern Aus- 
tralia, Indonesia and the Philippines westward through India 
to tropical Africa. They vary considerably in size, but the varia- 
tion may be continuous; in fact, all of these forms may well 
represent a single species. As mentioned above, the species de- 
scribed as Cryptopone rufotestacea by Donisthorpe belongs here 
and is a synonym of T. lainarki Santschi, which in all likelihood 
is only a size variant of darwini. It seems likely also that 
Motschulsky's Amhlyopone testacea belongs in this complex. 

The third group is the group of sharpi Forel, which consists 
of medium-small species, all castes blackish in color, with a dis- 
tinct elliptical mandibular pit; palpal segmentation (as seen in 
2 African specimens of hrunoi only) is 4, 4. Anal lobe lacking 
in 2 hrunoi females from Liberia, but present (though separated 
by a relatively shallow cleft) in a single male from Southern 
Rhodesia; perhaps in this group the character is variable or 
sexually dimorphic. Mid-tibiae without spine-like setae on ex- 
tensor faces. 

Some of the species {hrunoi, malayanus, katangana) were de- 
scribed originally in Ectomomyrmex, or were placed in that genus 
later. Actually, these species are all very close to sharpi — in 
fact, malayanus is almost certainly a straight synonym of sharpi. 
The form of sharpi from China and Japan has the propodeum 



8 BREVIORA NO. 190 

wider and more opaque above ; it was described by Wheeler as 
subspecies pilosior, but should be considered as a good species 
until more is known about the variation in this complex, which, 
if it matches the variation found in hrunoi in Africa, may be 
considerable. T. pilosior occurs westward to the scarp of the 
Tibetan Plateau, where I collected a worker on Mt. Omei, 
Szechuan Province, in 1945; Euponera (Trackymesopus) 
cJwsonensis Teranishi, 1940, from Osaka, Japan, is its new 
synonym. The Oriental and African forms of this group are 
also very close, and deserve careful comparison. The sharpi 
group does not really belong to Trackymesopus, and will eventu- 
ally have to be moved out. But in order to avoid one additional 
round of name changes, I am leaving it where it is until the 
limits of other closely related groups, especially Bracliyponera, 
are clarified. 

Trackymesopus brunoi comb. nov. 

Pachycondyla (Ectoviomyrmex) hrunoi Forel, 1913, Deutsche ent. Zeitsehr., 
beih., p. 20."), worker. Type loc: Bulawayo, S. Ehodesia. Syutype ex- 
amined. 

Euponera (Trachymesopus) nigeriensis Santschi, 1914, Boll. Lab. Zool. 
Portici, 8: 316, worker. Type loc: Olokomeji, Nigeria; also from 
Aburi, Gold Coast. Syntype examined. New synonymy. 

Euponera (Trachymcsopus) hayoni Menozzi, 1932, Ann'. Mus. Civ. Stor. Nat. 
Genova, 56: 97-98, worker. Type loc: Kome, Sesse Archipelago, Vic- 
toria Nyanza, Uganda. New synonymy. 

Ectomomyrmex nigeriensis var. Icatangana Santschi, 1933, Bull. Ann. Soc 
Ent. Belg., 73: 96, Type loc: Pweto, Elisabethville, Belgian Congo. 
Syntype examined. New synonymy. 

Euponera (Mesoponcra) dentis Weber, 1942, Proc. Ent. Soc Wash., 44: 43, 
fig. 9, dealate female. Type loc: Lotti Forest, Imatong Mts., Sudan. 
Type examined. New synonymy. 

Euponera {Trackymesopus) lamotiei Bernard, 1953, Mem. Inst. Fr. Afrique 
Noire, 19 (1): 195, fig. 2g, h, i, dealate female. Type loc: Keoulenta 
Savannah, French Guinea. New synonymy. 
This species occurs widely in Africa south of the Sahara. In 

addition to the types mentioned above, I have seen samples 

from Liberia, Cameroons and the Kalahari Desert. There is some 

small variation among these samples in relative head v.idth and 

in strength and opacity of the sculpture, but this appears to be 

both size-linked (allometric) and continuous. 



ANT CHARACTERS AND SYNONYMIES 9 

EcTOMOMYRMEX Mayr 

The taxonomy of this genus has been consideral)ly siniplitied 
at the species level by the partial revisions of Wilson (1958) 
and Yasumatsu (1962), but an even more fundamental change 
needed is the removal of the species hrunoi and malayanus. 
These, as has been demonstrated above, are members of the 
sharpi group of "Trachymcsopus." AVith these deletions, Ec- 
tomomyrmex is once more restored to the status of a strictly 
Indo-Pacific genus, ranging from India and northwestern China 
eastward to Japan and Korea, Samoa and northern Queensland. 
It is now also possible to define the genus. 

Ecfomomyt-mcx consists of medium-sized to large, usually 
basically black-colored forms, the workers and females of Avhich 
tend to have the posterior cranium somewhat prismatic, and the 
posterior face of the petiolar node strongly striate, or at least 
rugose-punctate. Worker-female palpi segmented, so far as is 
known, 4, 4; mandibles without basal pit, the oblique basal 
groove present and weak, or absent. The oblique mesepisternal 
suture is present and reasonably distinct in the worker. Com- 
pound eyes fairly to rather well developed and multifacetted in 
the worker. 

Centromyrmex Mayr 

Centromyrmex Mayr, 1866, Verb, zool.-bot. Ges. Wien, 16: 894. Type: 

Centromyrmex hohemanni Mayr, 1866, monobasic. 
Spalacomyrmex Emery, 1889, Ann. Mus. Civ. Stor. Nat. Gcnova, 27: 489. 

Type: Spalacomyrmex feae Emery, 1889, monobasic. 
Typliloteras Karawajew, 1925, Konowia, 4: 128. Type: Typhloteras hamu- 

latum Karawajew, 1925, monobasic. 
Glyphopone Forel, 1913, Kev. Zool. Afr. 2: 308. Type: Glyphopone he- 

quaerti Forel, 1913, monobasic. New synonymy. 
Glyphopone subgenus Leptopone Arnold, 1916, Ann. S. Afr. Mus., 14: 163. 

Type: GlypJwpone (Leptopone) rufigaster Arnold, 1916, monobasic. 

New synonymy. 

The type female of Glyphopone bequaerti has been compared 
directly with a winged female kindly sent by Dr. Arnold 
(Abercorn, Northern Rhodesia, 10-12-1943, Arnold leg. et det.). 
These two specimens are virtually identical and are surely 
conspecific, as indicated in the formal synonymy expressed below. 
The median lobe of the clypeus is hardly to be considered "bi- 
carinate," but the median portion of its surface is very slightly 
concave when viewed in the proper light. Although these are 



10 brevioka no. 190 

large females, with dark forebody and rufous gaster, they bear 
the general characters of Cenfromyrmcx, and in any ease, they 
are probably no larger or darker than the female of C. gigas, 
judging from the workers of gigas I have seen. It seems best 
to consider Glyphopone a synonym of Centromyrmex until the 
worker of G. hequaerti is found and we are able to determine 
whether it has the definitive characters. 

Centromyrmex bequaerti Forel comb. nov. 

Glyphopone Bequaerti Forel, 1913, Rev. Zool. Afr., 2: 308, fig. 1, alate 
female. Type loe. : Kibombo, Belgian Congo ; examined. 

Glyphopone {Leptopone) rufigaster Arnold, 1916, Ann. S. Afr. Mus., 14: 
163, figs. 10, 10a, alate female. New synonymy. 

Pro MYOPIAS silvestrii Santschi 

Myopias (Promyopias) Silvestrii Santschi, 1914: 324, fig. 10, worker. Type 
loc. : Mamou, French Guinea ; one syntype worker examined. 

Promyopias asili Crawley, 1916, Entomologist, London, p. 30, fig., alate 
female. Type loc: " Xyasaland, " [according to label on holotype: 
Mlanje, Nyasaland, 15-IV-1913, S. A. Neave leg.]. Holotype in 
British Museum, examined. New synonymy. 

REFERENCES 

Arnold, G. 

1951. The genus Hagensia Forel (Formicidae). Jour. Ent. Soc. S. Afr., 
14: 53-56. 

Emery, C. 

1901. Notes sur les sous-families des dorylines et ponerines. Ann. Soc. 
Ent. Belg., 45: 32-54. 

1911. Formicidae subfamily Ponerinae. Genera Insectorum, fasc. 118. 
Teranishi, C. 

1940. Teranishi Memorial Volume. Posthumous works, p. 8. 
Wheeler, G. C. and J. Wheeler 

1952. The ant larvae of the subfamily Ponerinae. — Part II. Amer. 
Midi. Nat., 48 : 604-672, 6 pis. 

Wheeler, W. M. 

1922. Keys to the genera and subgenera of ants. Bull. Amer. Mus. Nat. 
Hist., 45: 631-710. 
Wilson, E. O. 

1958. Studies on the ant fauna of Melanesia. IV. The tribe Ponerini. 
Bull. Mus. Comp. Zool., Harvard, 119: 320-371. 
Yasumatsu, K. 

1962. Notes on synonymies of five ants widely spread in the Orient. 
Mushi, 36: 93-97. 



BREVIORA 

Mnjiseitiinii of Cojmparative Zoology 



Cambridge, Mass. December 5, H)(5o Number 191 

THREE NEW SPECIES OF MANGORA 
(ARANEAE, ARGIOPIDAE) FROM CENTRAL AMERICA 

By Arthur M. Chickering 

In my paper dealing with the genus M angora in Panama (1954) 
I recognized eight species then known to exist in that country. 
This conclusion involved the recognition and description of three 
new species together with the establishment of considerable new 
synonymy. 

During my collecting trips back to the Panana Canal Zone and 
parts of Panama outside of the Canal Zone in 1954 and 1957-1958 
a considerable number of specimens belonging to this genus were 
taken but no new species were found among these. I have also had 
the opportunity to study all of the Mangora collections from 
Central America now in the Museum of Comparative Zoology at 
Harvard College. Some of this material has only recently been 
sorted out of general collections which have been awaiting atten- 
tion for many years. From all of this material I have been able 
to separate out what I believe to be representatives of three new 
species. These are described in this paper in accord with my 
usual formula. 

If recently established new synonymy is taken into considera- 
tion together with the new species recognized in this paper, the 
complete list of species in the genus Mangora 0. P. -Cambridge, 
1889, now definitely known from Central America, may be given as 
follows: M. bimaculata (0. P.-Cambridge) ; M. calcarifera F. P.- 
Cambridge; M. Candida Chickering; M. conspicua sp. nov. ; M. 
distincta sp. nov.; M. mobilis (0. P.-Cambridge); M. montana 
Chickering; M. passiva (0. P.-Cambridge) ; M. pia Chamberlin 
and Ivie; M. picta 0. P.-Cambridge; M. schnierlai Chickering; M. 
spinula F. P.-Cambridge; M. sufflava sp. nov.; M. trilineata 0. P.- 
Cambridge. Banks (1898) reported M. gibberosa (Hentz) from 
LowTr California. I have not been able to examine these specimens 
but I regard it as highly unlikely that the species exists in that 



2 BREVIORA No. 191 

part of Central America and, for this reason, the species is not 
included in the list given above. Kraus (1955) reported M. gib- 
berosa (Hentz) from El Salvador. Again, I think there is serious 
question regarding the validity of this identification. I am rather 
strongly inclined to believe it belongs to the species which I am 
regarding as M. conspicua sp. nov. but, in view of the uncertainty, 
I am omitting it from my list. Only males are known for M. 
distincta sp. nov., M. picta 0. P.-Cambridge, and M. sufflava sp. 
nov. Only females are known for M. passiva (0. P.-Cambridge), 
M. schnierlai Chickering, and M. trilineata 0. P.-Cambridge. 

All types established in this paper together with my entire collec- 
tion of Araneae are deposited in the Museum of Comparative 
Zoology at Harvard College. 

Genus Mangora O. P.-Cambridge, 1889 
Mangora conspicua sp. nov. 

(Figures 1-7) 

Female holotype. Total length 3.77 mm. Carapace 1.408 mm. 
long; .99 mm. wide opposite interval between second and third 
coxae where it is widest; .66 mm. tall at level of greatest width 
and, therefore, two-thirds as tall as wide; median fovea a narrow, 
longitudinal groove. 

Eyes. Eight in two rows as usual. LE on very slightly raised 
tubercles; AME protrude somewhat over clypeus. Viewed from 
above, anterior row strongly recurved, posterior row slightly so. 
Viewed from in front, posterior row procurved, anterior row re- 
curved, all measured by centers. Central ocular quadrangle wider 
in front than behind in ratio of about 20 : 17; slightly longer than 
wide in front. Ratio of eyes AME : ALE : PME : PLE = 8 : 5.5 : 
6.5 : 6. AME separated from one another by about 1.25 times 
their diameter, from ALE by slightly less than that distance. 
PME separated from one another by aljout ten-thirteenths of 
their diameter, from PLE by eleven-thirteenths of their diameter. 
Height of clypeus equal to about the radius of AME. 

Chelicerae. Typical of the genus. Promargin of the fang groove 
apparently with three teeth ; retromargin apparently with only two. 

Maxillae and Lip. Typical of the genus; with details regarded 
as unnoteworthy. 

Sternum. Quite convex; longer than wide in ratio of about 13 : 
11; sternal suture strongly procurved; not directly continued 
between fourth coxae but a sclerite is present there ; fourth coxae 
separated by nearly two-thirds of their width. 



1963 



MANGORA FROM CENTRAL AMERICA 




l ^il / / \ f-\\\ 4+\4l 



External Anatomy of Mangora 
Figures 1-7, M . conspicua 

Figs. 1-3. Epigynum from below, in posterior view, and in profile from 
right side, respectively. 

Fig. 4. Epigynum from a paratype, turned forward to reveal the dorsal 
surface. 

Fig. 5. Left second tibia of male allotype, showing ventral spines. 

Fig.s. 6-7. Two views of the male palpal tibia and tarsus. 



Legs. 1423. Width of first patella at "knee" .217 mm., tibial 
index of first leg 15. Width of fourth patella at "knee" .227 mm., 
tibial index of fourth leg 16. 



4 






BREVIORA 






No. 191 




Femora 


Patellae 


Tibiae 


Metatarsi 


Tarsi 


Totals 






(All measurements 


in mm.) 






1. 


1.474 


.440 


1.050 


1.320 


.638 


4.922 


2. 


1.350 


.440 


.924 


1.166 


.610 


4.490 


3. 


.748 


.330 


.528 


.638 


.438 


2.682 


4. 


1.474 


.450 


.968 


1.122 


.550 


4.564 



All legs with fairly robust spines. The long, branched hairs on the 
third tibiae appear to be arranged in two oblique rows of five and 
four, respectively, but short trichobothria are closely contiguous. 
Tarsal and palpal claws appear as usual in the genus. 

Abdomen. 2.795 mm. long; 1.826 mm. wide; ovoid; with at- 
tachment to cephalothorax only slightly anterior to middle ; other- 
wise as usual in the genus. 

Epigynum. General pattern resembles that of M. mobilis (O. 
P.-Cambridge) and M. passiva (O. P. -Cambridge) but the central 
tongue is longer than in either of these species and other differences 
are also visible. Features as shown in Figures 1-4. 

Color in alcohol. This is a very clearly marked species and all 
three females available are remarkably consistent in their color 
pattern. Carapace yellowish with a narrow, median, black stripe 
reaching from just behind PME through the median groove where 
it is extremely narrow; there is a marginal stripe beginning a little 
behind PLE and extending to about opposite the second coxae 
where it widens and continues nearly to the posterior border as a 
gray area. The sternum is dark brown with a light spot in the mid- 
dle just behind the sternal suture. The legs are basically yellowish 
but arc rather conspicuously spotted because each spine is sur- 
rounded at its base by a small black dot. The first and second fem- 
ora each have a ventral black dot on the membrane at the joint be- 
tween the trochanter and the femur, then an irregular black stripe 
reaches nearly to the distal end where there is a curved black band ; 
there is also an indistinct gray stripe on both of these femora just 
prolateral to the black stripe. The gray stripe is lacking on some 
paratypes but the fourth femur has a dark, gray, retrolateral stripe 
extending through the middle of the segment. The abdomen is also 
conspicuously colored by numerous small white flecks and dark 
brown or black stripes. Beginning about one-third from the base 
there is a narrow median, brownish stripe and on each side of this 
there is an irregular black stripe. White dominates the whole dor- 
sum except for these stripes. The lateral sides each have four nar- 
row, irregular, dark brown or black, oblique stripes extending to the 
venter. The latter part is yellowish anterior to the epigynum; 



1963 M ANGORA FROM CENTRAL AMERICA 5 

behind the epigynum there is a broad, median stripe with a group 
of white flecks on each side. There are also three small, white spots 
on each side of the group of spinnerets. 

Allotype male. Three males apparently belonging to this spe- 
cies are in the collection but they are all defective for one reason 
or another. For this reason the female has been chosen as the 
holotype and a male with normal palps as the allotype. The abdo- 
men of the allotype male was crushed and of little value. Total 
length of a paratypc male 2.134 mm. Carapace of allotype male 
1.122 mm. long; .718 mm. wide opposite interval between second 
and third coxae where it is widest; about .44 mm. tall. Otherwise 
essentially as in female. 

Eyes. Essentially as in female; details regarded as unnote- 
worthy. 

Chelicerae. Difficult to see teeth along fang groove ; apparently 
the number is the same as in female. 

Maxillae, Lip, and Sternum. Each maxilla has a tooth w^hich is 
in apparent opposition to a chitinized tubercle on the palpal femur ; 
otherwise these structures are essentially as in female. 

Legs. 1243. Width of first patella at "knee" .162 mm., tibial 
index of first leg 13. Width of fourth patella at "knee" .152 mm., 
tibial index of fourth leg 15. 

Femora Patellae Tibiae Metatarsi Tarsi Totals 
(All measurements in mm.) 



1. 


1.232 


.418 


.880 


1.056 


.594 


4.180 


2. 


1.056 


.360 


.682 


.990 


.550 


3.638 


3. 


.638 


.242 


.396 


.506 


.374 


2.156 


4. 


1.034 


.330 


.704 


.792 


.462 


3.322 


Palp 


.220 


.120 


.098 




.462 


.900 



Spines on legs are considerably more conspicuous and much 
longer than in females. The curled hairs on the third tibiae ap- 
parently are arranged in two oblique rows of four and three, re- 
spectively. Additional trichobothria are also present but their 
exact distribution has not been determined. The second tibia has 
specialized spines; those seen in ventral view are represented in 
Figure 5. The first coxa has the ventral hook and the second femur 
has the corresponding groove and ridge. 

Palp. Complicated; parts are small and generally inconspicu- 
ous but the clavis is clearly visible. Both patella and tibia are 
short; the latter is broad, angular, and very irregular in outline 
(Figs. 6-7). The femur has the chitinized tubercle as already 
stated. 



6 BREVIORA No. 191 

Abdomen. This part of the body is badly crushed and unsuit- 
able for study ; apparently it is essentially like that of the female. 

Color in alcohol. The color in general is similar to that of the 
female holotype but it is less conspicuous. The marginal stripes on 
the carapace of the female are merely indistinct grayish areas 
in the male allotype. The sternum is nearly surrounded by a dark 
gray margin with the greater part yellowish with small grayish 
areas. The legs are much as they appear in the female but the 
black dots at the bases of the spines are less conspicuous and the 
prolateral, grayish stripes on the first and second femora are absent 
in the male. Abdomen: there are numerous white, chalky flecks on 
the dorsum; there are two black dots in the middle line in the 
anterior half of the dorsum; the posterior half of the dorsum con- 
tains a dark, rectangular, fairly broad spot composed of a series of 
black marginal spots more or less connected by very narrow, black 
lines extending through a brownish area. 

Type locality. The holotype female is from Mexico, Nuevo 
Leon, El Potosi, Cerro Potosi, June 13, 1938 (H. Hoogstraal). One 
paratype was taken with the holotype. Another in the collection is 
from the same general region, Sabinas Hidalgo, 1000 ft. elevation, 
June 13, 1940. Three females in the collection are from Honduras, 
Escuela Agr. Panam., 27 km. S. of Tegucigalpa, San Antonio del 
Oriente. Two males accompany the females. One was mature; it 
had undergone severe injury to its abdomen but was selected as 
the allotype; the other was immature. The Honduran specimens 
were collected by A. and M. Carr at 3800-4000 ft. elevation, No- 
vember 17, 1945. Two other males in the collection are from 
Mexico: Nuevo Leon, Villa de Santiago, Hacienda Vista Hermosa, 
2000 ft. elevation, July 19, 1940 (H. Hoogstraal). Both of these 
males have their tarsal bulbs so distended as to be useless for 
description. 

Mangora distincta sp. nov. 
(Figures 8-11) 

Male holotype. Total length 2.535 mm. Carapace 1.202 mm. 
long, .902 mm. wide opposite interval between second and third 
coxae where it is widest, .484 mm. tall and, therefore, about .54 as 
tall as wide; gently arched with tallest point nearly at level of 
greatest width; median fovea a rather deep, narrow, longitudinal 
groove. 

Eyes. Eight in two rows as usual; LE on slightly raised tu- 
bercles; AME protrude i)rominently over clypeus. Viewed from 



1963 



MANGORA FROM CENTRAL AMERICA 



above, anterior row strongly recurved, posterior row slightly so. 
Viewed from in front, anterior row slightly recurved, posterior row 
moderately procurved, all measured by centers. Central ocular 
quadrangle wider in front than behind in ratio of 19 : 16; slightly 




External Anatomy of Mangora 

Figures 8-11, M. distincta 
Figure.s 12-15, M. sufflava 

Fig. 8. Left second tibia to show ventral spines. 
Figs. 9-10. Two views of the palpal tarsus. 
Fig. 11. Tarsal clavis. 

Fig. 12. Left second tibia to show ventral spines. 
Figs. 13-14. Two views of the palpal tarsus. 
Fig. 15. Tarsal clavis enlarged. 



8 BREVIORA No. 191 

longer than wide in front. Ratio of eyes AME : ALE : PME : 
PLE = 6:5:6:5. AME separated from one another by their 
diameter, from ALE by slightly less than this distance. PME 
separated from one another by slightly more than their radius, 
from PLE by their diameter. LE separated only by a line. Height 
of clypeus about equal to the diameter of AME (surrounding pig- 
ment makes it difficult to obtain exact measurements of eyes and 
their relative positions). 

Chelicerae. General features as usual in the genus. Unable to 
observe teeth along fang groove without injury to holotype. 

Maxillae and Lip. Maxillae with a sharply pointed tooth near 
their bases ; probably used in opposition to the chitinized tubercle 
on the palpal femur. In general, both structures appear to be typi- 
cal of the genus. 

Sternum. Scutiform; strongly convex; somewhat depressed in 
anterior portion; sternal suture quite procurved; widest between 
second coxae where it is nearly as wide as long; continued broadly 
between fourth coxae which are separated by 8/11 of their width. 

Abdomen. Ovoid; quite typical of the genus; details regarded 
as unnoteworthy. 

Color in alcohol. Legs and mouth parts generally yellowish; 
each chelicera has a small, irregular, grayish area in front near the 
base. The carapace is yellowish with a narrow central stripe from 
a little behind PME and extending two-thirds of the distance to 
the posterior border. The sternum is yellowish with a very irregular 
light grayish marginal area. Abdomen: yellowish with whitish 
flecks dorsally and dorsolaterally ; in the posterior half of the 
dorsum there are three narrow brownish stripes ; in the region of 
the spinnerets there are several black and white dots; on each 
lateral side there is a narrow oblique grayish stripe followed by an 
area of many small white flecks and finally a rounded irregularly 
grayish spot; the venter is generally yellowish but just in front 
of the genital groove there is an oval grayish spot and behind the 
genital groove there is an irregularly grayish, rounded spot con- 
taining a group of white flecks. 

Type locality. The holotype is from Honduras, Escuela Agr. 
Panam., 27 km. S. of Tegucigalpa, San Antonio del Oriente, Nov. 
17, 1945, 3800-4000 ft. elevation (A. and M. Carr). There are no 
paratypes and the female is unknown. 



1963 M ANGORA FROM CENTRAL AMERICA 9 

Mangora sufflava sp. nov. 
(Figures 12-15) 

Male holofype. Total length 2.6 mm. Carapace 1.3 mm. long, 
.99 mm. wide opposite interval between second and third coxae 
where it is widest, .638 mm. tall at about the level of its greatest 
width and, therefore, about half as tall as wide; median fovea a 
narrow, well defined, longitudinal groove; considerably overlapped 
by anterior end of abdomen. Other features as usual in the genus. 

Eyes. Eight in two rows as usual. Viewed from above, anterior 
row strongly recurved, posterior row moderately so. Viewed from 
in front, anterior row moderately recurved, posterior row moder- 
ately procurved, all measured by centers. Central ocular quad- 
rangle wider in front than behind in ratio of 7 : 6; slightly longer 
than wide in front. Ratio of eyes AME : ALE : PME : PLE = 
7.5 : 6 : 6.5 : 6. AME separated from one another by about their 
diameter, from ALE by about two-thirds of their diameter. PME 
separated from one another by about two-thirds of their diameter, 
from PLE by about 1.3 times their diameter. LE separated only by 
a broad line. Height of clypeus equal to about four-fifths of the 
diameter of AME. 

Chelicerae, Maxillae, Lip and Sternum. All apparently typical 
of the genus. 

Legs. 1243. Width of first patella at "knee" .200 mm., tibial 
index of first leg 13. Width of fourth patella at "knee" .162 mm., 
tibial index of fourth leg 14 

Femora Patellae Tibiae Metatarsi Tarsi Totals 
(All measurements in mm.) 



1. 


1.364 


.506 


.990 


1.100 


.649 


4.609 


2. 


1.320 


.484 


.902 


1.078 


.627 


4.411 


3. 


.792 


.308 


.506 


.528 


.418 


2.552 


4. 


1.292 


.418 


.770 


.880 


.528 


3.888 


Palp 


.286 


.119 


.119 




.440 


.964 



All legs with well developed spines and hairs. Second tibia with 
modified spines ; those seen in ventral view shown in Figure 12. The 
long, slender, branched hairs on the prolateral side of third tibia ap- 
pear to be arranged in two oblique rows of three each. The first coxa 
bears the usual hook and the second femur has the corresponding 
prolateral, proximal groove and chitinizcd ridge. Tarsal claws ap- 
pear to be as usual in the genus. 



10 BREVIORA No. 191 

Palp. The patella has the usual long, slender, terminal, dorsal 
spine. Both patella and tibia are short. Important tarsal features 
shown in Figures 13-15. The clavis appears to be different from 
any other found in the collection. The tooth is probably present 
on the maxilla together with the chitinized tubercle on the palpal 
femur but neither can be clearly seen without danger of damage 
to the holotype. 

Abdomen. 1.625 mm. long; 1.170 mm. wide near middle; ovoid. 
At the base are two long, slender, dorsal, spine-like bristles. Other 
features typical of the genus. 

Color in alcohol. Carapace yellowish with a grayish median 
stripe extending from AME to middle third of steep posterior de- 
clivity ; the center of the grayish stripe is a narrow, black line. The 
sternum is broadly grayish around the margin but yellowish in the 
center. The legs and mouth parts are yellowish with many varia- 
tions in the depth of color. Abdomen: this part of the body has a 
rather unusual color pattern; dorsolaterally there is a fairly broad, 
white stripe on each side extending nearly the length of this part 
of the body; there is a large, median, grayish yellow spot in the 
anterior half of the dorsum containing a pair of small white dots 
in the posterior half with another pair of small white dots at its 
posterior border ; the posterior half of the dorsum has a pair of dark 
gray stripes separated by a yellowish stripe; these stripes do not 
reach to the posterior end of the abdomen; the venter is yellowish 
with faint graj^ irregularities. 

Type locality. The holotype male is from Boquete, Chiriqui, 
Panama, August, 1950. This specimen was overlooked among 
specimens belonging to different genera at the time of completion 
of my previous paper on Mangora (1954) and only recently found. 
There are no paratypes and the female is unknown. 

BIBLIOGRAPHY 

Banks, Nathan 

1898. Ararhnida from Baja California and other parts of Mexico. Proc. 

California Acad. Sci., ser. 3, Zoology, 1(7) : 205-309, 5 pis. 
1929. Spiders from Panama. Bull. Mus. Comp. Zool., 69: 53-96, 4 pis. 

Bonnet, Pierre 

1957. Bibliograiihia Araneorum. 2. 3me partie. Toulouse: Les Artisans 
de rimprimerie Douladoure. 

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

1889- Arachnida-Araneida. /?* .• Biologia Centrali-Americana. 
1905. Dulau & Co., London. 



1963 MANGORA FROM CENTRAL AMERICA 11 

ChAMBERLIN, R. V. AND WiLTON IviE 

1936. New spiders from Mexico and Panama. Bull. Univ. Utah, 27: no. 5, 
biol. series, 3, no. 5:3-103, 17 pis. 

Chickkring, Arthur M. 

1954. The spider genus Mangora (Argiopidae) in Panama. Bull. Mas. 
Comp. Zool., Ill : 195-215, 28 fig.s. 

Kraus, Otto 

1955. Spinnen aus El Salvador (Arachnoidea, Arancae). Abhand. Senck- 
enberg. naturforsch. Ges., 493: 1-112, 12 pis. 

ROEWER, C. F. 

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

Simon, Eugene 

1892- Histoire naturelle des araignees. Deuxieme edition. 
1903. Librarie Encyclopedique de Roret, Paris. 



BREVIORA 

Miasemm of Compsiratlve Zoology 



Cambridge, Mass. December 5, 1963 Number 192 

A DESCRIPTION OF DINOPIS LONGIPES 
F. P.-CAMBRIDGE, 1902 

(Araneae, Dinopidae). 
By Arthur M. Chickering 

F. P.-Cambridge described this highly interesting species en- 
tirely by means of figures and a statement of the total lengths of 
the two sexes. It may be that one reason for this extreme brevity 
was due to the fact that fully mature individuals have extraordi- 
narily long and fragile legs and the body is usually badly distorted 
in the preservative. This results in considerable difficulty in getting 
specimens sufficiently intact to make description significant. It has 
usually been assumed that this is a rare species but this is probably 
because collecting activities have not been sufficiently intensive. 
At any rate, I now have in my collection several mature specimens 
of both sexes and numerous immature individuals. 

Because of the extreme brevity of the original description it has 
been thought advisable to prepare a detailed description of both 
sexes in order to make clear the fundamental features of the spe- 
cies. After a careful search through the collection, individuals of 
both sexes have been selected for detailed description in accord 
with my usual formula. The selected male is smaller than some 
other individuals of the same sex but its parts are quite well pre- 
served and its legs are much less fragile than in most of the other 
available specimens. It appears to have been captured soon after 
the last moult and before its full size had been reached. The female 
selected for description had probably reached her full mature size. 

Genus DiNOPIS Macleay, 1839 
DiNOPis LONGIPES F. P.-Cambridge, 1902 

Male. Total length 16.282 mm. Carapace 4.42 mm. long; about 
2.73 mm. Vv-ide; extremely flat; with a shallow median depression 
continued posteriorly by a long, very narrow groove ; with several 
very short spines. 



BREVIORA 



No. 192 



Eyes. Eight in three rows with arrangement typical of the 
genus; posterior centrals enormously enlarged and moved forward 
to dominate the optical area. Viewed from above, only posterior 
eyes seen in two rows. Viewed from in front, anterior row definitely 
procurved. Ratio of eyes AME : ALE : PME : PLE = 13 : 15 : 
32 : 16. AME separated from one another by 18/13 of their diame- 
ter, from ALE by 42/13 of their diameter. PME separated from 
one another by about 2/3 of their diameter, from PLE by nearly 
3/2 of their diameter. (All measurements are made from the inner 
margin of the iris and not the margin where the cornea becomes 
continuous with the cuticle, which is often difficult to determine.) 
The iris of the PME is purplish and the border of the cornea is 
bright red. The clypeus is lacking because of the marginal posi- 
tion of the first row of eyes. The AME are located on a common 
tubercle; each ALE is also mounted on a fairly prominent tubercle. 

Chelicerae. Rather small for the size of the animal ; basal seg- 
ment about .77 mm. long; distal half of medial surface with many 
long, curved, stiff, brownish bristles. Fang evenly curved, without 
special features. Fang groove fairly well defined; with 5 definite 
small teeth plus a terminal nodule along the promargin; with 10-12 
minute teeth along the retromargin, some of which are no more 
than nodules; some irregularities have been noted on left and 
right sides and in different specimens. 




mmmh 



liiii 



External Anatomy of Dinopis longipes 

Fig. 1. Lip. 

Fig. 2. Palpal tarsus of male showing coiled embolus and central apo- 
physis. 

Fig. 3. Di.stal end of embohis; removed from a second male. 

Fig. 4. Epigynum from below. 



1963 REDESCRIPTION OF DINOPIS LONGIPES 3 

Maxillae. Nearly parallel in general; massive in proximal 
halves; narrowed and somewhat diverging in distal halves; with 
well developed scopulae. 

Lip. General features shown in Figure 1. 

Legs. 1243. Width of first patella at "knee" .550 mm., tibial 
index of first leg 3. Width of fourth patella at "knee" .440 mm., 
tibial index of fourth leg 4. 

Femora Patellae Tibiae Metatarsi Tarsi Totals 
(All measurements in mm.) 



1. 


13.780 


1.950 


14.495 


20.150 


5.460 


55.835 


2. 


11.700 


1.755 


11.310 


12.675 


3.510 


40.950 


3. 


9.750 


1.280 


7.410 


7.280 


1.625 


27.345 


4. 


10.855 


1.690 


10.075 


10.725 


1.755 


35.100 


Palp 


2.015 


.845 


.520 




1.170 


4.550 



Spines: Numerous femoral spines are present but apparently 
all are very small and, perhaps, irregularly placed; no patellar 
spines have been observed ; tibial and metatarsal spines are numer- 
ous and of moderate size. Three tarsal claws are present with each 
upper claw provided with a single row of about six teeth ; the third 
claw is simple and without teeth ; there are also numerous spurious 
claws present. Numerous trichobothria have been observed on 
tibiae and metatarsi; they are not confined to "ventral surface at 
base of tibia" as stated by Dr. Petrunkevitch (1939). The cala- 
mistrum is moderately well developed on the proximal half of the 
fourth metatarsus of immature specimens; on the specimen here 
described in detail it is, apparently, much reduced. 

Palp. Both patella and tibia are short, simple and lacking 
apophyses or other special modifications; the cymbium is a deep 
cup-shaped structure containing the bulb which has an extra- 
ordinarily long, coiled embolus ; enclosed in the center of the circu- 
lar coils of the embolus is a somewhat distinctive structure noted 
by F. P.-Cambridge and again noted in this study. Figures 2 and 
3 show the chief features of the palp as observed by this author. 
The length of the free part of the embolus seems to vary in different 
individuals. In one example there were apparently ten circular 
coils making a total length of the embolus approximating two 
inches. In the specimen described here in detail the free part of the 
embolus is much shorter; when stretched out it resembles a clock 
spring. The median apophysis surrounded by the coiled embolus 
also exhibits different shapes in different specimens. The signi- 
ficance of these differences is not at all clear. 



4 BREVIORA No. 192 

Abdomen. Long, slender; of nearly uniform width; with a pair 
of very small dorsal tubercles slightly behind the middle. The six 
spinnerets and cribellum appear typical in males of the genus. The 
abdomen is well supplied with plumose hairs. 

Color in alcohol. Carapace: the basic color is a yellowish 
brown; with a narrow darker median stripe and a fairly broad 
and still darker dorsolateral stripe on each side; white plumose 
hair extends between the PLE up to the enlarged PME. The 
sternum is yellowish, with a very irregular, broken, narrow, brown 
border and several irregular whitish subchitinous patches. The legs 
are yellowish with sparsely located small black dots ; the coxae are 
much spotted with black ventrally. Abdomen: light brownish dor- 
sally and laterally ; with a darker, median, dorsal stripe ; the lateral 
sides possess several alternating, light and dark longitudinal 
stripes; the venter has a fairly broad, brownish, variegated, median 
stripe and many small, whitish, subchitinous, irregular spots on 
each side of the darker median stripe. 

Female. Total length about 24.7 mm. Carapace about 8.58 mm. 
long; 4.225 mm. wide op]:)osite interval between second and third 
coxae where it is widest; much narrowed immediately in front of 
first coxae where it is about 2.405 mm. wide; surface with nu- 
merous shallow depressions. 

Eyes. General features as in male. Ratio of eyes AME : ALE : 
PME : PLE = 9.5 : 11 : 44 : 12.5. 

Chelicerae. Essentially as in male except for teeth along the 
fang groove. The specimen from which this description is chiefly 
taken shows seven teeth along the promargin of which the second, 
fifth, and seventh are very small. Along the retromargin there are 
five or six teeth of moderate size and many minute teeth, little 
more than minute tubercles; immediately behind the last tooth 
of moderate size there is a series of five very minute teeth some of 
which are on the lower part of the larger tooth itself; some of the 
minute teeth occur in the fang groove. The fang groove in another 
female specimen showed the promargin with six teeth of which the 
fourth and sixth were small, the others of moderate size; the re- 
tromargin had eight teeth in an irregular row together with numer- 
ous minute tubercles. There is a well defined scopula along the pro- 
margin of the fang groove. 

Maxillae. Essentially as in male. 

Lip. This structure is abnormal in the specimen chiefly used 
for this description. Another specimen has the lip essentially as 
shown in Figure 1. 

Sternum. Also essentially as in male. 



1963 REDESCRIPTION OF DINOPIS LONGIPES 5 

Legs. 1243. Width of first patella at "knee" .88 inm., til)ial 
index of first leg 4. Width of fourth patella at "knee" .902 mm., 
tibial index of fourth leg 6. 

Femora Patellae Tibiae Metatarsi Tarsi Totals 
(All measurements in mm.) 



1. 


18.525 


2.925 


17.680 


18.850 


4.160 


62.140 


2. 


18.200 


2.860 


16.900 


16.900 


2.929 


57.789 


3. 


13.780 


2.015 


10.400 


10.725 


1.950 


38.870 


4. 


14.170 


2.665 


12.545 


11.895 


1.950 


43.225 



I have found difficulties in making leg measurements as accurate 
as usual because of fragility and breakage. The palpal claw has 
three prominent teeth ; the palpal tarsus bears many stiff bristles 
and numerous spines. The calamistrum is fairly well developed on 
the proximal third of the fourth metatarsus. All legs are richly 
provided with spines of two kinds; the usual type are numerous, 
short and, apparently, not regularly placed; the second type is a 
slender, highly branched, unusual kind. Trichobothria have not 
been observed in females, perhaps because of the multiplicity of 
spines. 

Abdomen. Badly distorted by the preservative. Apparently as 
in male in all essential general features except that the dorsal tu- 
bercles somewhat in front of the middle are moderately well de- 
veloped in contrast to the minute tubercles in the male ; each tu- 
bercle has its posterior surface covered by a coating of short, white 
hairs. The cribellum is well developed and undivided. 

Epigynum. Essentially as represented in Figure 4. 

Color in alcohol. Essentially as in male except that the whole 
coloration is darkened to a dull brown in general ; the dorsal sur- 
face of the abdomen bears a series of isolated, small, white dots 
caused by clusters of short, white hairs. 

Collection records. Both specimens chiefly used for this de- 
scription were collected on Barro Colorado Island, C. Z., July, 1954. 
Several other adults and numerous immature specimens are in the 
collection from many localities in the Canal Zone and Boquete, 
Chiriqui, Panama. The species seems to be fairly abundant on 
Barro Colorado Island. All specimens referred to in this paragraph 
are now deposited in the Museum of Comparative Zoology at Har- 
vard College. 



6 BREVIORA No. 192 

REFERENCES 

Bonnet, Pierre 

1956. Bibliographia Araneorum. 2. 2me partie. Toulouse: Les Artisans 
de rimprimerie Douladoure. , 

Cambridge, F. Pickard 

1897- Arachnida-Araneida. In: Biologia Centrali-Americana. Vol. II. 
1905 Dulau & Co., London. 

Petrunkevitch, Alexander 

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

South America, etc. Bull. Amer. Mus. Nat. Hist., 29: 1-809. 
1939. Catalogue of American Spiders. Pt. 1. Trans. Conn. Acad. Arts and 

Sci., 27: 51-248. 

Roewer, C. F. 

1954. Katalog der Araneae. Vol. 2, Pt. 2. Brussels. 



BREVIORA 

Mmseiinii of Comparative Zoology 



Cambridge, Mass. December 5, 1963 Number 193 

A MIOCENE TOAD FROM 
COLOMBIA, SOUTH AMERICA 

By Richard Estes^ and 
Richard Wassersug^ 



The fossil toad described here is part of the large vertebrate 
assemblage of the La Venta fauna (Fields, 1959), and was col- 
lected by R. W. Fields during the 1949 ITniversity of California 
field expedition to the upper Magdalena Valley, Huila, Colombia, 
South America. It is the same specimen identified by D. Savage 
(1951) as a leptodactylid, cited by Estes (1961) as a bufonid close 
to the living Bufo ah>arius and B. crucifer, and discussed briefly 
by Tihen (1962b, p. 14) as Bufo sp., near B. marinus. Tihen's very 
useful paper on bufonid osteology (1962a) now makes it possible 
to give a much more accurate and clear-cut assessment of the 
relationships of this animal. Few fossil Bufo have been recorded 
previously from South America (Tihen, 1962b). 

Both authors wish to express their gratitude to Drs. E. E. Wil- 
liams, R. F. Laurent, J. A. Tihen and J. M. Gallardo for their 
many helpful suggestions, and to Mr. Howard Hamman, who 
prepared the illustrations. This work was supported by National 
Science Foundation Grant NSF G-18905, held by the senior au- 
thor, and research time for the junior author was provided through 
the Thayer Academy Advance Studies in Science Program (also 
supported by the National Science Foundation). We are grateful 
for this support. 



' Department of Biology, Boston University, and Associate in Vertebrate Paleontology, Mu- 
seum of Comparative Zoology, Harvard University. 

^ Thayer Academy, Braintree, Massachusetts 



2 BREVIORA No. 193 

Class AMPHIBIA 

Superorder SALIENTIA 

Order ANURA 

Family BUFONIDAE 

BuFo MARiNus Linnaeus 

Referred specimen — University of California no. 41159, postor- 
bital portion of skull, eight articulated vertebrae, both scapulae 
with articulating proximal ends of the humeri, both distal ends of 
humeri in articulation with proximal ends of radioulnae, two frag- 
ments from the region surrounding the acetabulum of the pelvic 
girdle, distal end of right femur, proximal ends of both tibiofibulae, 
distal extremity of right tibiale-fibulare, and two unidentified 
fragments. Two bone shafts which were collected at the same time, 
and have the same specimen number, are probably mammalian. 

Horizon — University of California locality V-4517, Monkey 
unit, Honda group (Fields, 1959, p. 419, and fig. 2). 

Age — Late Miocene. 

Fauna — La Venta. 

Preservation — The skeleton was apparently complete before ex- 
posure and erosion disarticulated and destroyed parts of it. The 
matrix is a silty mixture of claystone and sandstone, and the cavi- 
ties have been filled either with this materialor with calcite. In 
some cases, calcite or matrix have remained as endocasts of parts 
of the bones. Breakage has apparently been the result of erosion, 
and has occurred at the weakest and thinnest points. 

The shoulder and elbow complexes are articulated in flexed posi- 
tions, and the vertebral column has a slight ventral curve, similar 
in both cases to their positions in life in a normal resting position. 
It is possible that the toad died and was buried in such a position. 

Description — The maximum width of the skull across the pos- 
terior arms of the squamosals is 39.5 mm. A height measurement 
of the skull taken from the dorsal extremities of the prootic part 
of the otoparietal (in Bvfo, paired processes directly dorsal to the 
foramen for the ninth and tenth cranial nerves) to the parasphe- 
noid-ptcrygoid suture is 10.0 mm. A ventral measurement along 
the median line from the exoccipital condyles to the jiostcrior 
border of the sphencthmoid is 1L7 mm. In occipital view, the 
skull roof appears flat, except for the strong postorbital crests 
and suggestions of the supraorbital crests. The foramen magnum 
is about 65 per cent wider tlian deep, and the foramina for ninth 



1963 



MIOCENE TOAD 



and tenth cranial nerves are prominent. Only the left columella 
is present and its tip was broken in handling. Its true length, as 
indicated in the figures, was taken from a j^hotograph made before 
the breakage occurred. The prootic parts of the otoparietal are 
strongly ossified laterally and ventrally and form the ventral bor- 
ders of strongly marked troughs for the columellae. The pterygoids 



^P^. foa 




opa 




Figure 1. Bufo marinus, U. C. no. 41159, ventral view of skull, anterior to 
the top; above, outline, and below, shaded drawing; X2. On the outhne, 
solid lines indicate sutures, dotted lines are contours, and cross-hatching 
indicates breakage. Abbreviations : ecc, endocranial cast; ex, exoccipital; 
joa, canal for occipital artery; joe, oculomotor foramen; jom, foramen 
magnum; jor, foramen for maxillo-mandibular branch of trigeminal nerve; 
fps. frontoparietal .suture; ma, attachment for nuchal muscles; oc, exoccipital 
condyle; oj, large opening into braincase for optic and trochlear nerves; 
opa, otoparietal; pas, parasphenoid ; pt, pterygoid; sph, sphenethmoid ; sq, 
squamosal; st, columella auris; tr, foramen for fifth, sixth and seventh cran- 
ial nerves; vj, foramen for ninth and tenth cranial nerves. 



BREVIORA 



No. 193 



are broken just medial to the point at which they would have given 
off their quadrate and palatal processes, and the descending quad- 
rate processes of the squamosals are broken off at approximately 
the same level. The cultriform process of the parasphenoid is 
broken off just posterior to its juncture with the sphenethmoid, 
but the bone shows nothing unusual. The squamosal-prootic and 
prootic-parasphenoid sutures are difficult to discern, especially on 
the right side, and it appears that almost complete fusion has oc- 
curred. 




opa 




Figure 2. Bufo marinus, U. C. no. 41159, posterior view of skull; above, 
outline, and below, shaded drawing; X2. Comments and abbreviations as in 
Figure 1. 



In dorsal view, the dermal ornamentation lacks strong protu- 
berances other than the postorbital and orbital crests, though weak 
parietal crests are present, and two small bumps occur near the 
midline. The all-over pattern of the dermal ornament is a pitted 
and wrinkled one, in which the wrinkles extend more or less trans- 
versely across the top of the skull. 

The frontoparietal portions of the otoparietals appear to be 
fused to each other, though an anterior groove, shown in Figure 



1963 



MIOCENE TOAD 



3, perhaps indicates part of their suture. Conjoined, these fronto- 
parietal areas are 21.7 mm. at their widest point. The anterior sec- 
tion of the frontoparietal area is broken off before its contact with 
the nasals. The occipital groove (Tihen, 1962a), is enclosed to 
form a canal. On the right side, breakage has removed the overly- 
ing bone and exposed the endocast of this canal. 

In ventral aspect, much of the brain cavity appears as an en- 
docranial cast. Small posterior fragments of the sphenethmoid, 
along with the anterior arms of the prootics and the cultriform 
process of the parasphenoid, completely surround an opening 
which was membrane covered in life and through which the optic 




opa 




Figure 3. Bufo marinus, U. C. no. 41159, dorsal view of skull, anterior to 
the top; above, outline, and below, shaded drawing; X2. Comments and 
abbreviations as in Figure 1. 



and trochlear nerves passed. This opening is somewhat squared- 
off , with notches in anterodorsal and posteroventral corners. Be- 
cause of greater ossification, especially on the posterior border of 
the sphenethmoid, the opening is relatively smaller than that of 



6 BREVIORA No. 193 

Ptecent Bufo. The lateral wings of the parasphenoid are broadly 
overlapped by rugose medial processes of the pterygoids. The fora- 
men for cranial nerves five, six, and seven, and the oculomotor 
foramen are as in Recent Bufo. The occipital grooves (here en- 
closed to form canals) open on the ventral surface of the orbital 
portions of the frontoparietals, and are continued for a short dis- 
tance as channels on the roof of the orbit. These paired canals, 
which in life transmitted the occipital arteries, diverge slightly 
as they pass anteriorly, but lie essentially parallel to the midline. 
Dorsal to the suture between the prootic and the pterygoid, im- 
mediately lateral to the foramen for cranial nerves five, six, and 
seven, and on a level with it, is a groove which passes through the 
prootic and emerges immediately lateral to the base of the colu- 
mella. The maxillo-mandibular branch of the trigeminal nerve 



opa 




for pt 



Figure 4. Bufo mnrinus, U. C. no. 41159, anterior view of right half of 
skull, outline only, showing posterior wall of orbit ; X2. Comments and 
abbreviations as in Figure 1. 



is found in this groove in Rana (Holmes, 1924, p. 296) and dissec- 
tion showed this to be the case in Bufo as well. The groove in Bufo 
terminates on the lateral borders of the prootic, leading into the 
supraptcrygoid fenestra (Tihen, 1962a, j). 160), but in this speci- 
men there has been extremely high ossification in this area and 
portions of the prootic, squamosal, and pterygoid have completely 
closed the fenestra, and enclosed the groove in a foramen. 

The thick and massive vertebral colunni is complete from the 
atlas back to about the level of the anterior third of the eighth 
vertebra, though all of the transverse processes are badly broken. 
The column was originally articulated with the skull, but was re- 
moved by the senior author in order to study the configuration of 
the exoccipital condyles and atlantal cotyles. An approximate 



1963 



MIOCENE TOAD 



overall measurement of the seven presacral vertebrae is 42.5 mm. 
The atlantal cotyles are separated by a shallow notch. The neural 
spines are flattened and capped with dermal bone that has a 
slightly pitted texture. None of the broken transverse processes 
are long enough to give positive information about their true 
length or orientation, though they seem to have been oriented as 
in Recent Biifo. The zygapophyses are robust, as are the centra. 






Figure 5. Biifo mannus, U. C. no. 41159, right, medial view of left scapula 
and humerus {h) , showing sutural contacts of scapula (sc), coracoid (c), 
and clavicle (cl) ; left, anterior view of distal end of left humerus and 
proximal end of radioulna (r), to show crista medialis (cm) ; below, ventral 
view of vertebral column, anterior to left; all X2. 



The right scapula is the better preserved. Its dorsal edge is 
broken parallel to, and near, the original natural border. The left 
scapula is broken at about the same point, but its posterior half is 
missing. In the glenoid fossa, the small band of cartilage that 
separates the scapula from the clavicle in the Recent species is 
completely replaced by a firm bony suture. In Bufonidae, the 
pars acromialis of the scapula (Proctor, 1921, p. 197) forms a 
strong prominence which is relatively larger in the fossil (on the 
right side ; it is broken on the left) than in the Recent species. The 



8 BREVIORA No. 193 

paraglenal cartilage (Ecker, 1889, p. 40), between the scapula and 
the coracoid, usually forms a noticeable portion of the floor of the 
glenoid cavity in modern anurans, but is absent in the fossil and 
replaced by a firm suture of these two bones. The large foramen 
in the glenoid fossa is, as a result of the heavy ossification in this 
region, slightly smaller than in the Recent forms. 

The deltoid crest of the humerus is quite prominent, and is 
about as high from base to peak as the circumference of the hu- 
merus at the same point. The shafts of both humeri are missing, 
distal to the crests. 

The radioulnar articulations are similar to those of Recent Bvfo. 
On the distal segment of the left humerus, the crista medialis is 
well developed, and has a strongly rugose muscular attachment 
surface. 

All that remains of the pelvic girdle is the ventroposterior cor- 
ner of the ischium and a ventral piece of the ilium from near the 
acetabulum. 

The proximal articulation surfaces and short segments of shafts 
of the tibiofibulae show no differences from Recent Bujo. The 
right tibiofibula has the longest shaft preserved. The grooves sepa- 
rating fused tibial and fibular components are deep but no more 
so than in the Recent species. The distal end of the right femur is 
no longer in natural articulation with the tibiofibula but still 
makes a perfect fit. 

Both proximal and distal extremities of the right tibiale-fibulare 
are present. The diaphyses of the bones have been broken so that 
a natural fit is no longer possible. 

Two small fragments may represent a posterior section of the 
ilial shaft and a section of tibiofibula, but are insufficient for posi- 
tive identification. 

Discussion — Tihen (1962a, p. 163) defines the valliceps species 
group of Bufo as follows: frontoparietals broad, usually pro- 
duced into crests; roofing bones ornamented; occipital groove en- 
closed to form a canal; frontoparietals and prootics fused. He 
then divides the valliceps group into three essentially geographical 
subgroups: the Mexican section in Central and North America; 
the South American section in South and Central America; and 
the Caribbean section, throughout the Neotropical Region, "par- 
taking to some extent of the characteristics of each of the others, 
besides developing its own features." 

This fossil is placed in the valliceps group on the basis of pres- 
ence of all of the above characters. Within this group, it is elimi- 
nated from the Mexican section by having a strong overlap of the 



1963 MIOCENE TOAD 9 

medial wings of the pterygoids onto the wings of the parasphenoid 
and complete closure of the suprapterygoid fenestra {ibid., p. 
168). The strongly-overlapping pterygoids are characteristic of 
the South American section, and do not usually occur in the Carib- 
bean section, but the occlutled sui)rai)terygoid fenestra is present 
in the latter (ibid., p. 171 ) . This single resemblance to the members 
of the Caribbean section will be discussed below. 

Bufo chilensis of the South American section and B. retiformis 
of the Caribbean section were the only species not available for 
this study. The dermal ornamentation of the fossil is most like 
that of the South American section, which has crests of only mod- 
erate extent and development and a lined or wrinkled sculpture; 
while crests in members of the Caribbean section are often exten- 
sive, exaggerated, and the sculpture is pustular. 

A final factor used in allocating the fossil to the South Ameri- 
can section was the width of the vertebral centra, wiiich are per- 
ceptibly narrower in proportion to their length in the Caribbean 
group. As might be expected, the results are correlated with size, 
so that only specimens of relatively large size appear to be well 
separated. Many specimens of the Mexican section resemble the 
Caribbean forms in also having the narrow centrum. 

The above considerations indicate that the fossil belongs to the 
South American section, but it differs from these forms in one 
characteristic of importance to Tihen's classification. He indicates 
(ibid., pp. 165-166) that the suprapterygoid fenestra is not mark- 
edly occluded in the South American section of the imlliceps 
group, while it is often nearly closed by flanges of pterygoid and 
squmosala in members of the Caribbean section. The fenestra is 
completely closed in the fossil, thus indicating a possible relation- 
ship to the latter group in terms of the classification based on 
skeletons of Recent species. However, the suprapterygoid fenestra, 
as a taxonomic character, may be weak in some cases (as Tihen 
realized), owing to its qualitative nature. Tihen points out {ibid., 
p. 174) that B. typhonius of the Caribbean section lacks an oc- 
cluded suprapterygoid fenestra. One specimen of B. peltocephalus 
of the Caribbean section (M. C. Z. no. 23564) also has an open 
fenestra. Within the South American section, one specimen of B. 
paracnemis (M. C. Z. no. 343) has the fenestra closed on one side 
of the skull. It is possible that closure of the fenestra in Recent 
specimens may be both variable within the species as a whole, 
and be partly a function of age of the specimen. In addition, since 
there has been a trend in many groups (including anurans) toward 
deossification, geologic age can be a modifying factor as well. The 



10 



BREVIORA 



No. 193 



latter is probably the most important with respect to this fossil, 
since a number of regions, e.g. shoulder girdle (see above) and 
prootic show a greater amount of ossification than comparable 
regions in any Recent specimen of either New or Old World Bufo 
seen by us. 

The crista medialis of the humerus is of some interest in this 
specimen, because it is a secondary sexual characteristic in some 
Recent frogs. The crista forms the attachment for the M. flexor 
carpi radialis, which aids in flexing the wrist and is important in 
amplexus. Ecker (1889, pp. 42-43) indicated that the crista was 
present in males of Rana esculenta, R. temporaria, and R. oxy- 
rhinus. Holmes (1924, p. 241), in discussing the same species, 
states that it is present in both males and females, but is more 
prominent in males. Inasmuch as the methods of embrace and 
courtship are more or less uniform throughout the Salientia 
(Noble, 1931, p. Ill), this characteristic may be of similar signif- 
icance in other anurans as well. Table I gives the results that 
were derived from specimens of Bufo related to the fossil. 



Taxon 



TABLE I 
MCZ no. sex 



crista medialis 



B. peltocepha 


ilus 


29600 


9 


absent 


23564 


S 


present 


B. ar en arum 




1263 


9 


absent 


30650 


S 


present 


B. bloinbergi 




29669 


9 


absent 


J^ )t) nriYt )iR hi 


irribilis- 


21439 


9 


present but small 




29028 


S 


absent — small specimen 


B. ictericus 




321 


s ? 


present 



Only a small number of the M. C. Z. collection of New World 
Bufo were sexed at the time of skeletonization, and of the others, 
only a few have the ridge to any great extent. B. marinus horribilis 
(M. C. Z. no. 29028) , a male which lacks the crista medialis, was 
the smallest individual of that species in the osteological col- 
lection and may not have reached maturity. The fossil is thus 
most probably a male, though exceptions such as the above render 
this uncertain. 



SPECIFIC ASSIGNMENT OF THE FOSSIL 

As indicated above, the specimen clearly belongs to the South 
American section. Tihen (1962a, p. 165) includes the following 



1963 MIOCENE TOAD 11 

species witliin this grouj): Bufo arenarum, B. blomhergi, B. chi- 
lensis, B. crucifer, B. ictericus, B. m. marinus, B. m. horribilis, 
and B. paracnemis. The fossil differs strongly from Bufo blombergi 
in having strong supraorbital and postorbital crests. B. arenarum 
has very high cranial crests, which are rugose and slightly flat- 
tened on their dorsalraost surfaces, and has a relatively smooth 
skull surface between the crests — both characters in contrast to 
those of the fossil. B. crucifer has a sculpture pattern much finer 
in texture and more deeply incised than in the fossil, has strong 
parietal crests, and has a relatively smooth skull surface between 
the crests, as in B. arenarum. The remaining taxa are very closely 
related, and all have been included in a single undifferentiated 
species, B. marinus, by some workers, or treated as subspecies or 
full species by others. We have followed Tihen's taxonomy 
(1962a) for convenience in discussion. M. C. Z. specimens of 
B. m. horribilis indicate that this primarily Mexican group tends 
to have thin, fine, almost pitted sculpture, and relatively low 
cranial crests, unlike the fossil. Specimens of B. ictericus contrast 
with the fossil in having prominent parietal crests, though this is 
a variable character. Both skeletons of B. paracnemis available 
to us resemble the fossil in sculpture and crest pattern, but on each 
side have a very strong anterior angulation of the posterior skull 
margin, in contrast to the fossil and the other species included in 
the South American section, though the consistency of this char- 
acter is not determinable at this time. Hence the strongest resem- 
blance between the fossil and any Recent group, on the basis of 
the available specimens, is with Bufo marinus marinus. The tend- 
ency toward obliteration of sutures, the relatively greater ossifica- 
tion in prootics, suprapterygoid fenestra, and shoulder girdle, as 
well as the general robustness of some of the bones themselves, do 
not, in our opinion, warrant nomenclatorial recognition at the 
specific level, but merely reflect the often greater ossification seen 
in many fossils (discussed above on p. 9) and probably indicates 
a stage in the evolution of Bufo marinus in the broad sense. In 
identifying this specimen with the Recent Bufo marinus, we real- 
ize, with Taylor and Smith (1945, p. 541) and J. Savage (1960, p. 
235) that the concept of B. marinus as currently used is a broad 
one, yet further refinement in the assessment of relationships of 
this fossil cannot precede re-evaluation of the Recent Bufo ma- 
rinus complex. 

The beginnings of such a re-evaluation have been provided by 
Bertini and Cei (1962), who studied the serological relationships 



12 BREVIORA No. 193 

between some of the species involved. Bufo marinus, a predom- 
inantly Amazonian aquatic species, appears to be the most 
primitive type, and has given rise to xerophilic continental, and 
moisture-loving coastal populations in the southern half of the 
continent. Identification of this fossil with the populations usually 
referred to as B. 7n. marinus is thus consistent with the northern 
occurrence of the fossil within South America, and with the 
primitive nature of B. m. marinus 

Blair (1963, p. 11) has discussed evolutionary patterns in Bujo, 
and has suggested that the ancestral stock of the marinus group 
"somehow crossed into South America during its time of isolation 
from North America in the Tertiary." There is still insufficient 
fossil material to make any positive statement, but most recent 
work on fossil anurans suggests that by the late Cretaceous or 
Paleocene, family groups were well differentiated, and lines re- 
sembling many modern genera were already present. It is thus 
quite possible that the ancestral stock for the marinus group 
entered South America by a land connection with North America 
which persisted until late Paleocene time. If this is so, then it is 
not necessary to propose that they reached South America by 
some sort of "sweepstakes" method after the connection was 
severed. 

SUMMARY AND CONCLUSIONS 

A late Miocene toad from the upper Magdalena Valley, Huila, 
Colombia, South America, is referred to the Recent species Bufo 
marinus. Within this poorly understood group, it seems to show 
closest resemblance to B. m. marinus from northern South 
America. It differs from B. marinus only in having slightly greater 
ossification in the suprapterygoid fenestra, lateral parts of the 
prootics, and glenoid region of the shoulder girdle. This condition 
is insufficient evidence for recognizing a taxonomic difference 
from B. marinus, at least until restudy of the Recent forms is 
efi'ected. The broad geograi)hic and altitudinal range of the Recent 
species precludes ecologic interpretation. The species is aquatic, 
and common today along large river courses and may have been 
so on the Miocene floodplains as well. The fossil, in combination 
with the lizards described by Estes (1961), is another indication 
of the modernity of the hcrpetological elements of the late Mio- 
cene La Venta fauna, and of the greater extent of the floodplain- 
aquatic habitat in northern South America during the late 
Miocene. 



1963 MIOCENE TOAD 13 

REFERENCES CITED 

Bertini, F., and J. M. Cei 

1962. Seroprotein patterns in the Bufo mnrinus complex. Hcrpetologica, 
17: 231-238, 3 figs., 1 table. 

Blair. W. F. 

1963. Evolutionary relationships of North American toads of the genus 
Bujo: a progress report. Evolution, 17: 1-16, 4 figs. 

ECKER, A. 

1889. The anatomy of the frog. Oxford, Clarendon Press, translated 
by George Haslam, pp. i-x, 1^49, 261 figs., 2 pis. 
ESTES, R. 

1961. Miocene lizards from Colombia, South America. Breviora, Mus. 
Comp. Zool., Harvard Univ., no. 123: 1-11, 5 figs. 
Fields, R. W. 

1959. Geology of the La Venta badlands, Colombia, South America. 
Univ. Calif. Publ. Geol. Sci., 32: 408^44, 2 figs., 4 pis., 2 maps. 

Holmes, S. J. 

1924. The biology of the frog. MacMillan and Co., New York, pp. 
v-ix, 1-370, 94 figs. 
Noble, G. K. 

1931. The biology of the Amphibia. McGraw Hill and Co., New York, 
pp. 1-577, 174 figs. 
Proctor, J. B. 

1921. On the variation of the scapula in the batrachian groups Aglossa 
and Arcifera. Proc. Zool. Soc. London, 1921 ; 197-214, 10 figs. 
S.WAGE, D. E. 

1951. Report on fossil vertebrates from the upper Magdalena Valley, 
Colombia. Science 114: 186-187. 
Savage, J. M. 

1960. Geographic variation in the tadpole of the toad, Bufo marinus. 
Copeia, 1960 (3) : 233-235, 3 figs. 

Taylor, E. H., and H. M. Smith 

1945. Summary of the collections of amphibians made under the 
Walter Rathbone Bacon Traveling Scholarship. Proc. U.S. Nat. 
Mus., 95 : 521-613, 4 figs., 15 pis. 

TlHEN, J. A. 

1962a. Osteological observations on New World Bujo. Amer. Midi. 

Naturalist, 67: 157-183, 38 figs. 
1962b. A review of New World fossil bufonids. Ibid., 68: 1-50. 



v^ 



BREVIORA 

Museiiim of Comparative Zoology 



Cambridge, Mass. December 31, 1963 Number 194 

A NEW SUBSPECIES OF TROPIDOPHIS GREENWAYI 
FROM THE CAICOS BANK 

By Albert Schwartz 

The species of small boa, TropidopJiis greenwayi, has hereto- 
fore been known only from Ambergris Cay.^ No additional speci- 
mens of T. greenwayi have been taken since the type and paratype 
were collected in 1936. At the time of their review of the Carib- 
bean snakes of the genus TropidopJiis, Schwartz and Marsh 
(1960) examined the two extant specimens and separated them 
from the Cuban and Bahaman species T. pardalis (with which 
they had been nomenclatorially associated) and T. canus (to 
which they might be expected on geographic grounds to be 
related). 

From January 11 to 22, 1961, the writer and David C. Leber 
collected on South Caicos and its adjacent Long Cay and visited 
the Ambergris Cays as well in order to secure additional speci- 
mens of T. greenivayi and to ascertain if this snake were more 
widely distributed on the Caicos Bank. We were unable to find 
the boa on the Ambergris Cays, but a series of fourteen individ- 
uals was collected for us on South Caicos and Long Cay by resi- 
dents of these islands. Comparison of this large series with the 
two specimens of topotypic T. greenwayi indicates that the South 
Caicos and Long Cay populations represent a different form, 
which may be called : 

Tropidophis greenwayi lanthanus, new subspecies 

Type: Museum of Comparative Zoology (MCZ) 69630, from 
0.5 mi. north of Cockburn Harbour, South Caicos, taken 22 Janu- 
ary, 1961, by a native for A. Schwartz. 

Paratypes: MCZ 69619, 0.5 mi. east of Cockburn Harbour, 
South Caicos, 13 January, 1961 ; MCZ 69632, same locality, 14 

lActually, "Ambergris Cay" is one of a pair of cays, both known as the Amber- 
gris Cays, which lie about 13 miles southwest of Cockburn Harbour on South 
Caicos. 



BREVIORA 



No. 194 



January, 1961 ; MCZ 69620, same locality, 19 January, 1961 ; 
MCZ 69621, 7 mi. northeast of Cockburn Harbour, South Caicos, 

20 January, 1961 ; MCZ 69622, 0.5 mi. east of Cockburn Harbour, 

21 January, 1961 ; MCZ 69623, Cockburn Harbour, 21 January, 
1961; MCZ 69624-25, Long Cay, off Cockburn Harbour, South 
Caicos, 21 January, 1961 ; MCZ 69626-28, same locality, 22 Janu- 
ary, 1961 ; MCZ 69629, 69631, same data as type. All specimens 
were collected by natives for A. Schwartz. 

Distribution: Known only from South Caicos and adjacent 
Long' Cay on the Caicos Bank. 

Diagnosis: A subspecies of T. grcenwayi differing from the 
nominate form in higher number of ventral scutes and in colora- 
tion and pattern. 

Description of type: An adult spurred male with the following 
measurements and counts: total length, 257 mm., tail, 30 mm.; 
ventral scutes 160, subcaudal scutes 28 ; supralabials 9/10 ; infra- 
labials 11/11; parietals in contact; preoculars 1/1, postoculars 
3/2 ; dorsal scales smooth, rows 25-27-19 ; dorsal paramedian 
blotches 42/49 ; tail blotches four. Coloration : Head uniformly 




Figure 1. Dorsal uiidbody pattern of Tropidoplils grcenwayi lanthanus 
(MCZ 69630, type). 



1963 NEW SUBSPECIES OF TROPIDOPHIS GREENWAYI 3 

dark brown dorsally; neck slightly paler, quickly grading pos- 
teriorly into a grayish-tan middorsal zone about ten scales wide, 
this zone including the two paramedian rows of dark brown dor- 
sal blotches, each blotch faintly outlined with pale gray, the dorsal 
/one continuing posteriorly onto the upper surface of the tail, 
where it merges imperceptil)ly with the yellow coloration of the 
tail tip. Sides darker brown, becoming more reddish ventrally, 
and enclosing three rows of lateral blotches, the uppermost two 
rows the least conspicuous (because of the closeness of their color 
to that of the ground color and the reduction of pale outlining), 
the lowermost row outlined by gray especially along the pos- 
terior margins. A^entral surface reddish-tan with two rows of dark 
brown blotches, which may extend dorsally onto the first two 
scale rows, do not coalesce medially on the venter, and are out- 
lined posteriorly in white. Supralabials dark brown, the pos- 
terior three somewhat flecked and blotched with cream. Ilemi- 
penes extruded, weakly bifurcate and weakly s]iinose distally. 

Paratypes: The paratypes include three males and ten females : 
I cannot discern any difference in coloration, pattern or scalation 
between the sexes nor between the snakes from South Caicos and 
Long Cay, and all are discussed as one unit. All three males are 
spurred, as is the type. Two of the males are juveniles, each with 
a total length of 140 mm. Males range in total length from 140 
mm. to 320 mm., whereas females vary between 250 mm. and 336 
nun. Ventrals range between 156 and 165 (mean 161.2), caudals 
between 26 and 30 (mean 28.0). All have 1/1 preoculars. Post- 
oculars vary from 2/2 (two snakes), 2/3 (six, including type), 
and 3/3 (five), thus showing a tendency toward having three 
postoculars in 85 per cent of the population ; one snake has the 
head scales so damaged that its data are omitted. Paramedian 
blotches vary from 27 to 49 in the series, the largest discrepancy 
between right and left counts on the same snake being seven 
blotches, as in the type and one other individual. The average 
blotch number is 39.9. One snake, as noted below, lacks blotches 
completely. Scale rows at midbody are either 25 (eleven snakes) 
or 27 (three snakes). The number of rows of blotches around the 
body are eight or ten, with only one snake showing the reduction 
to eight rows of blotches. The tail/total length ratio (X 100) 
averages 10.5 (9.4—12.1 ; the highest ratio is shown by one of the 
juvenile males). All of the paratypes have the parietals in con- 
tact, and have the dorsal scales smooth. Upper labials vary from 
9 to 10, and lower labials from 9 to 12. 



4 BREVIORA No. 194 

Coloration of the paratypes : All the adult paratypes except 
one agree very closely with the type. All have a lighter median 
zone containing two rows of paramedian blotches ; this middorsal 
zone was noted in life as matching in various individuals (color 
notations from Maerz and Paul, 1950) : PL MAS, PI. 15C7, PI. 
13A6, PL 14A4, PL 12A6, all of which are shades of buffy tan to 
grayish tan. The sides are somewhat darker (PL 15A9, PL 16 A6, 
PL 13A10, PL 15C8, PL 15C8), then becoming lighter on the 
lower scale rows and venter to a more red coloration (PL 14G10, 
PL 15A4, for example). The blotches themselves are always dark 
brown. The venter varies from a rich reddish tan to an almost 
chocolate brown, at times relieved by rather extensive white bor- 
ders to the brown ventral blotches. One snake, an adult female, is 
unusual in that the pattern consists merely of the paler mid- 
dorsal zone and darker sides, without any indication of lateral or 
ventral blotches ; the paramedian blotch rows are represented by 
a rather diffuse dark brown smudging along the middle of the 
back. The tail tips are dark in two snakes and light (yellow) in 
ten ; two are indeterminate. 

The two juvenile paratypes are very like the adults in colora- 
tion and pattern ; the pattern elements are not appreciably 
brighter than in the adults (in contrast to juvenile and adult T. 
canus where the juveniles show the dorsal pattern much more 
distinctly than the adults). There is also no evidence of the 
lateral nuchal stripe which is a common feature of both juvenile 
and adult T. canus (Schwartz and Marsh, op. cit.: 61) and espe- 
cially prominent in the juveniles. 

In addition, the young paratypes have white, rather than 
brown, venters; apparently the darker pigmentation comes with 
increase in size. 

Comparisons: T. g. lanthanus requires comparison only with 
the typical form. It is quite distinct from the adjacent Bahaman 
and Ilispaniolan forms of Tropidophis. The comparison is ham- 
pered, however, by the paucity of specimens of the nominate form. 
If the type and paratype represent a fair sample of T. grccnwayi 
on the Ambergris Cays (and it is possible that they do not), then 
laniliayius is certainly very distinct from the snakes on these out- 
lying cays. Comparison of tlie illustration (Figure 1) of T. g. 
lanthanus with that of the type of T. g. greenwayi (Schwartz and 
^larsh, op. cit.: fig. 7) at once demonstrates the pattern differences. 
In the nominate form there is no indication of the dorsal })ale 
zone, the entire dorsal surface has a mottled or stippled effect, 



1963 NEW SUBSPECIES OF TROPIDOPHIS GREENWAYI 5 

with blacks, browns, tans, and whites more or less intermixed ; the 
blotches are extremely obscured by the mottled and stippled mark- 
ings on the interspaces. Such is not the case in lanthanus where 
in all specimens but one (the blotchless female mentioned above) 
the blotches are quite distinct and there is no interspace stippling. 
It might be argued that the greenwaiji coloration is a peculiarity 
of extreme age or adulthood ; in fact the type of greenwayi is the 
largest specimen of the species presently available. However, in 
the largest specimen of lanthanus, the trend is obviously just the 
reverse of that in the nominate snakes, and the pattern is com- 
pletely obliterated except for the dorsal pale zone. Another fea- 
ture of greenwayi is the "salt-and-pepper" effect on the dorsal 
surface of the head ; such a condition does not occur in lanthanus, 
where the head is always uniformly dark brown. In number of 
ventrals, lanthanus embraces the 157-158 ventral counts known 
from the two greenwayi (both are males). However, the specimen 
of lanthanus which has the lowest count for that race (a female 
with 155) stands alone in the series; all other lanthanus have 
counts ranging from 160 to 165. I regard this single specimen as 
being somewhat aberrant. When large series of topotypic 
greenwayi finally become available, I suspect that they will have 
ventral counts low^er than those of lanthanus. 

Both specimens of greenwayi have 2/2 postoculars, w^hereas 
only two of thirteen lanthanus have such a count; the remainder 
have counts of 2/3 or 3/3, the third postocular being wedged be- 
tween the fifth and sixth supralabials. It is possible that a ten- 
dency toward 3/3 postoculars is characteristic of lanthanus. 

Remarks: Perhaps the most interesting feature of the new 
specimens of T. greenwayi here reported is their uniformity. For 
example, the species was partly defined (Barbour and Shreve, 
1936:2) by having the parietals in contact. Schwartz and Marsh 
{op. cit. :57) noted that this character, as with all scale charac- 
ters in this assemblage of small boas, was variable ; the same com- 
ment applies equally to dorsal scale earination and number of 
dorsal scale rows. But all new specimens of T. greenwayi do indeed 
have parietals in contact and smooth scales. In fact, T. greeyiwayi 
can be in addition characterized as being a Tropidophis with 
smooth dorsals usuallv in 25 scale rows and tvpicallv ten rows of 
blotches, all features which were uncertain at the time of the 
review of the Caribbean Tropidophis. Likewise, additional data 
are now available on tail/total length ratio in T. greenwayi ; the 
ratio averages 10.5, which is the lowest mean of any member of 



6 BREVIORA No. 194 

the assemblage, most closely approached by that of T. canus 
canus (10.7). 

Even with additional material available, I am unable to guess 
as to the affinities and origin of T. grccnwayi. With two juveniles 
now at hand, the relationships of this species to T. canus seems 
even more remote than it did previously (Schwartz and Marsh. 
op. cit. : 62). This snake appears to have been long isolated from 
its relatives, whatever they may have been, and other than to 
remark that it is a member of the pardalis-maculatuf; assemblage, 
little can be said. 

I suspect that T. greemvayi will be found to occur on the other 
major islands of the Caicos Bank. It is surprising that it has been 
so long overlooked on South Caicos where the natives knew of it 
as soon as approached about small snakes. The name lanthanum 
is an allusion to the fact that the species has been overlooked on 
South Caicos. This boa appears to be genuinely lacking from 
Grand Turk and probably other islands on the Turks Bank, which 
is separated from the Caicos Bank by the Turks Island Passage : 
natives there were aware of Epicrates on some of the outer cays, 
but did not know of any snakes at all on Grand Turk. No snakes 
of any species were encountered in the one week spent by us on 
Grand Turk. 

Figure 1 was executed by Ronald F. Klinikowski. I wish to 
thank him for his efforts on my behalf. 

LITERATURE CITED 

Barbour, Thomas, and Benjamin Shreve 

1936. New races of Tiopidophis and of Ameiva from the Balianias. 
Proc. New England Zool. Club, 16: 1-3. 

Maerz, a., and Rea Paul 

1950. A dictionary of color. New York, McGraw-Hill Book Co., ])p. 
i-vii, 1-23, 137-208, 56 pis. 
SrHWAKTZ, Albert, and Robert J. Marsh 

1960. A review of the paradalis-maculatus complex of the boid genus 
Tropidophis of the West Indies. Bull. Mus. Conip. Zool., 123(2) : 
r)0-84, 10 figs. 




BREVIORA 



Miisemnti of Comparative Zoology 

Cambridge, Mass. December 31, 1963 Number 195 

CAYMAN ISLANDS TR0PID0PHI8 (REPTILIA, 

SERPENTES) 

By Richard Thomas 



New material from the Cayman Islands, gathered principally 
through the efforts of Dr. Albert Schwartz during his collecting 
in the West Indies, has provided a large enough series of Tropi- 
dopJiis from each of the three islands so that a clearer picture 
of the relationships can be seen. Specimens examined have 
been under the curatorship of Dr. Doris Cochran, United 
States National Museum (USNM), Dr. Ernest Williams, Mu- 
seum of Comparative Zoology (MCZ), Dr. Albert Schwartz, 
field collection (AS), and in the author's private collection (RT). 
The illustrations are the work of Ronald F. Klinikowski. 

The Cayman group consists of three islands : Grand Cayman, 
Ca.yman Brae, and Little Cayman. Grand Cayman is situated 
about 180 miles west-north-west of Jamaica and 150 miles south 
of Cuba ; it is about 22 miles long and eight miles wide at maxi- 
mum width. Cayman Brae, 89 miles east-north-east of Grand 
Cayman, is about 12 miles long with an average width of a mile. 
Little Cayman, five miles to the west of Cayman Brae, is ten 
miles long with a maximum width of two miles. The long axis 
of each island lies in a generally east-west direction. They are 
low-lying islands formed of calcareous rocks. 

Battersby (1938) described Tropidophis mcJanurus ca^iman- 
ensis from Grand Cayman; Grant (1940) further recorded T. m. 
cayDiancnsis from Cayman Brae and described Trojjidophis 
parkeri from Little Cayman. Schwartz and Thomas (1960) on 
the basis of obvious affinities with melanurus called parlicri a 
subspecies of T. melanurus. 

The new material recently acquired from the Caymans indi- 
cates that, despite affinities of the Cayman Tropidophis with T. 
melanurus, these snakes are distinct and should be known as 



2 BREVIORA No. 195 

Tropidophis caymanensis caymanensis and Tropidophis cayman- 
ensis parkeri. Tropidophis caymanensis may be distinguished 
from Tropidophis melanurus by the following characters: 

1. Size : The average total length of caymanensis from all 
three islands (48 specimens) is 369.5 mm with a range of 202-564 
mm. The average size of melanurus from Cuba and the Isle of 
Pines (77 specimens) is 607.1 mm with a range of 253-1057 mm. 

2. Scale-rows : Caymanian Tropidophis have usual scale-row 
counts of 23-25-17 (caymanensis), 25-27-17 (parkeri), or 23- 
25-17 (Brae Tropidophis). Tropidophis melanurus has a usual 
scale-row formula of 25-27-19. 

3. Light tail tip : All specimens of Caymanian Tropidophis 
examined have a light tail tip ; of 16 specimens of parkeri seen 
in the field all but three (with incomplete tails) had yellow tail 
tips, and one of the three with incomplete tails showed traces of 
yellow pigmentation on the remaining tip. Old specimens show 
a slight darkening on the dorsum of the tip, but the rest of the 
tip is yellow. Specimens from Grand Cayman and Cayman Brae, 
which were not seen alive by the writer, have plain but dis- 
tinctly light tail tips, presumably as a result of the yellow fading 
in preservation; field color notes on one adult (AS16230) from 
Grand Cayman describe a yellow tail tip. In mclanurns the yel- 
low tail tip is a distinctly juvenile condition and in the adult 
becomes either unicolor with the ground color or black. 

4. Dorsal body pattern : In caymanensis, the two rows of 
paramedian dorsal spots are squarish (2-4 scales long by 2-3 
scales wide) and distinct from the ground color; these spots are 
more discrete than in melanurus, with pale edging frequently 
occurring on the median edges, and in addition they are distinct 
from the middorsal stripe. T. melanurus typically has para- 
median dorsal spots much less distinctly developed and they are 
apt to be broadly confluent at the midline and not distinct from 
the middorsal stripe, when the latter is present. The light edg- 
ing is confined to the outer edges of the spots. 

Tropidophis caytnavensis is, therefore, a species with closest 
affinities to melanurus but distinguished from that species by its 
dwarf size, lower number of scale rows, persistence of the yellow 
tail tip in adults and bolder dorsal spotting. 

Examination of the series of 16 specimens from Cayman Brae 
shows that they may be distinguished from the Grand Cayman 
population with which they had previously been considered 
identical. I take great pleasure in naming this new subspecies 



1963 



CAYMAN ISLANDS TROPIDOPIIIS 



for Dr. Albert Schwartz, who suggested this problem, in honor 
of his zoological work in the West Indies; it may therefore be 
known as : 

Tropidophis caymanensis schwartzi subsp. nov. 

Type: MCZ 69618, an adult male from The Creek, 8 mi. NE of 
West End, Cayman Brae, collected 25 August 1961 by a native 
for A. Schwartz. 

Paratypes: MCZ 69603-04, Cayman Brae, The Creek, 8 mi. 
NE of West End, 22 August 1961 ; MCZ 69605, Cayman Brae, 
6 mi. NE of West End, 23 August 1961 ; MCZ 69606, Cayman 
Brae, The Creek, 8 mi. NE of West End, 23 August 1961 ; MCZ 
69607-608, Cayman Brae, The Creek, 8 mi. NE of West End, 24 
August 1961; MCZ 69609-11, Cayman Brae, West End, 25 Au- 
gust 1961 ; MCZ 69612-13, Cayman Brae, Spot Bay, 25 August 
1961 ; MCZ 69614-17, Cayman Brae, The Creek, 8 mi. NE of West 
End, 25 August 1961. All specimens were collected by natives 
for A. Schwartz. 





I 



Fig. 1. Dorsal cephalic pattern of Tropidophis caymanensis sclnvartsi 
(MCZ 69618, type). 

Fig. 2. Dorsal cephalic pattern of Tropidophis caipnanensis cai/manensis 
(AS 16230). 



4 BREVIORA No. 195 

Diagnosis: a race of caymanensis with a normal scale-ro^\ 
count of 25-25-15, rather than 23-25-17 (T. c. cayma7icnsis) or 
25-27-17 {T. c. porkcri), differing also from T. caymanensis in 
the coloration of the head and in a higher mean ventral count 
and from T. c. parl-cri in a somewhat lower mean ventral count. 

Description of type: An adult male, total length 485 mm, 
tail 50 mm. Body relatively stout, slightly compressed, head 
distinct from neck; scales weakly keeled except for five lower- 
most rows on each side, smooth scales becoming reduced to three 
lowermost rows posteriorly ; middorsal row enlarged and bi- 
earinate on posterior fiftli of body, ending anterior to vent ; 
dorsal scale-rows 25-25-17 ; ventrals 200 ; subcaudals 39 ; anal 
single ; spurs present ; preoculars 1/1 ; postoculars 3/3 ; temporals 
3-4/3-4; supralabials 10/10, 4 and 5 entering the orbit; infra- 
labials 12/12. Dorsal pattern : two rows of 55 and 61 parame- 
dian dark spots, 2 to ^/U scales long by 2 to 2V> scales wide : 
light edging to spots mostly lateral, but some median edging 
occurs also ; lighter middorsal stripe present, broadly connected 
to median edges of spots. Sixth and ninth scale rows with fine, 
light centered, dark lines; row of small spots betw'een lines on 
sixth and ninth scale-rows ; two rows of staggered, dark spots on 
lower five scale-rows ; venter irregularly stippled with dark 
pigment (see Figure 3) ; lowermost row of lateral spots becom- 
ing enlarged and migrating to venter on posterior fifth of l)()dy: 
about seven spots on dorsum of tail ; subcaudal surface with two 
rows of al)()ut seven dark spots. Dorsal ground color of body 
light grey changing to cream below sixth scale-row and on ven- 
ter; tip of tail light. (Vphalic pattern: trapezoidal, dark, dor- 
sal ce]ihalic figure mucli invaded liy light stippling (see Figure 
1 ) ; posterior center of figure light ; anterior portion of cephalic 
figure separated off to form interocular bar; nipple-like, ante- 
rior projection of interocular bar not extending beyond frontal ; 
dark, oval spot on internasals and prefrontals narrowly joined 
on each side to preocular portion of lateral head stripe, which 
contiiuics posteriorly onto neck to level of ninth ventral after 
which it is broken u]i into a lateral row of spots. 

VaridHini : The paratypes include seven males and nine fe- 
males; the largest male is 435 mm (the type); the lai-gest fe- 
male is 358 mm. The males of tliis series average larger than the 
females, having an av(>rage total lengt'i of 362.8 nun (202-425) : 
females avei-a<?,'e 336.8 mm (297-358). Ventrals in males, including 
th(> type, average 195.6 (191-202) ; females 200 (195-205) : sub- 
caudals in males average 36.6 (34-39), in females 36.2 (31-39). 



1963 CAYAIAN ISLANDS TROPIOOPITIS 5 

All specimens have 1/1 preoculars and 3/3 postoeiilars ; all have 
the st-ale-row formula of 25-25-17 except for two with counts of 
25-25-15. The model supralabial count is 10/10 with other counts 
of 12/10 (1) and 9/10 (2). Tails of males average 10.6 per cent 
of total length with a range of 9.1-11.8 per cent ; tails of females 
average 11.2 per cent of total length (10.8-12.07f ). 

Pattern and color are fairly uniform in the paratype series. 
In all of the specimens the dorsal cephalic pattern is much in- 
vaded and broken up by solid, light areas and by heavy stip- 
pling- of light pigment ; in all examples but three the interoeular 
])igniented area is separated from the cephalic figure to form 
an interoeular bar; in the three exceptional specimens the inter- 
oeular bar is but narrowly joined to the dorsal cephalic figure 
by a median strip of dark pigment made hazy by light stippling. 
Typically there are several (usually 4) dark spots on each side 
of the lower labials. The dorsal pattern in the paratype series 
is much the same as in the type ; the average number of dorsal 
spots is 60.1 in males and 58.9 in females. Ventral coloration 
in the paratype series varies from a uniformly clear cream to a 
rather heavy, irregular, dark mottling over nearly the entire 
undersurface ; the typical ventral coloration for the series is like 
that described for the type. All specimens, except one with an 
incomplete tail, have light tail tips. 

Comparisons: A recently acquired series of 16 T. c. parkeri 
agree with Grant's (1940) original diagnosis of it as a 27 scale- 
row form with a ventral count of over 200; the ventrals of the 
nine males average 201.0 (199-204), those of the females 205.1 
(201-212) ; the normal scale-row count for the series is 25-27-17 
with variants of 25-27-16 (1), 25-27-18 (1), and 25-27-19 (1). 
Sell wart zi may be distinguished from parkeri by the possession 
of a normal scale-row count of 25-25-15. Chromatically there 
does not seem to be much difference in the populations; the 
fresh parkeri material has a richer, deeper coloration, whereas 
schivartzi is much lighter. The difference, however, is likely due 
to fading, since field color notes on one specimen of schwartzi 
("dorsal ground color orange tan; dorsal blotches brown, not 
outlined posteriorly with orange") agree with the typical parkeri 
coloration. Parkeri tail tips are yellow; those of scJiwartzi are 
definitely light and probably faded from the yellow condition. 
The cephalic figure in parkeri tends to be broken up and in- 
vaded with lighter pigmentation much as in sch^rarfzi, though 
the figure is frequently broken into two or three disjunct por- 
tions in parkeri as o]:)posed to simple invasion by light pigment 



6 



BREVIORA 



No. 195 



in schwartzi to form an erose margin and light center. The aver- 
age number of paramedian, dorsal spots in parkeri is 54.7 for 
males and 55.9 for females, slightly lower than the averages for 
schwartzi. In parkeri the females average larger than males; 11 
females average 400.3 mm (230-564) ; nine males average 373.3 
mm (306-444) ; the tails of male parkeri average 10.6 per cent 




Fig. 3. Dorsal midbody i)attern of Tropidophis caymanensis schwartzi 
(MCZ 69618, type). 



of the total length, those of females 9.4 per cent. The slight sex- 
ual dimorphism evident in average total length and average 
tail/total length ratios for schwartzi is present but reversed in 
parkeri. 

The ten specimens of caymanensis which were examined have 
a modal scale-row count of 23-25-17 with variants of 23-25-19 
(1) and 25-25-19 (1). These agree closely with Battersby's 
(1938) type and paratype which had scale-row counts of 23- 
25-17. Schwartzi is distinguishable from caytuancnsis by the 
possession of 25-25-17 scale-rows. Grant (1940) thought that 
the type of caymanensis Avas abnormal in having only 191 ven- 
trals ; he also mentioned a male with 183 ventrals. In his analysis, 



1963 CAYMAN ISLANDS TROPIDOPIIIS 7 

he apparently omitted both of these specimens from considera- 
tion and speculated that they might be members of a localized 
population along the north coast characterized by a reduced 
number of ventrals. The material at hand shows that similar 
low ventral counts are not restricted to the north coast; counts 
of 183, 187 and 191 are found on specimens from Georgetown 
in the southwest. Grant also mentioned five females and five 
males in his own collection which, along with the female para- 
type, average 205 and 193 ventrals, respectively. However, he 
only listed seven specimens in his own collection from Grand 
Cayman ; just what the provenance of the other three specimens 
is cannot be determined. Counts from the new material plus the 
counts on the type, paratype, and the individual with 183 ven- 
trals noted by Grant (Br. Mus. No. 1939.2.3.78) yield an av- 
erage of 192.6 (183-200) for seven females, 191.7 (183-195) 
for six males. Therefore, caymanensis is seen to average 
lower in ventrals than parkeri and schwartzi, there being 
almost no overlap between caymanensis (183-200) and parkeri 
(199-212). There are apparently no significant differences in 
body pattern and color between caymanensis and schwartzi. 
However, schwartzi can be distinguished from caymanensis on 
the basis of the cephalic figure. In caymanensis the cephalic 
figure is much more solidly pigmented with black and uniform 
in outline ; the edges are not highly indented and eroded by 
invasion of light pigment as in schwartzi, and the light areas 
within the figure are more restricted and well defined, not exten- 
sive and hazy in outline by virtue of light stippling (see Figure 
2). Seven female caymanensis average 392.0 mm total length 
(222-488) and five males average 368.8 mm (192-538) ; tails of 
males average 12.2 per cent of the total length, those of females 
average 10.8 per cent of the total length. Once again, apparent 
sexual dimorphism is the reverse of that in schivartzi. 

Remarks: The author was able to do some collecting on Little 
Cayman in December of 1962. Some impressions were gathered 
concerning Tropidophis c. parkeri and are probably applicable 
to the species generally. A series of sixteen parkeri was col- 
lected during the visit to Little Cayman. These snakes gave the 
impression of being more secretive than T. melanurns, which is 
most frequently found prowling abroad at night. Parkeri was 
never collected in the open ; the majority were found beneath 
fragments of limestone which litter the island. Two were taken 
within rocks themselves, being secreted in eroded chambers ; one 
was taken among the leaves of a dead Agave, and another was 



8 BREVIORA No. 195 

found three feet off the gTOuiid in the leaf base of an epiphyte. 
One specimen was found approximately eight feet above ground 
in the roof of an outhouse, where it was partly concealed beneath 
corrugation in the tin roof. One specimen was found beneath 
rocks and litter under a sea-grape tree; it was this "beach situ- 
ation" wherein the natives claimed the snake to be most often 
found. A large female specimen found resting in a tin can dis- 
gorged a large Hula septe7ifrwnaUs shortly after capture. Many 
specimens autohemorrhaged freely upon capture or at subsequent 
disturbance; the "shutter" effect noted by Hecht, Walters and 
Ramm (1955) for Bimini Tropidophis was also observed; this 
effect is produced by the sudden and transient appearance of 
blood under the brille during autohemorrhage. Most of these 
snakes coiled in a tight, defensive ball and would so remain for 
long periods with their heads completely hidden. This habit is 
not in keeping with melanurus, which might be said to be less 
timid; experience with melanurus in the field indicates also that 
Contparafivc material: Tropidophis caymanensis caymanensis : 
USNM 108042-43, Grand Cayman; MCZ 44862-4, Grand Cay- 
man; AS 16209-11, 16230, Grand Cayman, Georgetown; AS 
X4876, Grand Cayman, Georgetown. Tropidophis c. parheri: 
rSNM 108009, Little Cayman; MCZ 44865 (type). Little Cay- 
man; MCZ 44866-8 (paratypes). Little Cayman; AS X489b- 
X4904, Little Cayman, western end; RT 36, Little Cayman, 
western end. Tropidophis melanurus: 79 specimens, Cuba and 
the Isles of Pines. 

LITERATURE CITED 

Anonymous 

1961. Cayman Islands: Report for the rears 1959 and 196U. London: 
Her Majesty's Stationery Office, pp. 1-46: 10 figs., 1 map. 
Battersby, J. C. 

1938. Some snakes of the genus Tropidophis. Ann. ilag. Nat. Hist., 
(11) 1:557-560. 
Grant, Chapman 

1940. The herpetology of the Cayman Islands. Bull. Inst. Jamaica, 
Sei. Scr., 2:1-56, 13 figs. 
Hecht, Max K., Vladimir Walters and Gordon Ramm 

1955. Observations on the natural history of the Bahaman pigmy boa, 
Tropidophis pardalis, with notes on autolieinonliage. Copeia, 
Xo. 3:249-251. 
Schwartz, Albert and Richard Thomas 

1960. Four new snakes (Tropidophis, Dromiciis, Alsopliis) from the 
Isla de Pinos and Cul)a. Herpetologiia, 16('2) :73-90. 



^>euo 



BREVIORA 



MnaseTLaim of Compsirative Zoology 



Cambridge, Mass. December 31, 1963 Number 196 

A NEW GENUS AND SPECIES OF BATHYPELAGIC 

OPHIDIOID FISH FROM THE WESTERN 

NORTH ATLANTIC 

By Daniel M. Cohen 

U. S. Fish and Wildlife Service 
Washington, D. C. 

Through the kindness of Dr. Richard H. Backus of the Woods 
Hole Oceanographie Institution and Dr. Giles W. Mead of the 
Museum of Comparative Zoology I have had the opportunity 
to study the three fishes reported upon below. Although they 
were taken off the southern edge of Georges Bank, hardly a 
zoologically unknown area, they represent the first of their kind 
ever captured. And, indeed, they are so different from any other 
known ophidioid that still another name must needs be added to 
the already lengthy roster of genera in this group. 

ThaLASSOBATHIA gen. nov. 

Diagnosis. Chin barbel absent. Vertical fins continuoiLS; 
ventral fins each with two rays, originating close to the level of 
the posterior margin of the preopercle and a short distance behind 
the symphysis of the cleithra ; the ventral fins covered with thick 
skin, short, extending only to the level of the origin of the pec- 
toral fin; pectoral fin rounded, without separate elongated rays. 
Branchiostegal membranes separate ; gill rakers reduced in num- 
ber and size ; branchiostegal rays seven. Preopercular spine pres- 
ent beneath skin. Anterior nostril lacking a tube, placed high 
on snout. Teeth present on premaxillary, vomer, palatine and 
dentary, lacking on basibranchials. Maxillary not expanded pos- 
teriorly, not sheathed. Premaxillary not protractile, bound to 
the snout anteriorly by a broad frenum. Head compressed, its 
height greater than its width. Eyes well developed. Body 
compressed, relatively short and stubby. Two lateral lines. 



2 BREVIORA No. 196 

Peritoneum dark. Swimbladder reduced, its cavity completely 
occluded ; bladder not bolind to vertebrae. Neural spines on 
abdominal centra not depressed ; their tips not truncate. 

Type species. Thalassohatlua pclagica sp. nov. 

The name Thalassohathia is derived from the Greek GaXaaaa, 
the sea ; ^aO'jq, deep. The gender is feminine. 

Relationships. The main problem encountered in establishing 
this genus has been to determine its relationships rather than to 
distinguish it from any presently known genera. The curiously 
reduced ventral fins and the general shape and consistency of the 
body might suggest relationship with the liparid fishes ; however, 
there is not the slightest trace of either a suborbital stay or a 
sucking disc. In addition, the branchiostegal membranes are sep- 
arate from each other and from the isthmus, and the gill opening 
extends far up on the side of the body in typical ophidioid 
fashion. The last character also separates ThalassohafJiia from 
the Zoarcidae (not including Lycodapus) . The non-protractile 
premaxillaries and attached broad frenum suggest the condition 
of the upper jaw found in salariine blennnies ; however, very 
different dentition, branchiostegal membranes, ventral fins, caudal 
fin connections and other characters separate Thalassohathia 
from the Salariinae. In addition, the absence of a one-to-one 
correspondence between dorsal and anal fin rays and vertebral 
centra separates Thalassohathia from any of the blennioids. 

Among the ophidioids the closest known relatives of Thalasso- 
hathia appear to be the Bythites — Oligopus — Cataetyx group 
of genera recently discussed by Cohen (in press). Some reasons 
for deriving Thalassohathia from this group are the similarly 
compressed and foreshortened body, the typical bythitine denti- 
tion, and especially the double lateral line system, which greatly 
resembles that found in some of the species of Oligopus (Cohen, 
in press). 

Although at first glance Thalassohatlna does not appear to be 
a bathypelagic fish, closer examination reveals that many of its 
characteristics are adaptations to a midwater life. The skeleton 
is so poorly ossified that X-ray photographs are very indistinct. 
Scales are completely lacking. The swimbladder is a small, tissue- 
filled organ similar to those which have been found in many other 
species of bathypelagic fishes. The above characters and their 
significance in floating fishes have been treated by Denton and 
Marshall (1958) and Marshall (1!)()0). Other structures wbich 
may help to keep Thalassohathia aloft are its subdermal fatty 



1963 NEW OPTTIDIOID FISH 3 

layer and enlarged liver. Among the most characteristic features 
of ophidioid fishes are the naked ventral fin rays which probably 
serve as tactile organs in benthic species (Herald, 1953, and 
serve as tactile organs in benthic species (Herald, 1953 and 
Briggs and Caldwell, 1955, have confirmed this for two species of 
benthic Ophidiidae). These structures would of course be useless 
in midwater, and Thalassobathia has reduced ventral fin rays 
which are covered with thick skin. 

Because of its midwater habitat, Thalassohathia should also be 
compared with the ophidioid subfamily Aphyoninae (Nybelin, 
1957) which comprises a small group of bathypelagic species. 
Aphyonines are small, pale fishes with thin, scaleless skin, weak 
musculature, reduced ossification and no swimbladders. Although 
they are livebearing, they have a copulatory apparatus which is 
very different from that found in benthic livebearing ophidioids. 
They also lack well developed eyes and lateralis systems. I believe 
that neoteny has played an important part in the evolution of 
the aphyonines. Tlialassuhathia , with its large eyes, well devel- 
oped lateralis system and bythitiiie dentition, seems, on the other 
hand, to be one more offshoot of the highly adaptive Bythites 
group of genera, and although it shares a common environment 
with the Aphyoninae, the two are not at all closely related. 

Specimens examined in preparing the foregoing section are 
listed in Cohen (in press). In addition I have examined the 
aphyonine material described by Nybelin (1957), specimens of 
Aphyonus in the Museum of Comparative Zoology and in the 
British Museum (Natural History), and specimens of Bara- 
thronus in the U. S. National Museum and the Chicago Natural 
History Museum. I thank the fish curators in Cambridge, Chi- 
cago, Goteborg, London and Washington for their kindness. 

Thalassobathia pelagica sp. nov. 
Figures 1, 2 

Holotype. MCZ 42161, 221 mm standard length, collected 
by Richard H. Backus on board the "Capt. Bill III," haul RHB 
913; October 13, 1962; 39° 26' N., 71° 00' W. to 39° 32' N., 
71° 00' W.; depth to gear 390 fms. (approx. 713 m), depth to 
bottom 1400-1350 fms. (approx. 2561-2469 m) ; time 1020 to 
1505 hrs. EDT ; 64 foot open midwater trawl. 

Paratypes. MCZ 42162 (2) ; data as for the holotype. 

Description. Body compressed and foreshortened, greatest 
depth between three and four in total length. Head relatively 



BREVIORA 



No. 196 




S3 
O 

tB 

.9 

O 



t-i 






be 

a> 






a 

S 






a 

o 

o 



'Si 

Si. 

e 

e 
o 

=0 
=0 

e 

e 



2 



1963 NEW OPIIIDIOID FISH 5 

short, between five and six in total lengtli. Snout rounded, nioutli 
slightly inferior. Interoi-bital area flat or slightly convex. Out- 
lines of head converging anteriorly when viewed from above. 
Eye prominent, horizontal diameter of cornea about three and 
one-half in head. Posterior nostril an elongate slit immediately 
in front of ventral half of eye, anterior nostril small and round, 
closer to tip of snout than to orbit. Body quite flexible due to 
weak ossification, feeling much the same as icosteid fishes. 

All fins covered M'ith thick skin. The dorsal fin appears to 
originate a short distance behind the level of the posterior margin 
of the opercle ; it is preceded by rayless pterygiophores and ptery- 
giophores bearing very reduced rays. The exact number of these 
elements is in doubt as their weak ossification makes them difficult 
to count on an X-ray photograph. Pectoral fins inserted ver- 
tically, broadly rounded; in the holotype and smallest paratype 
the pectoral fin extends about two-thirds of the way from its 
origin to the vent, in the other paratype the pectoral fin extends 
only about one-half of the pectoral origin to vent distance. The 
ventral fins much reduced and covered with thick black skin (Fig. 
2C), the longer, inner ray extending only slightly beyond the 
level of the base of the pectoral fin. One pectoral fin was cleared 
and stained to verify the number of rays. 

Teeth short, needle-like, extending on the dentary in a single 
line from the angle of the gape to the symphysis, where a broader 
patch of teeth is present ; premaxillary teeth arranged similarly 
to those on the dentary ; vomerine teeth in a roughh- circular 
patch; palatine teeth in an elongate patch, about three or four 
times longer than wide. Suprabranchial tooth patches present; 
however, I have been unable to find any basi-branchial teeth. 
Tongue massive, lacking teeth and lacking an anterior prow-like 
extension. The first gill arch with rakers reduced to a few fleshy 
flaps and protuberances laterally (Fig. 2A), medially (Fig. 2B) 
with seven or eight small rakers. A prominent pseudobranch 
located lateral to the upper arm of the first arch. Gill filaments of 
moderate length, about equal to diameter of lens. 

Color brown-grey over the body, the grey due mainly to a 
mucous coat ; the fins and head darker. 

Prominent pores on the head. The lateral canal w4th a single 
pore near the upper angle of the opercle ; supraorbital canal 
with two or three pores, one above the upper nostril, another 
below and slightly ahead of the upper nostril, a median pore in 
the interorbital region of the largest paratype ; infraorbital canal 



BREVIORA 



No. 196 






Fig. 2. Thalassobathia pelagica. A, First gill arch from left side of 
smallest paratype, lateral view. B, First gill arch from left side of smallest 
paratype, medial view. C, Ventral fins of largest paratype. Drawn by 
Mildred H. Carrington. 



i 



1963 NEW OPHIDIOID FISH 7 

with nine pores, the anteriormost below the anterior nostril and 
immediately behind one of the supraorbital pores, six in an 
irregular row in the loose skin above the upper jaw, and two 
behind the eye ; preoperculo-mandibular canal with five or six 
pores located along the ramus of the lower jaw; in the smallest 
paratype the pair of pores located closest to the midline of the 
lower jaw have joined to form a single pore. 

Lateral line with an upper and lower section, the upper line 
originating above the opercle and arching upward and back to 
the level of the origin of the dorsal fin, where it extends pos- 
teriorly about midway between the dorsal profile and the midline ; 
the dorsal line disappears at about two-thirds of the distance 
between the posterior margin of the opercle and the base of the 
tail; the ventral line originates slightly behind the level of the 
vent and extends posteriorly to the tail in the midline of the body. 
Each of the two lateral lines along the body is marked by about 
30, small, dark, papillae ; similar structures are sparsely dis- 
tributed over the body and head. 

Each specimen has a broad, fleshy hood over the genital area. 
The largest paratype has a pair of large ovaries which contain 
small eggs at the bases of numerous villi. The smallest paratype 
contains small paired ovaries with developing eggs. 

The swimbladder of the smallest paratype is a white, bean- 
shaped organ about 14 mm long. The lumen appears to be com- 
pletely occluded by white tissue.^ 

The liver is prominent in both paratypes, and the thick skin 
has a subdermal fatty layer. 

The brain case is cartilaginous. The olfactory nerves extend 
anteriorly in the orbit and are separated from each other by the 
cartilaginous interorbital. 

All three specimens bear numerous round marks left by squid 
tentacles. The smallest paratype has the top of its head bitten off. 

LITEEATURE CITED 

Briggs, John C. and David K. Caldwell 

1955. The characteristics and distribution of the spotted cusk eel 
OtopJiidium omostigmum (Jordan and Gilbert). Quart. Journ. 
Florida Acad. ScL, 18(4) : 285-291. 



1 I am indebted to N. B. Marshall for his kindness in examining the swim- 
bladder. 



8 BREVIORA No. 196 

Cohen, Daniel M. 

(in press). A review of the ophidioid fish genus Oligopus with the descrip- 
tion of a new species from West Africa. Proc. U. S. Nat. Mus. 
Denton, E. J. and N. B. Marshall 

1958. The buoyancy of bathypelagic fishes without a gas-filled swim- 
bladder. Journ. Mar. Biol. Assoc. U. K., 37: 7.')3-767. 
Herald, Earl S. 

1953. Spotted cusk-eel, the strange fish that stands on its tail. Cali- 
fornia Fish and Game, 39(3): 381-384. 
Marshall, N. B. 

1960. Swimbladder structure of deep-sea fishes in relation to their 
systematics and biology. Discovery Eepts., 31 : 1-122. 
Nybelin, Orvar 

1957. Deep-sea bottom fishes. Eepts. Swedish Deep-Sea Expedition 2, 
zoology (20) : 250-345, pis. 1-7. 

Table 1 
Counts and measurements in mm of Thalassobathia pelagica 

Ilolotype Paratypp Parafyiic 

Dorsal rays 

Anal rays 

Pectoral rays 

Vertebrae'^ 

Caudal rays 

Standard length 

Head length 

Snout 

Orbit 

Interorbital 

Upper jaw length 

Preanal 

Greatest body depth 

Body depth at vent 

Pectoral fin length 

1 Not including hypural. 



79 


72 


74 


65 


58 


61 


23 


24 


27 


49 


48 


48 


10 


10 


10 


221 


173 


200 


44.0 


13.5 


42.0 


10.0 


— 


7.4 


11.9 


— 


11.0 


16.0 


— 


13.3 


30.0 


21.8 


22.2 


87.9 


71.0 


87.2 


59.4 


47.2 


64.0 


54.1 


45.4 


50.0 


28.2 


23.4 


25.4 



X^uJ 



BREVIORA 

MMsemim of Coeiparative Zoology 



Cambridge, jMass. Deckmbek 31, 1963 Number 197 

ANOLIS WHITEMANI, NEW SPECIES FROM 
HISPANIOLA (SAURIA, IGUANIDAE)i 

By Ernest E. Williams 

Miss Cochran in 1941 commented on the apparent extreme 
variability of Anolis cyhotcs cyhotes, mentioning among other 
points that "some of the specimens have heavy keeling on the 
ventrals ; in others from precisely the same locality, taken at the 
same time, the ventrals are perfectly smooth." Inspection of 
material referred to cyhotes in the Museum of Comparative Zool- 
ogy, the United States National Museum, and in the unreported 
collections of the American Museum of Natural History revealed 
that this phenomenon of sharp dimorphism in regard to keeling 
was curiously localized and that many of the localities were in 
the Cul de Sac Plain in Haiti and in its continuation in the 
Dominican Republic. A closer look at the specimens from this 
area showed further that there were indeed two kinds of cyhotes- 
like anoles represented, and that they were in fact reported as 
taken at exactly the same localities by the same collector on the 
same dates. There seemed, however, to be not a difference in a 
single character but in several independent characters. The 
suspicion thus arose that species difference and not intraspecies 
dimorphism was involved. The suspicion was sharpened when 
P. S. Humphrey, collecting for Yale University and the Univer- 
sity of Florida in 1959, lirought back only two cybotcsAike anoles 
from the Cul de Sac region, one of each type. The fresh material 
permitted also an increased confidence in a color difference which 
had seemed to exist in the specimens long in collections. When, 
therefore, in August 1959, E. E. Williams and A. S. Rand 
planned a visit to Haiti, one of the objectives was to obtain and 
examine alive a series of the keeled cyhotcs-like anole from the 
Cul de Sac. 

1 Notes on Hispaniolaii herpetology no. 9. 



2 BREVIORA No. 197 

This objective was achieved, and the keeled form in the Cul 
de Sac Plain proves indeed to be a new species, which we name 
in honor of M. Luc Whiteman of Port-au-Prince who has assisted 
in the recent Haitian explorations. 

Since 1959, more material of A. whitcniaiii has been obtained 
by A. S. Rand and J. D. Lazell, Jr. at Mole St. Nicolas on the 
northwest peninsula of Haiti — an arid area very similar to the 
arid portions of the Cul de Sac Plain in which ivhitemani was 
first found. In addition to these (MCZ) specimens and the ma- 
terial in the United States National Museum (T'SNM) and the 
American Museum of Natural History (AMNH), further speci- 
mens have been found in the collections of the Hamburg Museum. 
The latter, from Monte Cristi in the Dominican Republic, extend 
the range of ivhitemani considerably to the east but still along a 
dry coast. 

Anolis whitemani new species ^ 

Type: An adult male, MCZ 60055, road to Eaux Gaillees, 

HaitL E. Williams and A. S. Rand collectors, 13 August, 1959. 

Paratypcs: Haiti. MCZ 60056-9, same locality as the type, E. 
Williams and A. S. Rand coll. MCZ 62844, same locality as type, 
A. S. Rand and J. Lazell coll. YPM 3193, Eaiix Gaillees, P. S. 
Humphrey and S. Van Vleck coll. AMNH 70588, Eaux Gaillees, 
A. Curtiss coll. USNM 117217-8, Trou Caiman, A. Curtiss coll. 
USNM 54189, Thomazeau, A. Curtiss coll. MCZ 62827-43, M<>lr 
St. Nicolas, A. S. Rand and J. Lazell coll. Dominican Republic. 
AMNH 50147, 50200, MCZ 61843, Las Baitoas near Duvereje, 
W. G. Hassler coll. Hamburg Museum 5198 (3), Monte Cristi. 

Diagnosis: Very close to A. cybotes but differing in squama- 
tion (dorsal scales larger, the middorsals hardly larger than 
adjacent scales, rather than abruptly larger; the ventrals smaller, 
hardly larger than middorsals, narrow and keeled, rather than 
wide, smooth and cycloid), and in color (body color very pale, 
nearly grey instead of distinctly brown, dewlap unmarked Avhite, 
rather than grey with yellow streaks or yellowish or pinkish with 
grey streaks). 

Description. Head: Head massive, snout to posterior border 
of eye about as long as tibia. Head scales mostly smooth. Five 
scales across head between second canthals. A shallow frontal 



1 Named for M. Liic Whiteniaii, who has helpetl so iiiiuh iu ainassiug the recent 
collections from Haiti. 



1963 ANOLIS WHITEMANI 3 

depression. Naris in front of canthal ridge. Anterior nasal 
scale in contact with rostral. 

Supraorbital semicircles narrowly in contact, partly in con- 
tact with supraocular disks. Supraocular disks consisting of 
about 8 enlarged keeled scales separated by 4-5 rows of granules 
from the scales of the supraciliary rows. One to two elongate 
supraciliaries continued backward by a double row of moder- 
ately enlarged scales. Canthus distinct, canthal scales 4, the 
second largest, decreasing graduallj' forward. Loreal rows 6-7, 
the lower row the largest. Supratemporal scales somewhat en- 
larged, flat, grading into the enlarged scales surrounding the 
interparietal. Interparietal larger than ear, separated from 
supraorbital semicircles by 1-2 scales. 

Suboculars separated from supralabials by one row of scales, 
anteiiorly separated from the canthal ridge by one scale, pos- 
teriorly continued behind eye for a short distance. Six supra- 
labials to center of eye. 

Mentals somewhat broader than long, in contact posteriorly 
with 4 throat scales. Infralabials narrow, in contact with 3 
large, wide, nearly lunate sublabials. Throat scales small, swol- 
len, not keeled, only the anterior ones elongate. 

Trunk: Middorsal scales keeled, hexagonal, not much larger 
than adjacent scales which grade very gradually into granules 
on flanks. Ventrals not much larger than middorsals, relatively 
narrow, distinctly keeled, the keels in lines. Postanal scales small, 
poorly differentiated. 

Gular fan: Moderate, scales cycloid, keeled, larger than 
ventrals. 

Limbs and digits: Head and foot scales multicarinate. About 
17 lamellae under phalanges 2 and 3 of fourth toe. Largest scales 
of limbs unicarinate, those of arm as large as, of thigh larger 
than ventrals. 

Tail: Compressed. Each verticil surmounted by 4 keeled 
scales, ventrally 3 pairs of somewhat larger keeled scales per 
verticil. 

Color in life: Four topotypes taken on road to Eaux Gaillees: 
1. Type S . Dewlap white. Ground color grey with indistinct 
transverse bands. Scattered brown spots on flanks and on sides of 
belly. 2. 6 . Dewlap pure white. Dorsum grey and cream. 
Pattern very obscure. Venter pure white. 3. 9 . Dorsum pale 
grey. Belly pure white. 4. c^ . Juvenile. Dewlap pure white. 
Dorsum cream and grey. 



BREVIORA 



No. 197 



Notes by W. G. Hassler 1935 (Las Baitoas paratypes) : 
" Anolisf-like cybotes: in grass on mountain side above road and 
forest. These ancles very grey or grey-brown. Pattern faint. 
Sides slight!}' yellowish grey. Belly and throat white. S similar 
to 9 . Faint yellowish brown-grey longitudinal bars like ordi- 
nary cybotes but much lighter, less red and all very faint. Throat 
(very faint vermiculations), belly and fan WHITE." 

SPECIES STATUS AND CHARACTER DIFFERENCES 

The squamation and color differences between whitemani 
and cybotes in the Cul de Sac Plain are striking but hardly more 
striking than those between cybotes and " Audantia" armouri 
Cochran from the Massif de La Selle. New evidence, however, 
shows that the latter taxon, formerly considered a distinct genus, 
intergrades with cybotes at intermediate elevations in the foot- 
hills of the La Selle range, e.g. at Furcy, 10 miles S. of Port-au- 
Prince. Some of the s(iuamation differences between whitemani 
and cybotes are in fact rather similar to those between armouri 
and cybotes — although they are greater (Table 1, and Figs. 2-4). 
Thus the character differences are not in themselves great 
enough to establish species status for whitemani. 




X Anolis whitemani n. sp. in 
central and western Hispaniola 

Fig. 1. Localities for Anolu whitemani n. s]). 



1963 



ANOLIS WliriEAIANI 



Table 1. 
Differences separating' cybotcs-Viko anoles in the Port-au-Prince 



VEXTRALS 



TEMPORALS 



MIDDORSALS 



BODY 
PATTERN 



DEWLAP 
COLOR 



area. 



icliit( iiiaiii 



keeled, ii;irio\v. 



cybotes armouri 

(Port-au-Prince 
and vicinity) 
smooth, cycloid, smooth, cycloid, 
wide. very wide. 



granular, but finely granular, granular, but 

larger than ilank even smaller than larger than flank 
scales. flank scales. scales. 



enlarged but 
grading into flank 
scales. 



abruptly larger 
than flank scales. 



enlarged but 
grading into flank 
scales. 



BODY COLOR pale tan. 



patternless or 
with pale grey 
transverse mark- 
ings. 

white 



brown to reddish, olive. 

patternless as often with bold 

adults or with rhombs on flanks 

greenish lines on = persistence in 
flanks. adults of a 

juvenile pattern. 



pale yellow or 
rarely pale pink 
with white or 
greyish streaks or 
greyish with yel- 
lowish streaks. 



pinkish or yel- 
lowish Avith green- 
ish smudges 
centralh'.l 



SPECIES STATUS AND DISTRIBUTION 

In the final analysis the jndgment that whiteniani is a full 
species rather than a morph of cybotes, as Cochran (1941) be- 
lieved, has been based on observation of the habits and habitats 
of the two species in the Cul de Sac region. 

The topotypic series of A. whiicmani was taken in open dry 
scrub along the road to Eaux Gaillees in the full sun of late 
morning. The same day quite typical A. cybotes were obtained 
in the grove of trees at the spring in the village of Eaux Gaillees 
and in a similar grove of trees at Manneville further east on the 
same road, again at a spring. 



1 Color notes from Information provided by Dr. Albert Schwartz. 



6 BREVIORA No. 197 

In the Cul de Sac Plain such groves of quite large trees around 
springs are quite literally oases in a thorn scrub desert. The 
fauna, like the flora of these oases, is characteristically that of 
rather moist areas. 

The faunas of these oases are obviously disjunct, in effect 
island populations derived from the faunas of the moist forests 
of the mountain foothills to the south and to the north. The con- 
trasts between the fauna within an oasis and that outside it are 
sharp and very characteristic. Within each oasis are animals 
that could be seen also in a pension garden in Port-au-Prince 
itself, e.g. Anolis chlorocyanus, A. cyhotes, A. distichus, Celestus 
costatus. Outside, in the dry scrub, are Anolis ichitemani, 
Celestus curtissi, Leiocephahts scmilineatus. 

A. cyhotes and A. whitemani thus live in the same area but 
are sharply segregated by ecological preference and sharply dis- 
tinguished by color and morphology. It is very important that 
the morphological differences are greater between cyhotes and 
adjacent whitemani than between cyhotes and more distant 
armouri; this is a demonstration that these two populations 
are behaving as good species. 

The discovery of A. whitemani on the dry northwest coast at 
Mole St. Nicolas is a very valuable addition to the data on its 
distribution. When its occurrence seemed limited to the Cul de 
Sac Plain, whitemani was something of an anomaly, since this 
area has only relatively recently emerged from under the sea. 
However, with the realization that it is probably to be found all 
along the dry northwest coast of Haiti and is known as far east as 
the similar dry coast at Monte Cristi on the north coast of the 
Dominican Republic, its recent extension into similar arid habi- 
tats in the Cul de Sac Plain becomes very plausible. It is prob- 
ably an autochthon of the dry west and northwest coast of tlie 
"northern island" — the portion of Ilispaniola north of the 
former seaway through the Cul de Sac. (For a discussion of the 
importance of the northern and southern islands in His])ain()la 
zoogeography, see Williams, 1961.) 

It is noteworthy that Humphrey obtained large series of the 
species cyhotes on Gonave Id. (an island mostly dry but again 
with oases around the springs) but no ivhifemani. The latter 
may thus really be absent there. This requires further confirma- 
tion, but the fauna of Gonave has been derived by a limited ami 
erratic sampling of the faunas of the neighboring mainland. The 
absence of any individual species is therefore not in its(^lf sur- 
prising and need not imply oversight in collecting. 



1963 



ANOIJS WniTEMANI 



ACKNOWLEDGMENTS 

T am indebted to Dr. Albert Schwartz, Dr. Philip Humphrey, 
Dr. Doris Cochran, Mr. C. M. Bogert and Dr. W. Ladiges for 
the privilege of examining specimens. National Science Founda- 
tion Grant G 5634 and a grant from the American Philosophical 
Society provided funds for expeditions to Haiti. National Sci- 
ence Foundation Grant G 16066 continues to support research 
on Hispaniolan and other anoles. 

REFERENCES CITED 

CochrjVN, D. 

1941. The heipetology of Hispaiiiola. Bull. U. S. Nat. Mus., 177: i-vii, 
1-398. 
Williams, E. 

1961. Notes on HLspiuiiolau heipetology. 3. The evolution and relation- 
ships of the Anolis semilineatus group. Breviora, no. 136: 1-8. 






Fig. 2. Anolis whitemani n. sp. Paratype. MCZ 60056. Top: Side view of 
head. Middle : Dorsum at midbody. Bottom : Venter. 



8 



BREVIORA 



No. 197 






Fig. 3. Anolis cybotes cybotes. MCZ 59883. Top: Side view of head. Mid- 
dle: DcTSum at niidbody. Bottom: Veuter. 






Fig. 4. Anolis eyhotcs annouri. MCT. ClO.'il. Top: Wide view of head. Mid- 
dle: Dorsum at niidliodv. Hotloiu: Venter. 



BREVIORA 

MmseiMii of Comparative Zoology 



Cajjbridge, :\Iass. March 10, 19()4 Number 19S 

AMPHISBAENA SCHMIDTI, A THIRD SPECIES OF THE 
GENUS FROM PUERTO RICO^ (AMPHISBAENIA : 

REPTILIA). 

By Carl Gans 

Departineiit of Biology, State University of New York at Buffalo, 

Buffalo 14, New York 

The island of Puerto Rieo is iiiiiciue auiouy- the islands of the 
Greater Antilles in possessing two almost eertainly sympatrie 
species of Amph'shaena. One of them, A. caeca, ranges over 
the entire island, while the second, A. hakeri, is restricted to the 
central portion of the northwestern corner of Puerto Rico (Fig. 
1). .1. caeca has been shown to vary more widely in the western 
than in the eastern portion of its range (Grant, 1932; Gans and 
Alexander, 1962). 

A recent study of the systematics and variations of the sev- 
eral Antilles species (Gans and Alexander, 1962; here followed 
for terminology) disclosed three individuals from Puerto Rico 
that clearly diifered from both described forms in five character- 
istics, i.e. in more ways than the former differed from each other. 
The specimens were left incerfae sedis for three reasons : two 
specimens came from a rather old collection ; the third specimen 
had been collected on the diametrically opposite end of the 
island; and only these three out of more than 200 Puerto Rican 
siKM'hnens showed this character pattern. 

Dr. 11. lleatwole has now made available three additional speci- 
mens that stem from two coastal localities adjacent to that of 
the first two specimens. They make it desirable to call attention 
to the probable existence of yet a third species of Anrphishaena 
on the island, a situation that may have some interesting zooge- 
ographical implications. 

1 Notes on amphisbaenids No. 11. 



BREVIORA 



No. 11J8 























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19G4 AMl'lllSHAEXA SCIIMIDTI 3 

1 take o-reat plPHsurc in iiaiiiiim this t'onii for the late Karl 
Patterson Seliuiidt. Tlic spcciniciis IicIohl;' in tlic collections of 
the Carneo'ie Mnscuni (CM), tlic Tniversitetets Zoolop^iske Mn- 
senni of Kobenhavn (KM), and the TTniversity of Puerto Rico 
at Kio Piedras (PIMvliP). One of the individnals from Kio 
Piedras has been sent on cxchano-e to the ]\Insenni of Comparative 
Zoology (MCZ). T am ui-atefnl to tlie curators of the several 
institutions, ])articularly to Dr. F. W. I>raestru]) who went to 
considerable tronble to check the original catalogs, and to Dr. 
IT. Ileatwole and students who made an effort to collect addi- 
tional specimens. A. A. Alexander and E. E. Williams contrib- 
uted by discussing this situation. Dr. Virginia Cunnnings ])re- 
pared the drawings and Miss Charlyn Rhodes furnished technical 
assistance. These studies are supported by Grant G-21819 from 
the National Science Foundation. 

KEY TO PUERTO RICAN AMPHISBAENA 

1. Body aiinuli more than 205, caudal aiinuli fewer than 18, tail markedly 

shorter (Fig. -4), generally no enlarged parietats 2 

Body annuli 205 or fewer (198-202), caudal annuli 18 or more (20-22), 
tail markedly longer (Fig. 4), parietals very large sclimidti sp. nov. 

2. Body annuli 218-230, tail slightly longer (Fig. 4), internasal suture con- 
siderably shorter caeca 

Body annuli 239-255, tail slightly shorter (Fig. 4), internasal suture con- 
siderably longer taTceri 

Amphisbaena sciimidti s]). nov. 

Diagnosis: A form of Ai)tpJiish(i( iia without fusion of head 
segments ; with markedly enlarged parietals ; having 198 to 202 
body annuli along the ventral line; 20 to 22 caudal annuli; 14 
dorsal and 1(3 to 17 ventral segments to a midbody annulns; two 
rows of postgenial and one row of postmalar chin shields and 
4 precloacal pores. The tail is cylindrical and its end rounded. 
The autotomy constriction is noticeable at the seventh to eighth 
postcloacal annulus and autotomy takes place here. 

Holotype: MCZ 73115, a female collected by M. J. Yelez, Jr., 
at Orilla (Cunta) Carrcamo, Isabella, Puerto Rico, on 21 Febru- 
ary 1960. 

Parafypes: UPRRP 12!)(), a male collected with the holotype; 
CM 36277 from Salinas, Puerto Rico ; KM R-4414 and R-4416 
collected by Dr. Meinert at Aguadilla, Puerto Rico, in January 



4 BREVIORA No. 198 

1892; and UPRRP 2502 colloeted by II. IIeat\v()](> along high- 
way P.R. 681, 2 miles east of Cueva del Indio, Puerto Rieo, on 
14 January 1968. 

DiscKssiou: The demonstrated differenees indicate that the 
new form is very distinct morphologically. They do not show 
whether it is a separate species, a (coastal?) race of A. caeca, or 
of A. hokcri. A case may be made for each of these. I have de- 
cided to treat the form as a full species for the following reasons : 
(1) The degree of morphological difference is greater than that 
fouiul between consiiecific populations among related species. 
It exceeds that between the two presently recognized species on 
the island. (2) The sample from Salinas contains five specimens 
of A. caeca and one of A. schuiidti. The coastal record of A. 
caeca from Arecibo lies midway between two records of A. 
fichinidti. Both of these cases suggest a limited degree of symjiatry. 
(3) Specimens of A. schniidfi from opposite ends of the island in- 
dicate almost no geographic variation, in contrast to the situation 
of A. caeca. (4) Designation of the form as a full species does 
not prejudge the open question of its affinities. 

Description: Meristic characters are listed in the table. Fig- 
ure 2 shows the head scalation. Figure 3 the segmentation of 
cloaca and tail, and Figures 5 to 7 are photogi-aphs showing 
details of color i)attern and midbody segmentation. Figure 4 
shows the body proportions. 

Specimens are a dai'k bi-owii with vei-y slight ventral counter- 
shading. None of the sjiecimens show a droi)ping out of pigment 
on the midventral area. The smaller specimens have each seg- 
ment more or less uniformly ])igmented, but the two largest show 
a considerable additional darkening of the rectangular segmental 
centers. The intersegmental raphes are always lightened. Dorsal 
surfaces of head and tail are a uniform dark violet brown in 
freshly preserved specimens. 

The head scalation is characterized by lack of major fusions. 
An azygous rostral invisible in dorsal view is followed by a pair 
of contacting nasals, very large elongate prefrontals, frontals, 
and wide and long parietals. The latter may be divided (trans- 
versely) or not. There are three supra- and three infralabials, 
but the angulus oris lies very slightly anterior to the posterior 
edge of the third paii'. The second of each series is by far the 
largest ; indeed, the second infralabials are larger than any 
scales but the prefrontals. A small segment lies innnediately 
in line with the slit of the mouth ajid contacts the posterior half 



1964 



AMl'lllSBAEXA SCliAllUTi 






Fig. 2. AmphishaoKi .scliniidfi. Dorsal, lateral ami venti-al views of 
the head of the holotype, MCZ 73115 from Isal)ella, Puerto Rico. The 
line equals 1 mm to scale. (V. Cummings, del.) 



6 



BREVIORA 



No. 198 



of the enlarged lateral postinalar segment. The sutures between 
the supralabials run at angles of 45° to the slit of the mouth. 
The oeular is (luadrangular. 

The mental is T-shaped and nuich lai'gcr tlian the tiny first 
infralabials. The third infralahials are (piite narrow. The post- 
mental is almost as large as a second infralabial. The triangular 
tips of the two segments of the first ])Ostgenial row embrace it 
posteriorly. There are three second postgenials. The anterior 
tip of the median one contacts the postnu^ntal in some specimens. 
The malars are relatively small and contact the second and third 
infralabials, but are clearly excluded from contact with the 
postmental. The segments of the ])ostmalar row ai-e relatively 
long, the lateral ones are almost wedge-shaped. Only the anterior 
half of the lateral ones contacts the third infralabial ; the pos- 
terior portion reaches the small segment back of the angulus 
oris. The postmalar row is thus counted as the first body annulus 




Fig. 3. Aniphisbacna sclnnuUi. \'('iiti;il view of cloacM ;iihI tail of the 
holotype, MCZ ZHllf) from Isabella, I'licito Rico. The line e(|iia Is 1 nini to 
scale. (V. Cuniniings, del.) 



1UG4 



AMl'JllSUAENA SCIIMIDTI 



as well. Tlu' lateraliiiost segments of the row are of the same 
width as the mahirs, and give the impression tliat the hitter 
have been split. 

Dorsally, the large segments of the first body annnhis curve 
to contact the sides of the frontals. The dorsal poi'tion of the 
second annulus consists of elongate segments that increase in 



E 
E 



c 

0) 



22 



18 



14 



10 



  



o 

o o 



OD 
O O 
00 • 4 






 s chmid ti 
o caeca 

• baker i 



8 



16 24 

Snout - Vent Length - cm 



Fig. 4. Amphisbaena. Scatter diagram showing plot of tail length versus 
snout-vent length for all specimens of A. schmidti and A. haktri, as well 
as specimens of A. caeca from western Puerto Rico. Inclusion of eastern 
Puerto Rico material of A. caeca would mask the difference between A. 
caeca and A. bal-eri. hut not affect that between the former and A. 
scJimidti. 



length toward the middorsal line. The posterior edge of the 
second annulus shows no forward curvature. There is no dor- 
sal intercalated half-annulus, though the parietals are split into 
two pairs in four specimens and on one side of another. 

The head is pointed, depressed, of horizontally oval cross- 
section. The lower jaw is but slightly shorter than the upper. 



8 



BREVIORA 



No. 198 




Fig. 5. Ainp}ii.sb(t( H(i .scliiuulti . Dorsal, lateral and ventral views of the 
head of UCZ 73115, to show uniform coloration and the effect of the bulg- 
ing temporal musculature. 



11)64 



A.MI'HISBAENA SCHMIDTI 



9 



The bulji'c of the t('iii|)()i-;il itinsculature is very noticeable, pro- 
ducing a concave folding of tlie skin along the middorsal raphe 
and a clear distinction of the head from the narrower neck. 

There are lf»S to 20'2 body anmili from the back of the third 
infralabial, np to and inehiding the pore-bearing precloacals. 
The second throngh fifth annuli are ventrally nuu'h narroAver 
than the succeeding ones and their segments do not line nj) with 
those following. There is no pattern irregularity in the "pec- 
toral" region, nor are there intercalated dorsal half-annuli. 
There are ]4 dorsal and 16 to 17 ventral segments to a midbody 
annuhis. 




Fig. 6. Antplii.shaciia sclnuidfi. Dorsal and ventral views at midbody of 
MCZ 73115 to show segment proportions and coloration. 



The cloacal region is characterized by 4 round precloacal pores 
which are strongly expressed in males as well as females. There 
are 6 precloacal and 9 to 12 post cloacal segments and 3 (once, 
unilaterally, 4) lateral rows. The extreme lateral postcloacal 
segments lie almost directly laterad of the extreme lateral seg- 
ments of the curved precloacal shield, so that one is tempted to 
count them as precloacals. The autotomy annulus falls on the 
seventh to eighth postcloacal annulus and autotomy takes place 



10 



BREVIORA 



No. 198 




Fig. 7. Amphisbaena schmidti. Ventral view of cloaca ami tail of MCZ 
7.3115 to show pore size (in a female) and coloration. 

here. Specimens have 20 to 22 caudal anniili. The cross-section 
of the tail is circular throughout and the distal tip is capped by 
a hemispherical portion. 

The lateral sulci are distinctly marked from back of the first 
quarter of the trunk length to the level of the cloaca. At their 
widest they are narrower than the width of one of the l)order- 
ing segments. The dorsal and ventral sulcus and the lateral 
sulcus in the anterior quarter are indicated only b}- the align- 
ment of intersegmental sutures. 

The dorsal segments of a midbody annulus are approximately 
one and one-quarter times as long as wide, while the ventral 
ones are one and one-half times as wide as long. 

Habits: Examination of the KM catalogs (by Braestrujv) indi- 
cates that Dr. Meinert (an entomologist) excavated some termite 
nests on the days when the two paratypes were taken, and may 
have found the specimens at that time. The Cueva del Indio 
specimen was collected under a fallen ])alm leaf on liunuis. 

Range: Coastal Puerto Rico. 



LITEEATURE CITED 

Gaks, Carl and A. Allan Alexander 

19Hi2. Studies on amphisbaenids (Amphisbaenia, Reptilia). -. On the 
aniphisliaenids of the Antilles. Bull. ^ilus. ('(iiii]i. /ool., vol. ll!8, 
no. 3, pp. 65-1.58. 
(iRANT, Chapman 

lit.'!!'. A red(>scrii)tion of Antpltisbdcrid caeca with a discussion of its 
relationship to A. bakcri. .lour. Dcjit. Aurii-. rucrto l^ico. vol. 1(3, 
no. '1, i)p. 153-155. 



I!)(i4 



AMPIIISBAENA SCHMIDTI 



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BREVIORA 

Mmseiiiinii of Coeipsirative Zoology 

Cambridge, Mass. March 10, 19G4 Number 199 

A NEW SUBSPECIES OF VARANL\S EXANTIIEiMATICU^ 

(SAURIA, VARANIDAE) 

Bv R. F. Laurent 



Dr. Williams asked me recently to study a small series of 
voimq: Savanna Monitors collected bv C. J. P. lonides in south- 
eastern Tanganyika, where many novelties have already been 
discovered. He had been impressed by their striking color pat- 
tern which is quite ditit'erent from the pattern exhil)it('d ])y other 
juveniles of the same species. 

In this, he was quite right. However, the morphological fea- 
tures selected by Mertens (1942) as characteristic of the recog- 
nized species and races of the genus Varanus did not suggest that 
the six critical juvenile specimens belonged to an undescribed 
form. In squamation tbey agreed perfectly "well with Voranus 
{Empayusia) crantluinaticus microstictKs Boettger. Neverthe- 
less, I believe that it is a serious mistake to ignore coloration 
when this character has the same degree of constancy as is 
generally considered significant for morphological characters. 

The difficulty here .seems to be that the diagnostic juvenile pat- 
terns gradually fade during the life of the monitors so that the 
adults are far more similar, uniform and dull, than are the 
juveniles. As a result, the number of specimens available for 
comparison is greatly reduced, the adults not being distinguish- 
able. 

When I compared the lonides specimens with other juveniles 
of Varanus e.ranthcDiaticus, I came to the conclusion that the 
sul)species recognized by Mertens are themselves geographically 
varialjle as far as their juvenile color pattern is concerned. 

The six juveniles from southeastern Tanganyika are alike, 
but obviously different from two specimens from Kenya ; like- 
wise, among the so-called southern alhigularis, two juveniles 
from Zulidand have a striped jiattern quite different from the 



2 BREVIORA No. 19!) 

ocellated pattern of five other juveniles from Transvaal, Becliu- 
analand and Southwest Africa. 

The photographs of specimens here discussed replace advan- 
tageously- any clumsy and lengthy description. Feeling that six 
specimens disclosing the same color pattern are enough to infer 
that the population to which they belong has on the whole 
peculiarities patently diiferent from those of other populations, 
I regard them as a new subspecies of Vorantis e.rantJicmaticus: 




Kilwa 



Fig. 1. Map sliowing the known localities for Varan us exanihcmatictis 
ionidcsi n. subsp. 



Varanus exanthematicus ionidesi subsp. n. 

Holotype: A half-grown individual (MCZ 544:9-1), Kilwa, Tan- 
ganyika, 1956, coll. lonides. 

Paratypcs: 1 adult, 2 subadults and 1 juvenile (MCZ 52536- 
39), Liwale, Tanganyika, 1953; 1 juv. (MCZ 54495), Masasi, 
Tanganyika, 1956; 1 juv. (MCZ 55470), Liwale, Tanganyika, 
1957; 1 juv. (MCZ 56176), Kilwa, Tanganyika, 1958; 1 juv. 
(MCZ 56981), Liwale, Tanganyika, 1958. All collected by C. J. 
P. lonides. 

Diagnosis. A subspecies of Varanus exanthematicus similar to 
microstictus Boettger ( ty])e locality, Ethiopia) but differing 
from it by a very distinctive juvc^iile color pattern som(>what 
sinular to that of albiyularis. 



VJiJ-it -\i:\v srusi'KciKs ok VxVKANus 3 

The u'cucral cdloi" is li^litcf jiiid with less coiitrjist of li,L;'li1 and 
(lark colors than in two yonn*:- niicrosi id us from Kenya. 

Two l)la<-k lines start i'roni the jjostei-ior border of the eye 
and a])i)roaeh each other on tiie neck, stoi)])in<i' ahiMiptiy in llie 
scapular region where they t'oi'in an acute an<ile. (Tiie same lines 
exist in inicrostichis and dlhifjiihiris, hut they are liardl\- distinct 
in inlcr(jsticfi(s.) Between these lines there are thi-ee or four 
blackish marks (these art' invisil)le in iiiio'ostictus) . 

There are four or five transverse series of 6 lig-ht spots (8 to 
18 in the typical form, or disposition irregular) tienerally sur- 
rounded by a black border, extending on 4 to 20 scales (in micro- 
si id us they fill 1 to 4 scales only and they have no black border) ; 
the mediodorsal spots are especially large and they fuse on the 
vertebral line (except in one individual Avhere they are at least 
contiguous). 

On the tail, the transverse bars are arranged in ])airs Avhich, 
in iouidcsi, have the appearance of groups of four or even of 
ei"'ht, because the central zone of each bar mav become as light 
as the intervals between the pairs and, in addition, two secondary 
zones of a less pronounced ]ialing may still intervene on both 
sides of the central zone. These complications exist only for the 
first three or four pairs; the distal bands are plain black. These 
areas of paling of the central zones disappear gradually in 
ionidcsi, but they are not at all apparent in the young micyo- 
stidus available for comparison ; in the latter, the terminal bars 
are not darker than the proximals. Lastly, the number of pairs 
is 9-10 in i)iicrostidus, but only (>8 in ionidcsi. The throat is 
blackish, as is usual, accoi'ding to ^lertens. for the other races.^ 

MORPHOLOGICAL CHARACTERS 

As stated previously, the morphological characters are not 
different from microstidus. Table 1 gives scale counts as com- 
pared with Mertens' descriptions of microstidus, alhigularis and 
(Difjolensis plus some specimens of microstidus (Kenya) and of 
alhigularis (southwest Africa and Transvaal) and angohnsis 
(Lobito). 

Certain measurements have been studied on scatter diagrams. 
Xo significant differences have been observed between ionidcsi 
and microstictus, but it seems that aJbigularis and (ixgoJcnsis 
have the head broader and the snout longer than liotii micro- 

1 This is not the case for tlic two mii-rostii-tiis juveniles from Kenya. 



BREVIORA 



No. 199 



Subspecies 

ionidesi 

microst ictus (Mertens) 
mierostictus (Kenya) 
albigidaris (Mertens) 
albipiihiri.s (Zululand ) 
albigularis (Transvaal, 

SW Africa) 
aiuiolensis (Mertens) 
(iiifioJensis (Loliito, 

Angola ) 



TABLE I 

Scales Ventral scales Scales lietween 

around (from gular fold the commissures 

N the body to the groin) of tlie moutli 



5 


129-150 


1-i 


122-152 


o 


131-134 


19 


137-1(37 


O 


139-147 


5 


137-149 


3 


110-138 



120-141 



88-99 

88-94 

87-90 

8.1-110 

98-99 

88-107 
77-92 

94-98(!) 



53-63 
51-56 
50-52 
57-64 
54-57 

53-60 
53-55 

55-56 



These data on angoleiisis suggest tlint this alleged subspecies might lie 
clinally merged with albigularis. 

stictus and ionidesi; the differences are minor and would have 
to be checked on larp'e series. ]\Iore interesting- perhajis is the 
fact that the two Zululand specimens, different from all others 
by their transversely banded pattern, are also ditt'erent in tliat 
their tail is shorter than the snout-vent length. This strongly 
suggests that another neglected race exists in, South Africa, but 
it would be imprudent to describe this on two specimens Avith 
relatively inexact locality. The existence of this race will have to 
be confirmed by studies on larger samples of Vfiraims (.raiiflK- 
'})}afir)(s; from South Africa. 



Specimens utilized for comparison with 
Varanus exanthematicus ionidesi 

VarauHS exanthonaticus micfost ictus Boettger, MCZ 41112, 
Sokoki Forest, Kenya (coll. H. J. A. Turner) ; MCZ 59154, Ka- 
koneni on Sabaki River, Kenya (coll. M. J. Coe). 

Varanus exanihematicus alhigularis (Daudin), MCZ 45417-18, 
Zululand; MCZ 39906, Waterberg, Southwest Africa (coll. W. 
Hoesch) ; MCZ 41888, Saltpan, Pretoria Distr., Transvaal (coll. 
A. Roberts) ; AMNH 57600, Otjuiarongo, Waterberg, Southwest 
Africa; CNHM 17141, Gaberones, Beehuanaland (coll. H. Lang) ; 
CNHM 75712, Hiittenhain, Southwest Africa (coll. W. Hoesch). 

Varanus exanthematicus angolensis Schmidt, AMNH 47721-24, 
Lobito, Angola. 



1964 NEW SUBSPECIES OF VARANUS 5 

Acknowledgmc )tf . Tliis ^v()l■k has been performed thanks to a 
grant o-enerously allowed by Ihe National Science Foundation 
(BG-18987). I take i)leasure in thanking Mr. C. M. Bogert and 
Dr. R. F. Ing(M' for tlie loan of sj)eeiinens belonging to the col- 
lections of tlie American ^luseum of Natural History (AMNH) 
and the Chicago Natural History Museum (CNHM), respectively. 

EEFEEENCE 

Mertens, R. 

1942. Die fnmilio der Varane (Varanidae). Dritter Teil : Taxonomie. 
Abh. Senekenb. Naturf. Ges., 466: 235-391. 



BREVIORA 



No. 199 



.:r'?tx^-&re^ 




Fig. 2. Faranus exanthematicus micro.st ictus juveniles from Kenya. 



1964 



NEW SI'BSIMMIKS Ol' \AKAXUS 




I 



Fig. 3. Varanus exanthematicus ionidesi n. subsp. Holotype sunoimded 
by paratypes. 



BREVIORA 



No. 199 





^i^K^ 




Fig. 4. Varaniis exanthematicus albigularis juveniles, showing the typical 
pattern. 



1964 



NEW SUBSPECIES OF VABANUS 





Fig. 5. Varanus exanthematicus albigularis juveniles from Zululand, show- 
ing the banded pattern. 



w 



BREVIORA 

MiiseTjinti of Coimparative Zoology 



Cambridge. Mass. April 10, 1!)()4 Number 200 



AN ANGUID LIZARD FliOM THE LEEWARD ISLANDS 

By Garth Underwood 

Department of Zoology, University of the West Indies, Trinidad. 

Through the good offices of Mr. L. Kasasian of the University 
of the West Indies, I received from Mr. J. Phillips, Director of 
Agriculture in Montserrat, a lizard which I was amazed to see 
was an anguid of the genus Diploglossus; the family had not 
previously been known, living or fossil, from the Lesser Antilles. 
Scarcely less amazing was the observation that this lizard resem- 
bles the Central American D. monotropis in respect of several 
major differentiating characters, and differs conspicuously from 
the known Greater Antillean species. 

The specimen was collected in Montserrat by Mr. J. Kingsley 
Howes who tells me that he has only once before seen this lizard, 
and that twenty -five years ago. On the grounds that I am therefore 
unlikely soon to see another I describe for the second time 
(Underwood, 1959a) a new species of Diploglossus on the basis 
of a single specimen. Once again my description contains tedious 
detail, some of which may be eliminated when more specimens 
are examined. As explained in the earlier paper, I do not recog- 
nize the genus Celestus as distinct from Diploglossus. Li the 
condition of the claws, prefontals, nasals, and numerous scale 
organs on the back, this species resembles only D. ynonofropis. 
I therefore borrowed two specimens of monotropis and prepared 
the description with these before me ; the specimens are MCZ 
37139 Yavisa, Panama, and MCZ 15353 Limon, Costa Rica. 

The name of the island of Montserrat is Spanish, from the 
Latin moiis serratus. As this is quite the most remarkable reptile 
yet reported from Montserrat, and as no other reptile bears the 
name of this island, I call it Diploglossus montisserrati. 



BREVIORA 



No. 200 



Family ANGUIDAE Cope 
DiPLOGLOSsus MONTissERRATi iiew species 

Type and only known specimen: adult $ MCZ 76924, and from 
this specimen : left ramus of lower jaw; slide 1, prefontals, frontal 
(damaged), frontoj^arietals, interparietal (damaged), parietals 
and occipital scales ; slide 2, series of 10 scales back from occipital ; 
slides 3a, b and c, 12 longitudinal rows of 10 dorsal scales ; slide 4, 
transverse row of scales from middorsal to midflank ; slide 5, 
three rows of 10 scales forward from groin ; slide 6, transverse 
row of tail scales at level of ankle from middorsal to midventral ; 
slide 7, series of 20 scales back from mental ; slide 8, 25 midbelly 
scales. Collected by J. Kingsley Howes at Woodlands Spring, 
elevation about 600 feet, Montserrat, West Indies, 16° 45i// N, 
62° 13' W. 

Diagiiosis: A large strong-limbed DipJoglossus with sheathed 
claws ; three prefrontal scales ; contact between nasal and rostral 
scales ; numerous scale organs on dorsal scales but none on belly 
scales ; very numerous scale organs on dorsal tail scales ; more 
than 80 scales from occipital to base of tail ; back plain brown, no 
crossbands. 




Fi^. 1. Dorsal view, head of Diplofjlossus montis.serrati type. Area of 
posterior frontal aud frontoparietals may be a little inaccurate owing to 
damage. 



1964 AN ANGUIl) FROM MONTSERRAT 3 

Description: As this species rather closely resembles mono- 
tropis, I note in square brackets the features in which monotropis 
differs. Upon receipt of the tir.st draft of my manuscript Mr. 
Benjamin Shreve kindly examined an additional seven specimens 
of monoivopis: in the Museum of Comparative Zoolo<iy for the 
characters which I had thought useful. His data are incorporated 
here. Shreve 's counts on the type of montisscrraii, when they 
differ from mine, are cited in parentheses. 

Head broad ; Avidth : length to anterior margin of earhole = 
0.83:1. Rostral one and a half times as broad as high, in 
contact with 1st labials, nasals, and supranasals. Pair of supra- 
nasals each meeting other supranasal, rostral, nasal, upper 
postnasal and frontonasal. Pair of frontonasals, each meet- 
ing other frontonasal, supranasal, upper postnasal, canthal 
and median prefrontal. Pair of canthals each meeting fronto- 
nasal, 2 prefrontals, 1st and 2nd loreal and upper postnasal [the 
same or meeting both postnasals and only 1st loreal]. Three 
prefrontals (sutures separating them incomplete posteriorly). 
Median prefrontal meets frontonasals, canthals and lateral pre- 
frontals [and sometimes frontal] ; each lateral prefrontal meets 
median prefrontal, canthal, 2nd loreal [1st or 2nd loreal], 1st 
and 2nd supraoculars [1st and 2nd or 1st only] and frontal. 




Fig. 2. Lateral view, head of Diploglossus montisserrati type. 

Frontal tapers to a point anteriorly, wide posteriorly ; meets pre- 
frontals, 2nd and 3rd supraoculars, frontoparietals and inter- 
parietal; broadly in contact with frontoparietals, narrowly in 



4 BREVIORA No. 200 

contact with interparietal [narrowly or broadly in contact with 
interparietal], excluded from contact with parietals. Palpebral 
aperture large. Four supraoculars, 2nd largest; 2nd loreal just 
meets first supraocular [no loreal meets supraoculars], 3rd loreal 
separated from supraoculars by 1st supraciliary [2nd or 3rd 
loreal so separated]. One subocular intruding only slightly be- 
tween 6th and 7th labials. Six supraciliaries [6 or 7 supracili- 
aries], postoculars 4, 6 [4 or 5]. Five temporals between fronto- 
parietal and labials, anterior lower temporal meets 7th and 8th 
labials. Single nasal meets rostral, supranasal, 1st and 2nd labials, 
and 2 postnasals. Three loreals [two or three loreals] : 1st small 
between 2 postnasals, 3rd labial, 2nd loreal and canthal ; 2nd 
loreal large, between 1st loreal, 3rd and 4th labials, 3rd loreal, 
1st supraciliary, 1st supraocular and prefrontal. [1st loreal 
l)etween lower postnasal, 3rd and 4th labials, 2nd loreal, supra- 
ciliary, prefrontal and canthal] ; 3rd loreal between 2nd loreal, 
4tli and 5th labials, preocular and 1st supraciliary [2nd loreal 
has the same contacts]. Small mental two-third.s width of rostral; 
8, 7 [6 or 8] infralabials, prominent postmental in contact witli 
labials, 1st and 2nd followed by 4 pairs of chin shields, 1st pair 
meeting in midline. Transparent area in lower eyelid covered by 
about 4 vertical scales. 

Middorsal scales from posterior part of neck to root of tail with 
prominent median keel. Keels reduced on tail at about level of 
hindfeet [keels on tail a little more pronounced]. From occiput 
to end of tail, dorsal scales have longitudinal striae. Keels and 
striae fade on lower flank. Scales on underside of body quite 
smooth. Scales on forelimb and hindlimb with striae and slight 
keels on dorsal surface only [no keels on scales of limbs]. Soles 
of feet with swollen soft scales. Vertical ridge-like scale in groove 
between 3rd and 4th toes and soft fold of skin on proximal 
posterior face of 3rd toe [this condition of the 3rd and 4th toes 
is not developed]. Dorsal scales of fingers smooth, dorsal scales of 
toes striated [no striation on scales of toes]. 

Forty-one (43) scale rows at midbody [36-40, also confirmed by 
Boulenger (1885) and Taylor (1956)] ; mental to vent 88 scales 
[85, 90] ; occipital to point above vent 91 scales (occiput to base 
of tail 86) [72-79] ; from anal to chest on line joining axillae 55 
scales [53, 55] ; at level of ankles 20 scale rows around tail [23, 
21]. 

Third digit of forelimb slightly longer than 4th ; lamellae 3rd 
digit: 8, 8 [7 or 8] ; lamellae 4th digit 7, 8 [7 or 8]. Length of 
4th digit of hindlimb to length of 3rd as 1.2:1; 4th toe lamellae 
11,11. 



1964 



AN ANGUID FROM MONTSERRAT 



As indicated, slides were made of scales from various ])arls of 
the body; of the monofropis only ]\ICZ 37139 had sufficient scales 
for a parallel set of i)reparations. 

There are numerous scale organs on the prefrontal, frontal, 
frontoparietal, parietal and occipital scales; there are none on the 
interparietal [many]. 

Ten rows of ten dorsal scales (less three missing) had an 
average of 4.94 [4.71] scale organs per scale. The totals of scale 
organs for ten longitudinal rows of ten showed a variance of 
16.7 per cent ; the totals for ten transverse rows of ten showed 
a variance of 10.6 per cent. This suggests that a transverse or 
oblique row gives a more reliable count than a longitudinal row. 
Six of the scales lacked central keels ; for the others the central 
keel divided the scale organs into two groups, mesial and lateral 
[central keel does not separate two groups]. Counting the scale 
rows from the vertebral line, the majority of scale organs lie on 
the lateral side of rows 1 and 2, the mesial side of rows 3 and 4, 
and all on the mesial side of rows 5 and 6. 

Descending the flank the scale organs become reduced in num- 
ber; on the belly there are none [on belly 2.08 organs/scale]. 

Three rows of ten scales forwards from the groin had a mean 
of 3.43 organs/scale [4.8]. On the dorsal tail scales there are 
very many scale organs, 42 - 73/scale [9 - 11/scale], falling to 
none on the underside [3 - 4/scale]. 




Fig. 3. Mesial view, left ramus of lower jaw of Diploglossus montisscrrati 
type. 



Left dentary bears 20 teeth. Teeth cylindrical ; crowns of an- 
terior teeth laterally compressed, crowns of posterior teeth 
rounded; many of the teeth show signs of wear on the lingual 
side of the crown. The outer face of the dentary has six foramina. 
I am indebted to Dr. Robert Hotfstetter for drawing my attention 
to the existence of a foramen on the inner face of the surangular 
before the posterior branch of the coronoid in the Cordylidae and 
some Anguidae. This specimen has two foramina in this position. 



BREVIORA 



No. 200 



The osteoderms show the network of canals in the basal area 
(buried in the dermis), as described and figured by Hoffstetter 
(1962) for D. monotropis. The distal, epidermis-covered area 
shows radiating canals. The scale organs lie in the portion of the 
skin beyond the margin of the osteoderm. It seems unlikely 
therefore that scrutiny of the osteoderm would give an indication 
of their presence. 




Fig-. 4. Tjower, middorsal osteodcini ; upper left, middorsal scale; 
upper riglit, dorsal tail scale: Diploglossus montisserrnti type. 



General coloration medium bi-owii. Top of head uniform brown, 
upper labials white with a few brown speckles mostly on margins, 
lower labials and underside of head immaculate. Starting on 
neck indistinct darker brown lines; aci-oss width of 12 dorsal 
scale rows plain brown; on flanks and onto side of tail speckles 



1964 AN AXGUID FROM MONTSERRAT 7 

generally covering one or two scales consisting of a dark brown 
mark followed by white ; regenerated portion of tail darker and 
more uniform brown. Legs with speckles similar to those on 
flanks. Underside of body immaculate. D. monotropis has dark- 
edged, white crossbands which Boulenger notes may be broken 
into ocelli on the flanks. Dr. Ernest Williams notes a constant 
dark streak on the temples of moyiotropis as an additional dif- 
ference from montisserrati. 

Dimensions (contorted in preservation) : snout to vent c. 180 
mm, tail (half regenerated) c. 165 mm, forelimb 41 mm, hind- 
limb 45 mm, axilla to groin c. 95 mm. 

DISCUSSION 

We may well ask if the resemblance between monotropis and 
montisserrati indicates close relationship. Reference to the table 
(1959a, p. 13) in my previous paper shows that before the discov- 
ery of montisserrati the only species to combine the four charac- 
ters which I hesitatingly regarded as primitive was monotropis. 
The scale organs on the belly, which I did not notice in 1959, may 
represent an additional primitive character. The only differences 
which are greater than moderate differences of degree are the color 
pattern, the lack of scale organs on the interparietal and on the 
belly, and the very numerous scale organs on the dorsal side of the 
tail and, perhaps, the dorsal scale count. It is therefore difficult to 
suppose that the two species are not related and I suggest that it 
is plausible to assume that, widely separated as they are geograph- 
ically, they have diverged but little from a common ancestral 
stock. As I have previously suggested (1959a, p. 11) monotropis 
appears to be related to fasciatus and resplendens in South Amer- 
ica, to millepunctatus on lonely Malpelo Island and perhaps to the 
limbless Ophiodes. I have not examined material of bilohatus and 
lessonae but it seems possible that these species may be related to 
the monotropis stock with the difference that they have a single 
prefrontal scale (Boulenger, 1855; Schmidt and Inger, 1951; 
Taylor, 1956). In connection with this difference it may be noted 
that the sutures separating the median and lateral prefrontals of 
montisserrati are incomplete. The species of northern Central 
America and the Greater Antilles differ more markedly from the 
monotropis group. In the Greater Antilles, Jamaica and Hispan- 
iola have a diversity of forms while Cuba and Puerto Rico have 
onl}' one species each. The Puerto Rican species has not been 
reported in the Virgin Islands. 



8 BREVIORA No. 200 

It is interesting", therefore, to consider how montisserrati may 
have reached the Leeward Islands. There would seem to be two 
alternatives: either that monotropis-like Diploglossus was once 
widely spread in northern South America and from there reached 
Montserrat, or that monotropis-like stock reached the Greater 
Antilles from Central America and .sj^read eastward as far as 
the Leeward Islands. 

I regard arrival in the Leeward Islands via the Greater Antilles 
as unlikely. No forms are known in the Greater Antilles which 
resemble the monotropis group. We would have therefore to 
suppose that the monotropis stock passed through these islands 
but left no survivors, or that it underwent extensive evolutionary 
transformation there and gave rise to some of the now dissimilar 
modern forms yet survived with little change in the Leeward 
Islands. I have perhaps been too ready to assume in other cases, 
without detailed examination, that a genus which occurs in the 
Greater Antilles and the Leeward Islands has spread eastwards 
from Central America (Underwood, 1963). 

I consider the proposition more plausible that the monotropis 
group was once more widely distributed in northern South 
America. It is a long way from the mainland to Montserrat but 
the journey does not seem much more improl)able than the 
journe}' to Malpelo ; as there are many intervening islands the 
journey may not in fact have been made in one stage. As the 
ocean drift is westwards the take-off point would perhaps be 
somewhere in the Guianas. That there have been changes in the 
fauna of the coastal belt of northern South America is very 
likely. Phyllodactyliis has a distribution whicli is interesting in 
the present connection (Dixon, 1962). It occurs in Barbados, 
Grenada and Puerto Rico ; however, the easternmost record in 
northern South America is Patos Island in the Gulf of Paria, 
between Trinidad and Venezuela. The large and conspicuous 
Anolis richardii is common in Tobago, Grenada, and the Grena- 
dines and St. Vincent but none is known in Trinidad or on the 
mainland (Underwood, 1959b). Evidence of the former occur- 
rence or present survival of the monotropis group in the area 
from the Leeward Islands to the Guianas may yet be discovered. 
Diploglossus (lessonae) does in fact occur in eastern Brazil 
(Schmidt and Inger, 1951) south of the Amazon. 

What is the present status of Z>. montisserrati in Montserrat? 
Whatever may have been tlie manner of its arrival, it is now a 
relict, far removed geographically from relatives. I am inclined to 



]i*64 AN ANGUID F1{()IM IMONTSKRIJAT 9 

regard it as a survivor of the pluvial period, Avliicli lias persisted 
in a favourable habit<it in .spite of the recent climatic changes. 
llllUt barhudcnsis, on the other hand, is an example of a species 
wliieh Avas a])parently unable to survive the drying of the climate 
(Autt'enberg, 19.58). 

Although Montserrat is a small island (321/2 sq. mi.), it is quite 
mountainous with peaks high enough (up to 3000 ft.) to pre- 
cipitate rainfall, and has a good cover of moist forest and a con- 
siderable number of small rivers. Woodlands Spring is on the 
west side of the Island and, according to the 1 :50,000 survey map 
(Directorate of Overseas Surveys) it drains into a small stream 
called Runaway Ghaut. 

Mr. Kingsley Howes writes, "This was the second one that I 
have seen, so they are very rare on the island. The first one I 
saw was about 25 years ago. . . . They were both seen near fresh 
water. The first Avas at sea level near the estuary of a stream, 
the second at Woodlands Spring, the elevation being 600'. I did 
not see either of them feeding, but it is po.ssible that the second 
was feeding on young crayfish which were scurrying around the 
wet rocks where it was found." The stomach of the specimen in 
fact appears to be empty. The occurrence of both near fresh 
water would seem to fit in with the idea that it may be a relict 
of the pluvial period. The fact that there are many damp situa- 
tions in the island means that there may be other surviving 
populations. Taylor describes a specimen of monoiropis taken 
"in vegetation growing at the edge of Lake Bonilla . . . Costa 
Rica." 

Most of the reptile stocks of the Leeward and AVindward 
Islands range through several adjacent islands. The montisscrrati 
stock may well have been more widely distributed in the Leeward 
Islands and indeed may survive on others besides Montserrat. 
Survival on the limestone Leeward Islands, from Anguilla to 
Marie Galante, is unlikely ; they are low lying and dry. Some of 
the volcanic Leeward Islands, from Saba to Basse Terre (Guade- 
loupe), on the other hand, have some good wet forest on their 
higher slopes. Montserrat is one of the islands on which mongoose 
have not been introduced and this may have been a factor in its 
survival. Both St. Kitts, Nevis and Guadeloupe have mongoose; 
however, the mongoose may not extend up to the elevation at 
which the wet forest persists. Saba and St. Eustatius have no 
mongoose but they have very limited moist forest, in fact, only 
inside the crater on St. Eustatius. Several collections of bones 



10 BREVIORA No. 200 

have been made in the Leeward Islands and it would not be 
surprising- if the remains of DipJoglossus were found on islands 
on which it no longer occurs. For this reason I have figured the 
lower jaw and an osteoderm. 

I owe the specimen on which this paper is based to the gen- 
erosity of Mr. Kingsley Howes of Woodlands, Montserrat. I am 
indebted to Dr. Ernest Williams for the loan of the specimens 
of monotropis and for discussion of my manuscript, and to Mr. 
Benjamin Shreve for the report upon additional specimens of the 
mainland species. 

LITEEATUEE CITED 

AUFFENBERG, W. 

1958. A small fossil herpetofauna from Barbuda, Leeward Is., with 
the description of a new species of Hyla. Quart. J. Florida Acad. 
Sci., 21: 248-254. 

BoULEiNGER, G. A. 

1885. Catalogue of lizards in the British Museum (Natural History). 
2d ed., vol II, xiii + 497 pp. 
Dixon, J. 

1962. The leaf -toed geckos, genus Phyllodaclylus, of northeastern South 
America. Southwestern Naturalist, 7: 211-226. 

HOFFSTETTER, E. 

1962. Observations sur les osteodermes et la classification des Anguides 
actuels et fossiles (Eeptiles, Sauriens). Bull. Mus. Nat. Hist. 
Natur., Paris, 34: 149-157. 

Schmidt, K. P. and E. F. Inger 

1951. Amphibians and reptiles of the Hopkins-Branner expedition to 
Brazil. Fieldiana, 31: 439-465. 
Taylor, E. H. 

1956. A review of the lizards of Costa Eica. Univ. Kansas Sci. Bull., 
28: 3-322. 
Underwood, G. 

1959a. A new Jamaican galliwasp (Sauria, Anguidae). Breviora, Mus. 

Comp. Zool., No. 102: 1-13. 
1959b. The anoles of the eastern Caribbean (Sauria, Iguanidae). Part 
III. Eevisionary notes. Bull. Mus. Comp. Zool., 121: 191-227. 

1963. Eeptiles of the eastern Caribbean. University of the West 
Indies, Dept. of Extramural Studies, Jamaica, 191 pp. 



BREVIORA 

MnaseiLiei of Coimparative Zoology 



CvMBRiixiE, Mass. Ai'hil 10, 1<>()4 Number 201 

FOOD HABITS AND YOUNG STAGES OF 
NORTH ATLANTIC ALEPISAURUS (PISCES, INIOMI).^ 

By Richard L. Haedrich 

Mui^eum of Comparative Zoology, Harvard University 

INTRODUCTION 

Other than a few general statements (Clemens and Wilby, 
196U157; Gibbs and Wilimovsky, in press), there is very little 
information concerning food habits of the lancetfishes, genus 
Alepisaurus. During a recent exi)loratory fishing cruise (63-4), 
the U.S. Fish and Wildlife R/V DELAWARE obtained stomach 
contents from 36 Alepisaurus ferox and four .4.. brevirostris, taken 
by longline at 16 stations from the offing of New England to the 
Azores. These collections allowed an investigation of the food of 
Alepisaurus across a large portion of the North Atlantic over a 
relatively short period of time. In addition to the longline stations, 
four hauls were made with a modified Scharfe midwater trawl 
(Scharfe, 1960). Since the hauls were made in approximately the 
same levels as those in which the longlines fished, examination of 
this material provided some information concerning the selectivity 
of the lancetfish in its feeding. 

The longlines fished from the surface to about 80 fathoms. At 
most longline stations, sets were made at about 0600 hours and 
were hauled before 1500. Each midwater haul lasted less than an 
hour and reached a depth of 10 to 20 fathoms. All hauls were made 
at night, when vertically migrating animals would be expected to 
reach the upper limits of their depth ranges. The positions of the 
longline stations at which Alepisaurus were taken and the mid- 
water trawl stations of DELAWARE 63-4 are shown in Figure 1. 

Only five Alepisaurus smaller than 200 mm SL (standard 
length) were available to Gibbs (1960) for comparative purposes, 
and these could not be specifically identified. In the DELAWARE 



1 Contriliutioii Xo. 14-12 linin i\w Woods Hole Oa'anognipliic Institution. 



BREVIORA 



NO. 201 



material, AlepisaurKs was found to prey heavily on its own kind, 
and 42 young lancetfishes ranging from 50 to 585 mm SL were ob- 
tained from stomachs. Series of both species made specific identifi- 
cation of such small specimens possible. 







X ." 



ie«X 




Figure 1. Positions of longline stations at wliicli Alciiisauni.^ were taken 
( • ) and midwater trawl stations ( X ) of DELAWARE 63-4. 



ACKNOWLEDGMENTS 

Peter C. Wilson and personnel of the Bureau of Commercial 
Fisheries, Gloucester, made possible the participation in DELA- 
WARE 63-4 by representatives of the Museum of Comparative 
Zoology, Cambridge, and the Woods Hole Oceanographic Institu- 
tion. Jonathan L. Treible, representative of the Museum of Com- 
parative Zoology for the duration of the cruise, and Martin R. 
Bartlett, Woods Hole Oceanographic Institution, made the collec- 
tions and provided information concerning depths fished by long- 
lines and midwater trawls. Malcolm L. Clarke, National Institute 
of Oceanography, England, identified tentatively the cephalopods. 
Daniel M. Cohen, LTnited States National Museum, Washington, 
D.C., commented on the nomenclature of Lepidion. Richard H. 
Backus, Woods Hole Oceanographic Institution, and Giles W. 
Mead, Museum of Comparative Zoology, have read and criticized 
the manuscript. My work was supported by a Woods Hole Oceano- 
graphic Institution Summer Fellowship. Half the ship's costs for 
this cruise were paid for by the National Geographic Society. 
The material is housed in the Museum of Comparative Zoology. 



1964 FOOD AND YOUNG OF ALEPISAURUS 3 

IDENTIFICATION OF YOUNG STAGES 

Two series of Alepisaurus were present in the stomach contents. 
The largest fish in each scries was identifiable by characters pro- 
posed by Gibbs (1960), notably the shape of the head, the body 
proportions, and the melanophore structure. The largest A. bre- 
virostris was 585 mm SL; the largest A. ferox was 267 mm SL. 

As Gibbs (1960) observed, marked changes in proportions occur 
with growth, and the morphometric diff'erences which separate 
large specimens cannot be used to identify small lancetfishes. 
However, a distinctive melanophore pattern and the relative posi- 
tion of dorsal and pectoral fins do distinguish the young fishes. 
Examination of one series, the largest specimens of which were 
clearly Alepisaurus brevirostris, showed melanophores along the 
midventral surface of the body at all growth stages seen (38 speci- 
mens, 49-585 mm SL). These melanophores were absent in the 
other series (5 specimens, 40-267 mm SL), the largest specimen 
of which was clearly A. jerox. Since A. ferox greater than 500 mm 
SL do have a fine peppering of melanophores on the belly, these 
spots must appear at some stage in growth between about 270 and 
500 mm. The origin of the dorsal fin was in advance of the origin 
of the pectoral fin in all A. brevirostris, whereas the dorsal origin 
was over or behind the pectoral origin in all A. ferox. 

Alepisaurus from 260 mm SL down to at least 40 mm SL can 
be separated as follows: 

I. Melanophores present on belly, dorsal origin in advance of 
pectoral origin, snout profile convex 

AJrpisavrus brevirostris Gibbs 

II. Melanophores absent from belly, dorsal origin over or be- 
hind pectoral origin, snout profile generally straight (cf. Fig. 

2) Alepisaurus ferox Lowe 

Anal and pectoral fin-ray counts were taken from the young 

fishes. Many specimens were damaged, hence dorsal counts were 
impractical. The meristic data (Table 1 ) show a similar modality 
to that found by Gibbs (1960). 

TABLE 1 

Fin-ray count frequencies in young Alepisaurus 
Anal rays Pectoral rays 

13 14 15 16 17 18 12 13 14 15 16 

A. brevirostris 2 13 16 6 2 18 17 1 

A. ferox 13 1 112 1 



BREVIORA 



XO. 201 





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1964 FOOD AND YOrXO OF ALEPISAURUS 5 

Reported here for the first time is ti 40 nuu lancetfish, MCZ 
42377, taken l)y Isaacs-Kidd niidwater trawl during a shakedown 
cruise of R/V ANTON BRUUN; 39°10'N, 71°52'W; 1015-1420 
hr.; 9 Jan. 1963. The snout of this small fish is curved in profile, 
hut there are no melanophores along the belly and the origin of 
the dorsal is well behind that of the pectoral. There are 16 rays in 
the anal fin, and 14 rays in the pectoral. I identify this specimen 
as Alepisaurus ferox, and include it in the above analysis. The 
56 mm Alepisaurus figured by Maul (1946: 155, fig. 20) is refera- 
ble to A. brevirostris on the l)asis of head shape and fin positions. 

The three specimens of Alepisaurus brevirostris from 23^-'30'W 
(Table 2, Sta. 9, Coll. 2) constitute an eastward range extension 
for the species of some 1480 miles. Gibbs and Wilimovsky (in 
l)ress) report no specimens east of 57° W. 

FOOD OF ALE PI SAVE US 

(Table 2) 

Morning longline sets produced far more lancetfish than night 
sets. Twenty per cent of the longline stations were made at night, 
but only one of 45 Alepisaurus taken was caught on a set which 
fished primarily during the dark hours. This may indicate that 
Alepisaurus is a day-time feeder, but might also suggest that the 
bait is not readily seen by the lancetfish at night. In the dark, 
luminescent animals, such as the hatchetfish Sternoptyx, may be 
more easily detected. No difference, however, was noted between 
the stomach contents of the night-caught Alepisaiirus (Sta. 24) 
and those of fish from morning sets. 

Most animals from Alepisaurus stomachs were in excellent con- 
dition. Few had been digested to any extent. Rofen (in jiress) sug- 
gests that digestion proceeds almost entirely in the intestine of 
Alepisaurus, and that the stomach serves only for storage. This 
may be a mechanism allowing the lancetfish to dine at ojiportunity 
and to digest at leisure. 

TABLE 2 

Alepisaurus stomach contents, DELAWARE 63-4, 27 April to 7 June 1963. 
Each collection represents one jar, usually the stomach contents from one 
fish. In some cases the stomach contents from two or more fish were put in 
one jar; these appear as single collections with the number of stomachs in- 
dicated. The fork length (FL) of the fish from which the stomach contents 
were obtained is given when known. Ximiber of specimens and range of 
standard lengths of fishes in riglilhand column. 



6 



BREVIORA 



NO. 201 



Station 1. 40°09'N, 49°54'W. 0645-1510 hr. 27 Apr. 1963 



Collection 1, from A. ferox: 

Chiasmodon niger 
Diretmus argenteus 
JJrophycis sp. 
Alepisaurus breviroslris 
Paralepis coregonoides 
Carinaria lamarcki 
Pterotrachea sji. 
Phrosina sp. 
polychaete worm 



(Pisces, Percoidea) 
(Pisces, Berycomorphi) 
(Pisces, Anacanthini) 
(Pisces, Iniomi) 
(Pisces, Iniomi) 
(Mollusca, Heteropoda) 
(Mollusca, Heteropoda) 
(Crustacea, Amphipoda) 15 

1 



2, 19 & 38 mm 

1, 14 mm 
1,21mm 
4, 50-75 mm 

2, 41 & 45 mm 
7 

10 



Collection 2, from A. ferox: 
Schedophilus medusophagus (Pisces, Stromateoidea) 1, 120 mm 

Collection 3, from A. ferox: 
Carinaria lamarcki 



Pterotrachea sp. 
Phronima sp. 
Phrosina sp. 

Collection 4, from A. ferox: 

Paralepis coregonoides 
Pyrosoma sp. 
Carinaria lamarcki 
Pterotrachea sp. 
Phronima sp. 
Phrosina sp. 

Collection 5, from A. ferox: 

2 small pieces of scjuid 
Carinaria lamarcki 
Pterotrachea sp. 
Brachyscelus sp. 
Phrosina sp. 

Collection 6, from A. ferox: 

Alepisaurus breviroslris 
Notolcpis rissoi 
Paralepis coregonoides 
Argonauta sp. 
Carinaria lamarcki 
Pterotrachea sp. 
Phrosina sp. 
polychaete worm 



(Mollusca, Heteropoda) 3 
(Mollusca, Heteropoda) 39 
(Crustacea, Amphipoda) 2 
(Crustacea, Amphipoda) 1 



(Pisces, Iniomi) 2, 48 & 49 mm 

(Tunicata, Thaliacea) 1 

(Mollusca, Heteropoda) 21 
(Mollusca, Heteropoda) 65 
(Crustacea, Amphipoda) 1 
(Crustacea, Amphipoda) 7 



(Mollusca, Cephalopoda) 
(Mollusca, Heteropoda) 2 
(Mollusca, Heteropoda) 31 
(Crustacea, Amphijioda) 1 
(Crustacea, Amphipoda) 16 



(Pisces, Iniomi) 1,365 mm 

(Pisces, Iniomi) 1, 89 mm 

(Pisces, Iniomi) 1, 62 mm 

(Mollusca, Cephalopoda) 1 

(Mollusca. Heteropoda) 5 

(Mollusca, Heteropoda) 6 

(Crustacea, Amphipoda) 5 



1904 



FOOD AND YOUNG OF ALEPISAUHUS 



Collection 7, fioiu A. jerox: 

Alepisaurus brevirostris 
Paralepis sp. 
Octopodoteiithid? 
Carin a ria la m a rcki 
Pterolrachea sp. 
Cavolinia sp. 
Brachyscelus sp. 
Phronima sp. 
Phrosina sp. 
nemertean 
polychaete worms 



(Pisces, Iniomi) 


1, 585 mm 


(Pisces, Iniomi) 


1,50 mm 


(Mollusca, Cephalopoda) 


1 


(MoUusca, Heteropoda) 


9 


(Mollusca, Heteroputla) 


6 


(Mollusca, Pteropoda) 


2 


(Crustacea, Amphipoda) 


20 


(Crustacea, Amphipoda) 


2 


(Crustacea, Amphipoda) 106 

1 




X 

18 



Station 2. 39°20'N, 44°50'W 

Collection 1, from A. jerox : 

Brama brama 
Alepisaurus brevirost ris 
Paralepis coregonoides 
Paralepis sp. 
Stemoptyx diaphana 
fish larvae 
doliolid test 
octopod 

Pterolrachea sp. 
brachyuran 
Palmurus larva 
Brachyscelus sp. 
Phronima sp. 
Phrosina sp. 

Collection 2, from A. jerox : 

Lepidion lepidion 
Alepisaurus brevirost ris 
Notolepis rissoi 
Paralepis coregonoides 
Chiroteuthis sp. 
Taoniinae 
Carinaria lamarcki 
Pterolrachea sp. 
brachyuran 
small euphausids 
Brachyscelus sp. 
Hyperoche sp. 
Phronima sp. 
Phrosina sp. 



0650-1430 hr. 29 Aiir. 1963 




(Pisces, Percoidea) 


2, 34 & 101 mm 


(Pisces, Iniomi) 


1,223 mm 


(Pisces, Iniomi) 


1, 164 mm 


(Pisces, Iniomi) 


36, 23-60 mm 


(Pisces, Stomiatoidea) 


1, 34 mm 
13 

1 


(Tunicata, Thaliacea) 


(Mollusca, Cephalopoda) 


1 


(Mollusca, Heteropoda) 


2 


(Crustacea, Decapoda) 


1 


(Crustacea, Decapoda) 


1 


(Crustacea, Amphipoda) 


34 


(Crustacea, Amphipoda) 


2 


(Crustacea, Amphipoda) 


9 


(Pisces, Anacanthini) 


2, 25 & 45 mm 


(Pisces, Iniomi) 


3, 60-295 mm 


(Pisces, Iniomi) 


1, 105 mm 


(Pisces, Iniomi) 


5, 30-108 mm 


(Mollusca, Cephalopoda) 


1 head 


(Mollusca, Cephalopoda) 


147 


(Mollusca, Heteropoda) 


5 


(Mollusca, Heteropoda) 


1 


(Crustacea, Decapoda) 


1 


(Crustacea, Euphausiacea) 


4 


(Crustacea, Amphipoda) 


1 


(Crustacea, Amphipoda) 


1 


( Crustacea, Amphipoda) 


3 


(Crustacea, Amphipoda) 


10 



8 



BREVIORA 



NO. 201 



Station 4. 40°17'N, 36°07'W. 0630-1435 hr. 1 May 1963 



Collection 1, from A.jerox : 

Carinaria laniarcki 
Pterotrachea sp. 
Phrosina sp. 
Eunjdice sp. 

Collection 2. from A. brci 

Alepisaunis bre virost ris 
(loliolid test, containinp; a 

Phronima sp. 
Carinaria laniarcki 
Cavolinia sp. 
Brachyscelus sp. 
Phronima sp. 
Phrosina sp. 
])()l>-chaete worms 

Collection 3, from A. Iircviroslriti: 

Carinaria laniarcki 
Pterotrachea sp. 
Cavolinia sp. 
Brachyscelus sp. 
Phronima sp. 
Phrosina sp. 
Platyscelus sp. 
polychaete worm 



(Mollusca, Heteropo(ia) 7 

(Molhisca, Heteropo(la) 1 

(Crustacea, Ami)hi])0(la) 2 

(Crustacea, Isoiiotla) 3 

'irostris : 

(Pisces, Iniomi) 1,64 mm 

(Tunicata, Thaliaeea) 1 

(Mollusca, Heteropoda) 15 

(Mollusca, Pteropoda) 4 

(Crustacea, Amphi]ioda) 17 

(Crustacea, Amphijioda) 3 

(Crustacea, Ami)hipoda) 50 

2 



(Mollusca, Heteropoda) 17 

(Mollusca, Heteropoda) 1 

(Mollusca, Pteropoda) 6 

(Crustacea, Amjihipoda) 44 

(Crustacea, Amphipoda) 2 

(Crustacea, Amphipoda) 125 

(Crustacea, Amphipoda) 1 

1 



Collection 4, from .4. hrcvirostris: 

Anolopterus pharno 
Alepisaurus brevirost ris 
Carinaria laniarcki 
Pterotrachea sp. 
CdvoUuia sp. 
Brachyscelus sp. 
Phrosina sp. 
polychaete worm 



(Pisces, Iniomi) 


1,277 mm 


(Pisces, Iniomi) 


2, 48 & 56 mm 


(Mollusca, Heteropoda) ca 


.60 


(Mollusca, Heteropoda) 


3 


(Mollusca, Ptero])0(la) 


7 


(Crustacea, Amphipoda) 


69 


(Crustacea, Amphipoda) 


256 

1 



Station 6. 41 20'X, 2!) 20'\V. 1805-1050 hr. 3-4 May 1963 
Collection 1, from A. jerox 1365 mm FL: 



Paralepis coregonoides 
Carinaria lamarcki 
Pterotrachea sp. 

Phronima sj). 
Pln'osiiKi sp. 



iV 



isces. 



Iniomi) 



( M ollusca , Hct eropoda ) 
( Mollusca, Het(>rop()da) 
( Crust ac(>a, A ii 1 1 >! i i | loda ) 
( Crust acea , A m ph i | loda ) 



1, 123 mm 
23 
12 

1 

(I 



19(U 



FOOD AND YOrXO OF ALEPISArRUS 



Station 9. 36°21'N, 23°30"W. 0635-1530 hr. 8 May 1963 



Collection 1, from .1. jvrox: 

Urophycis sp. 
fish lar\a 

Carinana lammrld 
Pterotrachea sp. 
Brachysceltis sp. 
Phrosina sp. 

Collection 2, from A. fcrox: 

Lophius piscatorius 

Cubiceps gracilis 

Schedophilus medusophagits 

Alepisaurus brevirostris 

Stemoptyx diaphana 

fish larvae 

doliolid tests, one containing 

a, Phro7iimo 
unident. squids, 4 spp. 
Carinaria lamarcki 
Pterotrachea sp. 
Phronima sp. 
Phrosina sp. 



(Pisces, Anacantliini) 

(MoUusca, Heteropoda) 
(Mollusca, Heteropoda) 
(Crustacea, Amphipoda) 
(Crustacea, Amphijioda) 



(Pisces, Pediculati) 
(Pisces, Stromateoidea) 
(Pisces, Stromateoidea) 
(Pisces, Iniomi) 
(Pisces, Stomiatoidea) 



1, 9 mm 
1, 14 mm 
1 
29 
1 

9 



2, 40 & 48 mm 
1, 65 mm 

1, en. 60 mm 

3, 69-178 mm 

2, 24 & 27 mm 
8 

5 



(Mollusca, Cephalopoda) 4 
(Mollusca, Heteropoda) 9 
(Mollusca, Heteropoda) 220 
(Crustacea, Amphipoda) 5 
(Crustacea, Amphipoda) 5 



Station 14. 37°25'X, 29 lO'W. 0610-1335 hr. 16 May 1963 
Collection 1, from A. jernx 1585 mm FL: 



Schedophilus medusophagits 
Paralepis coregonoides 
Carinaria lamarcki 
Pterotrachea sp. 



(Pisces, Stromateoidea) 
(Pisces, Iniomi) 
(Mollusca, Heteropoda) 
(Mollusca, Heteropoda) 



3, 33-40 mm 
1,129mm 
11 
6 



Station 15. 36 55'N, 32°32'W. 0400-1110 la-. 17 May 1963 
Collection 1, from A. jerux 1437 mm FL : 



Lophius piscatorius 
Schedophilus medusoplwgiis 
Alepisaurus brevirostris 
Stemoptyx diaphana 
doliolid test 
Argonauta sp. 
Phronima sp. 
Phrosina sp. 



(Pisces, Pediculati) 
(Pisces, Stromateoidea) 
(Pisces, Iniomi) 
(Pisces, Stomiatoidea) 
(Tunicata, Thaliacea) 
(Molkhsca, Cephalopoda) 
(Crustacea, Amphipoda) 
(Crustacea, Amphipoda) 



2, 62 & 80 mm 
1, 35 mm 
1,98 mm 
28-30 mm 



10 



BREVIORA 



NO. 201 



Collection 2, from ^4. jerox 1342 mm FL: 



Paralepis atlantica 
Stemoptyx diaphana 
doliolid test 
Taoniinae 
Phronima sp. 
Phrosina sp. 
Platyscelus sp. 
nemerteans 
polychaete worm 



(Pisces, Iniomi) 

(Pisces, Stomiatoidea) 
(Tiinicata, Thaliacea) 
(Mollusca, Cephalopoda) 
(Crustacea, Amphipoda) 
(Crustacea, Amphipoda) 
(Crustacea, Amphipoda) 



1, 98 mm 

1,28 mm 

1 

1 

1 
24 

2 
18 

1 



Station 16. 33°20'N, 39°50'W. 0410-1110 lir. 19 May 1963 



Collection 1, from A. jerox 1265 mm FL: 



Cubiceps gracilis 
Alepisaurus brevirostris 
Macro paralepis affine 
Stemoptyx diaphana 
fish larvae 
doliolid tests 
salps 

Argonaata sp. 
Japetella sp. 
unident. scjuids 
Carinaria lamarcki 
Pterotrachea sp. 
Cavolinia sp. 
Limacina sp. 
Braclii/scchis sp. 
Phruiiiina sp. 
PJiromia sp. 
Platyscelus sp. 
]iolychacte worms 



(Pisces, Stromateoidea) 
(Pisces, Iniomi) 
(Pisces, Iniomi) 
(Pisces, Stomiatoidea) 

(Tunicata, Thaliacea) 
(Tunicata, Thaliacea) 
(Mollusca, Cephalopoda) 
(Mollusca, Cephalopoda) 
(Mollusca, Cephalopoda) 
(Mollusca, Heteropoda) 
(Mollusca, Heteropoda) 
(Mollusca, Pteropoda) 
(Mollusca, Pteropoda) 
( Crustacea, Amphipoda ) 
(Crustacea, Ami)liipuda) 
(Crustacea, Amphipoda) 
(Crustacea, Amphipoda) 



Collection 2, from A. jerox 1292 mm FL: 
Alepisaurus brevirostris (Pisces, Iniomi) 



Alepisaurus jerox 
Carinaria lamarcki 
Phrosina sp. 
Lanceola sp. 



(Pisces, Iniomi) 
(Mollusca, Heteropoda) 
(Crustacea, Amphipoda) 
(Crustacea, Isopoda) 



Collection 3, from A. jerox 1325 mm FL: 

Alepisaurus brevirostris (Pisces, Iniomi) 

Paralepis atlantica (Pisces, Iniomi) 

Stemoptyx diaphana (Pisces, Stomiatoidea) 
nemerteans 



2, 76 & 83 mm 

7, 71-439 mm 

2, 64 & 114 mm 
21, 8-31 mm 
17 
29 
146 

2 

7 

2 

2 
remnants 
15 

3 
36 
32 
14 
42 

5 



2, 113 & 124 mm 

1, 267 mm 

1 

1 

1 



1,486 mm 
1, 84 mm 
1, 22 mm 
15 



1964 



FOOD AND YOUNG OF ALEPISAURUS 



11 



Collection 4, from A.jerox : 
Alepisaunis brevirostris 



(Pisces, Iniomi) 



1, 437 mm 



Station 18. 31°00'N, 47°05'W. 0555-1320 hr. 21 May 1963 



Collection 1, from A. jerox 

Diplospinus multistriatus 
Cubiceps gracilis 
Alepisaunis brevirust ris 
Alepisaurus jerox 
Paralepis atlantica 
Paralepis elongata 
Sternoptyx diapliaiia 
fish larvae 
Pyrosoma sp. 
Abraliopsis sp. 



1166 mm FL: 

(Pisces, Scombroidea) 
(Pisces, Stromateoidea) 
(Pisces, Iniomi) 
(Pisces, Iniomi) 
(Pisces, Iniomi) 
(Pisces, Iniomi) 
(Pisces, Stomiatoidea) 

(Tunicata, Thaliacea) 



4, ca. 30 mm 
1, 84 mm 

1, 154 mm 
1, 91 mm 
1,96 mm 

5, 101-162 mm 
9, 18-30 mm 

35 
1 



(MoUiisca, Cephalopoda) 2 



Station 21. 32^00'N, 56°10'W. 0605-1320 hr. 24 May 1963 



Collection 1, from A. ferox 

Brama brama 
Alepisaurus brevirostris 
Paralepis sp. 
Sternoptyx diaphana 
Onychoteuthis sp. 
Carinaria lamarcki 
Cavolinia sp. 
Phrosina sp. 



1068 mm FL: 

(Pisces, Percoidea) 
(Pisces, Iniomi) 
(Pisces, Iniomi) 
(Pisces, Stomiatoidea) 
(Mollusea, Cephalopoda) 
(Mollusca, Heteropoda) 
(Mollusea, Pteropoda) 
(Crustacea, Amphipoda) 



Collection 2, from 3 A. ferox 

Bothus sp. 

Diplospinus multistriatus 
Alepisaurus brevirostris 
Alepisaurus ferox 
Paralepis coregonoides 
Lobianchia dofleini 
Sternoptyx diapliana 
fish larvae 
doliolid tests 
salps 

Argonauta sp. 
Japetella sp. 
Carinaria lamarcki 
brachyuran 
Brachyscelus sp. 
Phronima sp. 
Phrosina sp. 



941-1043 mm FL: 

(Pisces, Heterosomata) 
(Pisces, Scombroidea) 
(Pisces, Iniomi) 
(Pisces, Iniomi) 
(Pisces, Iniomi) 
(Pisces, Iniomi) 
(Pisces, Stomiatoidea) 

(Tunicata, Thaliacea) 
(Tunicata, Thaliacea) 
(Mollusca, Cephalopoda) 
(Mollusca, Cephalopoda) 
(Mollusca, Heteropoda) 
(Crustacea, Decapoda) 
(Crustacea, Amphipoda) 
(Crustacea, Amphipoda) 
(Crustacea, Amphipoda) 



2, 37 & 55 mm 
1,463 mm 
1, 75 mm 
11, 21-32 mm 
1 
1 
1 
1 



1, 19 mm 
7, 20-163 mm 

1, 81 mm 

2, 82 & 131 mm 
7, 68-99 mm 
1,31 mm 

5. 20-27 mm 

7 

4 

2 

4 

2 

3 

1 

1 

4 

3 



12 



BREVIORA 



NO. 201 



Lanceola sp. 
nemertean 
polyehaete worms 



(Crustacea, Isopoda) 



1 

1 

15 



Station 23. 32°05'N, 59 lO'W. 0605-1340 lir. 25 May 1963 

Colloctioii 1, from 4 .4. ferox 772-1276 mm FL: 

Diplospiiius DiullLst rial Its 
Anoplogaster cornuta 
Re go Ice us glesne 
leptocephalus larvae 
Alepisaurus brevirostris 
Pamlepis atlantica 
Paralepis coregonoicles 
Pamlepis eloiigoln 
Steriioptyx diaphana 
bits of Pyrosoma test 
unident. squids 
C armaria lamarcki 
Pterotrachea sp. 
Brachyscelus sp. 
Phroiiima sp. 
Phrosina sp. 
Platyscelus .sp. 
polyehaete worms 



(Pisces, Scombroidea) 


14, 27-131 mm 


(Pisces, Berycomorphi) 


1, 11 mm 


(Pisces, Allotriognathi) 


1, 160 mm 


(Pisces, Apodes) 


5, ca. 60 mm 


(Pisces, Iniomi) 


5, 82-383 mm 


(Pisces, Iniomi) 


2, 76 & 100 mm 


(Pisces, Iniomi) 


5, 87-ca. 110 mm 


(Pi.sces, Iniomi) 


2, 135 & 153 mm 


(Pisces, Stomiatoidea) 


12, 15-27 mm 


(Tunicata, Thaliacea) 




(Mollusca, Cephalopoda) 


2 


(Mollusca, Heteropoda) 


2 


(Mollu.sca, Heteroi)oda) 


1 


(Crustacea, Amphipoda) 


3 


(Crustacea, Amphipoda) 


1 


(Crustacea, Amphipoda) 


26 


(Crustacea, Ami:)hipotla) 


1 




15 



I 
I 



Station 24. 32°45'N, 64°36'W. 1855-0825 hr. 26-27 May 1963 



Collection 1. from ,4. jerox 1146 mm FL: 



Anoplogaster cornuta 
Paralepis elongata 
Sternopt yx diaphana 
Octopodoteuthis sp. 
Pholidioteuthis sp. 
polyehaete worm 



(Pisces, Berycomoriilii) 
(Pisces, Iniomi) 
(Pisces, Stomiatoidea) 
(Mollusca, Cephalopoda) 
(Mollusca, Cephalopoda) 



1. 45 mm 

2, 127 & 138 mm 
2, 27 & 30 mm 

1 
1 
1 



Station 29. 32'50'N, 68'15'\V. 0610-1340 hr. 1 June 1903 
Collection 1, from .4. ferox 1028 mm FL: 



Diplospin us ni utt isl rial u.^ 
St e rn ojit i/x dia pli a n a 
doliolid test 
Phronima sp. 
Phrosina sp. 
polyehaete worms 



(Pisces, Scombroidea) 
(Pisces, Stomiatoidea) 
(Tunicata, Thaliacea) 
(Crustacea, Amphipoda) 
(Crustacea, Anipliipoda) 



1, 116 mm 

2, 20 ct 22 mm 

1 

1 

1 

3 



1 i)iU 



FOOD AM) YOUNG OF ALKIMSAUHUS 



13 



Station 31. 34°45'N. 7ri5'\V. 

Coll(H'tion 1, rioni 5 .1. fcro-i 

Dij)l<)sj)iitiis Diiillisl ridtiis 
Brania bmma 
Anoplogaster coniula 
Alepisaiinif> brcvirosl ris 
Macro pamle pis ajjiiie 
Famlepis allantica 
Pa ra lepis coregonoidcs 
Pamlepis elongala 
Argyropelecus aculealus 
Slernoplyx diaphana 



0610-1515 hr. 2 June 


19G3 


:• 1039-1254 mm FL: 




( Pisces, iScoml)ioi(lca) 


1, 131 mm 


(Pisces, Percoiclea) 


2, 40 & 95 mm 


(Pisces, Berycomorplii) 


1, 75 mm 


(Pisces, Iniomi) 


2, 124 & 131 mm 


(Pisces, Iniomi) 


1, 150 mm 


(Pisces, luiomi) 


5, 93-lli) mm 


(Pisces, Iniomi) 


14, 79-120 mm 


(Pisces, Iniomi) 


2, 61 & 125 mm 


(Pisces, Stomiatoidea) 


1, 32 mm 


(Pisces, Stomiatoidea) 


6, 14-29 mm 



Station 33. 36°27'N, 67°50'W. 0610-1505 hr. 3 June 1963 
Collection 1, from A. jcrox 811 mm FL : 



D iplospin us tn ultist rial us 
Pa m le pis coregonoidcs 
Slernoptyx diaphana 
Pterotrachea sp. 
Phrosina sp. 
polychaete worms 



(Pisces, Scombroidca) 
(Pisces, Iniomi) 
(Pisces, Stomiatoidea) 
(Mollusca, Heteropoda) 
(Crustacea, Amphipoda) 



5, 61-194 mm 
3, 67-106 mm 
3, 20-29 mm 
2 

2 

2 



Station 37. 39°23'N, 68°33'W. 0620-1445 hr. 7 June 1963 
Collection 1, from A. brcvirostris 778 mm FL: 



Tetraodontidae 
Helicolenus dactyloplerii): 
Lepidion lepidion 
Syngnathus pelagicus 
Macroparalepis afjine 
Ommastrephidae 
Taoniinae 
Brachyscelus sp. 
Phronima sp. 
Phrosina sp. 



(Pisces, Plectognathi) 
(Pisces, Scleroparei) 
(Pisces, Anacanthini) 
(Pisces, Solenichthyes) 
(Pisces, Iniomi) 
(Mollusca, Cephalopoda) 
(Mollusca, Cephalopoda) 
( Crustacea, Amphipoda) 
(Crustacea, Amphipoda) 
(Crustacea, Amphipoda) 



1, 16 mm 

2, 24 & 25 mm 
2, 42 & 42 mm 
1,113 mm 

1, 138 mm 

2 

6 

3 

1 

2 



With regard to the size of its prey, Alepisaurus practices no 
discrimination. Adapted for swallowing large prey (Marshall, 
1955:330), it will also readily devour small creatures. A 1265 mm 
A. ferox (Sta. 16, Coll. 1 ) had eaten seven .4. brevirostris ranging 
from 71 to 439 mm in standard length; another A. ferox (Sta. 1, 
Coll. 7) contained a 585 mm A. brevirostris in addition to 106 
hyperiid amphipods, each about 20 mm long. 



14 BREVIORA NO. 201 

Lancetfish stomachs most commonly contained hyperiid amphi- 
pods, heteropods, paralepidids, Sternoptyx, and other Alepisaurus. 
At least one of these groups was found at every station, and three 
or more occurred simultaneously in 66 per cent of the collections. 
Of the 32 collections, amphipods occurred in 78 per cent, hetero- 
pods in 72 per cent, paralepidids in 59 per cent, Alepisaurus in 
56 per cent, and Sternoptyx in 44 per cent. Since these animals do 
occur so regularly in the stomachs, they must habitually freciuent 
the same depths as does Alepisaurus. 

Stomach contents reflected local abundances. Heteropods oc- 
curred in 17 out of 19 (89%) of the stomachs from stations 1 
through 15, and they were also a major constituent of the mid- 
water hauls. Heteropods were found in only 6 out of 13 (46%) 
of the stomachs from stations 16 through 37. In contrast, Sternop- 
tyx had been eaten by 77 per cent of the fish from stations 16 
through 37, but by only 44 per cent of the fish from stations 1 
through 15. On the southerly leg of the cruise, fishes predominated 
both in stomach contents and in the midwater haul. 

Myctophid fishes were notably absent from most stomach con- 
tents. Only one was found, a 31 mm Lobianchia dofleini (Sta. 21, 
Coll. 2). None occurred in the fish from station 9, although a one- 
hour midwater trawl at station 12 took 101 myctophids of a dozen 
species. The commonness of these small fishes in midwater hauls 
makes their absence from Alepisaurus stomachs particularly 
])uzzling. Myctophids are noted vertical migrators (Marshall, 
1960). If Alepisaurus does not migrate extensively, and if its prin- 
cipal depth differs from the day and night levels of myctophids, 
the duration of the lancctfish's encounter with myctophids would 
be brief during any diurnal cycle, and consumption of them there- 
fore low. 

Completely absent from the stomach contents were the charac- 
teristic middle-depth fishes Gonostoma, Stomias, and Chauliodus. 
These fishes are distributed mainly between 200 and 1500 meters 
(Haffner, 1952; Marshall, 1960). Their absence from Alepisaurus 
stomachs, and the presence of primarily epipelagic fishes such as 
Cubiceps, Brama, and Paralepis elongata strengthens the belief 
(Gibbs and Wilimovsky, in press) that the lancetfish frequents 
the upper layers. 

Little difference was noted between food of Alepisaurus ferox 
and A. brevirostris. A. brevirostris may favor invertebrates, par- 
ticularly the hyperiid amphipod Phrosina. All A. brevirostris 
stomachs contained these amphipods, but only 75 per cent of those 
from A. ferox did. There were 256 amphipods in an .4. brevirostris 



19G4 FOOD AND YOUNG OF ALEPISAURUS 15 

from Station 4 (Coll. 4), and only two in the A. ferox from the 
same station (Coll. 1). The maximum number of Phrosina from a 
single A. ferox was 106, and the average number per fish was 9.3. 
The maximum number of Phrosina from a single A. brevirostris 
was 256; 108.3 was the average. Fishes were more common in A. 
ferox stomachs. Sternoptijx and paralepidids occurred in 50 and 64 
per cent, respectively, of A. ferox stomachs, but were found in 
and 25 per cent, respectively, of the A. brevirostris stomachs. Since 
stomach contents were obtained from only four A. brevirostris, 
and since three came from one station, these comparisons are 
inconclusive. 

Rarely encountered fishes found in the stomach contents were 
Anotoptei'us pharao (Sta. 4, Coll. 4), Regalecus glesne (Sta. 23, 
Coll. 1), and four pelagic young of Lophius piscatorius (Sta. 9, 
Coll. 2; Sta. 15, Coll. 1). 

LITERATURE CITED 

Clemens, W. A. and G. V. Wilby 

1961. Fishes of the Pacific Coast of Canada. (Second edition). Fish. 
Res. Bd. Canada. Bull. No. 68, 443 pp. 
GiBBS, Robert H., Jr. 

1960. Alepisaurus brevirostris, a new species of lancetfish from the 
Western North Atlantic. Breviora, No. 123, 14 pp. 
GiBBS, Robert H., Jr. and Norman J. Wilimovsky 
In press. Family Alepisauridae, in Fishes of the Western North Atlantic. 

Sears Found. Mar. Res., Mem. 1, part 5. 
Haffner, Rudolph E. 

1952. Zoogeography of the bathjqjelagic fish, Chauliodtis. Syst. ZooL, 
1: 113-144. 
Marshall, N. B. 

1955. Alepisauroid fishes. Discovery Reports, 27 : 303-336. 
1960. Swimbladder structure of deep-sea fishes in relation to their sys- 
tematica and biology. Discovery Reports, 31 : 1-122. 
Maul, G. E. 

1946. Monografia dos peixes do Museu Municipal do Funchal. Ordem 
Iniomi. Bol. Mus. Municipal Funchal, 2(2) : 154-156. 
Rofen, Robert R. 
In press. Family Omosudidae, in Fishes of the Western North Atlantic. 

Sears Found. Mar. Res., Mem. 1, part 5. 
Scharfe, J. 

1960. A new method for one-boat trawling in midwater and on the 
bottom. General Fisheries Council for the Mediterranean, Studies 
and Reviews, ser. B, 21 pp. 



u^ 



v\^\^ 



BREVIORA 



Museeim of Coainparative Zoology 



Cambridge, Mass. April 10, 1964 Number 202 

THE BLIND SNAKES (TYPHLOPS) OF HAITI 
AVITII DESCRIPTIONS OF THREE NEW SPECIES^ 

By Neil D. Richmond 

Carnegie Museum, Pittsburgh, Pa. 

When the Herpetology of Hispaiiiola (Cochran, 1941) was 
published onlj^ 24 specimens of Typhlops were available from 
the entire island. Recent collecting (1950-1962) in Haiti has 
produced 93 specimens of blind snakes from that country alone, 
most of them now in the Museum of Comparative Zoology. 

Part of the field work was made possible by the following 
grants to the Museum of Comparative Zoology : National Science 
Foundation Grants NSF 16066, NSF 5634, and one from the 
American Philosophical Society. The six specimens from Gonave 
Island were obtained by Philip S. Humphrey collecting for the 
Yale Peabody Museum and the University of Florida. A descrip- 
tion of the various expeditions is given in Williams ct al., 1963. 

1 wish to express my appreciation to Ernest E. Williams who 
was responsible for getting this material together and making it 
possible for me to study it. Also I wish to thank Doris M. 
Cochran for permission to study the type of Typldops sulcatus. 
Miss A. G. C. Grandison supplied counts and descriptive notes 
on four specimens in the British Museum. Nicholas Strekalov- 
sky made the drawings of the new species herein described. 

The following abbreviations are used to designate the collec- 
tions where these specimens are deposited : AMNH, American 
Museum of Natural History; CM, Carnegie Museum; MCZ, 
Museum of Comparative Zoology; UF, University of Florida 
Museum; USNM, United States National Museum; YPM, Yale 
Peabody Museum. 



1 Notes on Hispanlolan herpetology no. 10. 



2 BREVIORA No. 202 

Examination of this material discloses that in addition to the 
two species previously known from Haiti, lumhricalis and piisil- 
lus, there are three forms new to science. All of the five species 
represented in this collection liave the following characters in 
common : relatively narrow strap-like rostrals ; completely di- 
vided nasals; preocular contacting the third upper labial only; 
scale rows either 20-20-20 or 20-20-18 ; at least one pair of 
enlarged parietal scales ; four upper labials ; three lower labials ; 
and clearly visible eyes. "With this great similarity they also 
show distinctive differences in the number of middorsal scale 
rows, number of preoculars, shape and size of ocular, shape of 
head, and color. 

In number of middorsal scales these snakes may be divided 
into two distinct groups : those with a relatively low number 
(240-330), two species, and those with over 385, three species. 
The three new forms are all in the group with the high number 
of middorsal scales. Two of these are further distinguished by 
having very small pointed heads. 

Typhlops capitulatus^ sp. nov. 
Figure 1 

Holotype: MCZ 62636. From Manneville, Haiti, at the north- 
west end of Lake Saumatre. Collected by A. S. Rand and J. 
Lazell, 10 August 1960. 

Paratype: MCZ 69006. From the same locality as the type. 
Collected by E. E. Williams and A. S. Rand, 13-14 August 1959. 





Fig. 1. Typhlops capiluhiius sp. iiov. Type MCZ 62636. From Manne- 
ville, Haiti. Dorsal and lateral views. 



iFroni tlip T^atin "having a small lipad." 



1964 Tvi'jiLoi's OF HAITI 3 

Diagnosis: A slender small-headed species of Typhlops char- 
acterized by the following combination of characters : high num- 
ber of dorsal scales, 385-400; scale rows 20-20-20; dorsal color 
extending on to venter, a white anal spot ; preocnlar almost 
triangular ; head small and pointed ; ocular almost as wide as 
high. 

Description: Sides of head tapering from about the level of 
the seventh middorsal scale. Rostral narrow, its width one-third 
that of the head, not extending to a line connecting the eyes ; 
nasals completely divided by a suture that extends from the 
.second upper labial to the rostral ; preocular almost triangular, 
approximately as high as wide and in contact with the third 
upper labial, its anterior edge curved but not elongated. Ocular 
broad, but little higher than wide, its posterior edge strongly 
convex (Fig. 1). Tavo pairs of enlarged parietal scales; two 
postoculars on each side. Eye small, scale rows around body 
20-20-20, dorsal scales from rostral to caudal spine about 400. 

Color: Light brown above, somewhat paler below although pig- 
mented, venter Avith scattered white scales, anal region and 
underside of tail white, dorsal color fades gradually on head 
to entirely unpigmented snout. 

Size: Total length 205 mm, head width at level of eyes 2.7 
mm, diameter at midbody 4 mm, tail slightly longer than wide. 

Variatio7i: MCZ 69006 is a very small specimen, total length 
93 mm, and is pale brown above and below although the charac- 
teristic pattern of pigmentation can be seen with magnification. 
This specimen has about 385 dorsal scales from rostral to spine. 
In detail of head scutellation it agrees with the type. 

Remarks: This species is most closely related to the species on 
Gonave Island, described below, from which it may be dis- 
tinguished by the shape of the preocular. 

Typhlops gonavensis sp. no v. 
Figure 2 

Holotype: YPM 3003. From Point a Raquette, on the south 
shore of Gonave Island, Haiti. Collected by Philip S. Humph- 
rey and Sarita Van Vleck, 9 April 1959. 

Paratypes: YPM 3004, and UF A943. With the same data 
as the holotype. 

Diagnosis: A slender small-headed species of Typhlops char- 
acterized by the following combination of characters : high num- 
ber of dorsal scales, 409-423 ; scale rows 20-20-20 ; dorsal color 



4 BREVIORA No. 202 

extending on to venter, a white anal spot ; may be distinguished 
from capitulatus which it most closely resembles by the dis- 
tinctive shape of the preocnlar which is wider than high and 
narrowest anteriorly. 

Description: Sides of head tapering from about the level of 
the seventh middorsal scale. Rostral narrow, its width one-third 
that of the head, not extending to the level of the eyes ; nasals 
completely divided by a suture that extends from the second 
upper labial to the rostral ; preocular wider than high, narrowest 
anteriorly (Fig. 2), in contact with the third upper labial; ocu- 
lar small but little higher than wide, its posterior edge strongly 
convex ; two pairs of enlarged parietals ; two postoculars on each 
side ; eye small ; scale rows around body, 20-20-20 ; middorsal 
scales from rostral to spine about 418. 

Color in preservative: Dark purplish brown above, somewhat 
paler below but heavily pigmented with an irregular white area 
in anal region. Dorsal color fades gradually on head to unpig- 
mented snout. Color in life (from notes by Sarita Van Vleck) : 
Reddish brown inferiorly, grading to burnt umber on tail, venter 
purplish gray anteriorly becoming pale gray-brown posteriorly. 

Size : Total length 189 mm, tail 4.2 mm ; head width at level 
of eyes 2.7 mm ; diameter at midbody 4 mm ; tail slightly longer 
than wide. 

Remarks: This species is most closely related to capitulatus 
on the mainland of Hispaniola and apparently has differentiated 
from that species following isolation on Gonave Island. It is 
easily recognized by the peculiar shape of the preocular. These 
two species have the smallest and the most pointed heads of any 
of the known Antillean Typhlops. 





Fig. 2. Typhlops gonavensis sp. iiov. Tyjje YPM 3003. From Point a 
Raquette, Gonave Island. Dorsal and lateral views. 



1964 TYPllLOPS OP HAITI 

Typhlops haitiensis sp. IIOV. 
Figure 3 

Holotype: MCZ 62635. From Manneville, Haiti. Collected 
by A. S. Band and J. Lazell, 10 August 1960. 

ParatijiJes (14) : MCZ 69007-12, from Manneville, Haiti. Col- 
lected by E. E. Williams and A. S. Rand, 13-14 August 1959. 
USNM il7273-74, 117276, from Trou Caiman, Haiti. Collected 
by A. Curtiss, 18 February 1943. CM 38804-8, from Manneville, 
Haiti. Collected by George Whiteman, 1963. Manneville is near 
the northwest end of Lake Saumatre, on the north edge of the 
Cul de Sac Plain; Trou Caiman is a small freshwater lake just 
west of Manneville. 





Fig. 3. Typlilnpn haitiensifi sp. nov. Type MCZ 62635. From Manne- 
ville, Haiti. Dorsal and lateral views. 

Diagnosis: A species of Typhlops characterized by a high 
number of dorsal scales, 400-452; scale rows 20-20-20; dorsal 
color extending on to venter, white anal spot; preocular higher 
than long, in contact with the third upper labial ; ocular narrow, 
more than twice as high as wide ; head broadly rounded as seen 
from above. 

Description: Snout, as viewed from above, truncate and 
broadly rounded, not tapering; rostral width about one-fourth 
the width of the head, rostral extending to the level of the eyes ; 
nostrils lateral ; nasals completely separated by a suture extend- 
ing from the second upper labial to the rostral. The preocular 
higher than wide in contact with the third upper labial only 
(Fig. 3). One pair of enlarged .strap-like parietals each as wide 
as two body scales ; a single postocular ; eye large and conspicu- 
ous. Scale rows around body 20-20-20, dorsal scales from rostral 
to tail spine about 435. 



6 BREVIORA No. 202 

Color: The head scales have dark brown centers and relatively 
wide light margins giving the top of the head a striped appear- 
ance ; dorsal brown color extends on to the venter where it is 
interrupted by irregular light areas along the midventral line. 
The dorsal scales are dark brown with light margins. The light 
margins are aligned giving the effect of fine light longitudinal 
lines. 

Size: Total length 240 mm, tail 5 mm, width of head at level 
of eyes 3.5 mm, diameter at midbody 5 mm, tail slightly longer 
than wide. 

Variation: The 14 paratypes agree in details of head scutella- 
tion, and in having 20-20-20 scale rows. The number of dorsal 
scales in this series varies from 400 to 452. The number of 
enlarged parietal scales also varies. All have one pair of greatly 
enlarged strap-like scales ; some have a second pair equally 
enlarged, while others show a graded change from this condi- 
tion to where the second pair is not distinguishable from the 
body scales. All have a single postocular. The color varies from 
dark to light brown in the large specimens. The three smallest 
specimens are very pale above and below, but under magnifica- 
tion it can be seen that the dorsal pigment extends on to the 
venter in the same pattern observed in the larger specimens. The 
extent of the light areas on the venter varies considerably ; the 
three specimens from Trou Caiman have little or no brown 
pigment on the venter, while MCZ 69012 has the venter almost 
entirely pigmented. 

Typhlops pusillus Barbour 

The 27 specimens examined from the mainland of Hispaniola 
agree with those reported by Cochran (1941) except that in this 
series all have two postoculars between the fourth upper labial 
and the parietal. 

The greatest variation is shown by the three specimens from 
Gonave Island. These agree with the mainland forms in pos- 
sessing divided preoculars, two postoculars and in other details 
of head seutellation. They differ from the mainland specimens 
in having a lower number of dorsal scales (258-268 compared 
with 285-319 for the specimens from Bombardopolis on the 
northwest peninsula). They also differ in color as they are 
brown above and pale below and in this respect resemble lum- 
hricalis. 



l!)64 TYPIlLOrS OF HAITI 7 

Typhlops lumbricalis (Linnaeus) 

This species has the most extensive range of any of the Antil- 
lean Typhlops. It has been found on Hispaniola, Cuba, several 
islands in the Bahamas, and in British Guiana, South America. 
Throughout this range, it is remarkably consistent in characters : 
scale rows 20-20-18, rarely 20-18-18 ; usually one pair of enlarged 
parietals and usually two postoculars. The middorsal scales 
range from 240-320 (Legler, 1959). 

The established lack of variation in the number of scale rows 
in various parts of its extensive range makes it very surprising 
to find a population of this species on the southwest peninsula of 
Haiti that is highly variable. 

There are 41 specimens of lumhricalis at hand from the south- 
west peninsula, representing several localities from Miragoane 
to Jeremie, and one specimen from Grand Cayemite Island. 
In number of rows of scales around the body they vary as 
follows: 22-20-20 (2), 20-20-20 (27), 20-20-18 (12). 'The num- 
ber of enlarged parietals also varies: IL-IR (14), 1L-2R (11), 
2L-1II (5), 2L-2R (10). All but one have two postoculars on 
each side. The exception has the upper postocular on each side 
fused to the parietal. The number of middorsal scales ranges 
from 273-324, average 801. Although this is within the known 
range of variation for lumlyricalis, it is higher than for most 
specimens of that species from other parts of Hispaniola. The 
shape and arrangement of the head scales is the same as in 
other populations. The color pattern is also the same. Although 
the number of rows of scales is both high and variable, this 
population can not be distinguished from lumhricalis by any 
character or combination of characters, and it appears to be one 
variable population rather than a mixture of two or more forms. 

In addition to the specimens from the southwest peninsula, 
there are five specimens from Port-au-Prince and Manneville. 
Tliese have the scales in 20-20-18 rows, middorsal scales ranging 
from 261 to 290, and are typical liunhricalis. 

Grant (1956) discusses four specimens of lumhricalis, three 
from Eaux Gaillees and one from Port-au-Prince. The number 
of middorsal scales ranges from 257 to 275 and indicates that 
these are lumhricalis as currently defined. There are also four 
specimens of lumhricalis in the British Museum from Haiti, two 
from Pont Beudet and two from Hinche. Miss A. G. Grandi- 
son supplied the counts for these and the middorsal scales 



8 BREVIORA No. 202 

range from 260-294 — well within the range of variation for 
lumhricalis. The average number of dorsal scales for all 13 
specimens from Haiti outside the southwest peninsula is 279 
with a range of 257-294, 



^&^ 



DISCUSSION 

The five species now known from Haiti are similar in having 
a maximum of 20 scale rows, preocular in contact with the third 
upper labial only, and a relatively narrow rostral. Yet with this 
apparent similarity the five forms represent three distinct 
phyletic lines of Typhlops. 

1. The group including lumhricalis and pusilhis is dis- 
tinguished by having a low number of middorsal scales, 
240-330. These two forms on Hispaniola are readily 
separated from each other by the presence of a single 
preocular in lunihricalis and by a divided preocular in 
pusillus. 

2. A slender, small-headed group with a high number of 
middorsal scales, 385-423 ; this includes two species, gon- 
avensis on Gonave Island, and capitulatus on the main- 
land. These are the most distinctive Typlilops in the 
area as no other species known from, the Antilles has 
such a small head. The two forms are obviously closely 
related and are probably insular populations of what 
was originally one form. The two may be distinguished 
from each other by the shape of the preocular. 

3. The third group is represented by one species, haitiensis. 
Like the small-headed forms it has a very high number 
of middorsal scales (400-454) but unlike the species of 
that group it has a broadly rounded truncate snout, as 
well as a distinctively shaped preocular and ocular. Of 
the five species known from Haiti it is the only one that 
typically has a single postocular. The others have two 
postoculars with one occurring as an uncommon variant. 

These three different groups do not seem to be closely re- 
lated to each other. The distributional data, although limited, 
seem to support the concept that only unrelated forms can 
occur sympatrically, and even then probably l)y occupying 
different ecological niches. For example, Iwnibricalis and pusil- 
lus appear very similar in body proportions and scutellation, 
and differ in the number of preoculars. A divided preocular is 
also known as a rare variant in lumhricalis (Legler, 1959). 



1!)G4 TYPIU.OI'S OF HAITI 9 

Probably pusiUus was derived from lumhricalis at sometime 
when Hispaiiiolfi was a series of islands. 

Todaj^ pusillus occurs on the northwest peninsula and ranges 
across the northern part of the island at least as far as the 
southern shore of Bahia de Samana. Cochran (1941) reported 
one specimen from Sanchez and I have examined four from 
Samana (AMNH 50353-56). 

The range of lumhricalis includes all of the southwest penin- 
sula and extends eastward along the southern coast where it is 
known from Barahona (Noble and Hassler, 1933)' and San 
Pedro de Macoris. On the northern coast it is known as far west- 
ward as Puerto Plata. The range of the two species overlap on 
the northeast coast. Whether they are ecologically separated in 
this area of overlap can not be determined from the available 
data. At Manneville, the one locality where three species have 
been found, each species represents a different group. 

In view of the large number of Typhlops in collections from 
the northwest peninsula and all of them pusillus, it would appear 
that lumhricalis does not occur there. The opposite is true of 
the southwest peninsula where extensive collecting has revealed 
only lumhricalis. 

Typhlops sulcatus Cope 

The finding on Haiti of species with a high number of dorsal 
scales raised the question of w^hat relationship these forms might 
have with sulcatus on nearby Navassa Island, the only other 
Typhlops known from the Antilles wdth both 20 scale rows and 
a high (397) number of dorsal scales. As no other specimens of 
sulcatus have been collected since Cope (1868) described it on 
the basis of one specimen, the type was examined. 

As the type appears today, it is a specimen that was pre- 
served just prior to shedding ; in fact, the old head shields 
have been shed (probably after preservation), and the new head 
shields are represented by soft raised areas outlined by deep 
sulci that mark the position of the original sutures. The exten- 
sions of the nasal sutures from the nostril to rostral are little 
more than faint lines but sufficient to indicate that this species 
does have completely divided nasals. Since Cope (1868) also 



iThe specimen with 385 micUlorsal scales from Alta Vela Island reported as 
Uunbricalis by Noble and Hassler (iy;j3) is not that species as currently defined. 
A preliminary examination discloses this specimen as more closely related to 
sulcatus than it is to the mainland forms. 



10 BREVIORA No. 202 

described this specimen as having completely divided nasals he 
may have seen it with the nasal plates in place Avhen the sntures 
might have been more distinct. Other characters of this specimen 
are : very long parietals extending down the posterior edge of 
the oculars to well below the level of the eyes, and a single 
postocular; both of these characters are well shown by Cochran 
(1941, p. 310, fig. 88). As stated by Cope the body is more 
slender than that of lumhricaUs ; it is also distinguished from 
that species by the high number of middorsal scales. From the 
species on Haiti with middorsal scales over 385, it is distin- 
guished by the shape and size of the ocular and preocular, and 
by the very long parietals. 

This specimen has the snout tapered in front of the eyes very 
much like monensis but not at all like the narrow-headed forms 
on Haiti which have the entire head tapered from well back of 
the eyes. Until such time as more specimens are available it 
seems best to recognize sulcatus as a distinct species. 

The other characters for this species mentioned in the litera- 
ture, trilobate snout and incomplete nasal sutures, are conditions 
that are associated with shedding (Richmond, 1961). The ex- 
tension of the nasal suture from nostril to rostral is very faint 
on most specimens of small Typhlops and difficult to see at best. 
As the nasal plate thickens, prior to shedding,, the nasal suture 
from labial to nostril becomes more conspicuous while the suture 
from nostril to rostral becomes fainter and may actually disap- 
pear as a suture although its presence can usually be detected as 
a fine dark line crossing an otherwise smooth area of the nasal 
plate. Also associated with shedding are the deep sulci repre- 
senting the sutures between the other head scales. 

Since sulcatus is known from only one specimen, and that one 
shedding, it is little wonder that it has been variously described 
as having complete nasal sutures (Cope, 1868), incomplete nasal 
sutures (Cochran, 1924, 1941), and complete on one side and 
incomplete on the other (Legler, 1959). 

TYPHLOPS 
Key to the forms occurring in Haiti 

A. Preocular divided pusillus 

AA. Preocular single B 

B. Ocular narrow and liigli, over twic-e as liigli as wide, posterior 
edge of ocular almost straight, preocular higher than wide (400- 
460 dorsal scales) (Fig. 3) haitiensis 



i 



li)64 TYPIILOl'S OF HAITI 11 

BB. Ocular wide, but little higher than wide, posterior edge strongly 

convex, preocular as wide or wider than high C 

C. Venter unpigmented, same color as underside of tail, less 

than 330 dorsal scales, head rounded lumhricalis 

CC. Dorsal dark color extending on to venter, underside of tail 
and anal area white, 380-430 dorsal scales, head tapering . D 
1). Preocular longer than high, anterior angle acute (Fig. 

2) gonavensis 

DD. Preocular almost triangular, anterior angle rounded (Fig. 
1 ) capitulatus 

EEFERENCES CITED 

Cochran, D. M. 

1924. Typldops lumhricalis and related forms. Jour. Washington 

Acad. Sci., 14(8): 174-177. 
1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus., 117: 
VII + 398 pp. 
Cope, E. D. 

1868. Additional descriptions of neotropical Eeptilia and Batrachia 
not previously known. Proc. Acad. Nat. Sci. Philadelphia, 
1868: 128. 
Grant, C. 

1956. Eeport on a collection of Hispaniolan reptiles. Herpetologica, 
12: 85-90. 
Legler, J. M. 

1959. A new blind snake (genus Typhlops) from Cuba. Herpetologica, 
15: 105-112. 
Noble, G. K. and W. G. Hassler 

1933. Two new species of frogs, five new species and a new race of 
lizards from the Dominican Eepublic. Amer. Mus. Novit., no. 
652: 1-17. (Eeceived Sept. 26, 1963) 

ElCHMOND, N. D. 

1961. The status of Typhlops silus Legler. Copeia, 1961 (2) : 221-222. 
Williams, E. E., B. Shreve, P. S. Humphrey 

1963. The herpetology of the Port-au-Prince region and Gonave 
Island, Haiti. Parts I-II. Bull. Mus. Comp. Zool., 129: 291-342. 

(Eeceived Sept. 26, 1963.) 



12 



BREVIORA 



No. 202 








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BREVIORA 

Miiseimi of Comparative Zoology 



Cambridge, Mass. April 10, 1964 Number '203 

A NEW CAPROMYID RODENT 
FROM THE QTTATERNARY OF IITSPANTOLA 

By Clayton E. Ray^ 

U. S. Nntioiial Museum 



During the early part of 1963, Mr. Robert Allen and I con- 
ducted field work on several Caribbean islands under the joint 
auspices of the Museum of Comparative Zoology at Harvard 
College and the Florida State Museum, supported in part by 
National Science Foundation grants GB 178 and G 16066. Our 
primary objective was the collection of vertebrates, both Recent 
and fossil. The period from 17 March through 4 April was spent 
in the Dominican Republic, during Avhicli time we received the 
cordial cooperation of Prof. Eugenio de Jesus Marcano F. and 
Ing. Emile de Boyrie Moya, Universidad Autonoma de Santo 
Domingo. The cave on the north flank of the Sierra de Neiba, 
from which Antillogale marcanoi Patterson 1962 was obtained 
in 1958, was revisited on 31 March through 2 April with the 
specific objective of securing supplementary material of this form. 
Although we had little success in this, meager evidence of two 
additional taxa was obtained : a single metatarsal of a tiny species 
of Acratocnus (Hooijer and Ray, in press), and a single cheek- 
tooth (MCZ 7675) of a large caviomorph rodent, described below. 
These specimens derive from the same thin reddish brown cave 
earth in the first chamber, from which the 1958 collections were 
obtained (see Patterson, 1962). This deposit contains a moder- 
ately rich accumulation of well-preserved remains (including 
skulls) of the common Hispaniolan caviomorphs, among which 
are interspersed a few fragments and isolated elements of other 
taxa, including Solcnodon, NcsopJiovUs, bats, and birds. 



1 Published by permission of the Secretary of the Smithsonian Institution. 



2 BREVIORA Xo. 203 

Through the courtesy of the various curators in charge, I have 
had at my disposal the rich collections of fossil and Recent cavio- 
morph rodents in the American Museum of Natural History, the 
Museum of Comparative Zoology (MCZ), and the U. S. National 
Museum (USNM). Comparison with relevant genera in these 
collections shows that the single tooth is that of a capromyid 
having its closest affinities with Plagiodontia," but referable to 
none of the described species of that genus (Johnson, 1948). 
The peculiarities of the tooth are sufficiently striking to provide 
adecfuate basis for characterizing the species which it represents. 
It is probaWe that the species will require a new genus for its ac- 
commodation when it is better known, but it may be assigned 
with (|uestion to Plagiodontia pending discovery of additional 
material. Tlie find provides yet another example of the novelties 
that reward exploration in Hispaniola, the least known, paleonto- 
logically, of the Greater Antilles. 

RODENTIA 
CAPROMYIDAE 

Plagiodontia? araeum-'' sp. nov. 

Type : MCZ no. 7675, isolated left upper cbeektooth, almost 
certainly the fourtli (deciduous) premolar; damaged slightly 
along the posterolabial wall ; the onlj- known specimen of the 
species. 

Horizon and locality -. Late Pleistocene or Recent ; first chamber 
of unnamed cave 2 kilometers SE of Rancho de La Cuardia, 
Municipio de Hondo Valle, Provincia de San Rafael, Republica 
Dominicana. The coordinates of Rancho de La Guardia as given 
in Gazetteer no. 33 of the United States Board on Geographic 
Names are 18°43'N, 71°39'W. 

Diagnosis: Differing from the upper cheekteeth of all other 
members of the genus Plogiodonfia in its absolutely greater 
maximum diameter of occlusal surface and in its extreme antero- 
lingual-i)osterolalnal compressioii (to which the specific name 
alludes). 



2 It mif^ht l>o snsiip<'tc(l on tlio basis of sizp and ^'oncral Htrufturc that the 
spf'cinicii is an upper clu'cl^todtli (as yet iinkiiown) of (Jncniisid. However, tlie 
walls of eacii rei'ntrant folil in tli(! upper elieckteetli of Qiirininia would un- 
doubtedly be tif;lill.v appressecl as they are in the lower cheekleelh. Thoui;!! the 
folds in .M("Z 707."i are narrow, tlieir walls are not appressed ( l''i>;. 1). 

"^ (ipOCtOV' narrow. 



1964 



NEW RODKXT FROM IIISrANIOLA 





S MM 



Figure 1. Occlusal views of (A) MCZ 7675, left DP^ of Plagiodoniia 
araeum, the holotype, aiul (B) MCZ 35314, right DP^ (reversed) of P. 
hylaeum. Fine stippling represents cenientuni ; conrse stippling, dentine. 
Drawn bv 8ue Hirschfeld under XSF GB 178. 



Description : Anterior margin of tootli strongly convex as in 
DP^ of Plagiodontia, not flattened as in M^- IVP; incipient anter- 
olabial fold absent, as in all Plagiodontia, excepting P. aedium 
(Johnson, 1948) ; reentrant folds relatively long and narrow, 
parallel- sided ; lingual reentrant with slight posterad flexure at 
internal extremity, not observed in other Plagiodontia; labial 
reentrant with concomitant flexure near its mouth ; posterolabial 
concavity in enamel wall shallow in comparison to other species ; 
longitudinal axis of tooth curved (convex anterolingnally) as 
in upper cheekteeth of other Plagiodontia. 

Measurements : All measurements of DP^ are given in mm in 
the following table. Anterolingual-posterolabial diameter of 
occlusal surface in MCZ 7675 is estimated owing to damage along 
posterolabial wall of crown. 



BREVIORA 



No. 203 



P. araeitm P. ipnaeum P. hylaeum 

MCZ 7675 USNM 254376, MCZ 35314, 
right DP^ right DP4 



Maximum diameter of 
occlusal surface (A) 

Anterolingual-posterolabial 
diameter of occlusal 
surface (B) 

Eatio of B/A 

Maximum diameter of crown 

perpendicular to longitudinal 

axis (C) 
Minimum diameter of crown 

perpendicular to longitudinal 

axis (D) 

Eatio of D/C 

Height of crown perpendicular 
to occlusal plane 



11.0 

5.2 
.47 

10.7 

5.0 

.47 

15.0 



8.2 

6.7 
.82 

7.6 

5.6 

.74 

19.5 



6.8 



5.0 
.74 



6.8 

4.1 

.60 

12.4 



EEFEEENCES 

Hooijor, D. A. and C. E. Eay 
[in press.] A metapodial of Acratocnus (Edentata: Megalonychidae) froni 
a cave in Hispaniola. Proc. Biol. Soe. Washington, 77, 1 fig. 
Johnson, D. H. 

1948. A rediscovered Haitian rodent, Plagiodontia acdium, with a 
synopsis of related species. Proc. Biol. Soc. Washington, 61: 
09-76. 
Patterson, B. 

1962. An extinct solenodontid insectivore from Hispaniola. Breviora, 
Mus. Comp. Zool., no. 165: 1-11, 4 figs. 



BREVIORA 

Mmseiim of Comparative Zoology 

Cambridge, Mass. May 15, 1964 Number 204 

THE STATUS OF PSEUDOGEKKO SHEBAE AND 

OBSERVATIONS ON THE GECKOS OF 

THE SOLOMON ISLANDS 

By Walter C. Brown^ 



INTRODUCTION 

Brown and Tanner (1949) referred a unique specimen (Brig- 
ham Young University [BYU], No. 7002) of a previously un- 
described geckonid lizard from Guadalcanal in the Solomon 
Islands to the new species shebae in the genus Pseudogekko 
Taylor (1922), thus establishing the second species known for 
the genus and extending the range to include a second peri- 
pheral group of islands, analogous in position to the Philippines. 
At that time we had not had the opportunity of examining any 
material of the type species of the genus, the Philippine species 
Pseudogekko compressicorpus. 

New material has now provided the opportunity to reassess the 
relationships of P. shehae and to redefine its differences from the 
other small geckos of the Solomon Islands. Dr. Ernest Williams, 
Museum of Comparative Zoology, recently called my attention 
to the difficulty of identifying certain specimens, which, on the 
basis of descriptions in the literature, were apparently referable 
either to Lepidodactylus guppyi or Pseudogekko shehae. The 
series of specimens in question (Museum of Comparative Zoology, 
Nos. 64152, 65862, 67122, 67124, 74517-19, and Stanford Uni- 
versity, No. 23720) were collected by Mr. Fred Parker on 
Bougainville Island, Solomon Islands, during 1961-62. These 
have provided the point of departure for the present paper. 

This study is part of the author's investigations on the 
herpetofaunas of the Islands of the Pacific area, supported by 



1 Division of Systematic Biology, Stanford University and Menlo College, Menlo 
Park, California. 



2 BREVIORA No. 204 

a grant from the National Science Foundation. Illustrations 
were prepared by Mr. Walter Zawojski, Stanford Research 
Institute. 

THE RELATIONSHIPS OF THE GENUS PSEVDOGEKKO 

Taylor (1922, p. 103), in erecting the genus Pseudogekko, 
suggested that it might have its closest affinities with Thecadac- 
tylus (I assume T. australis = Pseudothecadactylus austraUs: 
Brongersma, 1936, p. 136). There is reason, however, to believe 
that the closeness of this relationship is doubtful. Recently, five 
specimens of Pseudogekko compressicorpus became available : 
Stanford University Nos. 23548-49, from Zamboanga, Mindanao 
Island, and 23654-55 from Bohol Island, and Museum of Com- 
parative Zoology No. 44130, the latter collected by Taylor at 
Saub, Mindanao, and never previously reported. An examination 
of the foot structure of these specimens indicates probable close 
affinities with three other Oriental-Pacific genera (Lepidodac- 
tylus, Gekko and Luperosaurus) , which also belong to the sub- 
family Gekkoninae as defined by Underwood (1954). 

The six Oriental-Pacific genera, Gekko, Hcmiphyllodactylus, 
Lepidodactylus, Luperosaurus, Pseudogekko and Pseudothecadac- 
tylus^, all belong to that group with moderately to strongly 
dilated digits, with the distal joint relatively short, compressed, 
and arising from the tip or near the tip of the dilated part. 

If digital structure alone is considered, these six genera fall 
into four sections. Hemiphyllodactylus is rather sharply dis- 
tinguished by the greatly reduced first digit and the fact that 
the distal compressed phalanx is not attached all the way to the 
tip of the dilated portion, as pointed out by Stejneger (1899) 
and Smith (1933). Pseudothecadactylus forms a second section 
distinguished by the double series of lamellae which are widely 
separated distally. As they are presently understood, Gekko and 
Luperosaurus fall into a third section which may be distinguished 
from Lepidodactylus and Pseudogekko on the basis of the 
lamellae being entire throughout the length of the digit. In 
general the species of the genus Gekko are larger than are the 
species in the other genera, and the species of Luperosaurus 
exhibit more extensive webbing. However, a critical study of 



1 Pseudothecadactylus confined to the islands of Torres Straits and Australia is 
not properly Oriental-Pacific but is here considered because of Taylor's belief in its 
relationship to Pseudogekko. 



1964 



SOLOMON ISLANDS GECKOS 




Figure 1. PseudogelcTco compressicorpus. 



all known species in these last four genera and consideration of 
other characters than digital structure will be necessary before 
the generic limits and relationships can be clearly understood. 

This is borne out by the difficulties which have at times arisen 
in assigning certain species to one or the other of these genera. 
Thus Boulenger (1885a, p. 162) included in Lepidodactylus 
three species later placed in Hemiphyllodactylus by Stejneger 
(1899, pp. 788, 799) and Smith (1933, p. 15). Again, Boulenger 
(1885b, p. 473), in describing Gekko pumilus, noted that the 
species was very like a Lepidodactylus in many characters, being 
placed in Gekko on the basis of the undivided lamellae. Exam- 
ination of one specimen of pumilus (MCZ 69216) suggests that 



BREVIORA 



No. 204 



it may be more closely related to Liiperosaurus with reduced 
webbing and skin folds. Taylor (1915, p. 96), on first describing 
compressicorpus, placed it in the genus Luperosaurus but later 
(1922) separated this species from Luperosaurus and erected 




Figure 2. PseudogeTcTco shebae. 



the genus Pseudogekko for it. Again, an examination of the 
paratype of Luperosaurus rnacgrcgori Stejneger (Stanford Uni- 
versity No. 6263, a hatchling measuring 23.5 mm from snout 
to vent) reveals that the sub-terminal lamellae are divided in 
the midline, and hence that this specimen should be referred 
to Lepidodactylus species as that genus is presently understood. 
(The assignment of macgregori to the genus Luperosaurus is 
thus placed in doubt and the type .should be re-examined in 
this light.) 



1964 



SOLOMON ISLANDS GECKOS 



DISTINGUISHING CHARACTERS OF 

PSEUDOGEKKO SHEBAE AND 

LEPIDODACTYLUS GUPPYI 

Pseudogekko would appear to be distinguished from Lepi- 
dodactylus primarily on the basis of the more slender habitus 
and the more narrowly but uniformly dilated digits. Both P. 
shehae and P. compressicorpus are more slender in body and 
exhibit less broadly dilated digits (Fig. 1) than most of the 
several species of Lepidodactylns which I have had the oppor- 
tunity of examining. If the ratio ' ' breadth of head : snout-vent 
length" is used as a measure of habitus, the range for Pseudo- 
gekko shehae and Pseudogekko compressicorpus, based on the 
few adult specimen,s available, is from about 14-16 per cent ; for 
five species of Lepidodactylns {christiana, lugiibris, aureo- 
lineatus, planicaudus and giippyi) the range is 18-21 per cent. 




Figure 3. Lepidodactylns guppyi. 



6 BREVIORA No. 204 

(Lepidodactylus hrevipes from the Philippines is an exception 
with respect to both of these characters and is closer to Pseudo- 
gekko. A careful study of this species will probably show that 
it should be placed with P. shehae and compressicorpus.) 

The differences in foot structure of Lepidodactylus giippyi and 
Pseudogekko compressicorpus are illustrated in Figures 1 and 3. 
The condition of the terminal lamella — divided or entire — 
is not a generic character, since it is divided in Pseudogekko 
compressicorpus and Lepidodactylus lugubris and entire in 
Pseudogekko shehae and Lepidodactylus guppyi. 

The number of preanal and femoral pores in males and the 
size of the head scales, as illustrated by the number of scales 
between the eyes in the mid-orbital plane, will help to dis- 
tinguish Pseudogekko shehae from Lepidodactylus guppyi 
(Table 1). 

In addition to L. guppyi and P. shehae, members of the lugu- 
hris-woodfordi species complex occur also in the Solomon Islands. 
Individuals of this species or group of species, however, are 
readily distinguished from Lepidodactylus guppyi and Pseudo- 
gekko shehae on the basis of the divided terminal lamella on all 
toes but the first. This scale is undivided on all toes on specimens 
of L. guppyi and P. shehae. 

SUMMARY 

The Oriental-Pacific geckonid lizards of the genera Gekko, 
Lepidodactylus, Luperosaurus and Pseudogekko represent cat- 
egories which probably include closely related groups of species 
but, as they are presently understood, are not sharply and 
clearly delimited from each other. Two additional genera, Hemi- 
phyllodactylus and Pseudothecadactylus, although they probably 
represent lines of evolution distinct for a longer period of time, 
have by some authors been regarded as very closely related to 
these four genera. 

Superficial resemblances between species and the lack of sharp 
lines of demarcation between the genera not infrequently have 
made difficult the proper generic assignment of some of the 
species and even the determination of the species to which isolated 
individuals belong. A case in point is the identification of 
specimens of Pseudogekko shehae and Lepidodactylus guppyi, 
both known from the Solomon Islands. Their distinguishing 
characteristics and present generic assignment have been briefly 
discussed in the present paper. 



1964 SOLOMON ISLANDS GECKOS 7 

LITERATUEE CITED 

BOULENGER, GEORGE A. 

1885a. Catalogue of the lizards in the British Museum. London (Taylor 

and Francis), vol. 1, xii+436 pp., 32 pis. 
1885b. Descriptions of three new species of geckos. Ann. Mag. Nat. 

Hist., (5) 16: 473-475. 
Brongersma, Leo D. 

1934. Contributions to Indo-Australian herpetology. Zool. Meded., 
17: 161-251, 2 pis. 

1936. Herpetological note XIII. Zool. Meded., 19: 136. 
Brown, Walter C. and Vaso M. Tanner 

1949. Rediscovery of the genus FseudogeTcTco with description of a new 
species from the Solomon Islands. Great Basin Naturalist, 9: 
41-45. 
Smith, Malcolm A. 

1933. Remarks on some Old World geckoes. Rec. Ind. Mus., 35: 9-19. 

1935. The fauna of British India. Reptilia and Amphibia, vol. 2, 
XIII+440 pp., 1 pi. 

Stejnegek, Leonard 

1899. The land reptiles of the Hawaiian Islands. Proc. U.S. Nat. 
Mus., 21: 783-813. 
Taylor, Edward H. 

1915. New species of Philippine lizards. Philippine Journ. Sci., 10: 

89-109. 
1922. The lizards of the Philippine Islands. Bur. Sci. Manila, Publ. 
17: 1-269, 23 pis. 
Underwood, Garth 

1954. On the classification and evolution of geckos. Proc. Zool. See. 
London, 124: 469-492. 



8 BREVIORA No, 204 





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BREVIORA 



Miuseitiitni of Compsirsitive Zoology 



Cambridge, Mass. July 15, 1964 Number 205 

REDESCKIPTION OF AMPHISBAENA DUBIA MULLER 
(AMPHISBAENIA: REPTILIA)i 

By Carl Cans 

Department of Biology, State University of New York at Buifalo, 

Buffalo, N. Y. 14214 

This note provides a standardized (Gans and Alexander, 
1962) redescription of Amphishaena duhia Mliller, 1924, and 
adds many new records for the species, extending its range 
from Sao Paulo into Parana and Santa Catarina. I am grate- 
ful to Mr. W. C. A. Bokermann for the gift of two specimens. 
The following curators of institutions (referred to in abbrevia- 
tions throughout) placed me in their debt by loan of material: 
Miss Alice G. C. Grandison of the British Museum (Natural 
History), London (BM) ; Drs. Paulo E. Vanzolini and A. 
Stanley Rand of the Departamento de Zoologia, Sao Paulo, 
S. P., Brazil (DZ) ; Dr. Ernest E. Williams of the Museum of 
Comparative Zoology (MCZ) ; Dr. Konrad Klemmer of the 
Senckenbergischen Naturforschenden Gesellschaft, Frankfurt 
a. M., Germany (SMF) ; Dr. Joseph Eiselt of the Naturhistor- 
ischen Museums zu Wien, Austria (VM) ; and Dr. Heinz 
Wermuth [formerly] of the Zoologischen Museums der Uni- 
versitat, Berlin, Germany (ZMU). Specimens in the Gans col- 
lection are referred to by the letters CG. Dr. Virginia Cummings 
figured the specimens and Miss Charlyn Rhodes contributed 
technical assistance. The over-all project owes its support to 
Grant NSF G-21819 from the National Science Foundation. 



^Notes on amphisbaenids, 12. 



BREVIORA 



No. 205 



AmPHISBAENA DUBIA MtJLLER 

Amphishaena duhia Miiller, 1924, p. 86. Terra typiea: "Piracicaba, Staat 
Sao Paulo, Brasilien." HOLOTYPE: ZMU 26394. 

[Not=A. duhia Rathke, 1863; cf. Gans, 1961, p. 220; China, 1963, p. 197.] 
Diagnosis: A medium sized form of Amphishacna without 
major fusions of head shields; with one or more pairs of large 
parietals; with a blunt-tipped cylindrical tail without autotomy 
constriction or autotomy ; and with two clear round precloacal 
pores in males and none [or two very faint indications only] in 
females. Specimens have 213 to 231 body annuli ; 13 to 17 caudal 
annuli; 13 to 16 (generally 14 or 16) dorsal and 16 to 19 (gen- 
erally 16 or 18) ventral segments to a midbody annulus; and two 
rows of postgenials and no postmalars. The color of preserved 
specimens is a light brown faintly countershaded. Segments 
bear a light circular spot. 




Fig. 1. Amphinbaena duhia. Sketch map showing loc-alities mentioned 
in text. 



1964 



AMPIIISBAENA DUBIA 






Fig. 2. Amphisbaena dubia. Dorsal, lateral and ventral views of the head 
of the holotype, ZMU 26394. The line equals 1 mm to scale. (V. Cummings, 
del.). 



BREVIORA 



No. 205 



Discussion: The liolotype was available for examination and 
its assignment poses no problems. The specimen is slightly faded, 
but otherwise in excellent condition. 

The name A. dubia of Kathke (1863, p. 128), a senior 
homonym of A. dtihia Miiller, refers to AnipJiishaena fuUginosa 
ssp. (cf. Gans, 1961, p. 220) and has been suppressed under 
Opinion 664 (China, 1963, p. 197). 

The examination of the British Museum specimen confirms 
Vanzolini's (1949) statement that Boulenger (1885) had in- 
cluded an individual of A. dnhia in the series upon which he 
based his concept of A. verniicnlaris Wagier. 

It is interesting that the samples show no geographic variation. 




Fig. .3. Amphisbacna thtbia. Ventral view of cloaca and tail of tlie liolo- 
type. The line equals 1 mm to scale. (V. Cummings, del.). 



Description : Figure 2 shows views of the head, Figure 3 the 
ventral surface of the cloaca and tail, and 4, 5, and 6 photo- 
graphic details of the coloration and other aspects of the 
specimens. Figure 7 gives a scatter diagram of tail versus 
snout-vent length. Meristic data are listed in the table. 

This is a medium sized species of Amphishacna, of a light tan 
color in preservative wdth faint countershading. The pigment 
is evenly distributed across the segments and appears to fade 
out ventrally. The dorsal midbody segments shoM' a light cir- 
cular spot in the center of each segment. 

The head segmentation is characterized by lack of major fu- 
sions. An azygous rostral barely visible in dorsal view is followed 
by four or five pairs of enlarged cephalic shields in contact along 
the dorsal midline. The nostrils pierce the first pair (nasals). 
The second pair (prefrontals) are the largest segments of the 
head. There are two and one-half or three supra- and two and 



1964 



AMPIIISBAENA DUBIA 




Fig. 4. Amphishaena dubia. Dorsal, lateral and ventral views of SMF 
11813 from Curitiba, Parana. 



BREVIORA 



No. 205 



one-half (or three) infralabials with the third infralabial ex- 
tending considerably beyond the angulns oris. The supralabials 
are large, the second largest and the first next in size. The 
first two sutures incline anteriorly at an angle of approximately 
45°, the last ascends the snout almost vertically. The angulus 
oris lies anterior to the suture between frontals and parietals. 
The ocular is quadrangular. 

The mental is of approximately the same size as the first 
infralabials. The second infralabials are clearly the largest in 
the row, while the postmental is the largest segment on the 
lower jaw. Posteriorly its tip is inserted between the two large, 
tear-drop shaped first postgenials which in some specimens keep 
the postmental from even point-contact with the malars. The 
second postgenials are irregular. Occasionally a segment from 
this row extends forward to contact the postmental. The row 
back of the malars is counted as the first body annulus since it 
falls posterior to the angular oris ; there are thus no postmalars. 




Fig. 5. Ampliishaena duhia. Dorsal (left) and ventral (right) views of 
the holotype, ZMU 2(5394, at niidbody to show size of segments. The pig- 
ment has faded too uuu-h to be discernible. 



1964 



AMPHISBAENA DUBIA 



Dorsally, the first body annulus curves anteriorly, becoming 
wider to form the segments of the temporal-postocular row and 
abut to the lateral edges of the frontals. In a few specimens 
there is a splitting of this row^ giving a .semblance of a dorsal 
intercalated half-annulus. The second row includes the relatively 
large first parietals as its dorsalmost segments. The second 
parietals are ordinarily not elongate and their anterior and 
posterior sutures generally run in parallel, at right angles to 
the long axis of the trunk. Enlargement, if present, occurs 




Fig. 6 Amphisbaena didna. Dorsal, lateral and ventral views of tail 
of male specimen (CG 2093). Note clearly marked precloacal pores which 
are absent in females (ef. Fig. 3). 



8 



BREVIORA 



No. 205 



mainly by broadening (or fusion) of the middorsal segments 
into a pair of second parietals. 

The head is relatively blunt, flattened slightly dorsoventrally 
and oval in cross-section. The lower jaw is but .slightly shorter 
than the upper. The temporal muscles are only faintly indicated 
by swelling in these presumably adult specimens. The nuchal 
is verv faintlv con.stricted. 



region 



E 
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26 



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• • • 
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Snout - Vent Length - cm 



32 



Fig. 7. Amphisliacna dahia. Scatter diagram of tail versus snout-vent 
length for all specimens. 



There are 213 to 231 body annuli from the angulus oris up to 
and including the [pore-bearing] precloacals. The third through 
fifth or seventh are modified by being narrower and curving 
anteriorly on the ventral surface. There is often one inter- 
calated dorsal half-annulus or asjanmetric annulus both in the 
fifteen postcephalic, and the ten precloacal annuli. Otherwise 
the "pectoral" region shows no complexing of segments. There 
are 13 to 16 (generally 14 or 16) dorsal ;ind 16 to If) (generally 
16 oi- 18) ventral segments to a midbody annulus. 



1964 AMPlllSBAENA DUBIA 9 

The cloacal region is characterized by two small round pre- 
cloaeal pores in males and none or at best a very faint indica- 
tion in females. Five to eight precloacal and nine to 15 post- 
eloacal segments fringe the cloacal slit. There is no antotomy 
annnlns and the species appears incapable of autotomy. Caudal 
annuli number 13 to 17. The tail is wider than high in cross- 
section with a tendency toward ventral flattening. The distal 
half is faintly swollen and finally tapers very rapidly toward 
a faint vertical ridge on its tip. 

The lateral sulci are clearly marked, starting after the first 
fifth of the body length and proceeding to the level of the cloaca. 
At midbody each of them is as wide or wider than one of the 
adjacent segments, and filled with broken segments and diagonal 
folding lines that enter the inter-annular sutures at an angle 
that generally cuts off the corners of the bordering segments. 
The dorsal and ventral sulci are indicated only by alignment of 
the intersegmental sutures. 

The dorsal segments of a midbody annulus are approximately 
one and one-half times as long as wide, while the midventral 
segments range between one and a quarter to one and three- 
quarters times as wide as long. 

Hahits: The Curitiba specimen contained three elongate eggs 
each encased in a very thin leathery shell. The posterior and best 
formed one measured approximately 9 X 17 mm in situ. 

Range: Eastern portions of the states of Sao Paulo, Parana, 
and northern Santa Catarina, Brazil. 

Distribution records: BRAZIL: , (Boulenger, 1885; 

Vanzolini, 1949) ; BM 1961.2023. 8do Paulo: "Inland" , 

MCZ 20655-20657; VM 12335-4. Sao Jose do Rio Pardo, DZ 
6442. Cascata, DZ 6432. Aurora, DZ 6439. Limeira, DZ 6436. 
Piracicaba, (Miiller, 1924) ; ZMU 26394 (holotype). Sao Manoel 
do Paraiso (=Sao Manuel), DZ 1266, 1266B, 6520. Santo 
Antonio do Pinhal, DZ 6440. Belem, DZ 2425. Pirituba, DZ 
6438. Sao Paulo, DZ 7053, 7676. Osasco, DZ 7054. Barueri, 
CG 2092-2093. Mairinque, DZ 6433. Sao Bernardo do Campo, 
DZ 1284. Embu (=Embu Guacu), DZ 6461. Itarare, DZ 6443. 
Parana: "Wenceslao Braz. DZ 6667. Curitiba, SMF 11813. Santa 
Catarina: Tres Barras, DZ 6437. 



10 



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196-1 AMPHISBAENA DUBIA 11 

LITERATUEE CITED 

BOULENGER, GEORGE ALBERT 

1885. Catalogue of the lizards in the British Mu.seiiiii (Natural His- 
tory). Second edition. London, vol. 2, xiii-4-497 pp. 
China, W. E. 

19G3. Opinion 664. Amphishaena dubia Rathke, 1863 (Keptilia) : Sup- 
pression under the plenary powers. Bull. Zool. Nomencl., vol. 20, 
part 3, pp. 197-198. 
Gans, Carl 

1961. Dubia (Amphisbaena) Eathke, 1863 (Reptilia, Squamata). Pro- 
posed suppression under the plenary powers. Z. N. (S.) 1466. 
Bull. Zool. Nomencl., vol. 18, pt. 3, p. 220. 

Gaxs, Carl and A. Allan Alexander 

1962. Studies on aniphisbaenids (Amphisbaenia: Eeptilia). 2. On 
the aniphisbaenids of the Antilles. Bull. Mus. Comp. Zool., vol. 
128, no. 3, pp. 65-158. 

MilLLER, LoRENZ 

1924. Ueber neue oder seltene mittel-und siidamerikanischo Amphibien 
und Eeptilien. Mitt. Zool. Mus. Berlin, vol. 11, no. 1, pp. 75-93. 
Eathke, Heinkich 

1863. Untersuchungen iiber die Arterien der Verdauungswerkzeuge 
der Saurier. Abh. math.-phys. Classe konigl. bayr. Akad. Wiss. 
Miinehen, vol. 9, no. 1, pp. 125-183. 
Vanzolini, Paulo Emilio 

1949. Contribuic^bes ao conhecimento dos lagartos Brasileiros da fa- 
milia Amphisbaenidae Gray, 1825. 3. Sobre Ampliisbaena ver- 
micularis centralis Amaral, 1935. Anais Paulistas Medicina 
Cirurgia, vol. 58, no. 2, pp. 105-108. 



BREVIORA 



MuseiLim of Comparative Zoology 

Cammkidge, Mass. .IrLV 15, 1964 Number L'Od 

THE ALSTOPOD AMPHIBIANS SURVEYED 

By Donald Baird 

Depart nicnf of ( Joolosv. Princeton University, Princeton, N. J. 



INTRODUCTION 

In 1874 the Committee on the Structure and Classification of 
the Labyrinthodonts submitted to the British Association for the 
Advancement of Science a new and comprehensive chissification 
of the known Paleozoic amphibians. L. C. ]Miall, as secretary, is 
usually cited as the author of the several new group names whicii 
were published in the committee's report in 1875. Among these 
new taxa was the Ai'stopoda, proposed to include two genera of 
limbless, snake-like amphibians from the Coal Measures of Ire- 
land which committee-member T. H. Huxley had descril)ed in 
1866 and 1867. These genera, DolicJwsoma and Ophiderpeton, 
were included in a single family until 1908 when Hugo Schwarz 
established a separate family for (Jphulerpeton. Originally classed 
as a suborder of the order Labyrinthodontia, the Ai'stojioda are 
now recognized as an order of the subclass Lepospondyli. 

Knowledge of the grouj) was soon expanded through the work 
of E. D. Cope and Anton Fritsch: both families proved to be 
represented by identical or near-identical genera in the mid- 
Pennsylvanian cannel coals of Ohio and Bohemia. More recently 
J. T. Gregory and the Turnbulls have described a slightly older 
member of the Dolichoscvna group from the Mazon Creek nodules 
of Illinois. The oldest known ai'stopod was found in the Lower 
Mississippian of Scotland by Thomas Stock; the youngest is 
i-epi-esented by material from early Permian fissure deposits in 
Oklahoma, first reported by (iregory, Peabody and Price and 
currently being described by ]\IcGinnis. 

All the existing specimens of ai'stopods are imperfect or frag- 
mentary — only two complete skeletons are known — and most 
of the early descriptions were based on unpre]-)ared specimens. In 



2 BREVIORA No. 206 

these circumstances it is not surprising that the various nominal 
genera have been bandied about by taxonomists and confused 
with unrelated types of snake-like amphibians, or that the group- 
ing of these genera in families or even in a single order has 
sometimes been questioned (e.g. l)y Romcr, 1950A, p. 629). 

Difficulties in identifying from the literature my own collections 
of aistopods and other amphibians from Linton, Ohio, led me to 
examine Cope's type specimens from that locality at the American 
Museum of Natural History. A number of these specimens (but 
none of the aistopods) had been acid-etched by A. S. Romer in 
the course of his faunal revision (1930D) ; the rest had never 
been jirepared. Worse yet, many of them proved to be decompos- 
ing or disintegrating as a result of pyritc disease. To prevent the 
loss of this important collection I was authorized by curators 
G. G. Simpson and E. H. Colbert to undertake a salvage program. 
Accordingly, the decomposed bone and pyrite were dissolved away 
with hydrochloric acid and the remaining blocks of coal-shale, 
after thorough washing and drying, were reinforced with cellulose 
cement. Thus prepared, the specimens are preserved as high- 
fidelity molds of the skeletons. Red latex casts from these molds 
rejiroduce the originals in minute detail (Baird, 1955). 

SPECIT^IENS EXAMINED 

Stock's Mississipiiian ''Ophiderpetor' and specimens of Ophid- 
erpeton hrownriggii, 0. gramdosum and Phlegethontia mazoncn- 
sis at the Museum of Comparative Zoology (MCZ), as well as 
latex casts of the Turnbulls' Phlegethontia mazonensis and a suite 
of galvanotypes of Fritsch's Bohemian specimens, have been 
available for comparison with the American jNIuseum (AMNH) 
types and my own collections which are housed chiefly at Harvard 
and Princeton. A specimen of Phlegethontia linearis collected by 
Thomas Stock in 1888^ was loaned for preparation and study by 
the United States National Museum. 

ACKNOWLEDGMENTS 

I am pleasantly indebted to Drs. Simpson and Colbert and 
Mrs. Rachel H. Nichols for making available the Cope-Newberry 
collection, to Dr. Alfred S. Romer for the freedom of the Museum 
of Comparative Zoology's collections and great library, to Drs. C. 



1 Notes will l)c found on pago 13. following the text. 



1964 AISTOPOD AMPHIBIANS 3 

Lewis Gazin and Peter P. Vaughn for the loan of National Museum 
material, and to Dr. ^^'illiam D. Turnbull for supplying latex casts 
of his Braidwood Phlegethontia. My good friend Robert W. 
jMorris deserves especial thanks for guiding me to the Linton 
locality in 1950 and introducing me to the jileasures of Pennsyl- 
vanian herpetology. Two of my eight collecting trips were 
financed by the National Science Foundation and the W. B. Scott 
Fund of Princeton University, whose support is gratefully ac- 
knowledged. Dr. Joseph T. Gregory and JNIiss Helen J. IMcGinnis 
have helpfully criticized the manuscript; Miss McGinnis, further- 
more, has generously made available her unpublished monograph 
on Phlegethontia. 

SURVEY OF THE AlSTOPODA 

Our discussion of the ordinal characters of the Ai'stopoda had 
best be prefaced by a table of the known members with their 
source localities and stratigraphic ages. It should be noted that 
nearly every describer has acted on the time-honored principle 
of "new locality, new species" even though he was seldom able 
to compare his new species with a prepared specimen or an 
adequate description of any previously described species. On 
the other hand, the faunal similarities between the Westphalian 
coal-swamp faunas of Illinois, Ohio, Pennsylvania, Nova Scotia, 
the British Isles and Bohemia — similarities which current studies 
(Baird, 1963 I show to be even closer than Westoll (1944A, p. 106) 
indicated — should lead us to expect identical genera with closely 
related if not identical species. 

To determine specific differences in the fragmentary and 
variously preserved material at hand will be difficult at best. For 
present purposes, then, I have reluctantly followed prior authors 
in allowing each local fauna its indigenous nominal species, but 
I have recognized only one species per genus in each fauna. The 
latter restriction follows W. D. Matthew's principle that "in a 
collection of unified origin . . . congeneric animals will generally 
be of a single species, or, if they are of more than one species, 
the discontinuity between the groups will be large and evident. 
In such a collection, then, in the absence of fairly obvious dis- 
continuity, the variation within a genus should usually be taken 
as intraspecific." (Simpson in Matthew and Paula Couto, 1959, 
p. 51.) Thus the list which follows is essentially a record of 
generic distribution with the species of each genus arranged in 
approximate order of geologic age.- 



4 BREVIORA No. 206 

Class AMPHIBIA 

Subclass LEPOSPONDYLI 

Order AlSTOPODA IMiall, 1875A 

Family uncertain. 

"Ophiderpeton'' of Stock, 1882B. AVardie, near Edinburgh, Scot- 
land. Lower Oil-Shale Group, Calciferous Sandstone Series, 
Lower Carboniferous (early Mississippian). 
Family Ophiderpetontidae Schwarz, 1908B. [Syn.: Steenisauridae 
Huene, 1948G.] 
Ophiderpetoji Huxley in Etheridge, 1866A.'^ [Syn.: Oestocepha- 
lus Cope, 1868J; Thyrsidiiim Cope, 1875D;^ Steenisaurus 
Kuhn, 1938A.-'] 
0. broivnriggii Huxley in Wright and Huxley, 1866A (type 
species). Jarrow Colliery, Castlecomer, Co. Kilkenny, Ire- 
land. Leinster Coals, late Westphalian B (early Mid-Penn- 
sylvanian). 
0. naninn Hancock and Atthey, 1868A. Newsham, Northum- 
berland, England. Low ]\Iain Coal, late Westphalian B 
(early Mid-Pennsylvanian) . 
0. aranulomm Fritsch, ISSO (1883AV Nyrany (Niirschan), 
Plzen Basin, Bohemia, Czechoslovakia. Gaskohl, late West- 
phalian D (Mid-Pennsylvanian). 
0. cf. amphiuminuiti (this paper). I. F. Mansfield mine, Can- 
nelton, Beaver Co., Pennsylvania. Roof shale of Upjier 
Kittanning Coal, Allegheny Group, late AVestphalian D 
(Mid-Pennsylvanian) . 
0. amphiuminum (Cope), 1868J. Linton, Jefferson Co., Ohio. 
Canneloid shale below Upi^er Freei)ort Coal, Allegheny 
Group, late Westphalian D (late Mid-Pennsylvanian). 
0. vinnupi Frifpch, 1880 n88RA). Kounova, Kladno-Slany- 
Rakovnik Basin, Bohemia, Czechoslovakia. Late Stephan- 
ian (late Pennsylvanian).'"' 
Family Phlcgcthontiidac Cope, 187oD. |Syn.: Dolichosomatidae 
Lydekker, 1890A, a superfluous sul)stitute name proposed 
to include Cope's Phlegethontiidae and ^Nlolgophidac] 
Dolichosonia Huxley, 1867B. 
D. emersoni Huxley, 1867B (type species). Jarrow Colliery, 
Castlecomer, Co. Kilkenny, Ireland. Leinster Coals, late 
Westi)halian B (early ]Mid-Pcnnsylvanian). The specimen 
on which this genus was founded is obscured by a film of 



1964 AiSTOPOD AMPHIBIANS 5 

matrix: until it is prepared we hiiw no basis for distin- 
guishing Dolichosoma from Phlegethontia, with which it 
may w(^ll he congeneric. 
Fhlcgcthonfia Cope, 1871L. 

F. mazoncnsis Gregory, 1948(\ Mazon Creek, Grundy Co., 
and Brai(hvood, Will Co., Illinois. Francis Creek Shale 
above Morris (No. 2) Coal, Carbondale Formation, West- 
phalian C-D boundary (Mid-Pennsylvanian)." 
P. \r)olic}wso))ia] longissima (Fritsch, 1875A), new comb. 
Nyiaiiy, Plzen Basin, Bohemia, Czechoslovakia. Gaskohl, 

late Westphalian D (Mid-Pennsylvanian). The close simi- 
larity of this species to its near-contemporaries in America 
bespeaks a common genus. To group the four species to- 
gether as Phlegethontia seems more realistic than to seg- 
regate the Bohemian form as a species of the little-known 
genus DolichosoDia. If future work on Dolichosotna should 
confirm Fritsch's generic assignment, all the species of 
Phlegethontia would have to be transferred to that genus.** 

P. linearis Cope, 1871L (type species). Linton, Jefferson Co., 
Ohio. Canneloid shale below Upper Freeport Coal, Alle- 
gheny Group, late Westi:)halian D (late Mid-Pennsyl- 
vanian). The type and only specimen of P. serpens Cope, 
1871L, differs from P. linearis chiefly in its much larger 
size; its specific distinctness is very doubtful. 

P. new species, McGinnis MS, 1964. Quarry 6 miles north of 
Fort Sill. Comanche Co., Oklahoma. Fissure filling correla- 
tive with Arroyo Formation, Clear Fork Group, Autunian 
(early Permian). (Gregory, Peabody and Price, 1956, p. 3: 
"an aistopod.")^ 

Ordinal Characters 

Reserving for future discussion the undescribed Mississippian 
genus, we may examine the ordinal characteristics of the Ai'sto- 
poda as they appear in Ophiderpeton and Phlegethontia. In defer- 
ence to Miss McGinnis' forthcoming paper on the skeletal 
morphology of Phlegethontia. detailed osteological descriptions 
will be omitted here. The skull structure of Phlegethontia is ade- 
quately known from the descriptions by Fritsch, Gregory and the 
Turnbulls; that of Ophiderpeton has been interpreted by Steen on 
the basis of rather unsatisfactory specimens. Detailed comparisons 
between the skulls of these genera must therefore wait until the 



6 BREVIORA No. 206 

excellent type skull of Ophiderpeton amphiuminum (AMNH 
6857) can be fully prepared and analyzed. 

Different as the two genera are in skull and jaw configuration, 
in the more fundamental characters of the postcranial skeleton 
and skin they show many similarities — similarities beyond their 
common leposjiondyl heritage of holospondylous, ami^hicoelous 
vertebrae (Fig. 1). Among these characters in common are the 
following: 

1. Body long and snake-like. Vertebral counts are possible in a 
few specimens of aistopods : 

OphiderjjcloN brownriggii (type), po.sterior part missing c. 60 

O. granalosum (type of O. persuadens), complete juvenile lOO-f- 

Phlegethontia linearis (type), at least two loops missing c. 206 

P. longissima (type) ; Fritsch estimated 200+ in life 150 

P. mazoneitsia (Braidwood specimen), complete juvenile c. 140 

Number of vertebrae would appear to be a function of age in 
PhlegetJiontia. At least 230 must have been present in a presuma- 
bly adult individual such as the type specimen of P. linearis. 
Cervical, dorsal and caudal vertebrae are differentiated. 

2. Vertebrae with low, blade-like neural spines, transverse pro- 
cesses formed by parapophyses, hypapophyseal flanges on the 
caudals. and foramina for the spinal nerves. The neural spine ex- 
tends the full length of the vertebra, bifurcating where it merges 
with the posterior margin of the neural arch. (The artificial lack 
of neural spines on Miss Steen's plasticene squeezes of Ophiderpe- 
ton amphiuminum caused her to doubt their existence, but spines 
are well developed in this species and in 0. granulosuni.) 

The transverse process, which arises from the centrum and 
articulates with the capitulum of the rib, is accordingly interpreted 
as a parapophysis. Transverse processes of the cervical vertebrae 
slant posteroventrally; those of the long dorsal series gradually 
change direction from posterolateral to lateral to anterolateral, 
while those of the caudals slant anterolaterally. 

In the caudal vertebrae a pair of hypapophyseal processes form 
flanges which run the length of the centrum and fence the haemal 
canal. Basapophyseal processes may be present on the dorsal 
centra but are feebly developed at best. 

In Phlegethontia a foramen for the spinal nerve occurs below 
the anterior end of the posterior zygapophysis or slightly further 
forward, behind the transverse process. This foramen appears to 
be present also in Ophiderpeton although it is generally obscured 
by dorsoventral compression or concealed by the dermal armor 
in the specimens at hand. Small foramina for the spinal nerves 



1964 Ai'STOPOD AMPHIBIANS 7 

are clearly visil)le in a series of caiulals (MCZ 2165) from Linton 
which are attributed to Ophiderpeton amphiuminum on the basis 
of the associated osteoderms. Intra vertebral nerve foramina are 
not present in the Microsauria and Nectridea from Linton. 

3. Relatively straight, iinicipital, tetraradiate ribs. Their proxi- 
mal ends are K-shajied, the "tuberculum" (better termed the 
costal process) and the shaft forming the ui)riglit of the K, the 
capitulum forming the upper arm, and a pointed posteromedial 
process (the "ventraler Fortsatz" of Fritsch) forming the lower 
arm. The costal process, which in Ophiderpeton has a cupped head 
like the capitulum, api)ears to be a displaced tuberculum which 
has lost its vertebral articulation. In Ophiderpeton the shaft and 
posteromedial process are stout and stiletto-like; in Phlegethontia 
the shaft is slender and flexible and the posteromedial process is 
absent or weakly developed, perhaps in part because of retarded 
ossification. 

In articulated skeletons (which have usually been dorso- 
ventrally compressed) the ril)s tyiiically lie with their shafts par- 
allel to the column; the capitulum articulates with the transverse 
process while the costal process lies above or beside the overlap- 
ping shafts of the preceding ribs. In this manner three ribs overlap 
in Ophiderpeton, four in Phlegethontia. Gregory (1948C, p. 652) 
has interpreted the costal process ("tuberculum") as articulating 
with the transverse process in P. mazonensis, but this is definitely 
not the case in the specimens of Phlegethontia from Braidwood, 
Linton and Xyrany, nor in Ophiderpeton — if it were, the ribs 
would have to project laterally in a manner reminiscent of Draco 
volans! In life the ribs were probably aligned nearly parallel to the 
body axis with their distal ends not greatly dejiressed from the 
horizontal. In Phlegethontia the anterior dorsal ribs curved a little 
around the thoracic cavity, but in Ophiderpeton all the ribs were 
straight. Short dorsal and ventral muscles from the vertebrae 
presumably inserted on the costal process and posteromedial pro- 
cess, respectively, functioning to move the rib on the fulcrum of 
its capitulum. 

4. Hijoid shaped like a shalloivlij curved sickle. This element, 
clearly seen in the three local poiuilations of Phlegethontia,^'^ was 
first identified in Ophiderpeton by Fritsch (1901 A. ]). 88) but has 
not been mentioned in subsequent discussions of that genus. Prep- 
aration reveals its presence in the type specimen of 0. amphiumi- 
num (AAINH 6857 (. Gregory (1948C, p. 648) very i)lausibly in- 
terprets the sickle-shaped hyoid as the first ceratobranchial. 



8 BREVIORA No. 206 

5. Limbs and girdles absent. In none of the aistopod material 
is there anything which can be interpreted as the rudiment of a 
pectoral or pelvic girdle. The elements which Steen (1913A, p. 
876) identified (but did not illustrate) as the "clavicle and 
cleithrum" in a British Museum specimen of Ophiderpeton ani- 
phiuniinum are very jjrobably the liyoids. 

6. Ventral armor of needle-shaped gastralia arranged en chev- 
ron. Ophiderpeton has a stout thoracico-abdominal sheathing of 
fusiform osteoderms which are packed together in a herringbone 
pattern, the apex of the chevrons pointing forward. In the cervical 
region the spicules l)ecome smaller and grade into minute wheat- 
shaped ossicles which cover the gular area. This arrangement ap- 
pears to be the primitive condition in the A'istopoda and compares 
closely with that found in Paleozoic labyrinthodonts and reptiles. 
In contrast, the ventral armor of the specialized genus Phlege- 
thontia is reduced to a short series of widely spaced, thread-like 
gastralia in the anterior thoracic region. 

As dorsal armor does not occur in the Fhlegethontiidae it is not 
listed here as an ordinal characteristic, although it may well prove 
to be typical of the less highly specialized A'istopoda. In Ophid- 
erpeton the cheeks and the dorsal surfaces of trunk and tail are 
slieathed with oat- or pebble-shaped osteoderms (Fig. 1 C), and a 
similar dermal covering can be seen in the oldest known repre- 
sentative of the order, Stock's Mississippian aistopod from Scot- 
land (discussed below). 

DISCUSSION 

The foregoing tabulation of characteristics will, I trust, serve 
to establish the unity of the order Ai'stopoda and provide some 
basis for a discussion of its history and affinities. In contrast to the 
other lepospondyl grou])s of the Pennsylvanian coal-swamps, most 
of which make their first appearance — full-fledged but of un- 
known ancestry — in Westphalian time, the aistopods have an 
ancient pedigree. The oldest known member of this group is indeed 
the oldest known lepospondyl, and until the discovery of the late 
Devonian ichthyostcgids in Greenland it was the oldest known 
amphibian of any sort. 

The uniciuc skeleton of this eo-a'istopod (]\ICZ 2185) was col- 
lected by Thomas Stock in the Wardie shales of the Calciferous 
Sandstone Series of Scotland ; its age in American terms is early 
Mississi))i)ian. Although mentioned in a paper by its collector 
(Stock, 1882B I the sjiecimen remains uni^repared and midcscribed. 



1964 AISTOPOD AMPHIBIANS 9 

A long, yausagt'->'>ha|)C'd cont'ix'tiun of den^e irontstonc encloses the 
pyritized skeleton and promises formidable difficulties to the pre- 
parator. Fractures in the frontal plane at various levels, however, 
reveal aistopod characteristics: an elongate, apparently limbless 
body armored with pebble-like dorsal osteodcrms and needle-like 
gastralia; vertebrae with hourglass-shaped centra and long, low 
neural s})ines; and sharp, straight-shafted ribs. The bulbous, well- 
ossified calvarium recalls that of Phlegethontia. Although the 
proper classification of this genus must await preparation and de- 
tailed anatomical study, we may say that canny Thom Stock hit 
close to the mark in labeling it "Ophiderpeto7i." 

From the remarkable specialization already achieved by early 
Mississippian time we must infer that the history of the Aistopoda 
carries well back into the Devonian. Here the total absence of 
limbs and girdles in all known aistopods provokes speculation. 
Did the group originate from a tetrapod ancestor, or directly from 
some crossopterygian fish through the suppression of the fins? We 
have no proof either way. But the many features which aistopods 
share with the limbed leposj-jondyls, most obviously their basic 
skull structure and the division of the body into neck, trunk and 
tail, make it extremely unlikely that the forefather of the group 
was anything but a tetrapod. 

Affinities of the Aistopoda, as mentioned earlier, have been much 
debated. In surveying the literature we will do well to restrict our 
attention to those commentators who have had first-hand knowl- 
edge of the specimens. 

J. T. Gregory in 1948 assigned Phlegethontia to the "Dolicho- 
somidae" and classed this family with the Ophiderpetontidae as 
the suborder Aistopoda of the order Microsauria. Influenced by 
the reptilian features of Phlegethontia and by the belief that 
Cephalerpeton is a microsaur (which it is not) as well as a caji- 
torhinoid reptile (which it is ) , he placed the orders Microsauria 
and Captorhinomorpha together in the infraclass Captorhina of 
the subclass Eureptilia. In 1950, however, he conceded that the 
order Aistopoda should probal)ly be assigned to the amphibian 
subclass Lepospondyli. Gregory's 1948 paper convincingly demon- 
strated that the similarities of the Aistopoda to the Lysorophidae, 
Apoda and Amphisbacnidae are the result of convergence in ani- 
mals of similar ecotype, and that these groups are only remotely 

related. 

The Turnbulls in 1955 pointed out that Lydekker's family name 
Dolichosomatidae is objectively synonymous with Cope's earlier 
name Phlegethontiidae ; they assigned Dolichosoma to the latter 



10 BREVIORA No. 206 

family. They also noted without comment that both Lydekker and 
Schwarz had included the Alolgophidae in the Aistopoda." Stress- 
ing that the postcranial skeleton has more fundamental signifi- 
cance than the skull, they applied the same classification— class 
Amphibia, subclass Lepospondyli, order Aistopoda — used in this 
paper. 

Gregory, Peabody and Price (1956) carefully compared the 
vertebral structure of the various lepospondyl groups and con- 
cluded that the "Aistopoda are a highly specialized limbless group 
of great antiquity [which] may ]irovisionally be regarded as an 
early offshoot of the nectridian stock." 

Basing his observations in part on my preparations and the un- 
described Alississippian aistopod, Williams (1959) emphasized the 
antiquity of the tyj^ical lepospondylous centrum which he homol- 
ogized (quite rightly, in my opinion) with the pleurocentrum of 
other ancient tetrapods and the centrum of the living amphibians. 
Rejecting the theory that leposj^ondyls had an independent de- 
rivation from the crossopterygian fishes, he observed that "if then 
the lepospondyls are to be derived from the ichthyostegalians, 
rhachitomes, or anthracosaurs, the pleurocentral centrum speaks 
for anthracosaur affinities." 

Without questioning the morphological theory on which Wil- 
liams' hypothesis of anthracosaur affinities is based, I would like 
to point out some uncomfortable facts. The oldest true reptile 
which is adequately known, Cephalerpeton, is of mid-Pennsyl- 
vanian age and has composite vertebrae in which the relative sizes 
of pleurocentrum and hypocentrum ( = intercentrum) are the same 
as in Permo-Carboniferous mammal-like rejitiles and the living 
Sphenodon. In the same swamj) witli Cephalerpeton lived Di- 
plovertebron [Eusauropleura] , a form in which the hypocentrum 
is much larger in proportion to the ])leurocentrum; this genus may 
reasonably be taken as a morphological ancestor of the Seymouri- 
amorj^ha from which the true reptiles are believed to be derived. 
Diplovertebron, however, is so similar to the embolomerous 
labyrinthodonts in its skull and appendicular skeleton that it 
must be classed as an amphibian — an anthracosaur. Amphibians 
with similar vertebral proportions coexisted with true embolom- 
eres at least as far back as late ^Nlississippian time, so the com- 
mon ancestry of the diplovertebrontids and embolomeres can 
hardly be dated later than the middle Mississippian. That the 
ancestors of the reptiles had a relatively large hypocentrum, and 
that reduction of the hypocentrum characterized the transition 
from amphibians to rei^tiles, can hardly be doubted. Derivation of 



1964 a'istopod amphibians 11 

this pi'olo-ii'ptiliaii verti-hral structure from tlu' ichthyostegid 
pattern of late Devonian time is (juite undocumented but presents 
no great theoretical difficulties; the ichthyostegalian vertebra in 
turn is readily dei-ivahle from that of crossopterygian fishes such 
as Eusthcnopteron. 

In striking contrast to this plausible sequence of morphological 
stages stands the oldest known leposi)ondyl, Stock's early Missis- 
sippian ai'stopod. Its vertebrae are holospondylous without any 
trace of an ossified hypocentrum or of laterally paired pleuro- 
centra like those of the ichthyostegalians and crossopterygians. 
In vertel)ral form this eo-lepos))ondyl apjicars to be much further 
removed from the anthracosaur-reptile lineage than that lineage 
is from the rhachitome-stereospondyl line. Indeed, the evidence 
now available suggests rather that the dominance of the ])leuro- 
centrum evolved in convergent fashion in the lepospondyls and 
anthracosaurs. Thus in my opinion the evidence for leposjwndyl- 
anthracosaur affinities is too tenuous, and the i)oint of supposed 
common origin too remote in time, to justify speculation. 

Relationships of the Aistopoda to other lepospondylous amphib- 
ians are almost equally difficult to determine. The urodeles and 
anurans (Lissamphibia) need not be considered here, for their 
]X'rsistent four-leggedness and their relatively recent pedigree 
testify that any connection between Lissamphibia and Aistopoda 
must be indirect, by way of one of the Paleozoic lepospondyl 
groups. The apodans (Gymnophiona), though similarly devoid 
of limbs and girdles, show features of the scales and vertebrae 
which ally them rather with the microsaurs. 

Among Paleozoic Icjiospondyls the Nectridea (let us use the 
correct spelling if only for the sake of novelty) display similarities 
in vertebral structure which have led Gregory, Peabody and Price 
(1956) to consider them as distantly related to the aistopods. 
Tills liypothesis is the most plausible one yet advanced. But it 
should be emphasized that all known nectrideans had inter- 
vertebral spinal nerves and well-developed (though sometimes 
diminutive) appendicular skeletons, including the sculptured 
clavicles and interclavicle which characterize Paleozoic amphib- 
ians in general. Aistopods had intravertebral spinal nerves and 
lacked limbs and girdles, apparently since early iSIississippian 
time. Thus if these two groups had a common ancestor it must be 
sought in Devonian rocks — rocks in which amphibians of any kind 
are extremely rare. Until pertinent fossils can be found and de- 
scribed, therefore, the affinities of the Aistopoda must remain 
in doubt. 



12 BREVIORA No. 206 

The foregoing discussion will have served its purpose if it stimu- 
lates further research by clarifying the nature of this peculiar 
group of amphibians and pointing up some of the many unan- 
swered questions concerning them. Perhaps it is too much to hope 
that anything can remedy the common but unfortunate tendency 
to omit the diaeresis and pronounce the name as "ACE-topod" or 
''ICE-topod." Like the proverbial politician, the ai'stopods should 
be happy to be called by any name so long as they are recognized 
in print. 

OPHIDEEPETOX FROU CANNELTON, 
PENNSYLVANIA 

Seven associated vertebrae (PU 17293) whicli recently came to 
light in the Princeton paleobotanical collections provide a new 
horizon and locality record for the genus Ophiderpeton. This speci- 
men, collected by I. F. ]\[ansfield in 1877, comes from the richly 
fossiliferous roofing shale of the L'pper Kittanning Coal, a member 
of the Allegheny Group and late Westphalian D (^Middle Pennsyl- 
vanian ) in age. The sotu'ce locality is ]\Iansfiekrs mine near Can- 
nelton in Darlington Township, Beaver County, Pennsylvania. 
This is the first lepospondylous amphibian to be recorded from 
Cannelton. 

]\Iuch of the bone has flaked away and little remains but the 
neural arches and spines. Absence of the diagnostic ribs and gas- 
tralia makes generic identification difficult, as the neural arches 
of Ophiderpeton and Phlegethontia appear very similar when 
crushed. Dimensions of the vertebrae fall within the range of the 
pene-contemporaneous 0. amphiumvnum, but they are also com- 
patible with the exceptionally large Phlegethontia specimen 
named P. serpens by Cope. The most diagnostic feature available 
seems to be the medial notch in the posterior border of the neural 
arch: this is deep in the Cannelton aistopod and Ophiderpeton 
amphiuminum but shallow in PJdegethontia. As Ophiderpeton is 
common in the Linton fauna while only one Phlegethontia speci- 
men of e(iuivalcnt size is known, statistical probal)ilitics also favor 
assignment of the Cannelton ai'stoj^od to Ophiderpeton. Although 
the difference in age between the Linton and Cannelton deposits 
makes a specific difference possible, in default of morphological 
evidence the specimen may best be designated Ophiderpeton cf. 
nmphivmivinn fCope). 



1964 AisToi'on ami'hibians 13 

SUMMARY 

The order A'lstopoda is a natural gi'oiip of snake-like lepo- 
spondylous amphibians which inhabited the Paleozoic coal- 
swamps of Europe and North America. Two families, each es- 
sentially monotyi)ic, are distinguished: the Ophiderpctontidac of 
the early middle to late Pennsylvanian and the more highly spe- 
cialized Phlegethontiidac of the early Pennsylvanian to early 
Permian. The oldest known leposi)ondyl, an undescribed aistopod 
from the early Mississii)pian of Scotland, is of uncertain family 
position. 

Pending detailed comparison of the skulls, ordinal characteris- 
tics may be found in the postcranial skeleton : ( 1 ) body long and 
snake-like, reaching a vertebral count of about 230; (2) holo- 
spondylous, amphicoelous vertebrae with low, blade-like neural 
spines, transverse jirocesses formed by parapojihyses, hypapophy- 
seal flanges on the caudals, and foramina for the spinal nerves; 
(3) relatively straight, unicipital, tetraradiate ribs; (4) sickle- 
shaped hyoids; (5) limbs and girdles entirely lacking; (6) ventral 
armor of needle-shaped gastralia arranged en chevron, and (ex- 
cept in phlegethontiids) dorsal armor of pebble-shaped osteo- 
derms. 

The remarkable specialization already achieved by the early 
Mississippian implies an origin well back in Devonian time; a 
tetrapod ancestry rather than direct derivation from the crossop- 
terygian fishes is indicated. Relationships of the order are obscure. 
A proposed connection with the anthracosaur-reptile lineage is 
controverted by the available evidence, and among lepospondyl 
groups only the Nectridea show even distant affinities with the 
Aistopoda. 

NOTES 

'The man who actually collefted the superb skeletons of Linton amphib- 
ians which the U. S. National Museum acciuired from R. D. Lacoe, and 
whicli Moodie (1909B) attempted to describe, has not previously been 
identified. Circumstantial evidence points to Thomas Stock of Edinburgh 
whose great collection of Scottish Carboniferous fishes and amphibians was 
purchased by the Museum of Comparative Zoology in 1883. In the MCZ 
archives is a letter from Stock to Alexander Agassiz, sent from Wellsville, 
Ohio (the town just north of Linton) on April 30, 1888, which states in part: 
"I am staying about here . . . collecting the Linton fishes and labyrinthodonts. 
... I have been very .successful with the rare group of Carboniferous laby- 
rinthodonts and have already several nice ones that I should think would 



14 BREVIORA No. 206 

be a valuable addition to your collection." Unfortunately for Harvard, 
Agassiz declined to buy Stock's collection. Evidently it was sold to Lacoe, 
for se\-eral of the Lacoe specimens from Linton bear stickers inscribed 
"Stock #— ." 

" Current stratigraphic practice, followed in this paper, places the Mazon 
Creek fauna close to the Westphalian C-D boundary or in earliest West- 
phalian D and assigns the Cannelton fauna to late Westphalian D; the 
Linton fauna is very late Westplialian D. However, recent paleobotanical 
work by Cridland, Morris and Baxter (1963, p. 63) implies that the Francis 
Creek Shale (Mazon Creek fauna) is only slightly older than the Upper 
Freejiort Coal (Linton fauna) but considerably younger than the Kittanning 
Coal Group (Cannelton fauna). 

' Names of the Jarrow amphibians are usually cited as of Huxley, 1867. 
However, Ophiderpeton and four otiier genera were first validly published 
in January of 1866 by Etheridge, who credited their authorship to Huxley. 
Generic descrijjtions were pro\'ided although no species were named. As a 
genus coelebs (i.e. without named species) is valid under the International 
Rules of Zoological Nomenclature, the genus Ophiderpeton properly dates 
from January, 1866; its type sjiecies was first described in April of that year 
by Huxley in a paper by Wright and Huxley. Duiichusoma emersotti was 
not described until 1867. 

^Preparation of the type specimen (AMNH 6900) of the type species 
Thyrsidium jasciculare Cope confirms Romer's (1930D) identification of 
it as Ophiderpeton amphiumirinm. I mention this fact in order to resolve 
Mile. Dechaseaux's doubts (1955, p. 278) about tlie identity of Thyrsidium. 

' Steenimurus was proposed by Oskar Kuhn (1938A, p. 51) in a footnote 
to Ophiderpeton gmnulosuni Fritsch : "OpJiiderpeton Fritsch non Huxley 
(teste Steen) = Steeidsaiirus nov. nom." Steen's publications do not suggest 
any such generic distinction, nor do the specimens document it. 

"The specimen figured by Scliwarz (1908B, figs. 1-4) under the name 
Ophiderpeton vicinum is apparently from Nyrany. Because of the difference 
in age I prefer to assign the late Westphalian material from Nyrany to O. 
grduiilosiim and restrict the name O. vicinum to late Stejthanian material 
from Kouno\'a. 

' A si)ecimen (L^SXM 4313) which was assigned to PIdcgethontin mazonen- 
sis by Gregory (1950A, p. 867) proves on preparation to be Cocytinus, the 
first record of a lysorophid microsaur in the Mazon Creek fauna. 

'* D<dich<i.-:<nn<i sriilijrnnn Fritsch, 1901A, is liere s^'nonymized with Phle- 
gelhontia lungixsim". The type of "Dolirlio.somn" n)ig)islatu)n Fritsch is a 
ne(!tridean skull roof from which the postfrontal and postorbital bones have 
been displaced during fossilization: its i)roper name is Sauropleura [Uro- 
cordylus] scalaris (Fritsch). 

■'Two supposed aTstoi)Otls from Europe require some comment here. 
Palaeosiren beinerti Geinitz, 1864A, based on vertebrae from the LTpper 
Rotlicgende of Olivetin (Oelberg bei Braunau), Bohemia, has been inter- 
preted .us a gigantic aistopod l>y Fritsch (1883A, p. 125). So far as one can 



1964 Al.sToi'UU AMrillBlANS 15 

judge from Geinitz and Deichmiilh^r's lithograph (1SS2A, \A. 9) I he centra 
are hourghiss-shaped only externally: they lack the funnel-like amphicocly 
which characterizes the Aistoi)oda. Until preparation and rt^study of lliis 
.specimen reveal its affinities it remains inccrlae sedk. 

The iiuprinf of a small skeleton from Ihc late Stephanian or early Auluiiian 
of La Machine, Xicvre, France, has been tentatively identified as an aistopod 
by Thevenin (1910A, p. 39). Its preservation is so ])oor that little more than 
the gross niorjjhology can be made out. The skeleton looks more lysoroi)hid 
than aistopod to me, Ijut this suggestion must be tempered by the fact that 
lysorophids are known at present only from North America. Until further 
material is a\ailable tlie affinities of Thevenin's animal must be left snh 
j II dice. 

"The element in Phlegethontia longissimn ("Dolichosoma scKtifennn") 
which Fritsch (1901A, p. 88) called a grill-bar and Steen (1938, p. 226) called 
a rib is readily identifiable with the hyoid in P. linearis and P. nidzotiensis. 
Structures identified by Fritsch as gill-rays and shown forming external 
gill-tufts in his well-known reconstruction are actually gastralia, as Steen 
has pointed out. 

"Preparation of type specimens from Linton makes it abundantly clear 
that the Pennsylvanian genera Molgophis, Coeytiniis and Lysorophus and 
the Permian species commonly assigned to Lysorophus form a closely-knit 
family of microsaurian affinities. Although the family name Molgophidae 
Cojie 1875 has priority, I prefer to retain, at least for the present, the more 
familiar name Lysorojihidae Williston 1908. Whether Lysorophus Cope 1877 
is genericallj' distinct from Molgophis Cojie 1868 remains to be determined. 

REFERENCES 

References which are cited by date and letter (e.g. Cope, 1868J) will be 
found in the standard bibliographies of Hay, Camp and Romer wliich are 
listed below. 

B.AIRD, D. 

1955. Latex micro-molding and latex-plaster molding mixture. Science, 
vol. 122, no. 3161, p. 202. 

1963. Development of the Westphalian (early Pennsylvania) coal- 
swami) chronofauna. Geol. Soc. America Special Paper 73, pp. 
107-108. 
C.\MP, C. L., et al. 

1940-1961. Bibliography of fossil vertebrates, 1928-1933. Geol. Soc. 
America Special Paper 27, 1940; 1934-1938, Special Paper 42, 
1942; 1939-1943, Geol. Soc. America Mem. 37, 1949; 1944-1948, 
Mem. 57, 1953; 1949-1953, Mem. 84, 1961. 
Cridl.and, a. a., J. E. Morris, and R. W. B.axter 

1963. The Pennsylvanian plants of Kansas and their stratigraphic sig- 
nificance. Palaeontographica, vol. 112, Abt. B, Lief. 1-3, pp. 58-92, 
pis. 17-24, 4 text-figs. 



16 BREVIORA No. 206 

Dechaseaux, C. 

1955. Lepospondyli. In PIVETEAU, J., Ed., Traite de Paleontologie, 
vol. 5, pp. 275-305, 32 text-figs. Paris: Masson et Cie. 

Gregory, J. T., F. E. Peabody, and L. I. Price 

1956. Revision of the Gymnarthridae, American Permian microsaurs. 
Peabody Mus. Nat. Hist., Yale Univ., Bull. 10, ix + 77 pp., 
1 pi., 33 text-figs. 

Hay, O. p. 

1902. Bibliography and catalogue of the fo.ssil Vertebrata of North 

America. U. S. Geol. Survey Bull. 179, 868 pp. 
1929-1930. Second bibliographj' and catalogue of the fossil Vertebrata 
of North America. Carnegie Inst. Washington Publ. 390; vol. 1, 
viii + 916 pp. ; vol. 2, xiv + 1074 pp. 
M.\TTHEW, W. D. and C. De Pavla Couto 

1959. The Cuban edentates. Amer. Mus. Nat. Hist. Bull.; vol. 117, 
art. 1, pp. 1-56, 42 pis., 5 text-figs. 
McGiNNis, H. J. 

1964 MS. The skeletal moriihology of Phlcgcthontia, a Permocarbonif- 
erous aistopod. Unpublished MA thesis, Univ. California, 154 
pj)., 34 text-figs. 
RoMER, A. S., et al. 

1962. Bibliography of fossil vertebrates exclusive of North America, 
1509-1927. Geol. Soc. America Mem. 87, 2 vols., Ixxxix -^ 1544 pp. 
TuRNBULL, W. D. and P. F. Turnbull 

1955. A recently discovered Phlegethontia from Illinois. Fieldiana: 
Zoology, vol. 37, pp. 523-539, pis. 6-9, text-fig. 114. 
Williams, E. E. 

1959. Gadow's arcualia and the development of tetrapod vertebrae. 
Quart. Rev. Biol., vol. 34, no. 1, pp. 1-32, 22 text-figs. 



1964 



AISTOPOD AMPHIBIANS 



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Miuiseiuioii of Compsirsitive Zoology 



Cambridge, Mass. July 28, 1964 Number 207 



NOTES ON THE HORSESHOE BATS 
HIPPOSIDEROS CAFFER, RUBER AND BEATUS 

By Barbara Lawrence 

INTRODUCTION 

Attempts to identify a small series of Hipposideros, collected 
in the Belgian Congo by Alvin Novick in 1956, have made 
necessary a re-examination of the distribution and specific char- 
acters of caffer, ruber and beatus. This has brought to light 
some new characters of the nasal swellings and of the nose 
leaves, and emphasizes the importance of the nasal region in 
general for showing specific differences. 

Hipposideros ruber centralis Andersen 

Study of a series of six males and five females from Kivu 
Forest, Ituri Province, has led to a re-examination of the caffer- 
ruber group in the Museum of Comparative Zoology collections 
in the light of Verschuren's and Aellen's recent discussions of 
these small hipposiderids. The group is admittedly a confusing 
one. It is the more so because as competent a worker as Andersen 
(1906, pp. 281-282) interpreted his findings as evidence of a 
complicated distribution pattern which brought together in a 
single area different subspecies of the species caffer. Subsequently 
Hollister (1918, pp. 85-87) pointed out that two clearly distinct 
species, ruber and caffer, occur together in East Africa. Our mate- 
rial amply confirms this; we even have specimens of ruber and 
caffer taken in the same basement. This situation was apparently 
not clear to Aellen (1952, p. 76) when he followed Hollister 
in synonymizing centralis with ruber, although he failed to give 
ruber full specific status, considering it instead a subspecies of 
caffer. In addition he extended the range of what he called 
caffer ruber clear across the Congo to the Cameroons, whence 



2 BREVIORA No. 207 

he recognized three other races of caffer. Such a distribution 
pattern evidently did not satisfy him and subsequently (Perret 
and Aellen, 1956:435; Aellen, 1957:198) he stated that the whole 
caffer complex was in need of revision, though he did not further 
discuss the relation of ruber to centralis. 

Verschuren (1957, pp. 346-374), although he deals with a 
limited area, seems to have understood the situation better and 
it is significant that much of his work was based on field studies 
of the live bats. Aside from H. beatus which is easily distinct, he 
recognizes two species: caffer centralis, a forest form which 
readily adapts to life in houses, and nanus which is more char- 
acteristically a savannah form. According to him, the latter were 
far less numerous than the fonner and the two were only found 
together in one cave in heavily wooded savannah country. 

Verschuren's distinction between nanus and centralis is good 
and on a subspecific level our material agrees well with his. 
On a specific level, the evidence at hand for considering centralis 
a race of caffer rather than of ruber is not convincing. Length 
of tooth row, width across the molars at M^, and the relation 
of zygomatic to mastoid width, while useful as key characters, 
are not good indicators of specific relationships. Careful examina- 
tion of series of the caffer-ruber and beatus groups from a num- 
ber of localities in a belt across central Africa suggests very 
strongly that modifications of the nasal structures both internal 
and external are the best index to specific relationships. This 
is perhaps not surprising. Since hipposiderids make their sounds 
through their noses, small variations in the relative size of the 
diff"erent parts of the nasal swellings and the septa which in- 
completely divide them should be important. Before final decision 
can be made on the classification of this group, the functional 
significance of these morphological differences needs to be much 
better understood. 

As far as the material studied is concerned, attempts to 
separate ruber and caffer show that, whereas most of the differ- 
ences between the two species are of size not proportion, there 
is a real difference of kind in the compartments of the nasal 
swellings. In all forms, in dorsal view (strongly illuminated from 
below), three pairs of compartments are visible: 1. anterior, more 
or less dorsal and lateral to the external nares; 2. medial, be- 
tween one and the olfactory part of the brain; and 3. postero- 
lateral to the others, and the largest. Variation in size of these 
is a good diagnostic character. When caffer is viewed from above, 



1964 NOTES ON HIPPOSIDEROS 3 

the relatively very small size of compartment two and the some- 
what larger size of three are sufficient to distinguish it from 
ruber. No doubt sections and other more detailed studies would 
show further differences in this region. 

Present knowledge of the small hipposiderids, excluding beatns, 
of east and central Africa can then be summarized as follows. 
In the savannah country of eastern Africa two species, ruber and 
caffer, occur together, of similar proportions externally and 
cranially, but conspicuously different in size. They may be fur- 
ther distinguished by the jDroportions of the nasal compartments. 
Near topotypes of niber from Kilosa in southern Tanganyika 
and s])ecimens from Mtimbuka south of Lake Nyasa resemble 
each other closely, while to the north a series from Mt. Elgon 
is somewhat intermediate towards a more western race. 

This western race has the skull as long as, but slightly narrower 
than, typical ruber and small nasals which give it the appear- 
ance of a large caffer. The proportions of the nasal compartments, 
however, are of the ruber type. The specimens at hand from the 
Ituri Forest, two from Rutshuru and one from Avakubi all be- 
long to this western form, as does probably a specimen from 
Ulkerewe Island. Whether the name centralis is available for 
this western race or whether Hollister is correct in synonymizing 
centralis with ruber, cannot be determined until the type series 
is re-examined. Since the type locality of centralis, Entebbe, 
Uganda, lies west of Mt. Elgon whence we have intermediate 
specimens, it seems advisable for the time being to continue to 
use centralis for the Congo race. 

As for the specific status of centralis, the structure of the nasal 
compartments and apparent intergrading with ruber are evidence 
of a closer relationship with this form than with caffer. This is 
in accord with Andersen's (1906, p. 281) subdivisions of caffer, 
except that his group 2, "H. c. centralis" ( = ruber) , is raised to 
specific rank and two subspecies of this, ruber ruber and ruber 
centralis, are recognized. 

There also occurs in central Africa a bat with ca^er-like 
nasal compartments but of nearly the same size as centralis. We 
have one specimen from Beni of this apparent race of caffer 
which we take to be nanus and probably the same as the series 
so identified by Verschuren. The sympatric occurrence of rep- 
resentatives of two rather similar species of small Hipposideros 
recognized by Hollister in East Africa seems to be typical of 
the Congo region as well. 



4 BREVIORA No. 207 

Verschuren's habitat notes for caffer centralis ( = ruber cen- 
tralis) and nanus {— caffer nanus) suggest that caffer may be 
typically a savannah form which has spread west where suitable 
country is to be found, while ruber, a forest form, has moved 
into dry country fairly extensively, possibly because of its 
adaptability to roosting in buildings. 

HiPPOSIDEROS BEATUS MAXIMUS VeRSCHUREN 

Two males, collected at Mabali, agree with Verschuren's (1957, 
p. 364) description in being larger than beatus from the 
Cameroons, and extend the range of b. maximus considerably 
to the southwest. Until Verschuren published his account of the 
bats of Garamba, it had generally been accepted that nanus was 
a small relative of beatus and represented this species in the 
northeastern Belgian Congo. Actually, as Verschuren has said 
(1957, pp. 370-72), the eastern relative of beatus is a larger 
not a smaller race and nanus is quite a different animal. As 
stated above, it is very likely a forai of caffer. 

With the discovery of this larger race, Andersen's (1906, p. 
275) characters of tooth row and zygomatic arch no longer hold 
for the species. One of the specimens of beatus maximus from 
Mabali is almost identical in zygomatic-mastoid proportions 
with a similar-sized guineensis, a member of the caffer-ruber 
group, from Metet, while length of upper cheek teeth and width 
across the tooth rows taken outside IVP are exactly the same 
in both. As for the supposedly shorter tibia of beatus, the typical 
race is smaller than guineensis and so has a conspicuously shorter 
tibia. In the larger b. maximus, the tibia is only slightly shorter 
and the ratio of length of tibia to length of forearm while usually 
less than in guineensis occasionally shows no difference. 

A search for better diagnostic characters to separate the 
caffer-ruber races from beatus again has shown the importance 
of the nasal region. Cranially, these differences are hard to mea- 
sure though they are apparent enough to the eye. In beatus the 
rostrum bulks larger in proportion to the cranium with the width 
across the nasal swellings relatively greater in proportion to 
their total length. In spite of this, the inflation of the nasal region 
is less than in the caffer-ruber group. In particular, compartment 
three is relatively small as seen both in dorsal and in side view. 

Externally, width of nasals is usually reflected in width of 
nose leaf. While this character is often of subspecific value, 
specific relationships are better shown by differences in details 
of structure of nose leaves than by absolute size. Here again 



1964 NOTES ON HIPPOSIDEROS 5 

we find beatus sharply distinct from the ruber-caffer group. In 
beatus, the center of the horseshoe is shallower, and associated 
with this general difference are a number of particular char- 
acteristics. The septum slopes rather evenly inward to a point 
between the nostrils, whereas in caffer and ruber it has a very 
marked lumpy projection midway of its length, internal to which 
it makes a sharp angle where it meets the face between the 
nostrils. The nostrils themselves are less deep set and the pits 
ventral to them are shallower in beatus than in caffer and ruber. 
Further, in beatus, the two lateral accessoiy leaflets are of nearly 
equal length with a small warty growth dorsal to the end of the 
inner one, while in caffer and ruber the inner accessory leaflet 
is longer than the outer and the wart lies posterior rather than 
dorsal to its end. Verschuren's figures (1957, figs. 137, 141, and 
146) show well the differences in septum between the two groups, 
while the pale rim around the nostrils seems to be associated 
with their shallowness in beatus. The difference which he figures 
in surface of the sella is harder to see in alcoholic specimens and 
the development of the medial thickening is not a character 
which separates all the caffer-ruber group from beatus. Other 
external characters which help to confirm the distinct position 
of beatus are the average more distal attachment of the wing 
membrane and less funnel-shaped ears. 

LITERATURE CITED 
Aellen, V. 

1952. Contribution a I'etude des chiropteres du Cameroun. Mem. 

Soc. Neuchateloise Sci. Nat., 8, fasc. 1:1-121, figs. 1-26. 
1957. Les chiropteres africains du Musee Zoologique de Strasbourg. 
Rev. Suisse Zool., 64, fasc. 1, no. 6:189-214. 
Andersen, K. 

1906. On Hipposideros caffer, Sund. and its closest allies; with some 
notes on H. fuliginosus, Temm. Ann. Mag. Nat. Hist., (7) 
17 : 269-282. 

HOLLISTER, N. 

1918. East African mammals in the United States National Museum. 
Part I. Insectivora, Chiroptera and Carnivora. Bull. U. S. Nat. 
Mus., 99:1-194, pis. 1-55, figs. 1-3. 
Pereet, J. L. and V. Aellen 

1956. Mammiferes du Cameroun de la collection J. L. Perret. Rev. 
Suisse Zool., 63, no. 26:365-450. 

Verschuren, J. 

1957. Ecologie, biologie et systematique des cheiropteres. Institut 
des Pares Nationaux du Congo Beige, Exploration du Pare 
National de la Garamba, Mission H. de Saeger, fasc. 7:1-473, 
1 pi., 1 map, figs. 1-178. 



I 



cO 



BREVIORA 

Miasenam of Compsirative Zoology 



Cambridge, Mass. November 12, 19G4 Number 208 

THREE NEW SPECIES OF FROGS 

(LEPTODACTYLIDAE, ELEUTHERODACTYLUS) 

FROM HISPANIOLA 

By Albert Schwartz 

10,000 S.W. 84th Street, Miami, Florida 

Among specimens collected during the summer of 1962 in 
southwestern Haiti are representatives of two undescribed 
species of Eleutherodactylus, both of which belong to the ricordi 
group. A third new species from the Republica Dominicana was 
taken during the summer of 1963 ; the affinities of this third 
species are distinctly with species from Haiti, and it is ap- 
propriate to describe the three new forms in a single paper. In 
Hispaniola, I have had the capable assistance of Miss Patricia 
A. Heinlein, and Messrs. Ronald F. Klinikowski, Dennis R. 
Paulson, David C. Leber and Richard Thomas. For significant 
comparative material I wish to thank Dr. Doris M. Cochran, 
United States National Museum (USNM), and Dr. Ernest E. 
Williams, Museum of Comparative Zoology (MCZ), whose as- 
sistance in these studies is very deeply appreciated. The il- 
lustrations are the work of Mr. Leber, and I wish to thank him 
and the above mentioned persons for their assistance in the field. 
The types have been given to the Museum of Comparative 
Zoology. Paratypes of the new forms have been deposited in the 
American Museum of Natural History (AMNH) and the 
Museum of Natural History, University of Kansas (KU) ; 
material in my own collection is designated as the Albert 
Schwartz Field Series (ASFS). 

In July 1962, during a stay at Camp Perrin in Haiti, Mr. 
Leber took an especially interesting and rather large frog from 
a nearby cave during a diurnal visit. Nocturnal visits to the cave 
resulted in the taking of no more specimens of this frog, so that 



2 BREVIORA No. 208 

it is represented only by one individual. Although I am reluc- 
tant to describe a new species of Eleutherodnctylus on such 
limited material, the species is so distinctive that I have no 
hesitancy in naming it as 

Eleutherodactylus counouspeus new species 

Figure 1 



^o " 



Holotype: MCZ 43199, an adult male, from Grotte de Counou 
Bois, 1 mile (1.6 km) southwest of Camp Perrin, Dept. du Sud, 
Haiti, taken 30 July 1962 by David C. Leber. Original number 
X3266. 

Diagnosis: An Eleutherodactylus of the ricordi group charac- 
terized by a combination of large size (snout-vent length 48 
mm), large digital discs, a greenish yellow dorsal ground color 
with black markings consisting primarily of a dark interocular 
bar, a dark postocular V, and a scapular cross, a heavily mottled 
throat, and lacking inguinal glands. 

Description of type: An adult male, with the following meas- 
urements (in millimeters) : snout-vent length, 48.0; head length, 
18.5; head width, 17.2; diameter of tympanum, 3.1; diameter 
of eye, 7.1 ; naris to anterior corner of eye, 6.4 ; femur, 21.3 ; 




Fig. 1. Eleutherodactylus counouspeus, new species, type, MCZ 43199, 
snout-vent length 48.0 mm. 



1964 NEW FROGS FROM HISPANIOLA 3 

tibia, 22.5; fourth toe, 18.1. Head slightly narrower than dis- 
tance from snout to posterior border of tympanum ; snout de- 
cidedly truncate with nares prominent at anterior end of can- 
thus rostralis ; diameter of eye greater than distance from naris 
to anterior corner of eye ; diameter of tympanum about one-half 
diameter of eye, distance from tympanum to eye about one- 
third diameter of tympanum ; tympanum oval, the vertical 
diameter slightly greater than the horizontal. Interorbital dis- 
tance 5.5, less than diameter of eye. Digital discs present and 
well developed, those on fingers three and four distinctly larger 
than those on digits one and two, disc of finger three the largest 
and equal to about three-quarters size of tympanum. Fingers 
long and slender, unwebbed, 3-4-2-1 in order of decreasing 
length ; subartieular tubercles prominent, gray. Toes moderately 
long, with vestigial webs, 4-3-5-2-1 in order of decreasing length ; 
subartieular tubercles prominent, gray. Heels touch when fe- 
mora held at right angles to body axis. Inguinal glands absent. 
Dorsum smooth ; upper eyelids with low rounded tubercles. 
Throat slightly granular, belly smooth ; abdominal disc poorly 
developed but lateral margins of disc fairly conspicuous. Dorsal 
surfaces of forelimbs smooth, of hindlimbs granular, especially 
on dorsal surfaces of crura. Posterior face of thighs with many 
small, juxtaposed rounded granules. Tongue small, entire, free 
behind, its greatest width equal to about one-half that of 
floor of mouth. Vomerine teeth in two short, almost straight 
series, beginning well within the inner margins of the choanae 
and separated from the choanae by a distance equal to about 
three times the diameter of a choana, the two series separated 
from each other by a distance equal to the diameter of a choana. 
Coloration of type: Dorsal ground color in life greenish yellow 
(slightly brighter than PI. 21L1 of Maerz and Paul, 1950), with 
dorsal black markings as follows: a black interocular bar, a 
black median postocular V, a black scapular cross with a rounded 
black spot anterior and lateral to the side "arms" of the cross, 
a rather linear blotch on the sides posterior to the forearm in- 
sertion, and the balance of the dorsum and sides with lighter 
and irregular obfuscations of dark (Fig. 1). Fore- and hind- 
limbs diffusely spotted with dark with no apparent banding or 
barring; concealed surfaces pale purplish gray overlaid with 
brown suffusions. Throat yellowish green, heavily mottled with 
brownish; belly pinkish with admixture of yellow-green, and 
suffused with brown stippling ; underside of crura rather heavily 



4 BREVIORA No. 208 

marbled with brown. Lores yellow-green with an indistinct 
darker canthal line. Iris bronzy in life, with a reddish pupillary 
ring. 

Comparisons: No other Hispaniolan members of the ricordi 
group reach the size of connouspeus except E. schmidti Noble 
and its races rucillensis Cochran and limhensis Lynn, and pos- 
sibly E. femurlevis Cochran. The webbed feet in E. schmidti, 
along with the entirely different color pattern and the reddish- 
orange venter will distinguish it from counouspeus. E. schmidti 
is known only from the Cordillera Central and the Cordillera 
Septentrional in the Republica Dominicana and from the area 
about Limbe in Haiti and is thus not sympatric with counou- 
spejis. I have not examined the type and apparently only speci- 
men of femurlevis, which has a snout-vent length of 42 mm 
(Cochran, 1941:62); however, the detailed description of the 
pattern as well as the photograph of the specimen {ihid., pi. 
IOC) indicate that there is little similarity between it and 
counouspeus. E. femurlevis has a pale, wide, interoeular bar 
banded narrowly anteriorly with brown, a brown loreal spot, 
pale gray crossbands on the legs, immaculate venter — none of 
which corresponds to the pattern of counouspeus. The photo- 
graph of the type does not show a conspicuously large-eyed 
frog, and the legs appear shorter and the discs much less en- 
larged. It has seemed most pertinent to compare these two 
species not only because of some similarity in size but also 
because they both occur in the same general region on the 
southwestern Haitian peninsula, femurlevis having been de- 
scribed from Desbarriere, on the north and east foothills of 
the Massif de la Hotte. 

Observations: The type of E. counouspeus was taken by Mr. 
Leber as it actively hopped about on rocks about fifty feet back 
from the entrance of the Grotte de Counou Bois in the morning. 
No other specimens were secured on a later visit and no voice 
was heard which might be associated with this species ; E. 
pictissimus was taken fairly abundantly in the same cave. 

In general habitus, E. counouspeus resembles E. zeus from 
Cuba ; both are rather long-legged, large-eyed species with large 
digital discs. The pattern of the two is vaguely similar although 
they are not at all comparable in coloration. Since both are 
petricolous and/or cavernicolous forms, I feel that these simi- 
larities are due to convergence and do not show relationships. 
In Antillean Eleutherodactylus there is often a remarkable 



1964 NEW PROGS FROM HISPANIOLA 5 

similarity in pattern and coloration between unrelated forms 
which are petricolous (Schwartz, 1960:41-42). 

A second new species of the ricordi group from the western 
portion of the Tiburon Peninsula is named in honor of Dennis 
R. Paulson who helped collect the type series and whose interest 
and diligence in field work in the West Indies deserves special 
mention : 

Eleutherodactylus paulsoni new species 
Figure 2 

Holotype: MCZ 43200, a gravid female, from 4.5 miles (7.2 
km) northwest of Les Cayes, Dept. du Sud, Haiti, one of a 
series taken 7 August 1962 by David C. Leber and Dennis R. 
Paulson. Original number X3796. 

Parahjpes: ASFS X3797-98, AMNH 71987-89, same data as 
type; ASFS X2799, Camp Perrin, Dept. du Sud, Haiti, 23 
July 1962, collected by a native ; MCZ 43201, Grotte de Counou 
Bois, 1 mi. (1.6 km) SW Camp Perrin, Dept. du Sud, 26 July 
1962, Lucien Rigaud; KU 79811, 4 km (6.4 km) NW Les Cayes, 
Dept. du Sud, Haiti, 7 August 1962, D. C. Leber; MCZ 33825, 
Place Negre, near Jeremie, Dept. du Sud, Haiti, Luc and George 
\¥hiteman, 11 December 1960. 

Diagnosis: A species of Eleutherodactylus of the ricordi group 
characterized by a combination of rather small size (females 
snout- vent length to 26 mm), very small digital discs, a dorsal 




Fig. 2. Eleutherodactylus paulsoni, new species, type, MCZ 43200, snout- 
vent length 25.4 mm. 



6 BREVIORA No. 208 

pattern of dark brown marbling with a pair of butfy dorso- 
lateral stripes, scattered dorsal pinkish warts, without red or 
orange on the hindlimbs, and with inguinal glands. 

Description of type: A gravid female with the following 
measurements (in millimeters): snout-vent length, 25.4; head 
length, 10.5 ; head width, 10.0 ; diameter of tympanum, 2.1 ; dia- 
meter of eye, 3.6 ; naris to anterior corner of eye, 3.0 ; femur, 
11.9; tibia, 13.5; fourth toe, 11.5. Head slightly narrower than 
distance from snout to posterior border of tympanum ; snout 
truncate with nares prominent at anterior end of canthus ros- 
tralis; diameter of eye slightly longer than distance from naris 
to anterior corner of eye; diameter of tympanum greater than 
one-half diameter of eye, distance from tympanum to eye equal 
to about one-half diameter of tympanum. Fingers rather long 
and slender, unwebbed, 3-4-2-1 in order of decreasing length; 
subarticular tubercles prominent, gray. Toes moderately long, 
with vestigial webs, 4-3-5-2-1 in order of decreasing length ; sub- 
articular tubercles prominent, gray. Heels overlap when fe- 
mora held at right angles to body axis. Inguinal glands present. 
Dorsal surface, including eyelids, snout, and upper surfaces of 
all limbs heavily rugose ; a raised median dorsal line. Throat 
and belly smooth ; abdominal discs poorly developed. Posterior 
and ventral surfaces of thighs with many large, conspicuous 
juxtaposed granules. Tongue rather small, entire, free behind, 
its greatest width equal to about one-half that of floor of mouth. 
Vomerine teeth in two rather long, slightly bowed, series, be- 
ginning at outer margin of choanae and separated from the 
choanae by a distance equal to one-half the diameter of a cho- 
ana, the two series separated from each other by the same 
distance. 

Coloration of type: Dorsal ground color in life brown, with a 
rather irregularly marbled darker brown pattern, a dark brown 
interocular bar preceded by a paler tan snout, much marbled 
with dark brown. Scattered conspicuous warts from the level 
of the eyes onto the lower back, pinkish in life and strongly 
contrasting with the dark ground color. Dorsolateral stripes 
indistinct and buffy, almost completely obscured in the pre- 
served specimen by dark pigment. Dorsal ground color dis- 
tinctly more reddish posteriorly than anteriorly. Antebrachia 
with about three obscure crossbars, brachium marbled dark ; 
hindlimbs very obscurely crossbarred and dotted w^ith paler, 
their concealed surfaces dark brown, with some lighter stippling 



1964 NEW FROGS FROM IIISPANIOLA 7 

medially. Belly immaculate opalescent whitish ; throat with 
large to small discrete dots and stipples; underside of ante- 
brachia clouded with brown ; underside of thighs and crura 
rather heavily marbled and clouded with brown. 

Variation: Five adult females (including the type) have the 
following measurements: snout-vent length, 24.0 (21.5-25.8); 
head length, 9.6 (8.3-10.6); head width, 9.1 (8.1-10.1); tym- 
panum, 2.0 (1.9-2.1) ; eye, 3.5 (3.2-3.8) ; naris to eye, 2.8 (2.4- 
3.2); femur, 11.3 (10.0-13.6); tibia, 12.3 (11.3-13.9); fourth 
toe, 10.7 (9.8-12.2). A single adult male paratype measures: 
snout-vent length, 22.1 ; head length, 8.8 ; head width, 7.8 ; 
tympanum, 1.9; eye, 3.3; naris to eye, 2.3; femur 10.4; tibia, 
11.4; fourth toe, 10.7. Females reach a larger size than males; 
interestingly, the largest female (MCZ 33825) is from the north 
side of the Massif de la Hotte. The condition of this specimen 
is such that it is not readily compared with specimens from 
the south side of the Massif, but superficially it seems identical 
except for its slightly larger size. It is, of course, possible that 
such northern frogs may be found to be racially distinct from 
southern frogs. 

In color and pattern, the series shows variation principally 
in the distinctness of the dorsolateral stripes. When present, 
they are boldly outlined in dark brown and extend from the 
posterior portion of the upper eyelid to the groin. The brown 
dorsum with a more reddish (chestnut) tinge posteriorly was 
noted for all fresh material ; the pinkish warts are likewise a 
characteristic feature. The dorsolateral stripes vary from buffy 
to reddish; the interocular bar can be either tan or light buffy. 
The belly varied from white to translucent gray. The degree 
of throat spotting varies considerably, with the type showing 
the mean condition ; in some specimens the dots form almost 
a complex dark brown reticulum and in others they are larger 
and more uniform than in the type, but nonetheless discrete. 
All specimens, even the smallest juvenile (snout-vent 11.5), 
have the throat marked with dark spots. 

Comparisons: Shreve and AVilliams (1963:331) referred to a 
single specimen (MCZ 33825) under their discussion of E. 
furcyensis saying that "a single specimen from the vicinity 
of Jeremie . . . somewhat resembles this species and may be the 
La Hotte region representative." The specimen referred to is 
here designated as a paratype of E. paulsoni. Their comment, 
however, points out the comparison which it is most necessary 



8 BREVIORA No. 208 

to make — that between paidsoni and furcyensis: In coloration 
and pattern these two species are quite different. E. furcy- 
ensis has prominently banded crura and thighs, and lacks dorso- 
lateral lines. The anterior and posterior faces of the thighs are 
red (PI. 4D11) in furcyensis (these same areas are dark in 
paulsoni) and there is additionally a faint orange wash in the 
groin. Female furcyensis have the throat slightly purplish, 
flecked with black, whereas male furcyensis have the belly pale 
yellow; neither of these features occurs in paulsoni. Furcyensis 
has a dark brown dorsum with a pair of pale orange-buff re- 
versed parentheses, not a pair of dorsolateral lines. The two 
species do resemble each other in the throat spotting, which is 
a variable but always present feature in furcyensis, although 
no specimens at hand have the throat with a dark reticulum. 

Structurally, the two species differ in that paulsoni has in- 
guinal glands and furcyensis lacks them. In comparably sized 
specimens, the discs of furcyeyisis are larger than those of 
paulsoni. The differences in coloration and pattern are so strik- 
ing as to preclude the possibility that these two forms are 
related on a subspecific level. 

In size, furcyensis exceeds paadsoni in both sexes (largest 
female furcyensis snout-vent, 37.0; largest male, 28.0), although 
the small series of paulsoni makes comparison, difficult. At least 
fourteen adult female furcyensis have higher values in all 
measurements taken. 

The remaining ricordi group members in Hispaniola are : 
glandulifer, darlingtoni, weinlandi, rufifenioralis, schmidti, fe- 
murlevis, pictissimus, leoncci and counouspeus (list j^rincipally 
from Shreve and Williams but with some interpolations of my 
own). Of these species, schmidti and counouspeus are known 
to lack glands. These two forms are likewise much larger than 
paidsoni, are differently patterned, and schmidti has webbed 
feet. Of the forms with glands, paulsoni is exceeded in size 
by glandulifer, darlingtoni, weinlandi, pictissimus, and leoncei. 
None of these larger species have patterns which are in any 
way comparable to that of paulsoni, although the dorsum of 
weinlandi with its anteriorly and posteriorly differing dorsal 
ground colors is vaguely similar. E. darlingtoni has large digital 
discs, whereas leoncei, weinlandi and pictissimus have small 
digital discs. I have not examined specimens of either femur- 
levis or rufifenioralis; the latter is however much smaller than 
paulsoni (largest recorded snout-vent length 18) and is much 



1964 NEW FROGS FROM HISPANIOLA 9 

differently patterned. E. femurlevis, on the other hand, is much 
larger (snout -vent 42) and again with a different pattern. 

Observations : The type and paratopotypes were taken in leaf 
litter under large trees along the bank of an intermittent river 
(Riviere de la Grande Ravine du Sud) ; the adjoining area 
was thorn scrub. Another was taken at the cave entrance of 
the Grotte de Counou Bois, and a single individual from 4 km 
NW Les Cayes was taken on the ground among dead leaves 
in grass about 2 m high. At the type locality of paulsoni, E. 
pictissimus was the commonest frog encountered. 

Shreve and Williams (1963:338-9) discussed the Hispaniolan 
and Cuban components of the dimidiatus and varleyi groups. 
To the former they assigned the Cuban species alhipes, dimi- 
diatus, and emiliae and the Hispaniolan jugans, and to the latter 
the Cuban varleyi and cuhanus (which has customarily been 
placed in the dimidiatus group) as well as E. phyzelus Schwartz 
which I consider a synonym of E. varleyi Dunn. I concur in 
these assignments. Shreve and Williams also consider E. ventri- 
lineatus a member of the diynidiatus group (although it is pe- 
ripheral) in spite of its short vomerine series {dimidiatus group 
members are characterized by long vomerine series). E. ventri- 
lineatus is so obviously related to E. jugans that, except for the 
vomerine series, one might be inclined to regard them as con- 
specific. Thus, on Hispaniola, jugans and ventrilineatus are 
the only known members of the dimidiatus group. Both are 
restricted to higher elevations in the mountains of the south 
island, jugans in the Massif de La Selle, ventrilineatus in the 
Massif de la Hotte. 

A small series of a third Hispaniolan species belonging to 
this assemblage was taken by Richard Thomas in the Sierra 
de Neiba in the Republica Dominicana; since this form is the 
first recorded from the north island (and has thus "crossed" 
the Cul de Sac-Valle de Neiba plain), I propose that it be named, 
from the Greek word for "transgressor," 

Eleutherodactylus parabates new species 
Figures 3, 4 

Holotype: MCZ 43202, a gravid female, from 20 km south- 
west of Hondo Valle, 5950 feet (1800 m), Independencia Pro- 
vince, Republica Dominicana, taken 11 August 1963 by Richard 
Thomas. Original number V366. 



10 BREVIORA No. 208 

Parahjpes: ASPS V365, V367-70, same data as type. 

Diagnosis: An Eleutherodactylus, related to E. ventrilineatus 
Shreve, with short vomerine series but differing from both 
ventrilineatus and jugans in smaller size, in the presence of 
digital discs, in ventral pattern, and in relative size of tym- 
panum. 

Description of type: A gravid female with the following 
measurements (in millimeters): snout-vent length, 24.1; head 
length, 8.6 ; head width, 8.8 ; diameter of tympanum, 1.7 ; dia- 
meter of eye, 2.7 ; naris to anterior corner of eye, 2.4 ; femur, 
8.6; tibia, 9.8; fourth toe, 9.3; tympanum/tibia ratio 16.8. Head 
slightly broader than distance from snout to posterior border 
of tympanum ; snout rather pointed with nares conspicuous at 
anterior end of canthus rostralis; diameter of eye greater than 
distance from naris to anterior corner of eye ; diameter of 
tympanum greater than one-half diameter of eye, distance from 
tympanum to eye equal to about three-quarters diameter of 
tympanum. Interorbital distance 3.2 mm, greater than diameter 
of eye. Digital discs present and moderately well developed, 
those on fingers three and four larger than those on digits one 
and two, disc on finger three the largest and equal to about 
one-half size of tympanum. Fingers rather short, unwebbed, 
3-4-2-1 in order of decreasing length ; subarticular tubercles 




Fig. 3. Eleiitlierodactylus pardbates, new species, type, MCZ 43202, snout- 
vent length 24.1 mm. 



1964 NEW PROGS FROM HISPANIOLA 11 

fairly prominent, gray. Toes moderately long, nnwebbed, 
4-3-5-2-1 in order of decreasing length ; subarticular tubercles 
prominent, gray. Heels overlap slightly when femora are held 
at right angles to body axis. Inguinal glands absent. Dorsum 
smooth. Throat and belly smooth ; abdominal disc not prominent 
with only the pectoral fold moderately conspicuous. Dorsal 
surface of forelimbs smooth, of thighs smooth, but dorsal sur- 
face of crura studded with a few scattered low tubercles. Pos- 
terior face of thighs with many low rounded juxtaposed gran- 
ules. Tongue rather large, ovate, entire, free behind, its great- 
est width equal to slightly more than half of floor of mouth. 
Vomerine teeth in two short transverse patches, beginning at 
the midlevel of the choanae and separated from the choanae 
by a distance equal to about two and one-half times the diameter 
of a choana, the two series separated from each other by a 
distance equal to twice the diameter of a choana. 

Coloration of type: Dorsal ground color brown in a longitudi- 
nal band from snout to groin, enclosing a median tan dorsal 
line from snout to above vent, where it divides, each branch 
proceeding across the posterior of the thigh to behind knee; 
dorsal band separated from darker brown sides by a faint tan 
dorsolateral stripe which begins at the eye and proceeds to 
near the groin (Fig. 3). Sides darker brown. A tan canthal 




rig. 4. Eleutherodactylus paraiates, new species, ventral view, type, MCZ 
43202. 



12 BREVIORA No. 208 

line above a dark brown stripe which continues posteriorly as a 
supratympanic stripe to above the forelinib insertion; lores and 
cheek brownish, lips mottled. Dorsal surface of forelimbs brown, 
brachia slightly paler, antebrachia with remnants of two or three 
crossbars. Dorsal surface of hindlimbs dark brown, as are con- 
cealed surfaces, with a vague remnant of one darker crossbar 
on the crura ; a darker brown postanal triangle present. Throat 
and belly yellowish tan, heavily mottled with brown ; the mottling 
consists of dots of varying intensities, with the large dots gen- 
erally darker, the smaller lighter, giving an irregularly mottled 
appearance (Fig. 4). Undersurface of all limbs mottled with 
brown, heaviest on crura and pes. 

Variation: Four gravid females (including the type) have 
the following measurements (means followed by ranges) and 
ratios: snout-vent length, 23.8 (22.3-24.3); head length, 8.6 
(8.0-8.9) ; head width, 8.9 (8.5-9.4) ; tympanum, 1.6 (1.5-1.7) ; 
eye, 2.7 (2.6-2.8) ; naris to eye, 2.1 (1.9-2.4) ; femur, 8.6 (8.3- 
9.0) ; tibia, 9.6 (9.3-9.8) ; fourth toe, 9.2 (9.0-9.3) ; tympanum/ 
tibia ratio, 16.5 (15.6-17.3) ; tibia/snout-vent length, 40.3 (38.3- 
43.0). A single adult male measures: snout-vent length, 17.6; 
head length, 6.2; head width, 6.7; tympanum, 1.0; eye, 2.0; 
naris to eye, 1.5; femur, 6.7; tibia, 7.3; fourth toe, 7.2; tym- 
panum/tibia ratio, 13.7 ; tibia/snout-vent length, 41.5. 

In life the dorsal ground color of the entire series was like 
that of the type ; however, only one other paratype has the 
median dorsal line. In this specimen the dorsal band is some- 
what lighter than the type and there is conspicuous deposition 
of dark pigment adjacent to the median line and laterally along 
the region of the dorsolateral line, so that the dorsal band is 
rather conspicuously cut off from the sides, and the median 
line is also set off from the dorsal band. The four specimens 
which lack the median line also seem to lack a dorsal band. In 
these there is a dark interocular bar setting off a paler (more 
grayish) snout, and the sides are vaguely barred posteriorly. 
These specimens also have a single dark antebrachial bar and a 
single crural bar with a faint indication of a thigh crossbar as 
well. Even the smallest juvenile (snout- vent 11.0) shows the 
dark ventral mottling of the type, which is also a standard fea- 
ture of the balance of the series. One female paratype addi- 
tionally has a pair of tan scapular spots, each accompanied 
by a more laterally placed darker brown spot. 



1964 NEW FROGS FROM HISPANIOLA 13 

Comparisons: Only two known Ilispaniolan species of Eleu- 
therodactylus are comparable to E. parahates in squatty habitus 
— E. jugans and E. ventrilineatus. From both of these, para- 
hafcs differs in possessing- digital discs. From jugans, par abates 
differs as well in having a short vomerine series. I have not seen 
either jugans or venfrilincafus in life, and preserved specimens 
are all rather old and discolored. However, specimens of ventri- 
lineatus lack a middorsal line, which occurs in parahates. Jugans 
has a double crossbar on the crura and about three crossbars 
on the thigh, whereas ventrilineatus has a single bar as does 
parahates. The ventral coloration of the three species is very 
distinct ; the differences between jugans and ventrilineatus are 
well shown by Cochran (1941:34 and 35), the former having 
the belly heavily and irregularly pigmented, and ventrilineatus 
having the belly uniformly pigmented with a median ventral 
clear line. Thus, parahates has the least ventral pigmentation 
of the three. 

Of the three species, jugans reaches the largest size (females 
to snout-vent length of 33.3), with ventrilineatus smaller (fe- 
males to 30.6) and parahates smallest (females to 24.3). The 
single male of ventrilineatus (snout-vent length 24.9) is larger 
than the largest male jugans (22.5) or parahates (17.6). Para- 
hates and ventrilineatus may be separated by the tympanum/ 
tibia ratio; in the former, this ratio ranges (in females) between 
15.6 and 17.3, in the latter (females) between 20.0 and 21.9; 
the single male parahates has a ratio of 13.7, the single male 
ventrilineatus 19.6. Tibia/snout-vent length ratio likewise is 
diagnostic between these two species, varying in parahates be- 
tween 35.6 and 37.6 and in ventrilineatus between 38.3 and 
43.0 (all females). 

Ohservations and comments: The type series of E. parahates 
was collected under rocks and wood adjacent to a road through 
dense rain forest in the Sierra de Neiba (southern range). 
The forest in this area was composed of hardwoods and some 
ehano vercle (Magnolia doming uensis) and in the vicinity were 
low areas which supported small marshes with cat-tails, a 
rather unusual feature for such a high elevation. 

The members of the dimidiatus group, as presently understood 
are: 

dimidiatus Cope (with its race amelasma Schwartz) 

emiliae Dunn 

intermedius Barbour and Shreve 



14 BREVIORA No. 208 

alhipes Barbour and Shreve 

jugans Cochran 

ventrilineatus Shreve 

parabatcs Schwartz 
Of these, four are Cuban and three are Hispaniolan ; two of 
the Hispaniolan species occur on the south island, and the 
other just across the Cul de Sac-Valle de Neiba plain, in the 
Sierra de Neiba. Ve^itrilineatus and parabaies are slightly 
aberrant in that they both possess short vomerine series. Para- 
hates also has digital discs; as originally diagnosed (Dunn, 
1926 :210) this group was composed of members with very 
feebly developed discs. Aside from the color features which 
Dunn considered as diagnostic of the group, the structural fea- 
tures regarded as pertinent are : 1 ) discs present and small, 
or absent, 2) smooth belly, 3) long vomerine series (or short 
in two species), 4) no external vocal sac, 5) squatty habitus, 
and 6) inguinal glands absent. The last character may well 
not be valid ; I have examined only dimidiatus, jugmis, ventri- 
lineatus and parahates for this feature. 

Of the included species, E. dimidiatus, as Shreve and Wil- 
liams (1963:338) pointed out, is somewhat different in that it 
lacks the stocky build of the remaining forms. It too is the 
only known species which is not restricted to high elevations, 
although it does occur at high elevations in the Sierra de Trini- 
dad and Sierra de Grand Piedra in Cuba. As far as known, 
emiliae occurs only in the Sierra de Trinidad, and intermedius 
and alhipes in the Sierra Maestra. 

There is a possibility that E. unicolor Stejneger is the 
single Puerto Rican representative of the dimidiatus group. 
Relying on Stejneger 's (1904:597-598) description of the type, 
I feel that unicolor agrees with my concept of the group in all 
features except that the belly is granular rather than smooth. 
Additionally, it is a high mountain form, and thus agrees with 
most dimidiatus group species. No Jamaican species are obvi- 
ously associated with the dimidiatus group. 

I have had field experience only with dimidiatus. At least 
this species is vocal; the call is an insect-like twittering, males 
calling from the ground in mesic forested situations. No calls 
have been reported for any other species. 

Specimens examined: Eleutherodactylus ventrilineatus: MCZ 
19857-61, 19863 (type and paratypes), Mt. La Hotte (=Pic 
Macaya), Dept. du Sud, Haiti; Eleutherodactylus jugans: MCZ 



1964 NEW FROGS FROM HISPANIOLA 15 

21594-95, MCZ 19852-56, plus four unnumbered specimens (type 
and paratypes), near La Visite, La Selle Range, Dept. de 1 'Quest, 
Haiti; USNM 95423-27 (paratypes), La Visite, Dept. de I'Ouest, 
Haiti. 

LITERATURE CITED 

Cochran, Doris M. 

1941. The herpetology of Hispaiiiola. Bull. U.S. Nat. Mus, 177:i-vii, 
1-398, 120 figs., 12 pis. 
Dunn, E. R. 

1926. Additional frogs from Cuba. Occ. Papers Boston Soc. Nat. Hist., 
5:209-215. 
Maekz, a., and M. Rea Paul 

1950. A dictionary of color. New York, McGraw-Hill Book Co., pp. 
i-vii, 1-23, 138-208, 56 pis. 
Schwartz, Albert 

1960. Nine new Cuban frogs of the genus EleutJierodactylus. Reading 
Public Museum and Art Gallery, Sci. Publ., 11:1-50, 6 figs. 
Shreve, Benjamin, and Ernest E. Williams 

1963. The herpetology of the Port-au-Prince region and Gonave Island, 
Haiti. Pt. 11. The frogs. Bull. Mus. Comp Zool., 129(5) :302- 
342, 5 pis. 
Stejneger, Leonhard 

1904. The herpetology of Porto Rico. Rept. U.S. Nat. Mus., 1902:549- 
724, 1 pi., 196 figs. 



BREVIORA 

MiLiseiui.-nTi of Connparative Zoology 

Caimbridge, ]\Iass. .\()\ f.mhkr 12, 19()4 Number 209 



A XEAV SKATE. RAJA CERVIGONI, FROM VENEZUELA 

AND THE GUI ANAS 

By TTexkv T>. Bigelow and William C. SchroederI 

Raja cervigoni sj). iiov. 

Holofype: An immature male, 357 mm in total length, from 
10 miles (16 km) northeast of Carupano, in 20-30 fathoms 
(37-55 m), Museo Historia Natural La Salle, Venezuela, No. 
873. 

Paratjjpes: A male of 206 mm and a female of 229 mm from 
Punta Arava, Estado Sucre, in about 20 fathoms (37 m), from 
off the eastern part of Venezuela, and a male of 235 mm from 
off tlic Guianas, 07° 25' N, 54° 35' W, in 75-80 fathoms (137- 
U5 m), "Oregon" station 2289. 

Distinctive characters. Among rajids from the western Atlan- 
tic that have a pair of ocelli on the disc, cervigoni most ckjsely 
resembles Raja cyclophora Regan 1903, but it differs in lacking 
dark mucous pores on the under surface, in having orbital and 
nuchal thorns, and 3 rows of thorns on the tail, whereas cyclo- 
phora has prominent blackish streaks below marking mucous 
pores, lacks orbital and nuchal thorns, and has a single row of 
thorns on the tail. In cervigoni the more sharply rounded outer 
corners of the disc, which is also relatively wider (averaging 
73.2 per cent of the total length of the specimen on 4 indi- 
viduals), distinguishes it from texana Chandler 1921 (average 
disc width 63.8 per cent on 23 specimens) and also from ackleyi 
Garman 1881 (average 59.4 per cent on 3 specimens) ; and its 
fewer thorns (16-26) in the midrow on the tail, between the 
axils of the pectorals and the first dorsal fin, tcgether with the 
axis of greatest width further rearward (73-77 per cent, meas- 
ured between the tip of the snout and axils of pectorals), set 

1 Contrilnition No. 1522 from rhn Woods Hole Oceanograiihic Institution. 



2 BREVIOKA No. 209 

it apart from hahanu usis Hv^ehnv and Schroeder- on which the 
thorns number 34-47 (4 specimens) and the greatest disc width is 
64-69 per cent rearward. 

Description of Jioloti/jx'. Proportional dimensions in per cent 
of total length. 

Disc. — Extreme breadth 72.8 ; length 52.1. 

Snout length. — In front of oi-bits 12. 6 ; in front of mouth 14.6. 

Orbits. — Horizontal diameter 4.4; distance between 4.2. 

Spiracles. — Length 2.8 ; distance between 6."). 

Mouth. — Breadth 8.4. 

Exposed nostrils. — Distance between inner ends 8.2. 

Gill openings. — Length 1st 2.0; 3rd 2.0; 5th 1.5; distance be- 
tween inner ends, 1st 15.6 ; 5th 8.0. 

First dorsal fin. — Height 3.1 ; length of base 5.3. 

Second dorsal fin. — Height 3.1 ; length of base 4.8. 

Pelvics. — Anterior margin 13.2. 

Distance. — From tip of snout to center of cloaca 47.6 ; from 
center of cloaca to 1st dorsal 33.0 ; to tip of tail 52.4 ; from rear 
end of 2nd dorsal to tip of tail 3.7. 

Interspace. — 1st and 2nd dorsals 5.6. 

Disc 1.4 times as broad as long; maximum angle is front of 
spiracles 102°; snout pointed; anterior margins of disc nearly 
straight, outer corners shari)ly rounded; posterior and inner 
margins moderately convex. Axis of greatest breadth 73 per 
cent of distance back from tij) of snout to axils of pectorals. 
Tail rath<M- slender, the lateral folds low down, originating a 
little in advance of tips of pelvics, widening somewhat pos- 
teriorly as usual among rajids, reaching tip of tail ; length of 
tail from center of cloaca to origin of first dorsal fin 0.70 times 
as great, and to its tip 1.10 times as great as distance from 
center of cloaca to tiji of snout. 

A row of 7 thorns along the anterior and inner margins of 
each orbit and a tiny thorn opposite inner margin of each 
spiracle and 2 over tip of I'ostrum. Three jirominent thorns 
along the midline in the nuchal region, followed by a row of 
about 40 thorns which originates about midway between the 
rear margin of orbits and axils of pectorals, reaching to first 
dorsal fin, the first few thorns very small ; 4 thorns in the 
interdorsal space. A row of about 30 thorns low down each 
side of the midrow on tail, beginning ojiposite axils of ])elvics, 
reaching to opposite and second dorsal, very small rearward. 

2 In press (bull. .Mus. ('diup- Zdul.). 



IfUU NKW Sl'K(li;s OF SKATK 3 

i\Iust of tlu' thorns oil tail alternate in size. Tlie upper surface 

of disc and tail otherwise sinootli. A short i-ow of S oi- !) pale 

mucous pores eaeh side and close to the tii'st two nuchal thorns. 

Lower surface with small prickU's on end of snout and in a 

narrow band along margin of disc from snout to about opposite 

mouth. 

Snout in front of orbits 2.9 times as long as orbit ; its length 

in front of mouth 1.8 times as great as distance between exposed 

nostrils. Distance between orbits about equal to length of orbit. 

Orbit 1.6 times as long as spiracle. Nasal curtain and expanded 

outer margin of nostrils fringe:!. Jaws moderately arcluMl. T(n'th 

4'-' 

-T-, arranged in (junicunx, with ovate base ami triangular cusp. 

Distance between first gill openings 1.!) times as great as be- 
tween exposed nostrils ; between 5th openings about equal ; first 
gill openings O.-to times as long as longest diameter of orbit. 
Dorsals similar in shape, and nearly so in size, the interspace 
about equal to length of base of first dorsal. Second dorsal 
confluent with the caudal fin the base of which is shorter than 
that of the second dorsal. Pelvics deeply concave, scalloped 
alo)ig anterior side of excavation, weakly so rearward; anterior 
lobe slender, about % as long as distance from its own origin 
to rear tip of pelvic ; posterior lobe with sharply rounded tip, 
extending about % the distance from axil of pectorals to first 
dorsal. The claspers of this immature specimen extend only 
to the tips of the pelvics. Rostral cartilage firm, narrow, ex- 
tending nearly to tip of snout. Anterior pectoral rays reach- 
ing 58 per cent of the distance from axis through front margin 
of orbits to end of snout. 

Color. Upper surface, including dorsals, plain medium brown. 
A prominent ocellus on each side of disc, situated a little pos- 
terior to the greatest axis of disc, its center from the midline 
of disc a distance about equal to that from tip of snout to 
anterior ^ of orbit; distance between centers of ocelli 1.1 times 
distance to centers of orbits. The ocellus is formed by a narrow 
black-brown circle 14 mm in diameter, within which two dark 
spots are present on the left ocellus and one spot on the right. 
Below, plain whitish on disc and pelvics, with no dark mucous 
pores, the tail with pale brown blotches. 

The three paratypes agree closely with the holotype in most 
proportional dimensions and otherwise, the chief vai-iations 
being as follows: anterior angle of disc 104°-109° ; orbital thorns 
3, of which 2 are along anterior margin and 1 at inner rear 



4 BHEVIORA Xu. 209 

margin; 2 nuchal thorns; thorns in midline to first dorsal, 
beginning opposite axils of jiectorals, 15-16 ; 2 thorns between 
dorsals ; side row of thorns on tail, originating opposite axils 
of pelvies and ending opposite second dorsal, 16-21; these vary 
slightly in number from side to side ; few of the tail thorns 
alternate in size; sn(,ut in front of orbits 2.6-2.8 times as long 
as orbit, its length in front of mouth 1.8-2.0 times as great as 
distance between nostrils; space between dorsals about %-% 

as long as base of first dorsal. Teeth '— — — , with low tri- 

angular cusj). On the lower surface, the narroAv band of prickles 
along the anterior margin of disc is present, extending from 
near tip of snout to about opposite the axis midAvay between 
nostrils and mouth. 

The color above is plain brown, the ocelli each with one 
small roundish dark brown spot centrally, on two specimens, 
the spot being irregular in shape on one specimen. Below, the 
disc and pelvics are pale, the tail with faint brownish blotches 
as on the holotype. The distance between centers of ocelli is 
about 0.9 times the distance to centers of orbits. 

We thank Dr. Alwyne Wheeler of the British Museum (Na- 
tural History) for furnishing pertinent data on Regan's speci- 
mens of cyclopJiora and Drs. Paulo de Miranda Ribeiro and 
Alceu Lemos de Castro for checking certain characters on a 
number of cijclophora in the Museu Nacional collection, Rio de 
Janeiro; Harvey R. BuUis, Jr., of the U. S. Fish and Wildlife 
Service for the Guiana specimen of cervigoni; and Dr. Fernando 
Cervigoii of the Estacion de Investigaciones Marinas de Marga- 
rita for the opportunity to describe the Venezuela specimens 
and for whom we are pleased to name this species. 

EErEEENCES 

Chandler, Asa C. 

1921. A new species of ray fruni the Texas eoast, and report of the 
occurrence of a to]) minnow new to the fauna of eastern Texas. 
Proc. U.S. Nat. Mus., vol. .19, ]>]k G57-658. 
Garman, Samuel 

1881. Eeports on tlic results of <lredging under tlie supervision of 
Alexander Agassiz, along the Atlantic Coast of the United 
States. Eeport on the selachians. J^>ull. .Mus. Comp. Zool., vol. 8, 
no. 11, pp. 231-2.37. 
Regan, C. T. 

1903. On a collection of fishes made li\' Dr. (ioeldi at Eio .Janeiro. Pioc. 
Zool. Soc. Lomlon, jiai-t 2, pj). n9-68, 2 pis. 



1964 



NEW SPECIES OF SKATE 



5 




Haja cenigoni sp. nov., holotype, immature male, 357 mm long. 



i^MJ 



BREVIORA 

Mmseiuiini of Compsirsitive Zoology 



Cambridge, Mass. November 12, 1964 Number 210 

THE ANTS OF THE FLORIDA KEYS 

By Edward 0. Wilson 
Biological Laboratories, Harvard University 

The Florida keys should be of more than usual interest to 
zoogeographers on several grounds. They are an extensive sub- 
tropical archipelago adjacent to the United States mainland, 
easily reached by a main highway that runs for almost their entire 
length. They are in the hurricane belt, providing an opportunity 
to study the effects of severe periodic storms on population 
dynamics and dispersal. Finally, they are separated by the 
Florida current of the Gulf Stream — a narrow but formidable 
faunistic barrier — from the topographically similar Bimini 
Islands, with which they can profitably be compared. 

These advantages induced me to make a special collecting 
trip in June, 1958, to conduct an initial faunal survey. Time 
permitted only several islands to be investigated thoroughly. 
I selected Key Largo, Plantation Key, Big Pine Key, and Key 
West in order to insure the maximum geographic spread and 
diversity of habitats (see Davis, 1943, and Duellman and 
Schwarz, 1958). The ant fauna of the Bimini Islands had al- 
ready been surveyed by M. R. Smith (1954), while other studies 
of Floridian ants were available in the publications of Wheeler 
(1932) and Van Pelt (1956, 1958). 

The project was supported in part by a grant from the Na- 
tional Science Foundation. I am grateful to my wife Irene for 
her assistance in the collection of specimens and recording of 
ecological information, 

ZOOGEOGRAPHIC ANALYSIS 

In Table 1 the Keys fauna is classified according to nest site 
and probable origin. From a consideration of this partition plus 



2 BREVIORA No. 210 

other data given in the Systematic List later, the following 
generalizations are permissible. 

1. Elements introduced by human commerce are numerically 
more prominent in both species and individuals than on the 
Florida mainland. Eight of the 30 species recorded, or 26.7 per 
cent, are in this category. In the AVelaka Reserve in central 
Florida, according to published data by Van Pelt (1958), 8 of 
75 species, or only 10.7 per cent are in this category. The prev- 
alence of introduced "tramp" species is shared with most 
other small islands in the tropics, including Bimini, hence is 
a distinctly "insular" trait. (Of 12 introduced species occurring 
jointly in the Keys and Welaka Reserve, onlj^ four are held in 
common : Cardiocondyla emeryi, Tetramorium guineense, Mono- 
morium floricola, and Paratrechina longicornis.) 

2. The Antillean elements are mostly arboricolous, whereas 
the Floridian elements are mostly terricolous. 

3. Most of the arboricolous (hence Antillean) elements are 
very well adapted to life in the mangroA'e swamps, maintaining 
unusually dense populations there. The same species also oc- 
cur, for the most part, in other habitats in the interiors of 
the Keys. 

4. The arboricolous elements make up a disproportionate 
part of the Keys fauna. Twelve of the 30 species, or 40 per cent, 
are in this ecological category ; on the other hand, only 12 of 
75, or 16 per cent, are arboricolous in the Welaka Reserve of 
Central Florida. Judging from data by Smith (1954), the 
Bimini fauna resembles that of the Florida Keys in this respect ; 
of 23 species for which there is ecological information, 8 or 
34.8 per cent are arboricolous. 

Generalizations (2), (3), and (4) are probably interrelated. 
The mangrove swamps provide an excellent portal into such 
small islands as the Florida Keys. They form an extensive, per- 
sistent habitat around most of the margins of the islands. Dur- 
ing severe storms, such as the famous hurricanes of 1935 and 
1960 (Craighead and Gilbert, 1962), the mangroves were badly 
damaged but not exterminated. In 1960 many tree branches 
were torn loose and undoubtedly transported long distances by 
sustained winds of 140 mph over a period of nearly 36 hours. 
Such debris, unless submerged in water or literally torn to 
shreds, should provide vehicles of transport for intact colonies 



1964 ANTS OF THE FLORIDA KEYS 3 

of the arboricolous ant species found in the Keys, although the 
point has not been confirmed. The rapid regrowth of the man- 
grove swamps would provide the means for the reproduction of 
these propagules. 

The terricolous ant species, on the other hand, are not so well 
favored. Subterranean and log-dwelling colonies are not likely 
to be transported intact by the high winds. At the same time 
they are much more susceptible to mortality by drowning. In 
the hurricanes of 1935 and 1960, tides of 11 to 18 feet were 
reported. The effects on the ground ant fauna were not studied 
but must have been extensive. 

In sum, it is reasonable to postulate a higher immigration rate 
and lower extinction rate for arboricolous species in comparison 
with terricolous species in the Florida Keys. Whether or not 
the fauna is in equilibrium (see MacArthur and Wilson, 1963), 
it follows that it should consist disproportionately of arboricolous 
species. Hence generalization (4) of the present study seems to 
have a reasonable explanation in one aspect of the physical 
environment. But further field studies before and after tropical 
storms are clearly needed to illuminate this interesting situation. 

The faunas of the Florida Keys and Bimini Islands show 
some striking differences. There are, to begin with, the set of in- 
ferred ecological vicars listed in Table 2. In at least one case, 
Solenopsis geminata versus Pheidole megacephala, the two species 
are known to compete directly and replace each other on very 
small islands in the West Indies and Pacific. Whether this is 
also the case for the other pairs might be determined by ex- 
perimental introductions. According to the hypothesis, such 
introductions of one vicar into the territory of the other should 
result in few successful colonizations. 

Certain species appear to have no ecological equivalents. For 
the Bimini group this is true of the Antillean and endemic 
elements Macromischa pastinifera, Smithistruma nigrescens, 
and Brachymyrmex ohscurior. For the Florida Keys it is 
evidently true of the species Aphaenogaster miamiana and 
Xenomyrmex floridanus. Again, experimental introductions, 
conducted for comparative purposes with exchanges of the 
hypothesized ecological equivalents listed above, might prove 
instructive. According to the hypothesis, this second class of 
introductions should prove relatively successful. 



4 BREVIORA No. 210 

SYSTEIVIATIC LIST 
Platythyrea punctata (Fr. Smith) 

Center of Key Largo, winged queen at light, June 14. Ranges 
from northern South America to southern Texas, also through- 
out the West Indies to the Bahamas and southern Florida. 

Odontomachus ruginodis AVheeler 

Key Largo, Big Pine Key. 

According to W. L. Brown {in litt.) ruginodis is the correct 
name for the Floridian species, whereas insularis is correctly 
applied to the species referred to by Smith (1945) as insularis 
var. pallens Wheeler. 0. ruginodis ranges from the United States 
through the West Indies to South America as far south as 
Paraguay. Within the United States it is found throughout 
Florida and reaches extreme southern Georgia and southeastern 
Alabama. An isolated population, probably introduced, occurs 
within the city limits of Mobile, Alabama. 

This large ponerine, distinguished by its trap-like mandibles, 
was very abundant in undisturbed tropical hammocks in the 
center of Key Largo. Colonies were found nesting in cavities in 
the rather thick leaf litter. On Big Pine Key two colonies were 
found nesting beneath logs in open pine-palmetto woods. 

PSEUDOMYRMEX ELONGATUS Mayr 

Key Largo, Plantation Key, Big Pine Key, Key West (winged 
males in nest June 21). 

Ranges from Brazil north through the West Indies to the 
Bahamas and southern Florida. 

This typically tropical species was abundant on trees in open 
habitats, including mangroA^e swamps, lawns, and disturbed 
hammocks on Plantation Key. On Key Largo it did not pene- 
trate the denser hammocks in the center of the island. Colonies 
were found nesting in hard dead twigs attached to living trees. 

PsEUDOMYRMEX PALLiDUS Fr. Smith 

Big Pine Key. 

A species adapted primarily to savannas, P. pallidus ranges 
around the Caribbean from the West Indies through the coastal 
plain of the Gulf States into Mexico. 



1964 ANTS OF THE FLORIDA KEYS 5 

On Big Pine Key, F. pallidus occurred in both the mangrove 
swamps and on grass in the open pine-palmetto woods of the 
interior. Colonies were found nesting in dead hollow twigs at- 
tached to live mangrove trees. 

Cardiocondyla emeryi Forel 

Plantation Key, Key West. 

A pantropical "tramp" species that originated in Africa and 
has been introduced in the New World, including the warmest 
parts of the Gulf States, by human commerce. 

Workers were found foraging on hot, bare ground in open 
habitats during the day. The species is generally limited to the 
most disturbed habitats. 

Crematog ASTER ASHMEADi Mayr 

Key Largo (a series in Museum of Comparative Zoology col- 
lected in 1904), Plantation Key, Big Pine Key, Key West. 

Limited to the United States, from the Florida Keys north 
to North Carolina and southern Tennessee and w^est to Texas. 

The workers occur in trees in mangrove swamps, lawns, and 
disturbed hardwood forest. None could be found in the deep 
hammocks of Key Largo. Nests were found in dead, dry branches 
of living trees and (in one instance) under loose bark on the 
trunk of a small tree. 

Pheidole floridana Emery 

Big Pine Key (males in nest June 19-20). 

The above specimens compare well with floridana syntypes and 
are closer to other Florida material placed with this species than 
to the closely related (and possibly conspecific) flavens Roger 
of Cuba and the Bahamas. Both floridana and flavens are highly 
variable but can be distinguished by apparently consistent sculp- 
tural characters in the soldier. Floridana is known only from 
Florida, and the Florida mainland is therefore the inferred 
source of the Keys population. 

This small yellow species was abundant in open pine-palmetto 
woods, nesting in the soil beneath coral rock and rotting logs. 
Each colony appeared to contain between 100 and 200 workers. 



6 BREVIOBA No. 210 

Aphaenogaster miamiana Wheeler 

Key Largo (North). 

This native species, a typically Nearctic element, ranges over 
all or most of Florida, just reaching extreme southeastern 
Alabama. 

A single colony, containing between 30 and 50 workers, was 
found nesting in a rotting log at the edge of a clearing in a 
dense tropical hammock. 

Tetramorium guineense (Fabricius) 

Key Largo (Center). 

This species originated in Africa and has been spread by 
human commerce throughout the tropics of both hemispheres. 
It is a characteristic element of disturbed habitats everywhere, 
even in the smallest, most remote oceanic islands. 

Workers were found foraging at sunset on the trunk of a 
guava tree and fronds of a small royal palm in a lawn in the 
center of Key Largo. 

Tetramorium simillimum (Fr. Smith) 

Key West. 

Like T. guineense, this little species evidently originated in 
Africa and has been spread by commerce throughout the tropics. 
Whereas guineense is predominantly arboreal, simillimum is 
mostly or entirely terrestrial. 

Paracryptocerus (Cyathomyrmex) varians (Fr. Smith) 

Key Largo (North), Plantation Key (winged queens June 14), 
Big Pine Key, Key West. 

According to W. W. Kempf (1958), varians occurs in southern 
Florida, south of Miami, in the Bahamas (Andros, New Provi- 
dence, Bimini), and in Cuba and Jamaica. Previously published 
records of the species from Honduras, Trinidad, and northern 
South America were based on misdetermined specimens of P. 
pallens (Klug). 

The species is abundant in the Florida Keys in a variety of 
major habitats. The workers are exclusively arboreal and noc- 
turnal. Colonies were found nesting in hard, dead branches, 
one to two inches in diameter, attached to living trees of several 
species, including gumbo limbo, cocoplum, and mangrove. A 



1964 ANTS OF THE FLORIDA KEYS 7 

dealate queen, evidently in the act of nest-founding, Mas found 
in a dead branch on Plantation Key June 15. She was blocking 
the opening of a small cavity with her saucer-shaped head. 

Strumigenys GUNDLACHi (Rogcr) 

Key Largo. 

This characteristic Neotropical species also occurs in tropical 
Mexico, Central America, Trinidad, and the Greater Antilles. 
Brown (1959) records it from the Everglades National Park in 
Florida. 

Two specimens, a worker and a dealate queen, were collected 
in leaf litter in a dense, relatively undisturbed hammock in 
northern Key Largo. 

Xenomyrmex floridanus Wheeler 

Key Largo (North), Plantation Key (winged males June 14, 
winged males and queens June 16), Key West (males June 21). 

According to the recent revision of the genus by Creighton 
(1957), X. floridanus occurs in southern and central Florida, 
the Bahamas, Cuba, and Mexico. 

Colonies were found in abundance along the edge of forest 
and in mangrove swamps everywhere I collected. They nested 
exclusively in dead branches of trees. AVorkers were found 
foraging singly and in files during the day. They are apparently 
exclusively arboreal. 

MoNOMORiUM destructor (Jcrdou) 

Key Largo, Plantation Key. 

This aggressive little species originated in the Old World 
tropics, possibly Asia, and has been spread by human commerce 
throughout the tropics. 

Workers were abundant in lawns; a single colony was found 
nesting in the soil of a grassy roadstrip in North Key Largo. 

MoNOMORiUM FLORicoLA (Jcrdon) 

Key Largo (North), Plantation Key, Key West. 

M. fioricola is a pantropical tramp species that originated 
somewhere in the Old World tropics. 

Several colonies were located in dead branches of standing 
trees at the edge of hammocks. 



8 BREVIORA No. 210 

MoNOMORiUM PHARAONis (Linne) 

Plantation Key. 

This remarkable species is perhaps the ant most intimately 
associated with man. Originating in Africa, it has been spread 
by commerce throughout the world. In the tropics it nests out- 
of-doors in disturbed habitats, while in temperate zones it is 
abundant in greenhouses and dwellings. On Plantation Key it 
was discovered in both situations. Colonies were abundant in 
dead tree branches in disturbed native w^oods, and workers were 
foraging in the walls of a restaurant far from any native woods. 

SoLENOPSis (S.) GEMiNATA (Fabricius) 

Big Pine Key. 

This species, commonly referred to as the native fire ant, is 
found throughout the New World tropics, ranging northward 
well into the Gulf States. A reddish color phase, to which the 
Big Pine Key series belongs, occurs through much of this range 
and in addition has been carried by human commerce to many 
parts of the Old World tropics. It is especially successful in 
open habitats. 

On Big Pine Key workers were found foraging over the 
crushed coral surface of a parking lot. 

SoLENOPSis (Euophthalma) globularia (Fr. Smith) 

Plantation Key, Big Pine Key (winged queens in nest 
June 19-20) . 

aS^. globularia ranges from the extreme south of Alabama and 
Mississippi through Florida and the West Indies to Mexico, 
Central America, Brazil, and the Galapagoes (Creighton, 1930). 
Through much of this range it is limited to the coast. In colora- 
tion and propodeal sculpturing the Keys samples are closer to 
series from Florida (subsp. littoralis Creighton) than to series 
from Cuba, Haiti, and Puerto Rico. Hence the Keys population 
can be inferred to have originated from Florida. 

Near the center of Big Pine Key a single colony was found 
nesting in soil under a piece of coralline rock in pine-palmetto 
woods. 

SOLENOPSIS (Diplorhoptrum) longiceps M. R. Smith 

Key Largo (North). 

Workers collected at the above locality correspond well with 



1964 ANTS OF THE FLORIDA KEYS 9 

loncficeps paratypes in body proportions, pilosity, and color but 
are somewhat smaller in size. Longiceps ranges from Florida to 
Texas north to Tennessee (Creighton, 1950). 

A single colony was discovered nesting in a small cavity in 
firm leaf litter on the floor of an undisturbed tropical hammock. 

SoLENOPSis (Diplorhoptrum) picta Emery 

Plantation Key (winged queens and males in nest June 15, 
1958) ; Lower Matecumbe Key (collection by W. M. Wheeler, 
1930). 

The species is known from extreme southern Alabama through 
Florida to the West Indies and Central America. 

The Plantation Key colony was found nesting in a dead stem 
in disturbed native woods. 8. picta is characteristically an ar- 
boreal species. 

Cyphomyrmex minutus Mayr 

Key Largo (North) (winged queens and males in nest June 
16) ; Plantation Key. 

This common, primitive little fungus-grower ranges from ex- 
treme southern Alabama through Florida to the Bahamas, 
Greater Antilles, tropical Mexico, Trinidad, and South America 
as far south as Manaos. 

On Key Largo a colony was discovered in a small, crumbling 
log in a relatively undisturbed tropical hammock. On Plantation 
Key one colony was nesting in the soil beneath a piece of coral- 
line rock. Another was in rotting wood in a tree hole about 
three feet from the ground. Both were in disturbed, open na- 
tive woods. 

Tapinoma littorale Wheeler 

Key Largo (North), Plantation Key (males in nest June 15), 
Big Pine Key (males June 19-20), Windley Key (queens and 
males at light June 17). 

The species is distributed from extreme southern Florida to 
the Bahamas, Cuba, and Puerto Rico. 

This was one of the most abundant ant species in leaf litter 
on the floor of undisturbed tropical hammocks on Key Largo. 
Colonies were found nesting in small pieces of rotting wood 
buried in the litter. Colonies were also abundant in dead stems 
attached to living bushes and trees, especially in disturbed forest 



10 BREVIORA No. 210 

on Plantation Key and the mangrove swamp around Big Pine 
Key. Each colony appeared to consist of 100 to 300 workers 
and, in at least one case, up to several nest queens. 

CONOMYRMAPYRAMICA (Roger) 

Big Pine Key, Key West. 

This distinctive, terrestrial dolichoderine ranges from New 
Jersey to Florida west to Arizona and southward through the 
West Indies to Mexico, Central America, western South America 
and the Galapagoes. 

Colonies were found in open soil in elevated spots in the 
mangrove swamps. 

Paratrechina (P.) LONGicoRNis (Latreillc) 

Key Largo, Plantation Key. 

This species ranks with Tetramorium guineense as one of the 
most abundant and ubiquitous of all the pantropical "tramp" 
species of ants. It is characteristic of open, dry, highly dis- 
turbed habitats, from farmland to the centers of the largest 
cities. It originated somewhere in the Old World tropics. 

On the Keys listed above, P. longicornis was common in com- 
pletely open situations, especially around human dwellings. It 
was not found in the native woods. During a field trip to eastern 
Cuba in 1953, I noticed a similar distribution. P. longicornis 
workers abounded through sugar cane fields and along roads 
up to the very edge of the native forests that clung to limestone 
outcrop])ings ; inside the forests, over a distance of only a few 
feet, they were replaced by native Cuban ant species. 

Paratrechina (Nylanderia) bourbonica Forel 

Key West. 

This is another prominent pantropical "tramp" species of 
Old World origin. The Keys specimens have been compared 
with a syntype worker in the collection of the Museum of Com- 
parative Zoology. 

Workers were commonly seen foraging during the day at 
several locations on the streets of residential sections of 
Key West. 

Paratrechina (Nylanderia) parvula (Mayr) 

Key Largo (North). 

The workers of species in the parvula complex are too similar 



1964 ANTS OP THE FLORIDA KEYS 11 

to make tlie present identification more than tentative. P. par- 
vula, according to Creigliton (1950), ranges from soutliern New 
York west to Iowa and Texas and south to Florida. 

A single colony was found nesting in a small piece of rotting 
wood buried in leaf litter on the floor of a relatively undisturbed 
tropical hainmoek. 

Camponotus abdominalis ploridanus (Buckley) 

Key Largo, Plantation Key, Big Pine Key. 

C. floridanus is such a well-defined form with reference to the 
remainder of the abdominalis complex that it may well be a 
distinct biological species. I have used the trinomen in this case 
as an indication of the uncertainty of its biological status. 
Floridanus ranges from the east shore of Mobile Bay, Alabama, 
west through southern Georgia and south to the Florida Keys. 
The remainder of the abdominalis forms, which may or may not 
constitute a single species, range from southern Texas to Ecua- 
dor, the Amazon basin, and thence north again to the Lesser 
Antilles. The species is evidently absent from the Greater 
Antilles. 

Floridanus is one of the most abundant and adaptable ant 
species in the Keys. It occurs in undisturbed and disturbed 
native hammocks and on the lawns around human dwellings but 
is evidently absent from mangrove swamps. Nests are formed 
in leaf litter, under rotting logs, and in dead branches of living 
trees. Workers forage during the day, mostly on vegetation. 

Camponotus planatus Roger 

Plantation Key (winged queens and males in nest June 14-18), 
Key West. 

The species is found in extreme southern Florida and Texas, 
Cuba, Mexico, and Central America. 

C. planatus is a very abundant species in tropical hammocks 
but rare or absent elsewhere. On Big Pine Key one colony was 
found in a small, isolated hammock, but not a single specimen 
was collected in the extensive pine-palmetto woods that dominate 
the unsettled part of the island. Unlike floridanus, planatus is 
absent from the vicinity of human dwellings. Colonies were 
found nesting in dead tree branches, both attached to living trees 
and lying loose on the ground. The alert, swift-running workers 
forage during the day. 



12 BREVIORA No. 210 

Camponotus tortuganus Emery 

Plantation Key (winged queen June 14), Big Pine Key. 

As Creigliton (1950) points out, the status of this form, 
originally described as a subspecies of the widespread tropical 
species C. maculatus, will remain unsettled until a careful revi- 
sion can be made of the difficult species group to which it be- 
longs. Tortuganus was originally described from the Dry Tor- 
tugas. As currently delimited it occurs on the Keys and in the 
Florida mainland as far north as Lake Worth. 

A single colony was found beneath a rotting log in pine- 
palmetto woods on Big Pine Key. 

Camponotus (Colobopsis) sp. 

Big Pine Key. 

A single colony belonging to the subgenus Colohopsis was 
found nesting in a dead stem in a mangrove swamp on the south 
shore of Big Pine Key. It is similar to C. impressus (Roger) 
but differs sufficiently in the shape of the propodeum of the 
minor worker to warrant distinguishing it as a different, and 
possibly undescribed species. Additional materials considered in 
the context of a generic revision are needed to clarify the matter. 



SUMMARY 

Thirty ant species are listed from the Florida Keys. The 
disproportionate fraction of introduced species is cited as a 
typically insular feature of the fauna. Also, an exceptional 
prevalence of arboricolous (and Antillean) species is noted and 
is hypothesized to be the outcome of the periodic severe tropical 
storms that alter the Keys environment. Finally, comparisons 
are made with the nearby Bimini Islands fauna. 



1964 ANTS OF THE FLORIDA KEYS 13 

Table I 
Ecology and Zoogeographic Origin of the Keys Species 

Terricolous 

FbORiDiAN: Aphaenogaster miamiana, Phcidole floridana, Solenopsis 
glohidaria, S. longiceps, Paratredhina parvula. 

Antillban: Odontomachus ruginodis, Strumigenys gundlachi, Cyphomyr- 
mex minutus. 

Natives of uncertain origin : Solenopsis geminata, Conomyrma pyra- 
mica, Camponotus tortuganus. 

Introduced by man: Cordiocondyla emeryi, Tetramorium simillimum, 
Paratrechina longicornis, P. hourhonica. 

Arboricolous 

Floridian : Crematogaster ashmeadi. 

Antillean: Platythyrea punctata, Pseudomyrmex elongatus, Xenomyr- 
mex floridanns, Solenopsis picta, Paracryptocerus varians, Tapinoma lit- 
torale. 

Natives of uncertain origin: Pseudomyrmex pallidus, Camponotus 
(Colobopsis) sp. 

Introduced by man: Tetramorium guineense, Monomorium floricola, M. 
pharaonis. 

Both Terricolous and Arboricolous 
Floridian : Camponotus abdominalis floridanus. 
Antillean : Camponotus planatus. 
Introduced by man : Monomorium destructor. 



Table II 
Ecological Equivalents (with Zoogeographic Origins) 

Florida Keys Bimini Islands 

Odontomachus ruginodis Odontomachus insularis 

Pheidole floridana (Floridian) Pheidole flavens (Antillean) 

Solenopsis geminata (Neotropical) Pheidole megacephala (introduced) 
Crematogaster ashmeadi (Floridian) Crematogaster steinheili (Antillean) 
Cyphomyrmex minutus (Antillean) Trachymyrmex jamaicensis (Antillean) 
Camponotus bermudezi (Antillean) ^ (Camponotus floridanus (Antillean) 
C. lucayanus (endemic) U C. tortuganus (Antillean?) 

C. ramulorum (Antillean) J [c. planatus (Antillean) 

C. (Colobopsis) (mlmicola (endemic?) C. (Colobopsis) sp. (endemic?) 



14 BREVIORA No. 210 

EEFEEENCES 

Brown, W. L. 

1959. The Neotropical species of the ant genus Strumigenys Fr. Smith: 
group of gundlachi (Roger). Psyche, 66: 37-52. 

Craighead, F. C. and V. C. Gilbert 

1962. The effects of hurricane Donna on the vegetation of southern 
Florida. Quart. J. Florida Acad. Sci., 25 : 1-28. 

CREIGHTOlSr, W. S. 

1930. The New World species of the genus Solenopsis (Hymenop. 

Formieidae). Proc. Amer. Acad. Arts Sci., 66: 39-151. 
1950. The ants of North America. Bull. Mus. Comp. Zool., 104 : 1-585. 

1957. A studj' of the genus Xenomyrmex (Hymenoptera, Formieidae). 
Amer. Mus. Novit., no. 1843, 14 pp. 

Davis, J. H. 

1943. The natural features of southern Florida, especially the vegeta- 
tion, and the Everglades. Bull. Florida Geol. Sur., (Dept. 
Conservation), 25: 1-311. 

DuELLMAN, W. E. and A. Schwartz 

1958. Amphibians and reptiles of southern Florida. Bull. Florida 
State Mus., Biol. Sci., 3 : 181-324. 

Kempf, W. W. 

1958. New studies on the ant tribe Cephalotini (Hym. Formieidae). 
Studia Ent., 1 : 1-68. 

MacArthur, R. H. and E. O. Wilson 

1963. An equilibrium theory of insular zoogeography. Evolution, 17: 
373-387. 

Smith, M. R. 

1954. Ants of the Bimini Island Group, Bahamas, British West Indies 
(Hjnnenoptera, Formieidae). Amer. Mus. Novit., no. 1671, 
16 pp. 

Van Pelt, A. F. 

1956. The ecology of the ants of the Welaka Reserve, Florida (Hy- 
menoptera: Formieidae). Amer. Midi. Nat., 56: 358-387. 

1958. The ecology of the ants of the Welaka Reserve, Florida (Hy- 
menoptera: Formieidae). Part II. Annotated list. Amer. Midi. 
Nat.,59: 1-57. 

Wheeler, W. M. 

1932. A list of the ants of Florida with descriptions of new forms. 
J. New York Ent. Soc, 40: 1-17. 



BREVIORA 

Miasenainii of Comparsitive Zoology 

Cambridge, Mass. December 21, 1964 Number 211 

A NEW SPECIES OF THE SNAKE LEPTOTYPHLOPS 

FROM COLOMBIA 

By Benjamin Shreve 



In a shipment of snakes sent to me for identificaton by 
Hermano Niceforo Maria of the Instituto de La Salle of Bogota, 
Colombia, one appears to represent an undescribed species. An- 
other specimen, in the Museum of Comparative Zoology collec- 
tion, is apparently of the same species. 

Thanks are due the following individuals for lending compara- 
tive material from the collections in their charge : Mr. Charles 
M. Bogert of the American Museum of Natural Histoiy 
(AMNH), Dr. Doris M. Cochran of the United States National 
Museum (USNM), Miss Alice G. C. Grandison of the British 
Museum (Natural History) (BMNH), and Dr. William Duell- 
man of the University of Kansas Museum of Natural History 
(UKMNH). 

The new snake will be given a name suggested by the exceed- 
ingly low dorsal scale count. 

Leptotyphlops brevissima sp. nov. 

Holotype. Museo del Instituto de La Salle No. 1311, from 
Florencia, Caqueta, Colombia, collected by Niceforo Maria Feb- 
ruary 10, 1951. 

Paratype. MCZ No. 38950 (taken from the stomach of 
Micniriis tnipartitus (Dumeril, Dumeril and Bibron), MCZ No. 
21988, from Sonson, Antioquia, Colombia, collected by Niceforo 
Maria in 1925. This specimen has about a third of the head miss- 
ing on the right side. 

Diagnosis. Apparently most closely related to Leptotyphlops 
macrolepis (Peters) and L. anthracina Bailey, from which it 
differs in a lower dorsal and subcaudal count (Table 1), and in 



2 BREVIORA No. 211 

coloration (brown rather than black as in anfhracina and with- 
out the prominent white borders of the ventral scales of 
macrolepis) . Also related to L. dugandi Dunn to which it is 
nearest in longitudinal dorsal count (but without overlap, 
Table 1). L. dugandi, however, has still fewer subeaudals and a 
very different coloration : a whitish unmarked venter, a striped 
dorsal pattern, and a whitish fore part of the head. 

Description. (Paratype variation in parentheses.) Snout 
rounded ; supraocular present, small ; rostral extending nearly 
to level of eyes ; nasal completely divided into two ; ocular border- 
ing the lip ; three labials, two in front of, and one behind the 
ocular ; second labial not reaching the eye ; six lower labials ; 
14 scales around body at midbody ; dorsal scales from rostral to 
tail spine 152 (164) ; subeaudals is (14). 

Coloration in alcohol. Above, dark brown ; below, light brown ; 
scales, both above and below, narrowly bordered with whitish. 
The coloration resembles quite closely that of L. goudotii 
(Dumeril and Bibron). 

Measurements (in millimeters) 





head and body 


tail 


holotype 


63 


5 


paratype 


127 


11 



Remarks. The paratype was formerly identified as L. tnacro- 
lepis by the late Karl P. Schmidt, who found it during his work 
on the coral snakes. It may be that other specimens of hrevissima 
have been similarly misidentified ; longitudinal counts have been 
made all too infrequently. 

The dorsal coloration of the two species is quite similar. 

The ventral coloration of macrolepis gives the impression of 
a whitish ground color covered with brown spots, as each ventral 
scale has a brown spot in its center with much of the ground 
color visible around it. In contrast, the ventral coloration of 
hrevissima appears to be a uniform light brown, but on close 
examination it is seen that each scale has a very narrow light 
border. 

The coloration of anthracina appears to be even more similar 
to that of the new form. The pattern of spotting on each scale 
seems identical, but the brown coloration of hrevissima is repre- 
sented by black in anfhracina (or gray in some preserved ani- 
mals). 



1964 NEW SNAKE FROM COLOMBIA 3 

In order to shed more liglit on the relationships, material re- 
ferred to macrolepis was borrowed from various institutions. 
The following macrolepis were examined : V enezuela. Caracas : 
rSNM 62206, 107891. Chama, 200 meters: BMNH 1905.5.31.63. 
El Valle, Federal District: AMNH 59406. Merida, 1600 meters: 
BMNH 1903.4.28.12, 1904.6.30.2-5. Pauji, Acosta District : MCZ 
48752-3, 48918. Puerto Cabello : BMNH 1913.9.10.2. No definite 
locality: BMNH R.R.1964.33. Colombia . In swampy country 
of the Choco: MCZ 39705. New Grenada, Amazonas: AMNH 
17538 (in very poor condition). Brazil. Maues, Amazonas: 
AMNH 91642. British Guiana. Rupununi District, north of 
Acarahy Mts., west of New River : UKMNH 69821. 

E. R. Dunn (1944, p. 52) lists specimens of macrolepis from 
the following localities of Colombia : Norte de Santander : Ocana 
(1200 meters), two; Santander: Barichara (1336 meters), three; 
San Gil (1095 meters), four; Tolima: Mariquita (535 meters), 
one; Guamo (402 meters), one; Chaparral (800 meters), two. 

It can be seen from this list and the material mentioned pre- 
viously that there is no absolute evidence of sj'mpatry between 
the new form and what is regarded as macrolepis. In addition, 
there is no positive evidence that all of Dunn's material was 
properly identified ; he only reports the longitudinal dorsal 
scale count of one example and this is, of course, a critical point 
in separating the species. Unfortunately, all the Colombian ma- 
terial examined by Dunn was consumed by fire some years ago 
at the Museo del Instituto de La Salle. 

The dorsal and subcaudal counts of the specimens of macro- 
lepis examined, and those taken from the literature, are 202-246 
and 16-26, respectively. Those taken from the literature are 
those of J. A. Roze (1952, p. 153), E. R. Dunn (1944, p. 52) 
and J. R. Bailey (1946, pp. 3, 4). It may be that with such a wide 
range of extremes in these counts, more than one form is in- 
volved. At all events, the dorsal scale counts are considerably 
higher than those of hrevissima (152-164). 

Among the specimens received on loan as macrolepis was 
AMNH 35953 from Balzapamba, Ecuador. This has 183 dorsal 
scales from rostral to tail spine, 16 subcaudals, the usual 14 
scales around midbody, and a head and body measurement of 
113 mm, tail 12 mm. In these characters and in coloration the 
specimen matches anthracina, the only member of this complex 
known from Ecuador ; as anthracina, it represents the third known 
locality for this species. 



BREVIORA 



No. 211 



L. anthracina with scale counts (J. R. Bailey, 1946, p. 3) of 
182-189 dorsals and 15-19 subcaudals is closely related indeed to 
hrevissima. It may be that anthracina and hrevissima are only 
subspecifieally distinct. If so, it may be that the very low dorsal 
counts of hrevissima may represent "character displacement" 
in northern anthracina populations, sympatric with macrolepis. 

However, the difference in coloration between anthracina and 
hrevissima or macrolepis is not readily explainable on this hy- 
pothesis. It would be expected that anthracina, outside the 
range of macrolepis, would be as similar in coloration as it is in 
scales to macrolepis, instead of exhibiting a coloration which is 
more different from that of rnacrolepis than is that of hrevissima. 

KEFEEENCES CITED 

Bailey, Joseph K. 

1946. LeptotypMops anthracinus, a new blind snake from eastern 
Ecuador. Occ. Pap. Mus. Zool. Univ. Mich., No. 492 : 1-5. 
Dunn, Emmett Eeid 

1944. A review of the Colombian snakes of the families Typhlopidae 
and Leptotyphlopidae. Caldasia, 3 (11): 47-55. 
EozE, Janis a. 

1952. Contribucion al conocimiento de los ofidios de las familias 
Typhlopidae y Leptotyphlopidae en Venezuela. Mem. Soc. Cien. 
Nat. La Salle, 12 (32) : 143-158. 



head color: 



TABLE 1 
hrevissima anthracina macrolepis dugandi 



dorsal longitudinal 










scale count: 


152,164 


182-189 


202-246 


172-184 


subcaudal scale count : 


13,14 


15-19 


16-26 


9-12 


dorsal color: 


dark brown 


lilack 


brown 


striped 


ventral color: 


light brown 


Idack 


spotted 
(scales witli 
wide white 
borders) 


whitish 



dark brown 



black 



brown 



whitish 



BREVIORA 

MiaseMm of Comparative Zoology 

Cambridge, Mass. Dkckmbkk ."^0, 19G4 Xumbek 212 

A NEW SPECIES OP^ FRESHWATER GASTROPOD 

MOLLUSC OF THE GENUS SAULEA FROM THE 

MIOCENE OF KENYA 

Bv Thomas Pain^ and Dawn Beatty 



The genus Saulca Gray 1867 (family AmpuUariidae) has a 
thill, wholly corneous operculum, as found in the American 
genus Pomacca, and in this respect differs widely from the other 
African x\mpullariidae, all of which have a calcareous oper- 
culum. The corneous opercula are unfortunately not known in 
a fossil state. 

The only recent species of the genus Saiiha is *S'. vitrea 
{Born)=HeIi.r vitrea Born (1780, p. 383, pi. 15, figs. 15-16), 
originally described from an unknown locality, but now known 
to occur in the West African coastal regions of Liberia and 
Sierra Leone. The anatomy is at present unknown. 

The possibility of the occurrence of Saiilea in East Africa 
was first suggested by V. E. Fuchs (1936, p. 100) who con- 
sidered a poorly preserved fossil from the Kaiso (Pleistocene) 
beds of Lake Albert as possibly referable to this genus. Adam 
(1957, p. 48), however, pointed out that this strongly carinated 
shell was more probably a Vivipa)-us, perhaps a form of his own 
species V. coxi from the same deposit, and he later (1959, p. 15), 
correctly in our o]5iiiion, listed it in the synonymy of that species. 

In 1962, Dr. B. A'erdcourt very kindly sent to the senior 
author for examination a fossil shell, in the form of an internal 
cast in limestone, from Miocene deposits at Kirimon, east of 
Lake Bariiigo, Kenya, which, in spite of its poor state of preser- 
vation, he had no hesitation in referring to the family Ampul- 
lariidae (Pain, in Verdcourt, 1963, p. 7, tig. 10). This species 
is dextral in form, with a short, conical spire and somewhat 
barrel-shaped body whorl, features typical of the genus Saulea. 

1 47 Keyuolcls House, Millbank, Liunldu SW 1. 



2 BREVIORA No. 212 

It was not, however, sufficiently well preserved to make a 
description of it possible. 

Recently we have, through the kindness of Prof. Bryan Pat- 
terson of the Museum of Comparative Zoology, Harvard Uni- 
versity, received for study two further examples of this fossil 
collected at Kirimon by the Harvard University 1963 Kenya 
Expedition. These are both in a far better state of preserva- 
tion than the original specimen, and have convinced us that all 
represent a so far unknown fossil species of Saulea, which is 
described and figured herein. It is to be hoped that future col- 
lecting at Kirimon will produce further and perhaps even better 
preserved specimens of this unique fossil, which has established 
beyond reasonable doubt the existence of this genus in East 
Africa in Miocene times, and its probably far wider distribution 
during the Tertiary period. 

Saulea Gray 

Sduica Gia.v, 1867, ji. lOllU. Type species l).v moiiutypy: Hrli.r ritrrn 
Born. 

Saulea lithoides n. sp. 

Holotijpc. MCZ 28018, internal cast. 

Pamtypc. MCZ 28017, internal cast. 

Horizon and locality. Late( ?) Miocene, at Kirimon. approxi- 
mately 42 miles east of Lake Baringo and 38 miles SSE of Mara- 
lal, Kenya, East Africa. 

Diagnosis and description. An internal cast in limestone; 
51/4 whorls are preserved, the apex being decollate ; the spire 
short, somewhat convex, the body-whorl oval or barrel-shaped in 
outline and a little inflated at tbe periphery, % the total length ; 
umbilicus very narrow, columella erect, curving slightly toward 
the base. Aperture of moderate width, vertical, oblong. The 
outer lip and base of tlic columella are missing in all the ma- 
terial examined. 

Measurements fin millimetres: approx. only) 





Greatest 






-ciiKttl 


width 






u 


32 


MCZ 28017 


Paratyi)e 


39 


27 


MCZ 28018 


Ilolotype 


34 


:{() 


Coryndmi .Museum 


Topotype 



1!)(U NKW MIOCENE GASTROPOD FROM KENYA 3 

The authors" tlianks arc tluc to l*i'ot'. Bryan Patterson for 
the loan of material for study, to Mr. T. E. Crowley for much 
helpful adviee and to Mr. 1\. F. Cumberland for i)lioto»>rai)hin<i: 
specimens i'ov i-eproduction hei-ein. The collection of the speci- 
mens was made possible under National Science Koundation 
(Jrant CP-llSS. 



REFERENCES 

AiuM, W. 

lit.")?. Exjiloration dn Pare National Albert, ^fission J. de Heinzelin 
di' Braiifourt (1950). Inst. Pares Nat. Congo Beloe, 3; :M7l', 
pis. 1-9. 
19r)9. Molhisqucs Pleistocenes de la region dii Lae AUiert et de la 
Seniliki. Ann. Mus. Roy. Congo Beige, Sci. (Jeol., 25: 1-149, 
pis. 1-10. 
Born, I. 

1780. Testacea Mnsei Caesarei Vindobonensis. Vienna, 442 i)p., 18 j)ls. 
FucHS, V. E. 

1936. Extinct Pleistocene Mollusea from Lake Edward, Uganda, and 
tlieir liearing on the Tanganyika problem. Linn. Soe. J., Zool., 
40 (no. 269): 93-106, pis. 1-3. 
Gray, J. E. 

1867. Description of Sanlea, a new genns of Ani]nillariadae from 
Sierra Leone. Proe. Zool. Soe. London, 1867: 1000-1001. 
Verdcourt, B. 

1963. The ^Miocene non-marine Molhisca of Rnsinga Island, Lake 
Victoria and other localities in Kenya. Palaeontographica, 121 : 
1-37, 64 text-figs. 



BREVIORA 



No. 2] 2 




o 



c; 

a. 



o 






P-^'y;"-;- 



!)(U 



NEW MIOCENE GASTROPOD FIIOM KENYA 





{^ 






S 

c7 



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I— I 

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oo 

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. 


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f^ 2 



BREVIORA 

MiuiseiLMii of Coimparative Zoology 

Cambridge, Mass. .lAxrAUY 11, 1905 Number 21)^ 



A HITIIEKTO OVERLOOKED AXODOXTA 

(MOLLUSCA: UNIONIDAE) 
FROM THE GULF DRAINAGE OF FLORIDA 

Bv RiciiARn I. Johnson 



Li their reports on the inoUnsks of the Gulf drainaoe of 
Florida, Vaiider Schalie and Clench and Turner have included 
under Anodonia imhcciUs Say a number of records which belong 
to the species here described as distinct. 

I take pleasure in naming this species for my wife Peggy, 
who along with our two small daughters and S. L. II. Fuller, 
accompanied me during an extensive survey of Florida for 
Fnionidae during the summer of 1962. 

Anodonta peggyae new species 

Anodonta imhecilUs [sic] Vander Seluilie, H., 1940, Lluydia, 3:lS)(), pi. li, 

fig. 4; non Say 1829. 

Anodonia imhecilis Say 1S129, parliin. Clench, W. J. and Tuiiiur, E. I)., 

1956, Bull. Florida State Alus., 1:187, pi. 6, fig. 2. The figured 

sspeoimen becomes a ])aratypc of Anodonta prjimiae, :\1CZ 251U41. 

Holoixipc: MCZ no. 251040, from the SE shore of Lake 

Talquin [formed by a dam on the Ochlockonee River]. Leon 

County public fishing ground, Leon Co., Florida. 

Paratypes : Figured paratypes, MCZ 251041 ; additional para- 
types, MCZ 191840, all from the same locality as the holotype. 







Measurements 


jength 


Height 


Width 




mm 


mm 


mm 




71 


43 


24 


Holotype 


54 


33 


19 


Paratvpe MCZ 251041 


46 


28 


16 


Paratype MCZ 251041 



2 BREVIORA No. 218 

Description -. Shell small to medium, reaching a little over 80 
mm in length ; length to height ratio approximately 2 to 1. 
Outline subrhomboidal, valves slightly inflated, thin, fragile 
and smooth. Anterior end regularly rounded, posterior end 
more broadly rounded and slightly biangulate just above the 
base. Ventral margin broadly curved and obliquely descending. 
Dorsal margin straight or slightly curved, usually forming a 
distinct wing-like angle where it meets the obliquely descending 
posterior margin. Posterior ridge broadly rounded, posterior 
slojie sometimes slightly concave and located toward the anterior 
third of the shell. T'mbos low and broad, not extending above 
the dorsal margin, their scul])ture consisting of seven or eight 
low, delicate, slightly double-looped undulations. Periostracum 
smooth and shiny, except the posterior slope which may be 
slightly roughened, straw-yellow to yellowish green, sometimes 
very dark green, with numerous, generally fine, green rays 
over the entire surface. The rays are distinctly darker on 
the posterior slope. 

No hinge plate or teeth ; muscle scars inconspicuous and 
poorly defined. Nacre bluish white and iridescent. 

Habitat: Prefers sandy oi- mnddy bottoms of ]")onds and 
sluggish streams. 

Remarks: In the (!ulf drainage, Anodoitta, pajgiiav can be 
confused with ^1. inihecilis Say, in whose company it is often 
found, and with A. couperiaiiii Lea. These three species have 
umbos which do not extend above the dorsal margin, a character 
which distinguishes them from all other Anodonta of the (lulf 
drainage. 

A. imhecilis has an elongate' elliptical shell, with a ])osterior 
point which ends near the medial line, an almost straight 
ventral margin which is parallel to the dorsal one. and a perios- 
tracum which is rather uniforndy green. .1. peggyae differs 
from intlxcilis in that, with the hinge line held horizontal, it 
has a subrhomboidal shell with a less acute point located near the 
base, a ventral margin which sloi)es obliquely from the dorsal 
one, and a periostracum which has numerous green rays that 
are especially fine on the disk. 

A. couperiaria differs from both iitilx ciJis and peggyae in that 
couperiana has a shell which is much higher in ]iroiM)rtion to 
its length than either of the others, with a ])osterior j)oint which 
eiuls near the medial line as in imhecilis: and thougli the ven- 
tral margin is i-ouglily parallel to the dorsal one. it is hi-oadly 



1965 m:\v si'EciEs ok anodonta 3 

ami uiiifonuly curved. When c(nip( yiaixt luis i"iys at all, tlipv 
are usually even finer than those of pcg(j!j(i< . 

From the Choctawhatchee River system to the Oclilockonee 
River system (the southern tei-minus of imbccilis Say) A. 
imhecilis and A. pcggyac are often found together, while in 
peninsula!' Floi-ida .1. pi'iKJUac and .1. coiip( riaiid are geographi- 
eally separate. 

As the figured hoh)type of A)i(>do)ii<i c<)Hp< ruum eould not 
be located in the United States National Museum, the syntype, 
USNM 86673 is here selected as the lectotype and is figured on 
Plate 2, figure 4 of this report. The type of A7iodo7ita imhecilis 
Say, wliieh should be in the Academy of Natural Sciences of 
Philadel]ihia, is lost. 

Distribitfion : Gulf tlrainage of Florida, from the Chocta- 
whatchee River system to the Hillsborough River system. 

SPECIMENS EXAMINED 
CHOCTAWHATCTIEE RIVER SYSTEM 

Choctawhatchee River Drainage. — Alabama: Choctawhat- 
chee River, 8 mi. SW Abbeville, Henry Co. Bushy Point Lake, 
Choctawhatchee River ; Inlet Lakes, Choctawhatchee River ; both 
Walton Co. Horseshore Lake, Choctawhatchee River, Washington 
Co. 

Holmes Creek Drainage. — Florida: Holmes Creek, 1 mi. W 
(iraceville, -Jackson Co. 

APALACHICOLA RIVER SYSTEM 

Chipola River Drainage. — Florida: Spring Creek, Merritts 
Mill, 3 mi. E Marianna, Jackson Co. Chipola River, Scotts Ferr^^ 
Chipola River, Pole Bluff Landing ; Dead Lake of Chipola River, 
Chipola Park; Dead Lake of Chipola River, Chipola Pines, 
Idlewood; all Calhoun Co. 

Apalachicola River 'Drainaqe.- - Florida : Mosquito Creek, 
Chattfdiooclice, (iadsden Co. 

Flint River Drainage. — (icortjia : Flint River, Bainbridge ; 
Four Mile Creek, :] mi. SW Bain.bridge ; Blue Spring, Flint 
River, T'^^ mi. W Recovery; all Decatur Co. Spring Creek, 
2i/> mi. S Revnoldsville, Seminole Co. 



4' BREVIORA Xo. 21;^ 

OCHLOCKONEE KIVER SYSTEM 

Ochlockonee River Drainage. — Florida : Little River. 7 mi. 
SW Havana, Gadsden Co. Oehloi-konee River, 8 mi. \V Talla- 
hassee ; SE shore. Lake Talqiiin ; both Leon Co. Ochlockonee 
River, 71 o mi. E Hosford. Liberty Co. 

WAKT7.LA RIVER SYSTEM 

Florida: Wakidla River, In mi. S Tallahassee, \Vakulla Co. 

ST. MARKS RIVER SYSTEM 

Florida: St. Marks River, Natural Bi-idfre Sink, Leon Co. 

ST^WAXXEE RIVER SYSTEM 

Suwannee River Drainage. — Florida: Suwannee River, Old- 
town, Dixie Co. Suwannee River, Fannin Springs, (lilchrist Co. 

WITHLACOOCHEE RIVER SYSTEM 

Withlacoochee River Drainage. — Florida: Withlaeoochee 
River, 1 mi. N\V Lacooclice, Pasco Co. Little Withlacoochee 
River, Rerdell, Hernando Co. X shore, Lee's Lake, Panasott'kee ; 
Lake, 6 mi. XXW Panasolfkee ; both Sumter Co. SW shore. 
Lake Tsala Apopka, Floral City, Citrus Co. XW shore. Lake 
Tsala Apopka, Hernando Co. 

HILLSBOROT^GH RIVER SYSTEM 

Florida: Blackwater Creek. 8 mi. N Plant City; Hillsborough 
River. 4 mi. XE Tem])h' Ten-ace; IxdJi Hillsboroug'h Co. 



I 



l!)(ir) 



Xi:\V SI'KCIKS OF AXonoNTA 




I'lMtC 1 



The known distiiliution of Annthinta pri/c/yae, n. sp. in tlu' (uilf dijiinage 
of Florida is represented by a (dosed circle. 

The known distribution of Anodonta imbecilis Say in the southeastern 
United States is represented liy a partially filled circle. 

The known distribution of Anodonta coiipeiiana Lea is represented by 
an open circle. 



o 



1-^ 



c; 



BREVIORA No. 218 



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196.3 



XKW SI'KCIKS OK AXOnoXTA 




BREVIORA 

Miiseiim of Comtiparative Zoology 



Cambridge, Mass. February 15, 1965 Number 214 

A REVISED CLASSIFICATION 
OF THE DENDROCHIROTE HOLOTHURIANS 

By David L. Pawson 

U.S. National Museum 

and 

H. Barraclough Fell 
Museum of Comparative Zoology 

Recent studies on some unusual dendrochirote liolothurians 
and examination of available data on fossil liolothurians have led 
us to conclude that the hitherto accepted classification of the 
dendrochirote holothurians conceals some important evolutionary 
trends. In this article a new classification is proposed in which 
tentacle numbers are abandoned as criteria for diagnosing major 
taxa; instead, importance is attached to the shape of the cal- 
careous ring, the shape of the tentacles, and the calcareous de- 
posits of the body wall. The reasons for such changes in classifica- 
tion are given in detail elsewhere (Fell and Pawson, in press). 

NEW TAXA 

Certain genera which were hitherto grouped in the order 
Dendrochirotida are known to lack the richly branched tentacles 
which are so characteristic of other genera within that order. 
These genera with essentially unbranched tentacles also share 
other important features, which suggest a natural grouping of 
three families. We therefore propose a new order to accommodate 
these families. 

Order DACTYLOCHIROTIDA nov. 

Diagnosis: Tentacles 8-30 in number, not branched but digit- 
iform or digitate, the digits sometimes bifurcate. Body enclosed 



2 BEEVIORA No. 214 

in a test comprising imbricate plates. Calcareous ring simple, 
lacking complex posterior processes. 

Included families: Ypsilothuriidae Heding, 1942; Rhopalod- 
inidae Perrier, 1902 ; and a third family, diagnosed here as new : 

Family VANEYELLIDAE nov. 

Diagnosis: Anus and mouth at opposite ends of the body, which 
is U-shaped or fusiform. Plates in the body wall with or without 
small spires. Tentacles 10-20 in number. 

Included extant genera : Vaneyella Heding and Panning, 1954 
(type genus by tautonymy) ; Mitsukuriella Heding and Panning, 
1954. 

The order Dendrochirotida as now restricted may be diagnosed 
as follows : 

Order DENDROCHIROTIDA Grube, 1840 
(restricted herein) 

Diagnosis: Tentacles 10-30 in number, richly branched. Cal- 
careous ring simple or with complex posterior processes. Test 
sometimes well developed, or reduced to non-contiguous calcare- 
ous spicules. 

Morphological and paleontological evidence (Fell and Pawson, 
in press) suggests that this order includes some very primitive 
extant holothurian genera. These genera seem best assigned to 
new families ; one new genus and two new families are diagnosed 
here : 

Family PLACOTHURIIDAE nov. 

Diagnosis: Body completely enclosed in a test of imbricate 
plates without spires. Sole lacking. Calcareous ring long, slender, 
with long posterior processes. 

Included extant genus : Placothitria n.g. 

Placothuria n.g. 

Diagnosis: Tentacles 10, richly branched. Body U-shaped. Cal- 
careous ring long, with posterior processes composed of a mosaic 
of small pieces. 

Type-species: Stolus Juittoni (Dendy) (illustrated in Pawson, 
1963, pi. 7). 



1965 NEW HOLOTHURIAN CLASSIFICATION 3 

Etymology: The generic name is derived from the Greek plax 
(a plate) and Holothuria, and refers to the test formed by over- 
lapping plates. Gender, feminine. 

A second new family within the order Dendrochirotida is pro- 
posed : 

Family PARACUCUMIDAE nov. 

Diagnosis: Body completely invested in a test of imbricate 
scales with spires. Sole lacking. Calcareous ring simple, lacking 
posterior processes. 

Included extant genus : Paracuciimis Mortensen, 1925. 

A REVISED CLASSIFICATION 

The relationships of the orders Dendrochirotida and Dactylo- 
chirotida are discussed elsewhere, in the broad context of the 
inferred evolution of the Holothuroidea (Fell and Pawson, in 
press). From the new data the following revised classification of 
holothurians emerges : 

Subclass DENDROCHIROTACEA Grube, 1840 

(Nomen translatum herein, ex Dendrochirotae Grube, 1840) 
Introvert (with retractor muscles) always present. Tubefeet 
and respiratory trees usually present. Madreporite free in the 
body cavity. Mesentery of the posterior loop of the intestine in 
the right or left ventral interradius. Free tentacle ampullae lack- 
ing. Gonad in two tufts, one tuft to each side of the dorsal mesen- 
tery. 

Order DENDROCHIROTIDA Grube, 1840 

(restricted herein) 

Tentacles richly branched, 10-30 in number. 

Key to Included Families 

1 (6) Body partly or completely invested by plates. 

2 (5) Body enclosed by a test comprising conspicuous imbricate plates; 

sole lacking, 

3 (4) Calcareous ring complex, with long paired posterior processes. 

Placothuriidae nov. (Included ex- 
tant genus: Placoihuria nov.) 



4 BREVIOBA No. 214 

4 (3) Calcareous ring simple, lacking posterior processes 

Paracucumidae nov. (Included ex- 
tant genus: Paracucumis Mortensen, 1925) 

5 (2) Body invested dor sally by conspicuous plates; sole present 

Psolidae Perrier, 1902 (Included extant 

genera: Psolus Oken, 1815; StoUnus Selenka, 1868; Psolidium 
Ludwig, 1886; Lepidopsolus Bronn, 1860; Thyonepsolus Clark, 
1901) 

6 (1) Body more or less naked, not enclosed by a test; calcareous de- 

posits small, inconspicuous. 

7 (10) Calcareous ring complex, with paired or unpaired posterior proc- 

esses. 

8 (9) Processes composed of a mosaic of small pieces 

Phyllophoridae Ostergren, 1907 (emend, herein). 

(Included subfamilies: Phyllophorinae Ostergren, 1907; Sem- 
periellinae Heding and Panning, 1954; Thyoninae Panning, 1949). 

9 (8) Processes entire Sclerodactylidae Panning, 1949 (Nomen 

translatum herein, ex Sclerodactylinae Panning, 1949). (Included 
subfamilies : Sclerodactylinae Panning, 1949 ; Cladolabinae Heding 
and Panning, 1954). 

10 (7) Calcareous ring simple, lacking posterior processes 

Cucumariidae Ludwig, 1894 (emend. 

herein). (Included subfamilies: Cucumariinae Ludwig, 1894; Col- 
ochirinae Panning, 1949 [partim?] ; Thyonidiinae Heding and 
Panning, 1954). 

Order DACTYLOCHIROTIDA nov. 

Tentacles digitiforiii or digitate, the digits sometimes bifurcate, 
8-30 in number. Body enclosed by a test comprising imbricate 
plates. Calcareous ring simple, lacking complex posterior pro- 
cesses. 

Key to Included Families 

1 (4) Anus and mouth at opposite ends of the body, which is fusiform 

or U-shaped. 

2 (3) Plates with a prominent spire. Tentacles 8-10 in number 

Ypsilothuriidae Heding, 1942 (Included extant 

genera: Tpsilotlmria Perrier, 1886; Echinocucumis Sars, 1895; 
Ypsilocucumis Panning, 1949) 

3 (2) Plates with small spires, or none. Tentacles 10-20 in number. 

Vaneyellidae nov. (Included extant 

genera: Vaneyclla Heding and Panning, 1954; MitsuJcuriella Hed- 
ing and Panning, 1954) 

4 (1) Anus and mouth opening close together, body flask-shaped 

Rhopalodinidae Perrier, 1902 (Included 



1965 NEW HOLOTHURIAN CLASSIFICATION 5 

extant genera: Bhopalodina Gray, 1853; Rhopalodinopsis Heding, 
1937). 
The above arrangement of taxa is believed to reflect phylo- 
genetic relationships more satisfactorily than that currently em- 
ployed. 

Of the fossil holothurian families defined by Frizzell and 
Exline (1955), the Calclamnidae (including genus Thuroholia) 
and some members of the Stichopitidae and Priscopedatidae 
should be included within this subclass. 

OTHER HOLOTHURIAN GROUPS 

In view of the above proposed revision of the Dendrochiro- 
tida, it is desirable to consider briefly the arrangement of the 
remaining holothurian orders, the Aspidochirotida, Elasipodida, 
Apodida and Molpadida. 

Aspidochirotida and Elasipodida 

Both the aspidochirotes and elasipods have tentacles which 
terminate in an approximately circular disc. The body is usually 
bilaterally symmetrical in both groups, with the dorsal tubefeet 
modified into papillae or warts (Aspidochirotida) or into elongate 
sensory processes (Elasipodida). The two groups are distinguish- 
able on the basis of some anatomical features, but both may be 
conveniently placed into a single subclass, as follows : 

Subclass ASPIDOCHIROTACEA Grube, 1840 

(Nomen translatum herein, ex Aspidochirotae Grube, 1840) 
Diagnosis: Tubefeet present, tentacles shield-shaped, 10-30 in 

number. No introvert, hence retractor muscles lacking. Body with 

conspicuous bilateral symmetry. 

Key to Included Orders 

1 (2) Respiratory trees present. Mesentery of posterior loop of intestine 

attached to right ventral interradius 

Aspidochirotida Grube, 1840 

2 (1) Respiratory trees lacking. Mesentery of posterior loop of intestine 

attached to right dorsal interradius 

Elasipodida Theel, 1882 



6 BREVIORA No. 214 

Theel (1882) has discussed the possible antiquity of the elasi- 
pod holothurians and concluded that they are certainly not rep- 
resentative of an ancestral holothurian stock; rather they are 
secondarily adapted to deep-sea life. In elasipods, and in some 
aspidochirotes, the madreporite opens to the exterior, and does 
not hang free in the body cavity. This can be regarded, not as 
a primitive feature, but as a logical compensatory consequence of 
the absence of respiratory trees. The extremely fragile calcareous 
ring in elasipods is apparently secondarily reduced. 

Within this subclass should be placed some representatives 
of the fossil families Priscopedatidae and Theelidae as defined by 
Frizzell and Exline (1955). 

APOOroA AND MOLPADroA 

The important character shared by adopids and molpadids is 
the almost complete absence of tubefeet. Also both groups have 
simple digitate or pinnate tentacles. It is possible that the 
apodids and molpadids bear no close relation to each other, and 
the characters they have in common may have arisen through 
parallel evolution and convergence. 

Subclass APODACEA Brandt, 1835 

(Nomen translatum herein, ex Apodes Brandt, 1835) 
Diagnosis: Tentacles simple, digitate or pinnate. Tubefeet 
markedly reduced, or, more usually, lacking altogether. No intro- 
vert, hence retractor muscles lacking. Deposits may include 
anchors and anchor plates. 

Key to Included Orders 

1 (2) Body cylindrical. Eespiratory trees and anal papillae lacking. 

Deposits often include wheels Apodida Brandt, 1835 

2 (1) Body fusiform, often with tapering caudal portion. Eespiratory 

trees and anal papillae present. Wheels lacking 

Molpadida Haeckel, 1896 

It is remarkable that some members of both Apodida and 
Molpadida have anchors and anchor plates in the body wall. The 
anchors and their plates differ morphologically in the two orders, 
but presumably have the same functional significance as accessory 
locomotor organs, since the anchors project through the body 
wall. 



1965 NEW IIOLOTIIURIAN CLASSIFICATION 7 

The fossil family Achistridae and some members of the fossil 
families Stichopitidae, Theelidae, Synaptitidae and Calcanoridae 
as defined by Frizzell and Exline (1955) may be included in 
this subclass. 

LITERATURE CITED 

Fell, H. B. 

1965. The early evolution of the Echinozoa. Breviora (in press). 
Fell, H. B. and D. L. Pawson 

1965. Chapters on the Echinozoa; and evolution and phylogeny of the 
Holothuroidea. In Treatise on Invertebrate Paleontology, U, 
Echinozoa and Asterozoa. (In press.) 
Frjzzbll, D. L. and H. Exline 

1955. Monograph of fossil holothurian sclerites. Bull. Univ. Missouri 
Mines Met., 89:1-204, 11 pis., 21 text figs. 
Pawson, D. L. 

1963. The holothurian fauna of Cook Strait, New Zealand. Zool. 
Pubis. Victoria Univ. Wellington, 36:1-38, pis. 1-7. 
Theel, Hj. 

1882. Report on the Holothurioidea. I. Challenger Sci. Results: Zool- 
ogy, 4(13):1-176, pis. 1-46. 



BREVIORA 

Museimi of Comniparsitive Zoology 



Cambridge, Mass. February 15, 1965 Number 215 

TWO NEW SUBSPECIES OF AMPHISBAENA (AMPHIS- 
BAENIA, REPTILIA) FROM THE BARAHONA PENIN- 
SULA OF HISPANIOLA 

By RicHAED Thomas^ 

As a result of collecting sponsored and led by Dr. Albert 
Schwartz during the summers of 1963 and 1964 in which I was 
fortunate to be able to participate, 33 specimens of Amphishaena 
were collected in the low areas on and near the Barahona Penin- 
sula of the Dominican Republic. The general affinities of these 
specimens are with Amphishacna innocens Weinland. Compari- 
son of the Barahona Peninsula specimens with representatives 
of the forms assigned to the species innocens shows them to be 
distinct in themselves, but to resemble most closely A. i. gona- 
vensis Gans and Alexander. These authors, in their recent study 
of West Indian amphisbaenids (1962), examined the available 
Hispaniolan specimens and reviewed the forms. The combina- 
tion AmpJiisbaena innocens caudalis Cochran was first used by 
them, and the new name Amphishaena i7inoce7is gonavensis was 
proposed for the population on Gonave Island. Fourteen addi- 
tional specimens of gonavensis, recently acquired by Dr. Ernest 
Williams, allow a better comparison of the Gonave and Bara- 
hona populations than was possible previously. Although the 
differences between the two populations are perhaps on a level 
of specific separation, their relationship with one another is ob- 
vious. I feel that relationships are best expressed by regarding 
these two as conspecific and distinct from A. innocens. These 
related amphisbaenids of Gonave Island and the Barahona 
Peninsula should therefore be known as Amphishacna gonaven- 
sis Gans and Alexander. 

In lacking major fusion of the head scales, Amphishaena 
gonavensis is separable from all other West Indian Amphis- 
haena except innocens Weinland, caeca Cuvier, hakeri Stejneger 

1 10,000 S. W. 84th street, Miami, Florida .33143 



BREVIORA 



No. 215 



and fcnestrata Cope. From fcnestrata and bakeri it is imme- 
diately separable on the basis of lower number of body annuli, 
199-225 for gonavensis versus 236-249 for fcnestrata and 239- 
254 for hakeri; from fcnestrata it is additionally different in 
lacking the backward penetration of the rostral between the 







D 

Fig. 1. Ventral views of the heads of four forms of Amphishaena. The 
malar scales are stippled for points of reference. A, A. g. gonavensis (MCZ 
80296), note three rows of postgenials and anterior penetration of first 
postgenials between genial and second infralabial; B, A. g. leheri (ASFS 
V2596), note two rows of postgenials and lack of anterior penetration of 
first postgenials (position of first two postgenials in hyporissor is inter- 
mediate between A and B) ; C, A. innocens (ASFS X3114) ; D, A. caeca 
(ASFS X7382). 



nasals. From A. caeca, A. gonavensis may be distinguished in 
the possession of two scales in the first and second rows of 
postgenials (versus three for caeca). Gonavensis also differs 
from caeca in having enlarged precloacal scales forming a dis- 
tinct convex flap as opposed to the relatively undifferentiated 
precloacals of caeca, and in heraipenial structure (see discus- 
sion). 



1965 HISPANIOLAN AMPHISBAENA 3 

Employing the concept of accessory dorsal half -annul! in the 
nuchal region, as expressed by Gans and Alexander (1962: 79- 
80), the first two body annuli in innocens, gonavensis and caeca 
typically correspond to three dorsal half -annuli (Fig. 2). Gona- 
vensis (and innocens, typically) has the first complete annulus 
including (continuous with) the anteriormost of the first three 
dorsal half -annuli (temporals, postoculars, frontals) (Fig. 2, 
B-D). Exceptionally, the condition depicted in Figure 2 A 
occurs in innocens. Examination of 49 specimens of A. caeca 
shows the anteriormost half-annulus to appear split off from, in- 
stead of continuous with, the first complete body annulus (i.e. 
first ventral half-annulus) (Fig. 2, E-F) . 

A. gonavensis differs from its neighbor species A. innocens in 
the possession of a malar scale ^ (related to this are differences 
in shape and arrangement of the ventral head scales. Fig. 1), 
and in the condition of the first pair of parietal scales which nor- 
mally are in contact apically (thus only narrowly) with one 
another at the midline or may even be slightly separated from 
contact by the second pair of parietals (Fig. 2, C-D). In inno- 
cens by contrast there are typically two or four large squarish 
parietals (there may be other small supernumerary scales pres- 
ent, but these do not alter the basic appearance) formed by the 
median scales of the second and third dorsal half -annuli (Fig. 
2, A, B). Gonavensis is additionally distinguished from innocens 
in the shape of the tail which is rounded rather than tapered 
(Fig. 3, A, B), in the precloacal scales which form a more 
prominently convex flap, and in hemipenial structure (see be- 
low). 

A. gonavensis is broader and shorter headed, has a less 
sharply pointed snout, and is distinctly smaller than inno- 
cens. The largest of 55 specimens of gonavensis measures 
242 mm ; of 73 innocens 17 measure more than 242 mm, the 



1 The malar scale as defined by Gans and Alexander is a major and obvious 
diflference between those Amphisbaena which possess it and those which do not. 
With respect to the species discussed here, the malar, possessed by A. gonavensis, 
seems at least in part homologous to the two end scales of the second row of 
postgenials of A. innocens. The advisability of denoting this scale with a sepa- 
rate name seems somewhat dubious inasmuch as it implies an all-or-nothing 
diflference where there is really one of degree. Further, the designation of the 
row of scales behind the malar (the row just in front of the first body annulus) 
as postmalars in contrast to this row being merely the third row of postgenials 
in forms having no malar seems also inadvisable and misleading. The term 
"malar," however, is a very convenient one in description, and its usage is re- 
tained with the above comments duly noted. The postgenials are here used to 
apply to all rows of small scales hetween the genial and the first body annulus, 
including the "postmalar row" used by Gans and Alexander. 



BREVIORA 



No. 215 



largest being 279 mm. Gans and Alexander noted that a single 
specimen from Petite Gonave (MCZ 25549) was at that end of 
the range of values (Gonave population) of several characters 
which was closest to the Gul-de-Sac sample of innocens. On the 
basis of the new material, this specimen falls within the range 
of the Gonave population in those characters for which the trend 
was most noticeable (body annuli, dorsal segments, postcloacals). 
Numerical data used in above comparisons and in ' ' Comparisons 









Fig. 2. Diagrammatic illustrations of Amphisbaena lieads (dorsal view, 
half -heads) showing sealation differences. First two body annuli are shown 
"peeled out" from heads. A, atypical A. innocens (Camp Perrin) ; B, 
typical A. innocens (Camp Perrin) ; C and D, alternative common condi- 
tions of A. gonavensis ; E and F, A. caeca. Stippled scales are second 
dorsal half -annuli (complete annulus in F). In A-D the second half -annuli 



are accessory • 



in E and F the first are "accessory." 



1965 HISPANIOLAN AMPHISBAENA 5 

and Discussion" below are in i)art obtained from Gans and 
Alexander's paper and in part from personal observation (see 
' ' Specimens Examined ") . 

Amphisbaena gonavensis hyporissor new subspecies 

Holotype: MCZ 77149, an adult male, collected 13.1 mi. (20.9 
km) SW of Enriquillo, Pedernales Province, Republica Domini- 
cana, 30 July 1963, by David C. Leber and Richard Thomas. 

Paratypes: KU 79824-25, same locality as type, 22 July 1963, 
Richard Thomas ; ASFS X9974-76, AMNH 92792-95, RT 754-55, 
UIMNH 55600-02, same data as type; MCZ 77150, Republica 
Dominieana, Pedernales Province, 5 mi. (8 km) NE Oviedo, 30 
July 1963, Richard Thomas. 

Diagnosis: A subspecies of Amphishaena gonavensis charac- 
terized by lack of fusions of head scales, a high number of 
caudal annuli, more than six precloacals ; an occasional caudal 
autotomy; and a mottled but faded coloration. 

Range: Presently known from the southeastern portion of the 
Barahona Peninsula of Hispaniola. 

Description of type (Fig. 4) : (Methods of counting and ter- 
minology follow Gans and Alexander, except for differences in 
terminology already noted). Head scales not fused; prefrontals 
as broad or slightly broader anteriorly than posteriorly. First 
two body annuli correspond to three dorsal half-annuli. First 
pair of parietals border anteromedially on frontals and are oc- 
cluded from apical contact by an anterior extension of the left 
second parietal. Genial .81 times as broad as long. Three rows 
of postgenials present; first row with two enlarged scales (an 
abnormal, minute middle scale is present), apex of each scale 
projecting slightly beyond malar-second infralabial suture be- 
tween genial and second infralabial ; second row with three 
scales not including the two large triangular malars at each end ; 
third row with four scales not including the two enlarged, ter- 
minal " postmalars. " Body annuli 213, four laterals on each 
side, and 20 caudal annuli. Sixteen dorsal segments and 24 
ventral segments to an annulus counted at midbody. Four 
cloacal pores ; 8 precloacal scales and 13 postcloacals, including 
2 median, rectangular postcloacals. Snout-vent length 200 mm ; 
tail 20 mm. Overall coloration tan, becoming darker and slightly 
mottled dorsally due to increasing amount of central dark pig- 
mentation on more dorsal segments ; four ventralmost segment 
rows without dark pigment spots. 



BRE\nORA 



No. 215 



Variation: The paratypes are similar to the type in the con- 
figuration of the first pair of parietals. Two have the first 
parietals in broad contact and squarish ; in six they do not meet 
at the midline, the second pair of parietals contacting the 
frontals (Fig. 2C) ; in the balance of the paratypes the first 
pair of parietals are in apical or only narrow contact. Consid- 
erable asymmetry occurs in the degree of contact of the first 
pair of parietals with the midline. In all specimens of hyporis- 
sor the two scales of the first row of postgenials project not at 
all or only slightly beyond the malar-second infralabial suture 
and thus between the genial and second infralabial (Fig. 1). 
All paratypes have three rows of postgenials, the modal count 
being 2 + 3 + 4 but with variants of four and five in the second 
row and five in the third row. Body annuli vary from 199 to 
221; laterals two to five (mode four), most permutations of 
these counts occurring except 2/2, 5/5, 4/5, and 5/2 ; caudal 







Fig. 3. Ventral views of the caudal aud cloacal region of four forms. 
A, A. g. gonavensis (MCZ 80291); B, A. g. leberi (ASFS V2596, except 
for median postcloacals, is also typical of hyporissor) ; C, A. innocens 
(ASFS X3111); D, A. caecu (ASFS X940). Median postcloacals of A 
and B are stippled ; those of A are typical of gonavensis and hyporissor ; 
those of B are diagnostic of leberi. 



1965 HISPANIOLAN AMPHISBAENA 7 

aiinuli vary from 1!) to 21 (mode 19). Dorsal segments of a 
midbody ammlus are 16 (mode), 17 or 18; ventral segments are 
22 (mode), 23 or 24; totalled midbody segments 38-42. Pre- 
eloacal scales range from eight to eleven; postcloacals from 12 
to 14; totalled cloacals 20-24. There are two enlarged, median 
postcloacals; these are typically straight-sided (Fig. 3A). Clo- 
acal pores are four in every specimen. The smallest specimen 
measures 106 mm total length (tail 10 mm), the largest 231 
mm (tail 20 mm). Apparent caudal autotomy constrictions are 
evident between the fifth and sixth annuli of the tails of some 
of the juvenile specimens; adults do not possess obvious au- 
totomy constrictions. One medium size specimen possesses a 
broken tail, presumably autotomized, missing beyond the fifth 
annulus. The coloration is pale tan, darker dorsally and with 
some scattered dark mottling ; some specimens were noted as be- 
ing purplish dorsally in life. 

Specimens collected in the vicinity of the town of Pedernales 
in the northwestern section of the Barahona Peninsula represent 
a subspecies distinct from the other two known populations of 
this species. This form is named in honor of Mr. David C. 
Leber, for his enthusiastic participation in collecting in the 
Dominican Republic. 

Amphisbaena gonavensis leberi new subspecies 

Holotype: MCZ 77218, an adult male, collected 5 km N of 
Pedernales, Pedernales Province, Republica Dominicana, 25 
June 1964 by Richard Thomas. 

Paratypes: MCZ 77219, same data as type; ASFS V2595-96, 
8 km N of Pedernales, 26 June 1964, Richard Thomas; KU 
79855-56, UIMNH 55627-28, ASFS V2676-78, AMNH 92827-28, 
Pedernales, 29 June 1964, Richard Thomas; RT 985, same lo- 
cality as previous series, 3 July 1964, Richard Thomas. All 
specimens from Pedernales Province, Republica Dominicana. 

Diagnosis: A subspecies of Amphishaena gonavensis most 
closely related to A. g. hyporissor in the possession of a high 
number of caudal annuli, but differing from that form in con- 
tact of first pair of parietals with one another, in having typi- 
cally two rows of postgenials, and in the configuration of the 
median postcloacals. 

Distribution: Known presently from the low elevations of 
the northwestern portion of the Barahona peninsular region of 
Hispaniola. 



8 BREVIORA No. 215 

Description of type: Head scalation much like that described 
for the type of hyporissor. First pair of parietals in broad con- 
tact across the midline giving a squarish appearance to the 
parietals. Two rows of postgenials present, two scales in the 
first row and five in the second. Anterior penetration of first 
two postgenials between the genial and second supralabial very 
slight. Body annuli 212 ; four laterals on each side ; 16 caudal 
annuli. Sixteen dorsal segments and 22 ventral segments to an 
annulus counted at midbody. Four cloacal pores, nine pre- 
cloacal scales and 14 postcloacal scales. Enlarged median post- 
cloacal scales four. On each side of the midline, two median 
postcloacals separated by a transverse suture which curves pos- 
teriorly and laterally giving a rounded outline to the posterior 
two of these four scales (Fig. 3B). Total length 223 mm, tail 
19 mm. Coloration generally like that of the type of hyporissor, 
but the tan is more uniform with less dark mottling. 

Variation: Head scalation of the paratypes is much like that 
of the type. The first pair of parietals are in broad contact 
medially in all but one specimen in which the contact is slightly 
more than apical. First two postgenials typically have none or 
only slight anterior penetration between the genial and the 
second inf ralabial ; the characteristic condition is that illustrated 
in Figure 1 B. Body annuli vary from 207 to 220 ; laterals two 
to five with the same variation noted for hyporissor; caudal 
annuli range from 16 to 19 (mode 19). Many specimens have 
the tips of their tails scarred ; were this not so, higher caudal 
counts would probably be obtained. Dorsal segments of a mid- 
body annulus are 15 (two specimens) or 16; ventral segments 
22-25 (mode 24) ; totalled midbody segments 38-41. Precloacal 
scales range from 9 to 11; postcloacals 11-14; total cloaeals 
20-24. Median postcloacals are essentially the same as described 
for the type in all except one specimen in which they are very 
abnormal and frgamented and another in which the posterior 
scales reach the cloacal border between the two anterior scales 
which are reduced. Apparent caudal autotomy is found in one 
of the paratypes of leheri (UIMNH 55628) and in another speci- 
men from Pedernales which is currently being kept alive. In 
both the tail is missing beyond the fifth annulus. Coloration is 
generally darker and more uniform than that of hyporissor. 
Hemipenes are everted or partially everted in four specimens; 
the organs are attenuated and naked ; apparently they are only 
slightly bilobcd (the most completely everted organs are not 



1965 HISPANIOLAN AMPHISBAENA 9 

bilobed, but terminal bifurcation of the dissected retractor penis 
indicates that they may be slightly bilobed when completely 
everted). 

Comparisons and discussion: To facilitate comparisons of the 
three races of A. gonavensis the following table showing scale 
count data is presented. 

Table 1. Scale count ranges for the three races of 

A. go7iavensis 





Body 
annul! 


Cauflal 
annul! 


Total 
niidbody 
sosments 


Pre- 

cloacals 


Post- 
cloacals 


Total 
cloacals 


gonavensis 


207-225 


10-12 


36-41 


6-7 


11-14 


16-20 


hyporissor 


199-221 


19-21 


38-42 


8-11 


12-14 


22-24 


leberi 


207-220 


16-19 


38-41 


9-11 


11-14 


20-24 



As shown by the table, gonavensis differs strikingly from the 
Barahona populations in the low number of caudal annuli. It 
further differs in scalation in the low number of precloacals (the 
condition of seven precloacals occurs in only one specimen), and 
in total cloacals. The specimens of gonavensis examined by me 
(18) and the type (not seen by me but illustrated by Gans and 
Alexander) are rather uniformly characterized by the first pair 
of parietals meeting in apical contact at the midline ; in but 
three specimens was there a short suture between the parietals. 
As described above, hyporissor is generally, but much more 
variably, characterized by narrow contact of the first pair of 
parietals, while in leheri the parietals with one exception are in 
comparatively broad contact with one another. Gonavensis and 
hyporissor uniformly have three rows of postgenials, while 
leheri is typically missing a segment in the gular region and 
consequently has only two rows. Of the three exceptions noted, 
one is abnormal in having the "missing" segment partly inter- 
calated between the second postgenial row and the first body 
annulus, a postgenial count of 2 + 4 + 2 being the result. The 
two scales of the first row of postgenials penetrate very far for- 
ward between the genial and the second infralabial in gonaven- 
sis (with 35 to 69 per cent of the length of the scale lying an- 
terior to the malar-second infralabial suture) ; in hyporissor the 
penetration is less (10-30 per cent, four scales have no penetra- 
tion) ; and in leheri even less (6-18 per cent, 11 scales have no 
penetration). The corresponding differences in the configura- 
tion of these scales can be seen in Figure 1 A, B. In the con- 
dition of the median postcloacals, gonavensis and hyporissor 



10 



BREVIORA 



No. 215 



agree in having but two roughly rectangular, undivided scales 
(small marginal scales which are normally folded inside the 
cloaca are not considered). The condition which characterizes 
leheri and the one exception have already been noted. One speci- 
men (ASFS V2507) from approximately 10 km NW of Oviedo 





B 





Fig. 4. A and B, dorsal and ventral views of the head of the type of 
A. g. hyporissor (MCZ 77149) ; C and D, dorsal and ventral views of a 
specimen of A. innocens from Camp Perrin (ASFS X3123). 



on the road to Pedernales has the leheri configuration of the 
median postcloacals ; it also agrees more with leheri in the con- 
dition of the parietals, while in having three rows of postgenials 
it agrees more with hyporissor. More specimens are necessary 
from intermediate regions before the status of this specimen can 
be determined. 

The presumed caudal autotomy noted for some specimens of 
hyporissor and leheri has not been observed in gonavensis, nor 
does it occur in A. innocens (both have relatively short tails). 



1965 HISPANIOLAN AMPHISBAENA 11 

Gans and Alexander noted that caudal autotomy is variable in 
some forms, e. g. A. caeca. 

If the hemipenial structure noted for leheri holds true for the 
species as a whole, this will serve as an additional distinction 
from A. innocens and A. caeca. In contrast to the simple, naked 
structure of leheri, the hemipenis of innocens is strongly bilobed, 
very heavy and fleshy; the sulcus spermaticus bifurcates at the 
fork of the organ and each branch proceeds to the non-sulcate 
side and thence to the apices of the lobes which are flattened 
and disk-like. Proceeding from each branch of the sulcus sper- 
maticus over each lobe and onto the distal sulcate surface of the 
organ are very fine but regular flounces. The non-sulcate sur- 
face is naked (from ASPS X3112). The hemipenes of caeca 
are of the same general appearance as those of innocens, but 
the sulcus spermaticus is much more prominent (thick edged) 
and forks slightly before the bifurcation of the organ itself ; the 
apices are rounded not flattened, and there is no evidence of 
flouncing (from ASFS X937, X4111). 

The distribution of the races of Aniphishaena gonavensis 
appears zoogeographically a bit strange at first glance (see 
Fig. 5). The possibility exists that it is in reality not so strange. 
The Cul-de-Sac plain, a channel (in places below sea level) be- 
tween the north and south ''islands" of Hispaniola (Williams, 
1961) bordered on both sides by mountainous regions, debouches 
to the west at the angle between the Tiburon Peninsula and the 
main part of the island and to the east at the northeastern 
corner of the Barahona Peninsula just east of the Sierra de 
Baoruco. A. gonavensis may occur (or have occurred in the 
recent geologic past) up the eastern coastal fringe of the Bara- 
hona (presently almost nonexistent in places) and into the 
Cul-de-Sac plain. This being so, the island of Gonave would not 
be an unlogical extension of its range. The close affinities of 
hyporissor with gonavensis seem to support such a distribution. 
It is interesting to note that at least two other Hispaniolan 
lizards have a similar distribution. Anolis hrevirostris Bocourt 
occurs along the eastern Barahona into the Cul-de-Sac, east to 
the Golfe de Gonave and on Gonave. Diploglossus curtissi 
Grant, though of wider distribution to the east and north 
(Schwartz, in press), also occurs along the eastern Barahona 
coast and is channelled to the west through the Cul-de-Sac (lo- 
cality records not continuous) and occurs on Gonave. 



12 BREVIORA No. 215 

It is interesting to note that despite the geographic continuity 
of the Sierra de Baoruco with the mountain mass of the Tiburon 
Peninsula, Amphishaena innocens is not known from the Bao- 
ruco. On the contrary it is A. manni^ which has been collected 
there. Gans and Alexander record four specimens of manni 
from near Paraiso at 1800 feet (about 600 m) (Fig. 5). In the 
summers of 1963 and 1964 we obtained four additional speci- 
mens (ASFS X9907-09, V2911) from the eastern end of the 
Sierra de Baoruco (see Fig. 5) at an elevation of 2600 feet. 
(790m). Although further collecting may prove otherwise, it ap- 
pears that manni is geographically the nearest Amphishaena to 
hyporissor, possibly even interposed between it and innocens. Com- 
parable geographic relationships are seen in the species men- 
tioned above as having distributions similar to the one predicated 
for A. gonavensis. Diploglossus ciirtissi and Anolis hrevirostris 
are "replaced" in the highlands of the Sierra de Baoruco by 
other related species (Diploglossus cost at us Cope and Anolis 
distichus Cope). 

The type and most of the paratypes of A. g. hyporissor were 
collected in one of those occasional but not altogether rare situa- 
tions where fossorial creatures such as Amphishaena and Typh- 
lops are found concentrated near the surface. Such localities 
are generally but not invariably characterized by a rather 
friable soil, a scattering of moderate to large sized trees, a cover- 
ing of plant litter, and an abundance of rocks. This particular 
locality is a region of xeric woods inland from the western 
mangrove margin of the Laguna de Oviedo. The locality gener- 
ally satisfied the above conditions ; an abundance of limestone 
rocks was present. Aside from the amphisbaenids, two species 
of Typhlops were found in this region, Typhlops cf. sulcatus 
and another species (Thomas, MS). The localities to the north 
of Pedernales where A. g. leheri was collected were similar to 
the locality just described, at least in degree of aridity. The 
locality at Pedernales, however, was somewhat more xeric; the 
tree cover was primarily Acacia with an undergrowth of scat- 
tered clumps of Opuntia growing among outcroppings of lime- 
stone rock; the soil was sandy. Typhlops haitiensis, Typhlops 



1 A further note of comparison between gotiarensix and manni may be in order. 
A. manni, though distinct from {/onavcngis in having the nasals fused with the 
rostral, differs further in having the condition of the half-aniiuli in the nuchal 
region as described for caeca ( ride xiipra) . 6-9 (versus 4) cloacal pori's, a com- 
paratively stout tail with a very prominent autotomy constriction, and a deeply 
bilobed heniipenis. In meristic characters there are no striking differences. 



1965 HISPANIOLAN AMPHISBAENA 13 

sp. and Leptotyp/ilops sp. (Thomas, MS) were found in the 
same macrohabitat with A. g. leberi in the Pedernales region. 
In both of the localities where the two largest series were ob- 
tained, numbers of shed skins of these amphisbacnids were seen 
while collecting. Little can be said about the habits of these 
lizards from our encounter with them. They w^ere mostly col- 
lected under rocks; their passageways were frequently evident. 
One grasped and held onto a rootlet with its mouth in a pos- 
sible attempt to resist capture. None were found above ground 
in the open, and their abundance did not seem to be correlated 
with any weather phenomena. 

Specimens examined 

Amphisbaena gonavensis Jiyporissor : As listed for type and paratypes. 

Amphishaena gonavensis leberi: As listed for type and paratypes. 

Amphisbaena gonavensis hyporissor x leberi: Repiibliea Dominicana; 
Pedernales Province, ca. 10 km NW Oviedo, ASFS V2507. 

Amphisbaena gonavensis gonavensis: Haiti: Gonave Island: Pointe-a- 
Eaquette, PM 3385 (allotype), PM 3386, 3388 (paratypes), MCZ 80289; 
Ti Palmiste, 6 km from Pointe-a-Eaquette, MCZ 80290-302. 

Amphisbaena innocens innocens: Haiti: Department de 1 'Quest, Manne- 
ville, MCZ 62511, MCZ 8748; Thomazeau, MCZ 37595-97; Purey, MCZ 
51417, ASFS X3862; Department du Sud, Camp Perriu, ASFS X3109-34, 
X3240-41, DEP 2403. 

Amphisbaena innocens caudalis: Haiti, Grande Cayemite Island, MCZ 
25550 (type), MCZ 25551 (paratype). 

Amphisbaena caeca: Puerto Eico: Isla Verde, ASFS X937-43, X4104-25, 
X7381-98; 2.2 mi. SW Sabana, ASFS X7433-34. 

ACKNOWLEDGMENTS 

I wish to express my appreciation to Dr. Albert Schwartz, 
who allowed me to study this material gained as a result of his 
West Indian collecting; to Dr. Ernest E. Williams, Museum of 
Comparative Zoology at Harvard ; Dr. Charles A. Reed, Yale 
Peabody Museum (PM) ; and Mr. Dennis R. Paulson (private 
collection, DRP) for loan of specimens in their care. I also wish 
to thank Mr. Ronald E. Klinikowski for executing some of the 
illustrations for this paper. 

Types and paratypes designated herein now reside in the 
following collections: Museum of Comparative Zoology at Har- 
vard (MCZ), American Museum of Natural History (AMNH), 



14 



BREVIORA 



No. 215 



University of Kansas Museum of Natural History (KU), Uni- 
versity of Illinois Museum of Natural History (UIMNH), Al- 
bert Schwartz Field Series (ASFS), Richard Thomas private 
collection (RT). 

LITEEATURE CITED 

Gans, Carl and A. Allan Alexander 

1962. Studies on amphisbaeiiids (Amphisbaenia, Eeptilia). On the 

amphisbaenids of the Antilles. Bull. Mus. Comp. ZooL, Vol. 

128. no. 1, pp. 65-158. 
Schwartz, Albert 
(In press.) Biploglossus cosiatus Cope (Sauria, Anguidae) and its relatives 

in Hispaniola. Reading Public Museum and Art Gallery, Sci. 

Pub. 
Williams, Ernest E. 

1961. The evolution and relationships of the Anolis seinilineatus 

group. Breviora, Mus. Comp. Zool., No. 136, pp. 1-7. 



-18* 




O 20 40 
   I I 

km 



74* 

I 



Fig. 5. Map of southwestern Hispaniola showing Gonave Island, the 
Tiburon Peninsula and the Barahona Peninsula with localities for the spe- 
cies of AfnpMshaena (some localities are from Gans and Alexander, 1962). 
Hollow circles, A. innocens; circles with crosses, A. gonavensis ; solid 
circles, A. Ttmnni. 



BREVIORA 



Meseum of Comparsitive Zoology 

Cambridge, Mass. February 15, 1965 Number 21() 

THE GEOGRAPHICAL VARIATION OF THE FROG HY- 

PEROLIUS MAEM0EATU8 (FAMILY HYPEROLIIDAE) 

IX RHODESIA, NYASALAND AND TANGANYIKA 

By R. F. Laurent 



Mr. Yesey-Fitzgerald has generously provided the Museum 
of Comparative Zoology with several very interesting series of 
tree frogs collected in various localities of Rhodesia, Nyasaland 
and Tanganyika. Some of them belong to the marbled tree frog 
Hyperolius marvvoratus Rapp, the geographical variation of 
which is so striking and has already required the recognition of 
many subspecies. This new material permits a summary of our 
knowledge of this species in eastern Africa plus some significant 
additions, including one very distinct new subspecies. 

Thus far, the marmoratus subspecies recognized in the present 
area are as follows : 

orgentovittis Ahl : Shores of Lake Tanganyika. 

rhodoscelis (Boulenger) : Luapula River drainage and Lake 
Rukwa, Tanganyika. This is apparently a disjunct range 
and suggests that Lake Rukwa might have been part of the 
drainage of the Chambeshi River, which is itself loosely 
connected with the Luapula drainage through a swampy 
area south of Lake Bangweolo. 

lestagei Laurent (a doubtful form) : Lake Bangweolo. 

mclanolcucus Laurent: Primarily the Lufira basin in the 
Congo, but samples from Sakania (near the Rhodesian 
border) show conclusively that its range exceeds the limits 
of this drainage in a southward direction. 

nlhorufus Laurent: Recently described (Laurent, 1964) from 
the Moxico Province, Angola, but also present in the Lua- 
laba District (southwestern Katanga). This race is likely to 
be found in the extreme northwestern part of Rhodesia. 



2 BREVIORA No. 216 

aposematicus Laurent : Only known to me from Lealui. Prob- 
ably restricted to Barotseland. Possibly intergrading with 
alhornfus to the north. 

rlwdesianus Laurent: Described from Matetsi, Southern Rho- 
desia and presumably ranging into the Wankie region. 

swynnertoni FitzSimons : Described from Chirinda, eastern 
Southern Rhodesia. 

tneniatus Peters: Lowlands in Mozambique southward to 
northern Natal and eastern Transvaal. 

aJhofasciatus Hoffman : Southern Nyasaland. 

nyassae Ahl : Described from Langenburg. The precise loca- 
tion of this locality appears controversial. Loveridge has 
stated (1957, p. 329) that it is Manda, Lake Nyasa. Manda 
is on the eastern shore at 10°28' S. In the Gazetteer No. 1 
(British East Africa) published by the United States 
Board on Geographic Names, no Langenburg is found, but 
there is a New Langenburg, which is Tukuyu. Tukuyu is 
near Rungwe where many other specimens of nyassae and 
filllehorni (which is a synonym) have been collected. This, 
therefore, sounds more probable as a type locality than 
Manda which is rather far from the Rungwe Mountains. 
Lastly, according to Stieler's Atlas (1905), Langenburg is 
on the eastern shore of Lake Nyasa, but almost at its north- 
ern tip. It would require fresh material from the Rungwe 
region and the northern part of Lake Nyasa to resolve the 
point. 

Vesey-Fitzgerald's material includes good series from Lusaka, 
Seremje, Mazabuka and Lake Chilwa, in addition to series from 
Lake Tanganyika and Lake Rukwa, which merely confirm that 
argcntovittis Ahl and rhodoscclis (Boulenger) are respectively 
present there. 

Hypeholius marmoratus melanoleucus Laurent 

Hyperolius melanoleucus Laurent, 1941, Kev. Zool. Bot. Afr., 34: 157, pi. 
VIII, figs. D, E, r. — Lukaf u, Upper Katanga, Congo. 

New material. 3 $6,1 9 (MCZ 37362-65), Lusaka, North 
Rhodesia, 7-9-III-1962 ; 1 9 (MCZ 38824), River Mwambeshi, 
north of Lusaka, North Rhodesia, 14-III-1963 ; 2 9 9 (MCZ 
38807-08), Mkushi District, North Rhodesia, 3-IV-1963. 



1965 HYPEROLIUS MARMORATUS 3 

Color pattern. Two males (31-34 mm) have still the juvenile 
dull color pattern. The third male which is small (26 mm) and 
the female (34 mm) have the vivid " melanoleucus" markings: 
black with white bands and a few white spots. The pattern com- 
prises essentially one mediodorsal and two laterodorsal main 
elements with some irregularities (branches, lumbar inflexions 
and isolated spots) ; in the female, the center or the axis of 
these white spots and bands shows red spots or lines. In the 
male with the adult pattern, these red markings exist only on 
the sides. The belly also shows red spots. In live specimens (I 
saw them very often when I was in Katanga) these red ventral 
markings stand out on the paler pink general coloration. 

The female from Mwambeshi River (30 mm) has a somewhat 
disrupted pattern with red only on the sides, the belly and the 
throat. The two females from the Mkushi District have the 
usual pattern, with the remarkable feature that the red dorsal 
lines are very broad. 

Range. The older record of this subspecies at Sakania was 
already a proof that its range was not limited to the upper 
Lufira drainage. The specimens from North Ehodesia suggest 
that this range actually overlaps the Zambesi basin, at least in 
the Kafue drainage. It must be remembered, however, that two 
specimens with an adult pattern are far from an adequate 
sample and that a situation similar to that which I shall describe 
for the Serenje population is surely not excluded. 

HyPEROLIUS MARMORATUS NYASSAE Ahl 

Eyperolius nyassae Ahl, 1931, Daa Tierreich, 55: 339, fig. 213. "Langen- 
burg, " northeastern shore of Lake Nyasa, Tanganyika. 

Eyperolius fiilleborni Ahl, 1931, Das Tierreieh, 55: 349, fig. 224. "Langen- 
burg, ' ' northeastern shore of Lake Nyasa, Tanganyika. 

New material. 13 S S,2 $ $ (MCZ 38809-38823), Serenje, 
Chikoli River, North Rhodesia, 10-III-1963. 

Color pattern. Three males have the juvenile pattern. One 
has a " melanoleucus" adult pattern. The nine others have 
irregular black spots, a few large ones or many small ones, with 
a general predominance of the light ground color. Red lines 
in the middle of the light network are generally distinct, but 
they are less conspicuous when the black spots are small and 
numerous. Three of the males have still a white mediodorsal 
band free of black spots (this is a remnant of a " melanoleucus''' 



4 BREVIORA No. 216 

pattern) as well as the red markings. One of the males has red 
spots on the throat. One of the females has a moderately dis- 
rupted " melanoleucus" pattern, with heavy red markings; in 
the other, the pattern is strongly disrupted and, like most males, 
still with clear red lines. 

Size. Males with juvenile pattern : 31, 32, 35 mm. Males with 
adult pattern: 31 (2 specimens), 32 (1), 33 (4), 34 (1), 36 (2) 
mm. Females: 35, 36 mm. 

Discussion and range. The pattern displayed by the majority 
of the specimens is quite different from the "melanoleucus" pat- 
tern. On the other hand, it appears identical with that of 
filllehorni Ahl over which the name nyassae Ahl, based on 
specimens with a juvenile pattern, has page priority. This 
judgment is confirmed by the comparison with two fiillehorni 
paratypes. I still have misgivings about the identification of 
the new material from the Chikoli River because nyassae and 
fiillelorni are based on populations coming from the northern 
part of the Lake Nyasa basin, while the Chikoli River seems to 
belong to the Luangwa drainage. However, there are already 
several examples in Hyperolius marmoratus of ranges which 
overlap two adjacent drainages while, on the other hand, ranges 
can also be separated by dividing crests or highlands as well as 
by escarpments and falls. Therefore, we must now admit that 
the range of the nyassae subspecies should be extended to in- 
clude the Luangwa basin. 

Hyperolius marmoratus albofasciatus Hoffman 

Hyperolius alhofasciatus Hoffman, 1944, Soolog. Navors. Nas. Mus. Bloem- 

fontein: 178, fig. 8. Limbe, Nyasaland. 
Hyperolius marmoratus alhofasciatus, Loveridge, 1953, Bull. Mus. Comp. 

Zool., 110: 350. Limbe, Mtimbuka, Euo River, Nyasaland. 

New material. 3 $ $,1 $,6 juv. (MCZ 46186-95). Lake 
Chilwa (= Lake Shirwa), Nyasa, 3-X-1943. 

Size. Males : 20.9, 22.7, and 23.9 mm. Female : 22.4 mm. 

Color pattern. The four adults have the typical albofasciatiis 
pattern ; in the smallest male, however, the dark coloration is 
brown rather than black, reminiscent of the juvenile color. The 
juveniles show a strong tendency towards a longitudinal pattern 
similar to the ''hayoni" pattern; this is not surprising, since 
this pattern is established in taeniatns, a Mozambican subspecies, 
which occurs not far from southern Nyasa. The shift to a longi- 
tudinal pattern occurs by the elongation in a backward direction 



1965 HYPEROLIUS MARMORATUS 5 

of the hour-glass spots which are located between the eyes and 
in the scapular region in the juvenile " undidatus" pattern. 
Similarly, the lumbar undulations of the undulatus pattern are 
also elongated, but in a forward direction. 

Discussion. Loveridge (1953) rightly objected to my synon- 
ymizing melanolcucus with alhofasciatus (Laurent, 1947b). 
Indeed, the red elements characteristic of the "melanoleucus" 
pattern are not apparent in alhofasciatus. Another difference 
is that the sides are predominantly white in alhofasciatus. This 
is similar to the situation in the southern subspecies {marmor- 
atus and verrucosus) where the dark dorsal pattern is generally 
restricted to the back, showing no encroachment on the flanks. 
In melanolcucus, there is still a large black elongated spot on 
the sides or several smaller black markings. In addition, the 
juvenile pattern does not show any kind of elongation in melano- 
lcucus. 

The size of the specimens from Lake Chilwa is unusually small 
for the species. This striking peculiarity is not shared by the 
series collected at Limbe (type locality of alhofasciatus), Mtim- 
buka and the Ruo River (Loveridge, 1953). Apparently the 
populations from Lake Chilwa, which seems completely isolated 
(i.e. without any efferent river), are dwarfed. An additional 
problem derives from the fact that the type of alhofasciatus is 
an adult female of only 24 mm, which does not match the size 
of the frogs collected at Limbe by Loveridge, but corresponds 
to our Lake Chilwa series. Consequently, I wonder if the type 
was not actually collected on the shores of Lake Chilwa. 

HyPEROLIUS MARMORATUS PYRRHODICTYON^ SUbsp. U. 

Holotype. 1 9 (MCZ 46182), Mazabuka, in the water-grass 
fringe of the Kafue River, 31-1-1963, North Rhodesia, Vesey- 
Fitzgerald coll. 

Paratypes. 2 $ $, 1 juvenile, same data (MCZ 46183-85). 

Diagnosis. A race of Hyperolius marmoratus, characterized 
by the absence of any spots or marbling on the back, and the 
presence of a dark red network on the belly and throat. 

Color pattern. The absence of any definite dorsal pattern is 
not quite unique in the marmoratus group : it is characteristic 
of some well differentiated subspecies such as rhodoscelis (Bou- 
lenger) of the Luapula drainage (Congo and North Rhodesia) 

1 From the Greek for "red network." 



6 BREVIORA No. 216 

and Jcarissimhiensis Ahl of the region of the Karisimbi volcano. 
These two races differ in their juvenile patterns and their color- 
ation in life : belly vermilion red with bluish flanks and a 
white laterodorsal streak in rJiodoscelis, belly purple red with 
back almost entirely dark blue in karissimhiensis. I don't know 
the colors in life of pyrrhodictyon, but a red network on an 
otherwise white belly is a quite unknown feature in any other 
member of the genus Hyperolius. 

On close examination under the binocular, it appears that 
some black spots may be present on the flanks, around the vent, 
and on the limbs along the boundary between the dorsal gray 
or brown (in alcohol) and the ventral red which becomes a 
solid color (not reduced to a network) on all the parts of the 
limbs that are hidden in the normal resting position of the 
tree frog. Therefore, as generally in the genus, the thigh is 
almost entirely vividly colored (red in this case) except for a 
dorsal stripe which is gray or brown. In the smallest of the 
two female paratypes, the black spots are also distinct on the 
throat between the meshes of the red network. 

The juvenile is almost uniformly greyish. 

Discussion. Since the juvenile, as sometimes happens in the 
marmoratus group, has no distinct pattern, specific identifica- 
tion may seem questionable. However, the morphology appears 
to be that of H. marmoratus which is different enough from 
that of other species to be relied upon at least tentatively. 

Size. The holotype measures 32 mm, the two adult paratypes 
30 and 26 mm respectively. 

Range. Since there are only the four specimens from Maza- 
buka, we do not know the range of this form. Mazabuka is not 
far from the right bank of the Kafue River. We already know 
that the left part of the Kafue drainage is inhabited by melano- 
leucus, that aposcmaticus lives in the Upper Zambezi region 
(Barotseland), and rhodesianus exists to the south of the Zam- 
bezi in western Southern Rhodesia. Poynton {in litt.) believes 
that aposematicus cannot belong to the same species as rho- 
desianus since they do not form hybrid populations where they 
meet; if he is correct, we can expect similar discoveries in the 
future and the species Hyperolius marmoratus would then have 
to be split into several species. However, if the dividing line is 
the Zambezi itself or the Victoria Falls, the lack of a hybrid belt 
has no meaning at all, reflecting as it does mere physical in- 
ability to meet. 



1965 HYPEROLIUS MARMORATUS 7 

The real relations of pyrrJiodictyon with the surrounding 
populations attributed to //. marmoratus will thus remain un- 
certain until more material is collected. 

Acknowledgment. This work has been supported by National 
Science Foundation Grant NSF G-1342. 

BIBLIOGRAPHY 

Laurent, R. F. 

1941. Contribution i la systematique du genre Hyperolius Rapp 
(Batraciens). Rev. Zool. Bot. Afr., 34: 149-167. 

1943. Les Hyperolius (Batraciens) du Musee du Congo Beige. Ann. 
Mus. Congo, Zool., (1) 4: 61-140. 

1947a. Two new forms of the genus Hyperolius. Ann. Mag. Nat. 
Hist., (11) 14:294-296. 

1947b. On some misuses of Hyperolius names. Ann. Mag. Nat. Hist., 
(11) 14: 288-294. 

1957a. Catalogue des rainettes africaines (genres Afrixalus et Hypero- 
lius) de la collection du Museum National d'Histoire Naturelle 
de Paris. Ann. Soc. Roy. Zool. Belg., 82: 23-50, figs. 1-2. 

1957b. Apergu des formes actuellement reconnaissables dans la super- 
espece Hyperolius marmoratus. Ann. Soc. Roy. Zool. Belg., 82: 
379-397. 

1964. Reptiles et amphibiens de 1 'Angola (troisieme contribution). 
Publ. Cult. Museu do Dundo, No. 67 : 1-165. 

LOVERIDGE, A. 

1953. Zoological results of a fifth expedition to East Africa. IV. 
Amphibians from Nyasaland and Tete. Bull. Mus. Comp. 
Zool., 110: 325-406. 
1957. Check list of the reptiles and amphibians of East Africa 
(Uganda; Kenya; Tanganyika; Zanzibar). Bull. Mus. Comp. 
Zool., 117: 151-362 -f i-xxvi. 



8 



BREVIORA 



No. 216 




Fig. 1. Ventral aspect of Hyperolius marmoratus pyrrJiodictyon n.sp. Type, MCZ 46182, 



\ 



1965 



IIYrEROLIUS MARMORATUS 




A 


ALBORUFUS 


A 


APOSEMATICUS 


e 


ARGENTOVITTIS 


• 


RHODOSCELIS 


N 


NYASSAE 


A 


ALBOFASCIATUS 



TAENIATUS 

PYRRHODICTYON 

RHODESIANUS 

SWYNNERTONI 

LESTAGEI 

MELANOLEUCUS 



Fig. 2. Map of the localities for the subspecies of Ryperolius marmoratus in East 
Africa. 



BREVIORA 

Miiseiim of Comparative Zoology 

Cambridge, Mass. March 1, 1965 Number 217 

THE AUDITORY REGION OF THE 
BORHYAENID MARSUPIAL CLAD0SICTI8 

By 
Bryan Patterson 



Since the appearance of Sinclair's memoir on the marsupials 
of the Santa Cruz formation of Patagonia (1906), it has been 
known that certain members of the Borhyaenidae, such as 
Borhyaena and Prothylacynus, were peculiar among marsupials 
in lacking a tympanic process, or bulla, of the alisphenoid. So 
far, indeed, had reduction of this bone been carried in 
Borhyaena that all trace of the foramen ovale had disappeared, 
the mandibular nerve evidently having passed out between ali- 
sphenoid and tympanic without so much as a notch to mark its 
passage. Sinclair was, however, able to show that lack of a 
tympanic process of the alisphenoid was not characteristic of 
the family as a whole, this structure being present in at least 
two genera, Cladosictis and " Aniphiproviverra" (=TJiylacocUc- 
tis). His observations have been confirmed by others (e.g. Wood, 
1924). Recently, the surprising statement has been made by 
McDowell (1958, p. 173) that "the borhyaenines show no clear 
relationship to other marsupials (they have no alisphenoid bulla, 
otherwise characteristic of marsupials, for example)." Even if 
this claim were correct, these animals would not be unique in 
this respect within their order. As has long been known (e.g. 
van Kampen, 1905, pp. 406-407), Phascolomis lacks an alisphe- 
noid bulla, although it is remarkable in possessing a process from 
the squamosal that fulfills the same function. 

In the course of a recent visit to Princeton University, I was 
able once more to examine the borhyaenid material described by 
Sinclair and reassure myself that his descriptions were correct. 
In the course of this examination I came across an undescribed 



2 BREVIORA No. 217 

specimen that adds to knowledge of the borhyaenid auditory 
region. This is Princeton University no. 15705, consisting of 
various fragments, among which is an incomplete cranium. It 
had been identified, presumably by Sinclair himself, as Clado- 
sictis histratus Ameghino, a determination which there is no 
reason to doubt. From the fact that Sinclair made no mention of 
the specimen in his memoir, I suspect that it was not prepared 
prior to publication. Professor Glenn L. Jepsen very kindly 
loaned me the cranium for description. The accompanying fig- 
ures are the work of Mrs. Dorothy Marsh. 

DESCRIPTION 

The cranium is broken off anteriorly in the vicinity of the 
postorbital constriction and has been considerably damaged by 
weathering, which has removed the condyles, the bases of the 
zygomatic arches together with the glenoid cavities, and much 
of the bone of the right side. Compensating for these losses is 
the almost complete lack of crushing, something that cannot be 
said for the specimens of this species that Sinclair described 
and figured. 

As in other borhyaenines, and unlike the crushed specimen 
figured by Sinclair (pi. 55, fig. 1), the basisphenoid and basi- 
occipital progressively increase in width posteriorly. As he 
noted, neither bone has a distinct median keel and both are 
relatively flat transversely; at the suture they form a fairly 
prominent transverse ridge. No carotid foramen is to be seen in 
the basisphenoid ; in fact the only foramen visible in this bone 
is a minute one immediately antero-internal to the foramen 
ovale. Between basioccipital and periotic there is a conspicuous 
foramen antero-internal to and almost as large as the foramen 
lacerum posterius. This is the posterior carotid foramen of 
Gregory (1910, p. 233) ; a wide, deep groove in the postero- 
lateral portion of the basioccipital leads forward to it. The 
significance of this foramen is discussed below. The pars petrosa 
of the periotic, as is clearly seen on the left side, is rather sharply 
pointed anteriorly and extends far forward. Except for the 
opening of the foramen just described, the medial border of the 
pars petrosa abuts very closely against the basioccipital and 
basisphenoid. 

There is a long and low, slightly curved and narrow auditory 
bulla that does not project ventrally below the basicranium. The 



1965 



AUDITOEY REGION OF CLADOSICTIS 



anterior two-thirds of this structure is formed by the alisphe- 
noid. Antero-externally, this portion is suturally united with 
the squamosal (part of this area is broken away in the specimen). 
Medially, there is a long, fairly wide and deep depression be- 
tween the basicranium and the alisphenoid, which is roofed by 
the pars petrosa. The Eustachian tube opened into the posterior 
portion of this depression and passed forward within it. On the 



1 u /^^ 



r.er-- 



•fen. r.'' 




Fig. 1. Cladosictis lustratus Ameghino. Ventral view of incomplete 
cranium. Princeton University no. 15705, X2. Abbreviations: al., alisphe- 
noid; al. b., alisphenoid bulla; Eu., opening for Eustachian tube; f.c.p., 
posterior carotid foramen; f.l.p., foramen lacerum posterius; /. ov., foramen 
ovale; fen. r., fenestra rotunda; p. mas., pars mastoidea of periotie; p. p., 
paroccipital process; p. pet., pars petrosa of periotie; r. e., epitympanic 
recess ; sq., squamosal ; ty., tympanic. 



broken right side of the specimen a natural section of the ali- 
sphenoid portion of the bulla is preserved. This is revealed as a 
deep, oval chamber situated almost entirely within the lateral 
wall of the cranium. It is separated from the rather large, 
circular epitympanic recess by a low ridge formed by the alis- 
phenoid anteriorly and the squamosal posteriorly. The remain- 
ing third of the bulla is formed by the periotie, which meets the 



BREVIORA 



No. 217 



alisphenoid in a suture anteriorly and is clasped posteriorly by 
the heavy, squat and blunt paroccipital process. The portion of 
the periotie involved is not the pars petrosa, which, as in other 
borhyaenines, exhibits no trace whatever of a tympanic process, 
but the pars mastoidea, which, in this form at least, had enlarged 
downward and forward from the ventro-lateral corner of the 
occipital surface. 




— a.t.c.f: 



mas. 



f. ov. I 



p.pet 



Fig. 2. Cladosictis lustratus Ameghino. Lateral and slightly ventral view 
of incomplete cranium. Princeton University no. 15705, X2. a.t.c.f., aper- 
tura tympanici canalis facialis; /. pgl., postglenoid foramen; /. sub., sub- 
squamosal foramen; VII, grooves for facial nerve; other abbreviations as in 
Figure 1. 



The meatus — a meatus spurious — is large, very short, and 
deeper than long; it is bounded anteriorly and dorsally by the 
squamosal, posteriorly by the squamosal above and the pars 
mastoidea below (the suture between these bones is unfortunately 
not distinct), and ventrally by the alisphenoid in front and the 
pars mastoidea behind. The tympanic is present on the left side. 
It is free of the bulla and gives the appearance of having been 
situated wholly or almost wholly within it, although evidently 
lying close to the bony meatus. The bone has the primitive 



1965 AUDITORY REGION OP CLADOSICTIS 5 

horseshoe shape so frequently encountered in the order ; it is, so 
far as can be seen, slightly enlarged ventrally, and has a short, 
dorsally tapering posterior crus. Due to slight forward dis- 
placement, the anterior crus is concealed by the squamosal ; this 
displacement is evidence that, as is usual in marsupials, the 
tympanic was not fused to the skull. The internal surface of 
the pars mastoidea is grooved anteriorly for the passage of the 
facial nerve, and there is a corresponding slit in the ventral face 
of the tympanic. 

DISCUSSION 

This is the first specimen of a borhyaenine in which a fully 
formed auditory bulla has been preserved, but it is not unique in 
the family. A complete bulla, superficially similar to that of 
the machairodontines, is known in Thylacosmilus. In a brief 
description that I gave of this structure (m Riggs, 1934, p. 13), 
the alisphenoid was identified as a component, although the 
extent of its participation could not be determined, and the pars 
mastoidea and paroccipital were recorded as "overlapping" the 
bulla; uncertainty was expressed as to whether or not the 
tympanic and a tympanic wing of the pars petrosa also partici- 
pated. With Princeton University no. 15705 in hand I have 
recently gone over the Thylacosmilus material in the Chicago 
Natural History Museum, hoping for a clearer interpretation 
in the light of this new evidence. It now seems virtually certain 
that the tympanic was situated within the bulla, as in Clado- 
sictis, but, for the rest, cracks, absence of recognizable sutures 
and slight distortion prevent any assurance regarding the ele- 
ments contributing to the posterior portion of the bulla. I am 
now very dubious as to the existence of a tympanic wing of the 
pars petrosa and suspect that only the pars mastoidea was in- 
volved in the bulla, but certainty on these points can only come 
with additional evidence. 

As has long been known, the borhyaenids show very definite 
resemblances to two marsupial groups, the Didelphoidea and the 
Dasyuroidea, and there has been controversy as to whether they 
were definitely dasyuroid, even thylacinine, or an ofi'shoot from 
the didelphoid stock. The evidence has been summed up by 
Simpson (1941), who, in general agreement with such earlier 
authors as Winge, Ameghino, and Matthew, concluded that while 
all three groups are clearly related, and very probably had a 
common ancestry, the Borhyaenidae were descended from the 



6 BREVIORA No. 217 

Didelphoidea, the resemblances between them and the larger 
members of the Dasyuridae being due to parallelism. With this 
view I am in complete accord, and the structure of the auditory- 
region contributes a few additional points to its support. In 
the Dasyuridae there is a large depression or recess in the 
squamosal, posterolateral to the small recessus epitympanicus, 
which is covered by the pars flaccida of the tympanic membrane ; 
no such depression is present in the Didelphidae and Borhyae- 
nidae. The facial nerve in dasyurids, after leaving the middle 
ear cavity, runs through a bony canal formed mainly by the 
pars mastoidea with the squamosal contributing antero-ventrally 
{Thylacinus, in which there is only a groove in the pars mas- 
toidea, is an exception) ; no such canal occurs in didelphids and 
borhyaenids. Borhyaeuids, so far as known, lack a tympanic 
process of the pars petrosa. In the Didelphidae this process is 
less developed than in the Dasyuridae, in which it usually con- 
tributes largely toward the formation of the bulla {Thylacinus, 
which lacks the process, is again an exception). Two borhyaenine 
genera are known to lack an alisphenoid bulla. This structure 
is smaller in a number of didelphids than is the rule among 
dasyurids. Cladosictis is peculiar among marsupials in the par- 
ticipation of the pars mastoidea in the bulla, but it is not unique 
in this respect. The remarkable little Dromiciops, which, with 
Reig (1955), I believe to be a surviving microbiotheriine 
didelphid, has a large, complete bulla in which, in addition to a 
tympanic wing of the pars petrosa, the pars mastoidea is involved 
posteriorly. Sinclair (p. 410, pi. 62, fig. 7) has figured a speci- 
men of Microhiotherium with a fully formed bulla, to which this 
element may well have contributed, and, as noted above, the 
same may well have been true of Thylacosmilus. 

THE INTERNAL CAROTID ARTERY IN THE THERIA 

Shortly after preparing this description I had the opportunity 
to make a field dissection of the head of an opossum (Didelphis 
marsxipialis) . In the course of this, I noted a small branch leaving 
the internal carotid artery at the level of the foramen laeerum 
posterius, passing into the cranial cavity through the posterior 
carotid foramen and there joining the circle of Willis. The 
observation was subsequently confirmed in injected specimens 
by Miss Suzanne Kreinbrook in the Biological Laboratories, 
Harvard University. This condition is widespread in the Mar- 
supialia, to judge from the nearly universal occurrence of the 



1965 AUDITORY REGION OF CLADOSICTIS 7 

posterior carotid foramen. Among the Recent genera I have 
been able to examine, the foramen appears to be lacking only in 
the microbiotheriines, and in Acrohates — and possibly Phasco- 
larctos — among the Phalangeridae. Even in these cases it is 
impossible to be certain short of dissection, for the foramen may 
be little more than a slit between the basioccipital and the peri- 
otic, as in caenolestids, or, apparently, confluent with the foramen 
lacerum posterius, as in some macropodids. 

The internal carotid artery passes through the basisphenoid 
in pelycosaurs and therapsids (e.g. Romer and Price, 1940 ; Olson 
1944), and also in non-therian mammals so far as known — 
Monotremata, Triconodonta (Kermack, 1963) and Multituber- 
culata (Simpson, 1937). The marsupials have retained this prim- 
itive condition, but have an alternate route in the small branch 
of the artery that passes through the posterior carotid foramen. 
Cladosictis, so far as I am aware, is the only member of the order 
in which this alternate route was fully followed, and in which 
the anterior portion of the artery and the foramen in the basi- 
sphenoid was lost. Cladosictis had, in fact, essentially and inde- 
pendently achieved the placental condition. 

Retention of the primitive condition in marsupials suggests 
that the placental arrangements were not attained until after 
the eutherian-metatherian dichotomy, presumably at some time 
or times during the Cretaceous. Just how these arrangements 
came about is, of course, unknown. The posterior carotid foramen 
alternate route may have been followed by all or by some pla- 
cental groups, but the possibility would appear to exist that a 
small, more anterior branch may have reached the cranial cavity 
via an opening antero-internal to the tympanic region (a fore- 
runner of the foramen lacerum medium), and that this even- 
tually became the anterior portion of the internal carotid. In 
either case the artery would have run forward along the medial 
edge of the periotic and could thus have readily become enclosed 
in a groove or canal between basicranial and auditory elements. 
The stapedial artery presumably came into existence quite early 
in placental history. Matthew (1909), who recorded its presence 
in arctocyonids, creodonts and miacids, suggested, rightly I sus- 
pect, that the internal carotid early became divided into medial 
and lateral (stapedial) divisions. With the establishment of these 
main branches, and given the relative ease with which blood ves- 
sel patterns may change, a variety of combinations became pos- 
sible. Independent acquisitions of similar patterns could have 



8 BREVIORA No. 217 

occurred and almost surely did occur. Since the supply of blood 
rather than the route taken by it is the important thing, it is 
doubtful if there is any selective advantage in one vessel pattern 
over another. How varied these patterns may be has recently 
been demonstrated by Guthrie (1963) for the rodents. Within 
this one order almost all possible variations are present ; in some 
groups both internal carotid (medial) and stapedial (lateral) 
branches occur ; in others one of these is absent, and in yet others 
both are wanting. 

Details of the carotid circulation appear to be of dubious 
value as an item of evidence for determining higher affinities 
within the Eutheria. 

EEFEEENCES 

Gregory, W. K. 

1910. The orders of mammals. Bull. Amer. Mus. Nat. Hist., 27: 1-524. 
Guthrie, D. A. 

1963. The carotid circulation in the Eodentia. Bull. Mus. Comp. Zool., 
128: 457-481. 
Kampen, p. N. van 

1905. Die Tympanelgegend des Saugetierschadels. Morph. Jahrb., 
34: 321-722. 
Kermack, K. a. 

1963. The cranial structure of the triconodonts. Phil. Trans. Koy. Soc. 
London, (B), 246: 83-103. 
McDowell, S. B. 

1958. The Greater Antillean insectivores. Bull. Amer. Mus. Nat. Hist., 
115: 115-214. 
Matthew, W. D. 

1909. The Carnivora and Insectivora of the Bridger Basin, middle 
Eocene. Mem. Amer. Mus. Nat. Hist., 9: 291-567. 
Olson, E. C. 

1944. Origin of mammals based upon cranial morphology of the 
therapsid suborders. Geol. Soc. Amer. Spec. Pap. no. 55 : i-vi, 
1-136. 
Eeig, O. a. 

1955. Noticia preliminar sobre la presencia de microbiotherinos vivi- 
entes en la fauna sud-americana. Investigaeiones Zool. Chilenas, 
2: 121-130. 
ElGGS, E. S. 

1934. A new marsupial saber-tooth from the Pliocene of Argentina 
and its relationships to other South American predacious mar- 
supials. Trans. Amer. Phil. Soc, n.s., 24: 1-31. 



1965 AUDITORY REGION OF CLADOSICTIS 9 

ROMER, A. S. AND L. I. Price 

1940. Eeview of the Pelycosauria. Geol. Soc. Amer. Spec. Pap. no. 28: 
i-x, 1-538. 

Simpson, G. G. 

1937. Skull structure of the Multituberculata. Bull. Amer. Mus. Nat. 
Hist., 73: 727-763. 

1941. The affinities of the Borhyaenidae. Amer. Mus. Novit., no. 
1118: 1-6. 

Sinclair, W. J. 

1906. Marsupialia of the Santa Cruz Beds. Repts. Princeton Univ. 
Expeds. Patagonia, 4: 330-460. 
Wood, H. E. 

1924. The position of the "sparassodonts": with notes on the rela- 
tionships and history of the Marsupialia. Bull. Amer. Mus. Nat. 
Hist., 51: 77-101. 



(Received 15 December, 1964.) 



BREVIORA 

Mnaseem of Cooiparsitive Zoology 



Cambridge, Mass. ^r.w 7, 1965 Number 21S 

NEW FK()(;S OF THE ({ENUS CORNUFER 
(RANIDAE) FROM THE SOLOMON ISLANDS 

By AV ALTER C. Brown ^ 

INTRODUCTION 

Large collections made by Mr. Fred Parker on Bougainville 
and neighboring small islands are providing very consider- 
able additions to onr knowledge of the fauna of this area. The 
present paper reports three new species and one new subspecies 
of the genus Cornufer discovered by Mr. Parker. Future papers 
in this series will describe other novelties in both frogs and rep- 
tiles and will record important ecological and behavioral obser- 
vations. 

PLATY3IANTIS SYNONYM 1 ZED WITH CORNUFER 

In my revision of the amphibians of the Solomon Islands 
(Brown, 1952), I followed Boulenger (1918, p. 372), Noble 
(1931, p. 522), and Deckert (1938, p. 148) in maintaining Platy- 
mantis and Cornufer as distinct genera. My separation of the 
two genera was based primarily on the structure of the digital 
pads, as emphasized by Boulenger (1918, p. 372). On the basis 
of this character, the Solomon Islands representatives of this 
grou]i of ranid frogs, which were known at that time, fitted 
rather readily into one or the other of the two categories. Inger 
(1954, p. 348), on the basis of his experience with the Philippine 
ranid frogs, again placed Platymautis in the synonymy of Cor- 
nufcv. In so doing, he pointed out the difficulty of maintaining 
a separation of these two genera when a majority of the species 
is considered and the apparent evolution of the digital pads is 
taken into account. ITowevei-, he did note that the species of 



1 Division (if Systi'iiia tic Biolnj^.v. StJiiil'onl I'lii \ nsit y ami Mi'iilo ('(illrf;c. Mciilo 
Parlv, Califitrnia. 



BREVIORA 



No. 218 



this group do have a number of characters in common, which 
separate them from Rana {sc7isu stricto). As to their relation- 
ships with other ranid genera, it has been noted by Noble (1931, 
]). 520) and Brown (1952, p. 86) that as a group they are prob- 
ably more closely related to the genus Discodcles, which is some- 
what intermediate in position when the Bana-Discodeles-Cornu- 
fer-Batrahylodes series is being considered. 

Since 1952 I have worked exteusively with this group of ranid 
frogs (I have now examined 23 of the 28 species referable to the 
genus), and I agree with Inger that it is indeed difficult to 
maintain the two genera as distinct entities on the basis of our 
present knowledge of the digital or other known characters, or 
on the basis of any great difference in ecological adaptations. 
Therefore, in the following list I have assigned all of the species 
previously placed in PJatymantis to the genus Cornufcr and 
noted their distribution. However, a thorough study of these 
frogs in terms of their morphology and life histories is much 
needed in determining relationships within the group. 

Cornufer acrocliordiia new species, PI. 2 Soloiuon Islands (Bougainville) 
Cornufcr acuhodactylus (Brown), PI. 1 Solomon Islands (Bougainville, 

Choiseul ) 
Cornufcr bcauforii (Van Kanipen j 
Cornufer boulengeri Boettger 
Cornufcr cliccsniani (Parker) 
Cornufcr coriiutus Tayloi' 
Cornufer corrugatus (Dumeril) 
Cornufer dorsnli.'i Dumeril 
Cornufcr {lilliardi (Zweifel) 



Cornufcr gucntlicri Boulenger 
Cornufer guppyi Boulenger, PI. 1 
Cornufcr hazelac (Taylor) 
Cornufer ingeri Brown and Alcala 
Cornufer macrops new species, PI. 1 
Cornufer mo.szl-ou-ski (Vogt ) 
Cornufer mycrsi (Brown), PL 2 



Waigeu Island (not seen by me) 

Bismark Island (not seen by me) 

New Guinea 

Philii»pine Islands 

l'liilipi)ine Islands 

l'hili])pine Islands 

Bismai'k and Adiiiiialty Islands 

(not seen by me) 
IMiilippine Islands 
Solomon Islands 
Philippine Islands 
Philippine Islands 
Solomon Islands (Bougainville) 
New (luinea (not seen by me) 
Solduion Islands (Bongainville ) 



Cornufer necheri Brown and Myers, PI. 1 Solomon Islands 



Cornufer p. papucnsis (Meyer) 
Cornufcr papuen.sin wchcri (Sclunidt), 

PI. 2 
Cornufcr p. porkcri new species, PI. 1 
Cornufcr parJ.cri huluK itsix new 

subspecies, 
Cornufer pelewensis (Peters) 



Xew (Uiinea, Bismarks 

Solomon Islands 

Solomon Islands ( P>ongain ville i 

Solomon Islands (Bukaj 
Palau Islands 



1<J65 NEW FKOOS KUOM TllK SOLOMON ISLANDS 3 

C(>niiif< r polillciisis (Taylor) Pliilil)|iiii(' Islands 

Cornufcr punctata (Peters and Ddi-ia) New (iuinea (not seen ]>y me) 

Connifrr ruhififriatus (Bari)our) Koou Island 

('(>r)uif( r sdlonioni.s Boiilentjer, I'l. 2 Solomon Islands 

Cuntiif( r suht('rr(!<fri.'< Taylor Philippine Islands 

Cornufcr unicolor Tsehudi New Guinea 

Cornufcr ritianun (Dunieril) Fiji Islands 

Cornufcr riticnsis (Gir.ard) Fiji Islands 

When considered from a zoo<i'eo<>'raphical point of view, as 
shown by this list of species, Cornufcv appears to represent a 
relict, peripheral group of ranid frogs with the greatest number 
of species occurring at present in the fringing Philippine and 
Solomon archipelagos, and a smaller remnant in New Guinea 
and related islands, as well as in the Palau and P'iji Islands in the 
outer Pacific. 

NEW SPECTES 

The new species collected by Mr. Parker are ail relatively 
small and have been compared directly with the tyjx' of C. 
andvodartijlnx ( USNM 119769) and with a paratype of C. chccs- 
mani (MCZ 26501). (The relative sizes at maturity of the sev- 
eral species known from the Solomon Islands are given in 
Table 1.) 

CORNUFER PARKERI I'ARKERI ^ sp. and Subsp. IIOV. 

This diminutive frog with its rough, tuberculate skin sui)er- 
ficially resembles some of the small Oriental bufonids. However, 
its firmisternal girdle, well developed omosternum, undilated 
sacral diapophyses, teeth only on the upper jaw, reduced webs 
and united outer metatarsals place it in the ranid genus Coruu- 
fer. 

Holofjipc: MC'Z 86928, a mature female collected at Kunua 
area, Bougainville Island, Solomon Islands, on July 10, 1962, 
by Fred Parker. 
' Paratypes: MCZ 86911-18, 36921-22, 38194. 41860, 41866-69, 
42524-81, 48741-44, Stanford University Nos. 21778-74. AMNII 
70069-71, collected from the same general locality as the holo- 
type, during 1962-1968. 

Diagnosis: A diminutive Cornufcr, largest available mature 
female measuring 18.5 mm, and largest male 15.9 nnn from snout 

1 Niinicil f(ir Mr. Fred Parker. 



4 BREVIORA No. 218 

to vent; skin witli numerous warty tubercles on dorsum, lateral 
surfaces and limbs ; first finger shorter than the second ; tips of 
fino'ers blunt or slio-htly pointed, occasionally a eircummarp:inal 
o-roove faintly indicated; tips of toes scarcely dilated, rather 
pointed, a shallow groove, most prominent laterally, separating 
the dorsal from the ventral portions ; diameter of eye usually 
greater than, rarely equal to, length of snout. 

Description: A very small Cornufer, snout-vent length of 16 
mature females 15.1 to 18.5 mm, of 8 mature males 14.0 to 15.9 
mm, habitus slender; hind limbs long, the snout-vent length 
ranging from 60 to 72 per cent of the length of the hind liml) 
for 10 specimens ; head about as broad as long ; snout round- 
pointed, upper jaw scarcely protruding ; eye large, its diameter 
somewhat greater than the length of the snout (rarely equal 
to) ; tympanum distinct, large, its diameter about 50 to 70 per 
cent of the diameter of the eye, and usually almost 25 per cent 
of the breadth of the head ; canthus rostralis rounded ; loreal 
region concave, only slightly obli(iue; a moderately to faintly 
distinct, oblique fold dorsal and posterior to the tympanum ; 
forelimbs well developed, fingers relatively uniformly slender, 
bluntly round or slightly pointed, occasionally with a very faint 
groove separating a ventral ]iad laterally from the dorsal por- 
tion ; witliout webs; subarticular and metacarpal tubercles large 
and well developed ; first finger shorter than the second which 
is about equal to the fourth (PI. 1, fig. 3) ; hind limb long; toes 
very slender without web, rather pointed with the ventral pad 
delimited by a shallow groove except at the tip (the fifth toe is 
more blunt and usually lacks the groove) ; subarticular and both 
inner and outer metatarsal tubercles well developed. 

Skin of dorsum and dorsolateral surfaces marked by scattered, 
moderate-sized, round, oval or oblong tubercles ; ventral and dis- 
tal posterior part of the thighs granular; the proximal posterior 
part of thighs marked by elongate folds. 

Color (in ])reservative) : dorsum and lateral surfaces from 
grayish brown, through brown, to brownish black, the lighter 
shades with irregular darker blotches; several of tlie sj)ecimens 
exhibiting a tan, silvery or whitish middorsal band, wider 
anteriorly than posteriorly, and beginning anteriorly on the 
posterior jiart of the head or tlie pectoi-al region ; lower limbs 
and (Hh^r of jaws with wide, dai-k ti-ansverse bars; venter rather 
hea\ih- mottled with dark brown. 



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6 BREVIOEA No. 218 

Measurements of holotype (in mm) : Snout-vent length 18.5; 
length of head to posterior edge of tympanum 6.8 ; breadth of 
head 6.9 ; diameter of eye 2.4; diameter of tympanum 1.7 ; length 
of snout 2.4 ; length of hind limbs 27.5 ; length of tibia 8.1. 

Ecological Jinfc: Parker (personal eommunieation ) states that 
the specimens of this small frog were found under stones and 
logs in lowland secondary growth areas. 

Comparisons: This Cornufer is much smaller at maturity than 
any other known species of the genus. In the warty nature of 
the skin it is most similar to C. acrochordus. 

Cornufer parkeri bukaensis subsp. no v. 

Holotype: MCZ 35777, a mature female, collected in lowland 
forest at south end of Buka Island, Solomon Islands, on January 
28, 1962. l)y Fred Parker. 

Paratypcs: AMXII 69814-15, same locality as holotype. 

Diagnosis: The Buka population is distinguishable from the 
nominate subspecies by the much less warty skin (both dorsal 
and ventral surfaces) ; the larger eye as measured by the ratio of 
eye diameter to breadth of the head and the relatively broader 
head as measured by the ratio of the length of the head to its 
breadth (Table 2). ' 

Because of their obvious close affinities, and the fact that the 
observable morphological differences between individuals of these 
populations of diminutive Cornufer are based upon a very small 
sample of the Buka population, I prefer to regard these two 
populations as geographic subspecies of a polytypic species. 
AVere these populations overlapping in range and wer(^ there 
no intergradation, they would be recognized as full species. As 
pointed out by Mayr (1963, pp. 481-515), geographieally isolated 
populations such as these island populations are certainly incip- 
ient species wliether or not marked by pronounced morphologi- 
cal differences. If the isolation is relatively complete for a suf- 
ficiently long period of time, true reproductive^ isolates f distinct 
species) may arise. 

Color (in preservative) : Dorsum and iippci- hiteral surfaces 
more or less uniforml\- piifplisli bi'owu or with lighter blotches; 
h)wer fore limbs and to some (Icgi'cc the thighs uiMi'kcd with dai-k 
transvei-se bands; venter with a i-cticuiate pattern of bi-own and 
grayish white. 



196.') 



NEW FROGS KKOM THE SOLOMON ISLANDS 



Measnrcmcnts of liolotypc (in mm) : Snoiit-vcnt lon^tli 15.9; 
length of head to posterior ed<i-e of tympanum ^).\) ; l)readth of 
head 6.4; diameter of eye 2.1; diameter of tym])annm 1.2; 
length of snout 2.8 ; length of hind limb 2:]..") ; U'ugtli of tibia 7.2. 

(Table 2 

Ratios of dianu'ti'i- of eye to hieadth of head, and length of head to 

breadth of liead, for C. parkeri parkeri and C. parkeri bukaensis 

(R ^ range ; ^f ^ mean ; X ^ ninnl>er) 





iJiameter of eve 


Length of head 




Breadth of head 


Breadth of head 




R = 0.339-0.405 


R = 0.966- 1.050 


('. parkeri parkeri 


M = 0.376 


M = 1.003 




X = 20 


X = 20 




R = 0.318-0.328 


R = 0.922-0.952 


C. parkeri hukaensis 


M = 0.323 


^r = 0.937 




X = 3 


x = a 



CORNUFER MACROPS^ Sp. nOV. 

Hulutype: MCZ 41864, an adult female, eolleeted at 3U00 to 
4000 feet, in mountains of Aresi area, south of Kunua, Bougain- 
ville Island, Solomon Islands, 1963, by Fred Parker. 

Faratijpes: MCZ 38195-96 and 43740 in mountains of Kunua 
area, Bougainville Island ; Stanford University Xo. 21795, Kieta 
area, Bougainville Island, collected by Fred Parker, 1962. 

Diagnosis: A small Cornufcr, largest available mature female 
measuring 28.5 mm, and largest male 25.9 nnn from snout to 



1 From the Greek meaning "large eye." 



8 BREVIORA No. 218 

vent ; second finger longer than first ; slightly dilated disks at 
the tips of the fingers and toes ; the ventral pad separated from 
the dorsal by a circummarginal groove ; eyes relatively large, 
diameter of eye greater than length of snont (Table 3), about 
4U per cent of head breadth. 

Description: A moderately small Cornufer, snout-vent length 
26.0 to 28.5 mm for the two females; 23.2 to 25.9 mm for the 
three adult males ; habitus slender, tapering from head to groin ; 
hind limbs long; the snout-vent length about 65 per cent of the 
length of the hind limb ; head about as broad as long ; snout 
rounded, upper jaw not protruding; eye very large, its diameter 
about 16 to 17 per cent of the snout-vent length, greater than 
the length of the snout (Table 3) ; tympanum distinct, its diam- 
eter slightly more than 20 per cent of the breadth of the head ; 
canthus rostralis rounded ; loreal region oblicpie, concave ; a rela- 
tively inconspicuous fold above and ]iosterior to the tympainim ; 
fingers relatively long, slender, without web ; finger tips slightly 
dilated and more or less rounded, the ventral pad separated from 
the dorsal portion by a shallow marginal groove, first finger much 
shorter than the second which is shorter than the fourth ; distal 
subarticular tubercles large and strongly protruding, basal and 
metacarpal tubercles less protruding (PI. 1, fig. 5) ; hind limb 
long ; toes slender, without web ; tips of toes slightly dilated, 
round or round-pointed, the ventral portion separated from 
the dorsal by a circummarginal groove ; subarticular tubercles 
moderately large and strongly protruding; inner metatarsal 
tubercle large and broadly oval, the outer small and round. 

Skin of dorsal and dorsolateral surfaces of body and u])per 
surfaces of limbs without pronnnent tubercles or folds ; skin of 
ventral abdominal region with faint small granules. 

Color (in preservative) : Dorsal and lateral surfaces blotched 
light and dark bi'own ; hind limbs with dark crossbars; ventral 
surfaces heavily fiecked with brown. 

Mcasnrcmoiis of holoiijpv (in mm) : Snout-vent length 26.0; 
length of head to posterior edge of tympanum 10.7; breadth of 
head 11.1; diameter of eye 4.9 ; diameter of tympanum 2.3 ; 
length of snout 3.9; lengtli of liind limb 44.9; length of tibia 
13.6; length of third finger 4.3; diameter of third finger disk 0.8. 

Comparhons: Cor nu fry ))iac)'ops is distinguished from C. 
myersi by its snudler size at maturity ami relatively lai-ger eye, 
diameter of eye greater than length of snout (not less as for 
/y///r/'.s('). It is distinguished fi-oni ('. pdrhivi by its larger size. 



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lU BREVIORA No. 218 

In size, C. Diao-ops is most similar to the Papuan-Solomon 
species ('. acnleodactylus, C. acrochordus, n. sp., C. checsmmiae, 
C. (jilliardi and C. unicolor. It differs from C. aculeodactylus in 
its much less pointed fint^ers ; the first finger shorter than the 
second (not longer) and the fingers longer relative to other 
measurements, length of third finger to base of second subarticu- 
lar tubercle about 40 per cent of breadth of head as compared to 
25 to 32 per cent in (\ aculeodactylus (Table 8); smaller, 
rounded, outer metacarpal tubercle ; and eye larger relative 
to length of snout (Table 3). It differs from C. (jiUiardi in that 
the first finger is shorter than the second (not longer) ; head 
narrower ; dorsal folds absent. It differs from C. acrochordus 
in having tlic skin much less warty; the fingers less pointed; 
the fingers longer ( ditf'erences for third finger length relative to 
breadth of head are shown in Table 3) ; and the eye larger rela- 
tive to length of snout (see also Table 3). C. macrops diflPers 
from C. checsmanae in the larger eye (diameter of eye greater 
than length of snout, not less than, and more than 35 per cent 
of breadth of head, not less than as for chcesmanae) ; and the 
more granular posterior venter. It differs from C. unicolor in 
the absence of a web at the base of the toes, tlie relatively longer 
hind limbs; and the relatively broader head. 

CORNUPER acrochordus' sp. IIOV. 

_^. llolutypc: MC'Z 442(i4, a mature female collected at Aresi 
Mountain region, south of Kunua, between 2()()()-4000 feet, 
Bougainville Island, Solomon Islands, on 6 September 1!)63, by 
Mr. Fred Parker. 

Paratypcs: MCZ 44256-63, 44265-66, same general area as 
the holotype; MCZ 41871-2 and Stanford University 21832, 
Aresi area south of Kunua (elevation about 30()()-4()0() feet), 
Bougainville Island, Solomon Islands. 

Diagnosis: A moderate-sized Cornufer, largest available 
mature female measuring 39.3 nun and largest male 27.1 nnii 
from snout to vent ; dorsal surfaces of limbs and body with 
scattered, pi'ominent. i-oundish tubercles, dorsum also with some 
elongate folds, venter with coarse, rounded granules; fingers 
short ; first finger longer than the secoiul ; fingers and toes dis- 
linctly |)()inted; subarticulai- and innei' metacarpal and meta- 
tarsal tubercles very large and sti-ongly piv)1 i-nding. 

1 From the (Iicck lUr "wmiIv." 



1 !)(;,■) MEW KKOCiS FKO.M THE SUEOMON ISLANDS 11 

Description: A moderate-sized Cornufcr, siiout-veiit l('n<:tli 
about 25 to 28 mm for mature males (4 measured) ; 37.0 to 40.0 
mm for mature females (6 measured). (Two females about :5() 
mm in len<i'th have undilated, straiji'ht oviduets. ) Habitus mod- 
erately broad and depressed; hind limbs lon^-, snout-vent len<>-th 
60 to "to per cent of the length of the hind limb; head broader 
than long; snout broadly rounded; ujiper jaw not or scarcely 
protruding:; eye moderately large, its diameter slightly less to 
slightly greater than the length of the snout and about 15 to 16.5 
per cent of the snout-vent length ; tympanum distinct, its diam- 
eter about 50 to 70 per cent of the diameter of the eye and 19 to 
24 per cent of the breadth of the head ; canthus rostralis broadly 
rounded ; loreal region strongly oblique and only slightly con- 
cave ; a prominent fold above and posterior to the tympanum ; 
fingers slender, round-pointed to pointed, ventral pad lacking, 
without web or lateral fringe; first finger usually longer than 
second (rarely equal to) ; second finger about ecjual in length 
to the fourth; subarticular tubercles very large and protruding 
but not pointed (PI. 2, tig. 4) ; metacarpal tubercles large, the 
inner protruding laterally; hind limb relatively long; toes slen- 
der, tips of toes slightly dilated, pointed, the ventral portion 
separated from the dorsal by lateral grooves; subarticular tuber- 
cles moderate, strongly protruding, distally pointed ; inner meta- 
tarsal tubercle large, strongly protruding; the outer a rounded 
cone; solar and palmar tubercles small but prominent; skin of 
dorsal and lateral surfaces of head, body and limbs with numer- 
ous small to moderate, prominent, rounded or elongate tubercles ; 
dorsum also marked with relatively short folds, the longest pair 
forming an urn-shaped pattern between the post-orbital and the 
axillary levels; venter posterior to the fore limbs, and the pos- 
terior surface of the thighs marked by prominent, relatively 
large, rounded tubercles. 

Color (in preservative) : Dorsum variable, grayish to black, 
mottled usually with a broad occipital blotch ; fore and hind 
limbs marked by light and dark transverse bars of about e(iual 
width; lips with dark bars; venter with brown flecks, heavily 
concentrated anterior to the fore limbs ; areolated light-dark pat- 
tern on inner and usually lower surface of thighs. 

Measurements of holotype (in mm) : Snout-vent length 37.7; 
length of head to posterior edge of tympanum 15.1 ; breadth of 
head 16.5; diameter of eye 5.6; diameter of tymi)anum 3.3; 
length of snout 5.9 ; length of hind limbs 60.5 ; length of tibia 



12 BREVIORA No. 218 

18.0 ; length of third tinger to base of second subarticular tu- 
bercle 5.0. 

Eggs: A small clutch of 10 eggs, stated to be of this species 
by Parker, were measured. In the preserved state they measure 
3 or 4 mm in diameter. They are creamy white, without any indi- 
cation of pigment. 

Comparisons: Cormifer acrochordus is intermediate in size 
at maturity between C. macrops and C. papuensis weheri of 
species known from the Solomon Islands, and closest to C. 
macrops (Table 1). AVhen compared with extraterritorial species 
it is of about the same size as C. dorsalis from the Philippines 
and slightly smaller than C. gilliardi from New Britain. The 
sharply pointed and relatively short fingers distinguish C. 
acrochordus from known species of the genus other than 
C. aculcodactylus and possibly C. giUiardi. It differs from C. 
aculeodactyhis in its larger size (Table 1), much more warty 
skin, color pattern, larger eye relative to length of snout, and 
the broader head relative to snout-vent length (Table 3). It dif- 
fers from C. gilliardi in its smaller size, more pointed fingers, 
more warty skin, broader head, and larger eye (Table 3) ; the 
differences from C. macrops have been discussed in the section 
on that species (p. 10). 

AKTIFICIAL KEY TO SPE(^IES OF COBNUFEK 
KNOWN FEOM THE SOLOMON ISLANDS 

1. Tips of fingers liroadl.v dilated, hreadth of disk of tliird finger more 
than 30 per cent of the length of the third finger as measured to the 

base of the second subarticular tubercle 2 

Tips of fingers not or scarcely dilated, breadth of disk of third finger, if 
dilated, less than 20 per cent of the length of the third finger as 
measured to the base of the second subarticular tubercle ?> 

'2. Head relatively narrow, its breadtli usually less than 40 i)er cent snout- 
vent h'Ugth ; loreal region slightly or moderately oblique; eye large, 

its diameter nearly equal to length of snout nrcl'cri 

Head relatively broad, its breadth usually greatei- than 40 per cent 
snout-vent length; loreal region strongly obli(|ue; eye moderate, its 
diametei' ciiual to oi' slightly greater than the distance fi'oiu eye to 
nostril .'/"/'/'.'/' 

3. First finger distinctly shorter than the second 4 

First fiugei' loiigci- than (occasionally about ('(lual to) the second ;") 

4. Snout-vent length of adults L'l) to .''II unn ; tips of fingers aiul toes idund, 

witli moderately dilated disks; fourtli finger longer than second ... 
mormps 



lill),") XKW FK()(iS FKOiSl THE SOLOMO.N ISLANDS I'S 

SiKiiii-vt'iit li'iijitli (if mliilts less tli;iii l^n nun; tijis of finjicrs .-iihI toos 
lilunt or slifjlitly i)oiiite(l, scarcely dilated; fonrtli fiiit^ei' usually 
shorter than oi- ahout eriual to second parlceri 

o. Tijjs of tintieis sliaiply jiointed; fourth finger usually shorter than the 

sc-iind when ad]iressed (i 

Ti])s of fingers lilunt or rounded; fourtli finger usually kinger than the 

second when adjiressed 7 

0. Skill with nunu'rous itroiuinent warts and dorsal folds; solar area with 

numerous tubereles acltrucliordus 

Skin relatively smooth; solar area without tubercles acideodactylufi 

7. Tills of fingers lilunt ly swolk'u, la(d\ing a niai'ginal groove (kdiiniting a 

veiiTial ]iad ; length of tiliia usually less than .111 jier cent of snout-vent 

length solonionis 

Tips of fingers with slightly dilated disks, a nuirginal groove delimiting 
a, ventral pad; length of tibia usually greater than 50 per cent of 
snout-vent length 8 

8. Solar area with prominent tubercles; weli at base of toes not reaching 

])roximal edge of suliarticular tuliercle on inner margin of second toe; 

dorsum, especially of adults, with numerous narrow folds 

l)apuensis weh,eri 

Solar area lacking prominent tuliercles; web at base of toes reaching 
midpoint of subartieular tul)ercle on inner margin of second toe; 
dorsum lacking numerous narrow folds myersi 



ACKNOWLEDGMENTS 

I wish to thank Dr. Alan Leviton, California Academy of 
Sciences, Dr. Doris Cochran, United States National Museum 
(USNM), Dr. Richard Zweifel, American Museum of Natural 
History (AMNII), Dr. Alice Grandison, British Museum (Natu- 
ral Hi.story), Drs. L. D. Brongersma and M. Boseman, Leiden 
Museum, for the opportunity of examining pertinent materials in 
the collections of their institutions; and Dr. Ernest Williams, 
Museum of Comparative Zoology, for suggesting that I describe 
these interesting frogs. 

The study of this genus of frogs is part of the author's pro- 
gram concerned with herpetofauna of the Pacific Islands. This 
program is sponsored by the National Science Foundation grant 
no. GB-409. 

Drawings are by Mr. AYalter Zawojski, Stanford University. 



14 BREVIORA No. 218 

LITERATURE CITED 

BOULENGER, G. A. 

1918. Reinark^i uii tlii' liatracliiaii s'-'n^^'i':' Coniiifer Tsc-hudi, Plattj- 
mantis Giinther, Simomantis g. ii., and Staurois Cope. Ann. 
:\raR. Nat. Hist., (it) 1: 372-375. 

Brown, Walter C. 

1952. Aiii])hil)ians of the Solomon I.slands. Bull. ^lus. Conij). Znol., 
107: l-()4, pis. 1-8. 

Deckert, Kurt 

1938. Beitrage ziir Osteologie und Systeniatik lanider Fruschluiche. 
Sitz.-Ber. Ges. Naturforscdi. Fr., Berlin, Jahrg. 1938; 127- 
184. 

Inger, Robert F. 

1954. Philippine zoological e.xpedition 194(5-1947. Systematies and 
zoogeogra))liy of Pliili])])ine Anijiliiliia. Fieldiana : Zool., 33: 
183-531. 

Mayr, Ernst 

19()3. Animal species and evolution. Harvard University Press, Cam- 
bridge, Massaehu-setts, xiv + 797 pp. 

Noble, G. K. 

1931. Biology of the Aniiihihia. Mcdiaw-Ilill Hook Co., New York, 
xiii + 577 i)p. 

ZwEiPEL, Richard (J. 

19fin. Results of the 1958-1959 Gillian! New Britain expedition. 3. 
Notes on the frogs of New Britain. Amer. Mus. Novit., No. 
2023; 1-27. 

(Received 17 Decemlier, 19(i4.) 



196.') 



XKW FROGS FROM THE SOLOMON ISLANDS 



15 





■^•*fef 



FIGURE I 



FIGURE 2 






FIGURE 3 



FIGURE 4 



FIGURE 5 



PLATE 1 

Fig. 1. Cornufer (/uppyi, inferior view of hand. 

Fig. 2. Cornufer necT:eri, inferior view of hand. 

Fig. 3. Cornufer p. parkeri n. sp., inferior view of hand. 

Fig. i. Cornufer aculeodactyJus, inferior view of hand. 

Fig. 5. Cornufer maerop-s n. si)., inferior view of hand. 



16 



BREVIORA 



Xo. 218 





FIGURE I 



FIGURE 2 





FIGURE 3 



FIGURE 4 



I 'LATH -2 

Fiii. 1. Coniitfcr mi/crsi, inferior view of li.-iiid. 

Fig. 2. Cornufcr papurn^is inhrri, iiift'rioi' \ic\v of IkiikI. 

Fig. 3. Cnrnufcr .solo)iioni.s, infcMior view of IiiiikI. 

Fig. 4. Cornufcr acrocliordus n. sp., inferior view of hand. 



BREVIORA 

Meseenra of Coeipsirative Zoology 



Ca mbridge, Mass. May 7, 191)5 Number 219 

THE EARLY EVOLUTION OF THE ECHINOZOA 
By H. Barraclough Fell 

Museum of Oomiinrntive Zoology 



INTRODUCTION 

The phylum Echinodermata is customarily considered to 
embrace two contrasted siibphyla. One of these, the Pelma- 
tozoa, comprises forms which are attached to the substrate for 
part or the whole of the life history, and which have a U-shaped 
gut, with the mouth and anus both directed upwards. The other, 
the Eleutherozoa, comprises free-living forms in which the mouth 
is directed downwards, and the anus (if present) is normally 
placed on the upper surface. The best known members of the 
Eleutherozoa are the sea urchins, the sea cucumbers, the star- 
fishes and the brittlestars. However, these included forms 
differ so widely that is has been a difficult task to elucidate their 
interrelationship and the probable nature of their presumed 
common ancestor. Further, there are strong grounds for sus- 
pecting that the subphylum Pelmatozoa includes some forms 
which are really related more closely to certain Eleutherozoa 
than they are to other members of the Pelmatozoa. These grounds 
are here set out, and it is proposed to abandon the Pelmatozoa 
as a formal classificatory division, and to adopt patterns of 
body symmetry as the main criteria for defining subphyla, in- 
stead of using habit and attitude. 

Recent morphological and paleontological studies have led 
to the conclusion (Fell, 1962, 1963a, 1963b) that the star-shaped 
members are interrelated and comprise a single grouping which 
may be regarded as a subphylum, and for which the name 
Asterozoa is already available. Similarly, other evidence implied 
that the globoid members were probably interrelated, and these 
have been associated as another subphylum, Echinozoa. Conse- 
quently, the so-called Eleutherozoa proved to be a polyphyletic 



2 BREVIORA No. 219 

assemblage, and the name should therefore be abandoned. The 
present contribution is aimed principally at clarifying the in- 
ferred interrelationships between the various classes of Echino- 
zoa, and adapting the current classification to reflect these rela- 
tionships. 

The discovery of the hitherto unknown class Helicoplacoidea 
(Durham and Caster, 1963) has shown that primitive, free-living 
echinoderms, with characters intermediate between those of Echi- 
noidea, Holothuroidea and Edrioasteroidea, had already dif- 
ferentiated in early Cambrian times. The morphological char- 
acters of the Helicoplacoidea suggest a relationship to some 
common ancestral stock from which arose, on the one hand, the 
eleutherozoan Echinoidea and Holothuroidea, and on the other 
hand, the pelmatozoan Edrioasteroidea. Hitherto, the pelmato- 
zoan echinoderms have generally been thought to represent 
a single natural assemblage, the subphylum Pelmatozoa, but 
this concept now becomes suspect. Further grounds for 
doubting the validity of subphyla based on eleutherozoan or 
pelmatozoan habit are provided by recent work on Paleozoic 
sea-stars, from which it has become evident that the subphylum 
Eleutherozoa, erected to comprise the free-living echinoderms, 
consists actually of two entirely separate stocks, the Echinozoa 
and Asterozoa (Fell, 1962, 1963a). The Echinozoa represent an 
ancient, pre-crinoid stock, of which the modern representatives 
are the Echinoidea and Holothuroidea ; whereas the Asterozoa are 
of relatively late origin, derived from a pinnulate pelmatozoan 
stock, provisionally identified with Crinoidea. Some analogous 
results emerge from recent work by Ubaghs (1961), from which 
it is apparent that the lower Paleozoic Homalozoa comprise a 
stock of asymmetrical, or bilaterally symmetrical, echinoderms, 
some members of which were free-living (i.e., eleutherozoan), 
while others were stalked and attached to the subtrate (i.e., pel- 
matozoan). Lastly, data given later in this paper imply the 
essentially archaic character of the dendrochirote orders of Holo- 
thuroidea, and point to possible relationships between these 
forms and the Cambrian Helicoplacoidea. Certain parallels be- 
tween the dendrochirote psolid holothurians, on the one hand, 
and the Edrioasteroidea, on the other, serve also to reinforce 
suspicions that the Edrioasteroidea should be classified with the 
echinozoan echinoderms, and not with the so-called Pelmatozoa, 
where they are commonly placed. Indeed, this inference is 
already implicit in a phylogenetic diagram published by Fell 
(1962). 



1965 EARLY EVOLUTION OF THE ECIIINOZOA 3 

PATTERNS OF SYMMETRY 

Four structural patterns may be contrasted in echinoderms; 
these are : 

(1) Homalozoan pattern, seen in those early Paleozoic echino- 
derms in which the skeletal plates are arranged either asym- 
metrically, or with more or less bilateral symmetry. These forms 
have been assigned to a separate subphylum, the Homalozoa 
(Whitehouse, 1941; Ubaghs, in press). 

(2) Echinozoan pattern, seen in the Helicoplacoidea, Holo- 
thuroidea, Echinoidea, Ophiocistioidea and Edrioasteroidea, all 
essentially globoid forms lacking arms, with meridional sym- 
metry. The Echinoidea and Holothuroidea were placed by Zit- 
tel (1895) and Jaekel (1918) in a subphylum Echinozoa, and 
the same name may be retained in a more extended sense, to com- 
prise all the classes listed here. 

(3) Crinozoan pattern, seen in the pelmatozoan classes Eocri- 
noidea, Paracrinoidea, Cystoidea, Blastoidea, Edrioblastoidea, 
and Crinoidea, initially globoid forms with partial meridional 
symmetry, but acquiring radially divergent systems of ambula- 
cral feeding appendages (brachioles or arms). These groups, 
together with the Edrioasteroidea and some dendrochirote Holo- 
thuroidea, exhibit a sessile habit, involving certain morphological 
features normally utilized in defining a subphylum Pelmatozoa. 
However, although the sessile holothurians have never been 
grouped, the so-called Pelmatozoa cannot be defined so as to in- 
clude the one without the other. It is evident that two categories 
of diagnostic criteria have been intermingled, and a more critical 
definition is required. 

(4) Asterozoan pattern, seen in the Somasteroidea, Asteroi- 
dea and Ophiuroidea, in which radial divergent axes of sym- 
metry produce arms, and the earliest morphological features of 
the arms correspond to those seen in pinnulate Crinoidea. These 
taxa fall within the subphylum Asterozoa, as defined by Zittel 
(1895), Jaekel (1918), and at greater length by Fell (1963a). 

It will be noted that whereas categories (1), (2), and (4) 
above appear to be natural groupings, and offer no diagnostic 
difficulties, some unsatisfactory features arise under category 
(3). These are now examined, in the light of evidence supplied 
by the other three groupings. 



4 BREVIORA No. 219 

BLEUTHEROZOAX AND PELMATAZOAN TRENDS 

In each of the subphyla Homalozoa, Echinozoa and Asterozoa, 
irrespective of the pattern of symmetry adopted, two mutually 
opposed evolutionary trends may be observed, fundamentally 
governed by the attitude which the animal adopts with respect 
to its habitat. These are: 

(a) Qeutherozoan tendencies, that is, adoption of a free- 
living habit, in which the animal acquires locomotor mechanisms 
permitting it to seek out food wherever it is to be found, by 
browsing on available algae, preying upon other animals, or 
swallowing large quantities of mud for the sake of its slight 
organic content. Such tendencies are invariably accompanied 
by the evolution of jaws, or of some special oral appendages 
adapted to gross (macrophagous) feeding. The anus, if de- 
veloped, tends to lie on a part of the body remote from the 
mouth. 

(b) Pelmatozoan tendencies, that is, adoption of a sessile 
habit, by which the animal becomes attached more or less per- 
manently to the substrate, either by the aboral surface itself or 
by an aboral stalk. Locomotor organs are atrophied or lost alto- 
gether, and the animal is then dependent upon such planktonic 
sources of food as the sea-currents may provide. It secures the 
food by some ciliary or comparable advective mechanism medi- 
ated by the tube-feet, the food particles being conveyed to the 
mouth by food grooves on the upper surface, the nutrition being 
therefore, of the microphagous type. The mouth and anus neces- 
sarily both lie on the upper surface, and the alimentary canal 
is consequently bent into a U-shape in the vertical plane. Al- 
though the modifications are here considered only in the context 
of echinoderms, analogous features, of course, occur in other 
phyla with sessile members. It may be noted here that radial 
symmetry is by no means a consequence of the adoption of sessile 
habits. On the contrary, echinoderms which already possess 
radial symmetry, if they adopt a sessile habit, may acquire a 
strongly marked bilateral symmetry, very similar to that acquired 
by the sessile tunicates, for example, with which the psolid 
holothurians were once confounded. Further, the discovery of 
Hclicoplacus (Durham and Caster, 1963) implies that the echino- 
zoan echinoderms were already free-living forms hefore radial 
symmetry was fully developed, and that no subsequent sessile 
stages supervened between the Cambrian Helicoplacoidea and 



1965 EARLY EVOLUTION OF THE ECIIINOZOA 5 

their presumed Ordovician successors, which include the earliest 
known echinoids and holothurians. 

It would appear that eleutherozoan and pelmatozoan ten- 
dencies are not directly related to the pattern of symmetry of 
the body in echinoderms, and that the two categories of evolu- 
tionary change, namely body symmetry and habit, have operated 
as simultaneous variables. This may be illustrated by reference 
to the echinozoan classes. 

EVOLUTION OF THE ECHINOZOA 

The oldest known echinozoan is the lower Cambrian Helico- 
placus, in which the body is fusiform in shape, with the mouth 
at a broad anterior end, and the anus at the tapering opposite 
extremity (Fig. 1). The skeleton comprises numerous quad- 
rangular or lozenge-shaped plates, sometimes bearing a rigid, 
erect spine, and disposed in counter-clockwise helical series. A 
single band of smaller platelets winds in a helix (sometimes bi- 
furcated) around the body, and evidently indicates the position 
occupied by a single external ambulacral water-vessel. The 
symmetry would appear to be bilateral, therefore, but distorted 
by the counter-clockwise torsion, and combined with an apparent 
radial symmetry displayed by the arrangement of the thecal 
plates. Similar torsion is observable in the earliest Echinoidea, 
notably Eothuria in the Ordovician; here, however, the sym- 
metry is overtly radial, or more correctly meridional, for there 
are now five ambulacra, disposed at regular intervals of 72°, 
forming twisted meridians. Analogous torsion is seen in the 
Edrioasteroidea, but has not yet been reported from Holothuroi- 
dea or Ophiocistioidea. The torsion was eventually lost in the 
echinoid line, but it persisted in the edrioasteroids until their 
extinction in the Carboniferous. 

As already noted above, eleutherozoan and pelmatozoan trends 
have arisen independently from time to time in the various 
groups of echinoderms. Within the Echinozoa, the classes Echi- 
noidea and Ophiocistioidea are not yet known to have produced 
any sessile forms. The initial echinozoan stock, to judge by the 
Helicoplacoidea, was itself free-living, too. Helicoplacus must 
have been a motile, bottom-feeding echinoderm, resembling a 
plated dendrochirote holothurian, as suggested below. The skele- 
tal plates formed a complete, robust though flexible test. The 



6 BREVIORA No. 219 

varying degrees of expansion and contraction reported in fos- 
sils by Durham and Caster (1963) imply an underlying muscula- 
ture, able to operate concertinawise. Thus, Helicoplacus prob- 
ably crept over the sea floor, like an annelid. The fossils occur 
in a fine clastic matrix, implying that the habitat was mud; 
Helicoplacus was probably a gross mud-swallower, like many 
aspidochirote holothurians. The primitive state of the ambula- 
crum suggests that the organ may have carried, at best, only 
rudimentary tube-feet, which could hardly have been more than 
respiratory organs, and probably Avere only sensory tentacles, 
like the dorsal tube-feet of many holothurians. The complete 
plating of the body-wall, and the probably rudimentary form of 
the tube-feet, imply the lack of an effective respiratory mech- 
anism on the outer surface of the body. If this is correct, then 
it may be inferred that rectal respiration was required, either of 
the pulsatory crinoid type, or by means of respiratory trees, 
as in holothurians. Study of the distribution of respiratory 
trees among holothurian orders suggests a direct relationship 
to the habits of these animals, and also implies that the earliest 
holothurians had already developed these structures. It therefore 
seems likely that respiratory trees of a rudimentary type were 
present in Helicoplacoidea. 

The earliest Echinoidea, such as £^o#/i?(na, possessed a multi- 
plated, flexible, spirally twisted body wall, similar to that of the 
Helicoplacoidea, and perhaps inherited from a helicoplacoid 
ancestry. They, however, had five well-developed ambulacra, on 
which the meridional water-vessels lay as external structures, 
though with internal ampullae for the tube-feet. Structural de- 
tails of the ambulacral pores show that the tube-feet were 
large, and probably suctorial — certainly extensile and muscu- 
lar. They would, therefore, serve the double function of loco- 
motor organs and respiratory organs, as in modern echinoids. 
The fossils exhibit a developed jaw mechanism, showing that the 
early echinoids were already capable of feeding in the manner 
of their extant endocyclic descendants, that is to say, by biting 
and grinding organisms in the substrate, and chewing algae. 
This type of feeding demands an eleutherozoan habit, and con- 
traindicates any pelmatozoan tendencies, since an animal with 
such feeding mechanisms would rapidly starve if it adopted a 
sessile manner of life. 

The Ophiocistioidea developed a rigid skeleton by soldering of 
adjacent plates of the test, in much the same manner as in the 



1965 



EARLY EVOLTTTION OF TITE ECfllNOZOA 




Figures 1-5. Archaic types of placoid echinozoans. 1, Helicoplacus lat- 
eral aspect, X 3. 2, Ypsilothuria lateral aspect, X 3. 3, Placothuria lateral 
aspect, X 2. 4, Isorophus actinal aspect, X 3. 5, Lepidopsolus actinal aspect, 
X 2. Fig. 1 from Durham and Caster, 1963; Figs. 2, 3, 5, drawn by D. L. 
Pawson, Fig. 4, from Kesling and Mintz, 1960. 



later echinoids. Locomotion, however, was effected by the use of 
the grossly enlarged and plated oral tube-feet, so characteristic of 
the class. The enlarged oral tube-feet on the lower surface would 
also subserve the function of nutrition, by sweeping up detrital 



8 BREVIORA No. 219 

material, and cramming it into the mouth, which was directed 
downwards. The anus, as in the endocyclic echinoids, lay on the 
upper surface, though not at the apical pole. Here, as in the 
endocyclic echinoids, the feeding and locomotor habits imply an 
eleutherozoan mode of life, and no sessile forms are known to 
have developed. 

The early Ilolothuroidea are known at present only from 
isolated skeletal plates. However, on the basis of recent studies 
by Pawson (1965) it would appear extremely probable that the 
Ordovician and later Paleozoic holothurians resembled the extant 
Ypsilothuriidae (Fig. 2). Further, when once the dendrochirote 
tentacle had been evolved, they would resemble the extant genus 
Paracucumis, or PlacotJiuria (Fig. 3). These are all heavily 
plated forms, with a complete test, flexible, made up of large 
plates with or without rigid spinous processes. The early holo- 
thurians would also be comparable with Helicoplacus, and with 
the flexible-bodied Ordovician echinoids, such as Eofhuria (the 
latter genus having originally been regarded as a holothurian). 
Suctorial tube-feet may have been lacking from the earliest 
holothurians, to judge by their rudimentary state in extant 
plated genera, though this is uncertain. If they were initially 
lacking, then locomotor movements would have been effected by 
contraction and expansion of the body wall and its flexible test. 
Once suctorial tube-feet had developed, locomotion on the echi- 
noid plan would be possible. There is morphological evidence 
that some kind of jaw apparatus, resembling the echinoid lan- 
tern, was developed early in the holothurians. Apparently it was 
abandoned once the dendrochirote tentacle had developed, but 
the skeletal elements of the presumed lantern acquired a new 
purpose — the attachment of the radial (and retractor) muscles 
— and hence the organ persisted in later holothurians as the 
calcareous ring on the pharynx. 

It is probable, especially from data given by Pawson (1965), 
that the dendrochirote tentacle evolved from a formerly simple 
oral tube-foot. Repeated dichotomy led to the complex dendritic 
tentacle of the Dendrochirotida. This is an efficient collecting 
organ for planktonic material which is conveyed to the mouth 
by the contractions of the tentacles, ciliary action, and the 
"spooning" action of two ventral tentacles. Dendrochirote holo- 
thurians, whether motile or not, are able to trap sufficient nour- 
ishment by filtering the surrounding sea water, provided there 
are currents replacing the surrounding water, and bringing fresh 



1965 EARLY EVOLUTION OP THE ECUINOZOA 9 

supplies of food particles. From such deudrochirote holo- 
tliurians more than one line of evolution is possible, for they 
have the means of adopting either eleutherozoan or pelmatozoan 
habits. If the locomotor system is retained, the oral tentacles 
can be adapted to serve as food-collecting- organs operating in 
various ways. In the Cucumariidae, for example, the body may 
be held erect, attached only by the posterior tube-feet, whilst the 
tentacles are spread out in a ring around the mouth, which is 
directed upwards, so that the habit of the animal resembles that 
of a sea anemone. In holothurians which adopt the horizontal 
attitude (lying on the ventral side where the tube-feet are re- 
tained), the tentacles can readily evolve from deudrochirote to 
aspidochirote forms, thus permitting gross mud-swallowing, and 
a markedly eleutherozoan habit. 

A further possibility is for the locomotor system to fulfill a 
purely adhesive role, leading to a sessile (pelmatozoan) habit. 
This is illustrated by the psolid dendrochirotes, in some genera 
of which the body is converted into a flattened, limpet-like form, 
adhering by a broad, flattened ventral surface, applied to a firm 
substrate (Fig. 5). The exposed dorsal and lateral surface is 
covered by a test of imbricating, robust plates. The mouth and 
anus lie on the upper surface, often protected by valvate plates, 
similar to those of cystoids or edrioasteroids. The whole body, in 
fact, is comparable to that of an edrioasteroid, the only dis- 
tinction being the lack of external ambulacral plates (Figs. 4, 
5). Thus the psolids are actually closer to edrioasteroids in 
morphological features than to many holothurians, or even to 
eehinoids, and the only character by which they differ from 
edrioasteroids is the same as that which distinguishes them from 
eehinoids — namely, the fact that the water-vessels are internal, 
and the test consequently does not form ambulacral plates. 
Hence a comparison of Psolus with an edrioasteroid illustrates 
two important features: (1) Edrioasteroids have the same pat- 
tern of symmetry as Echinozoa, and have evidently arisen from 
an early echinozoan stock similar to that which gave rise to the 
deudrochirote holothurians, of which the psolids are obvious 
members. (2) The pelmatozoan characters of edrioasteroids and 
of psolid holothurians have arisen as a direct habit response to 
adoption of a sessile mode of life, and do not indicate any near 
relationship to such pelmatozoan groups as are customarily in- 
cluded in the subphylum Pelmatozoa. 

Comparative study of the internal skeleton of the pliaryngeal 



10 BREVIORA No. 219 

region of dendrochirote holothurians suggests to me that the 
original holothurians must have had external ambulacra formed 
by modified plates of the test and similar to those seen in the 
fossil edrioasteroids and illustrated by Kesling and Mintz (1960) 
(Fig. 7). Apparently, when once evolved, the large dendrochi- 
rote tentacles required a protective mechanism, by which they 
could be withdrawn into the body. The protection was achieved 
by telescoping of the anterior end (termed the introvert), 
capable of retraction under the action of muscles derived from 
the radial muscle group. The evolution of the introvert, in turn, 
implied the conversion of the original external ambulacral areas 
of the test into internal structures, surrounding the pharynx, 
and serving for the insertion of the retractor muscles. In primi- 
ive dendrochirotes the pharyngeal skeleton is still recognizable 
as equivalent to the ambulacral plates of an edrioasteroid, but 
in most surviving holothurians the mechanism is very reduced 
or vestigial. Stages in the reduction are illustrated in Figures 
6 and 8 to 12. Inferred homologies of edrioasteroid and dendro- 
chirote skeletal elements are indicated in the captions to these 
figures. 

The Edrioasteroidea (Figs. 4, 7) adopted a similar habit to 
that of psolid Holothuroidea, but the ambulacra remained ex- 
ternal, instead of sinking inwards, and this permitted an alter- 
native method of feeding, suited to a pelmatozoan way of life. 
The feeding mechanism was provided by the whole complex of 
tube-feet. The five ambulacra grew outwards from the mouth 
as meridians, but only on the upper surface. Each ambulacrum 



Figures 6-12. Ambulacral plating systems and calcareous ring elements 
in holothurians (6, 8-12), and an edrioasteroid (7), considered in this 
paper to be homologous structures. 6, Pentadactyla (Dendroehirotida), X 3. 
7, Isorophus (Edrioasteroidea), X 4. 8, Placothuria (Dendroehirotida), X 
4. 9, Neothyonidium( Dendroehirotida), X 2. 10, Psolus (Dendroehirotida), 
X 3. 11, Euthyonidiella (Dendroehirotida), X 3. 12, Mitsukuriella (Dendro- 
ehirotida), X 4. Abbreviations: a.n., anterior notch of radial plate; g.d., 
gonoduct; ir.p., interradial plate; mad., madreporite; viad. d., stone canal; 
m.f., radial water-vessel; oes., esophagus; p.p.r., posterior process (considered 
in this paper to be the distal ambulacral plates) ; ph., pharynx; p.v., polian 
vesicle; r.m., retractor muscle; r.p., radial plate (here considered to be an 
ambulacral element). Figs. 6, 8, 9, from Pawson, 1963; 7, from Kesling and 
Mintz, 1960; 10, drawn by D. L. Pawson; 11, 12, redrawn from Heding 
and Panning, 1954. 



1965 



EARLY EVOLUTION OF THE ECIIINOZOA 



11 



an 





nm. 






10 




12 BREVIORA No. 219 

carried a median groove, bordered on either side by the tube- 
feet. The latter must have functioned in a manner similar to 
the tube-feet of sea lilies, that is, they would wave about in the 
water, trapping small plankton and organic particles on their 
mucous surface, and sweeping them inwards towards the mouth, 
along the food-groove on the ambulacrum. Evidently no intro- 
vert evolved, and this implies that dendrochirote tentacles never 
developed. 

It may be inferred that the Edrioasteroidea, soon after their 
differentiation from the initial echinozoan stock, adopted the 
sessile habit but, unlike the dendrochirote holothurians, had no 
oral tentacles on which to rely for nutrition. The ambulacra as 
a whole, therefore, took on the function of nutrition, mediated 
by the tube feet; and the advective food grooves, and the ambu- 
lacral plates on which they were carried, were a natural conse- 
quence. In the holothurian line, the radial water vessels were 
early converted into internal canals, as also occurred in the post- 
echinocystitoid echinoids. Hence the adoption of pelmatozoan 
habit inevitably demanded a pre-existing dendrochirote nutritive 
mechanism, and never involved external ambulacra in holo- 
thurians. As Bassler (1935) has pointed out, the earliest (i.e. 
mid-Cambrian) edrioasteroids retain a fully-plated ventral sole, 
unlike their later derivatives, and this may be taken as evidence 
that the test was originally spherical in edrioasteroids. The am- 
bulacra in the early edrioasteroids were also more simple than 
in later forms, and most of the so-called pelmatozoan features, 
such as cover-plates, may well be later specializations, analogous 
to the development of cover-plates in crinoids and somasteroids. 

Comparison of dissections of psolids with edrioasteroids sug- 
gests certain inferences as to the internal anatomy of edrioaster- 
oids. In the absence of any evidence to the contrary, we may 
assume that the edrioasteroids had a gonad placed in the dorsal 
interradius. In psolids the gonopore lies on the introvert, just 
posterior to the mouth. The corresponding position on an edrio- 
asteroid would be that in which a pore is known to occur, but 
the pore has hitherto been supposed to be a hydropore. Psolids, 
however, respire (at least in part) by rectal respiratory trees. 
It seems probable that respiratory trees would be required by 
edrioasteroids also, and that the hydropore would have been 
internal, as it is in dendrochirote holothurians. 

Irrespective of these latter inferences, the main conclusion 
emerges that edrioasteroids should be treated as members of 



1965 EARLY EVOLUTION OF THE ECIIINOZOA 13 

the Echinozoa aud that their pelmatozoan features are no more 
fundamental than the same features in psolid holothurians, being 
purely secondary responses to the demands of sessile habit. 

The Echinozoa have here been selected to illustrate an argu- 
ment which could also be developed on the basis of evidence pro- 
vided by other groups of echinoderms. The hitherto puzzling 
features of Homalozoa, for example, some being apparently pel- 
matozoans, others eleutherozoans, would appear to be no more 
unusual than the circumstances found within the Echinozoa. 
Without prolonging the discussion at this stage by reference in 
detail to other subphjda, the inference may be drawn that the 
eleutherozoan and pelmatozoan characters have arisen independ- 
ently, and at different times, in various echinoderm groups, 
and it is not possible to devise a natural classification on the basis 
of such characters alone. A revised classification now emerges in 
which the Edrioasteroidea are transferred to the subphylum 
Echinozoa. When this transfer is made, the residual pelmatozoan 
classes prove to comprise a much more uniform assemblage which 
may be defined, not in terms of their habit, but in terms of 
their morphological symmetry, as follows : 

Subphylum Crinozoa Matsumoto, 1929 (redefined) 

Fundamentally globoid echinoderms with partial meridional 
symmetry tending to produce an aboral calyx, the ambulacra de- 
veloping as aboral semi-meridians later forming radially diver- 
gent systems of ambulacral feeding appendages w^hich take the 
form of brachioles or arms. 

Included classes : Cystoidea, Eocrinoidea, Paracrinoidea, Blas- 
toidea, Edrioblastoidea and Crinoidea. 

The following extended diagnosis of the Echinozoa also 
emerges : 

Subphylum Echinozoa Zittel, 1895 (redefined) 

Fundamentally globoid echinoderms which never develop 
arms. In earliest members mouth and anus lay at opposite ends 
of the body, but in some later forms these have become sec- 
ondarily displaced. Meridional w^ater-vessels traverse the body 
wall in the direction of the anus, the vessels lying originally 
on the surface, but sinking into its substance in later Paleozoic 
and all post-Paleozoic groups. Skeleton, nervous system, repro- 
ductive organs and muscular system tend to differentiate into 
meridional systems, although an underlying bilateral symmetry 
is always evident. 



14 



BREVIORA 



No. 219 



Included classes : Helieoplacoidea, Edrioasteroidea, Ophiocisti- 
oidea, Holotliuroidea, Echinoidea. 

We can now set out a general classification of the phylum 
Bchinodermata, incorporating the proposals relating to the 
eleutherozoan groups already made in an earlier paper (Fell, 
1963a), together with the results of the present investigation. 
These are given in Table 1. It will be noted that uniform ter- 
minations in -zoa are adopted for subphylum categories, whilst 
class names end in -oidea. 



Conventional Classification 



Classification Herein Adopted 



Subphylum Pelmatozoa 
Class Carpoidea 

Class Cystoidea 
Class Eocrinoidea 
Class Paracrinoidea 
Class Blastoidea 
Class Edrioblastoidea 
Class Crinoidea 



Class Edrioasteroidea 
Subphylum Eleutherozoa 
Class Ophiocistioidea 
Class Echinoidea 
Class Holothuroidea 

Class Asteroidea 
Class Ophiuroidea 



Subphylum Homalozoa 
Class Carpoidea 

Subphylum Crinozoa 

Class Cystoidea 

Class Eocrinoidea 

Class Paracrinoidea 

Class Blastoidea 

Class Edrioblastoidea 

Class Crinoidea 
Subphylum Echinozoa 

Class Helieoplacoidea* 

Class Edrioasteroidea 

Class Ophiocistioidea 

Class Echinoidea 

Class Holothuroidea 
Subphylum Asterozoa 

Class Stelleroidea (including 
the subclasses Somasteroidea, 
Asteroidea and Ophiuroidea) 



^Helieoplacoidea unknown before 19G3. 

Table 1. Comparison of the conventional classification (left) of Echino- 
dermata with that herein adopted (right). The new arrangement of the 
classes in four subphyla is believed to avoid the polyphyletic categories 
which have been demonstrated in the conventional classification, and it at- 
tempts to reconcile the systematic treatment with the evidence of the fossil 
record. 



ACKNOWLEDGMENTS 

In preparing this paper I was assisted by having access to then 
unpublished data on the morphology of dendrochirote holo- 
thurians assembled bv Br. David L. Pawson, at the time when 



1965 



EARLY EVOLUTION OF THE ECHINOZOA 



15 



we were working together on South Pacific echinoderms; these 
data are presented elsewhere (Pawson, 1965). A revised classifi- 
cation of Holothuroidea recently proposed (Pawson and Fell, 
1965) synthesizes the sj'^stematic outcome of our separate con- 
tributions. A further contribution, incorporating some material 



<^^W?i^ 




z 
< 

< 



Volchovio 
V 
\ 
\ 

OPHIOCISTIOIDEA 



BothriocidarisEothuria 

\ \ / 

ECHINOIDEA 



Thuroholia Cystaster 
/ / 

HOLOTHUROIDEA / 
/ 




Edriooster 



Walcottidiscus 



Helicoplacus 
HELICOPUkCOlDEA 



/ 

(^ 

Stromctocystltes 
EDRIOASTEROIDEA 



Figure 13. Approximate inferred phylogeny of the Echinozoa as deduced 
from evidence discussed iu this paper. 



16 BREVIORA No. 219 

added by Professor Raj'mond C. Moore, University of Kansas, 
will be given in the forthcoming Treatise on Invertebrate Paleon- 
tology, volume U, where the material of this paper is given a 
formal text-book presentation (Fell and Moore, 1965) ; for assist- 
ance in re-lettering Figure 13, I am grateful to Dr. Moore. I 
have to thank Dr. Ernst Mayr for reading and criticizing the 
manuscript, and for suggesting the inclusion of Table 1 ; Dr. 
David Pawson for assistance in illustrating holothurians, and 
for permission to use Figures 2, 5, 6 and 8 ; Dr. Porter M. Kier 
for some suggestions ; and other colleagues and students who 
have discussed the interpretations offered here, and who have all 
to some extent influenced the manner of presentation here 
adopted. 

SUMMARY 

The morphology of fossil and extant Echinodermata implies 
that pelmatozoan and eleutherozoan characters must have arisen 
independently in the various classes at several different times 
since the Pre-Cambrian. Hence it is not possible to base a natu- 
ral classification on these two categories of contrasting characters 
alone ; for sessile and free-moving echinoderm assemblages each 
comprise two or more unrelated stocks, the similarities of which 
are due only to convergent evolution. At least four subphyla may 
be defined on the basis of four recognized patterns of growth and 
symmetry, mediated by the hydrocoel, evidently as innate trends 
initially independent of the environment. At least three of the 
four subphyla include pelmatozoan and eleutherozoan stocks, 
which are here regarded as arising as simple habit responses 
to the habitat. 

Edrioasteroidea are interpreted as members of the subphylum 
Echinozoa, and should be removed from their present conven- 
tionally accepted position among the so-called Pelmatozoa. The 
latter assemblage, after removal of the Edrioasteroidea, is re- 
defined and assigned the rank of subphylum, under the name 
Crinozoa. The so-called calcareous ring of holothurians is here 
interpreted to be the homologue of the ambulacra! plating system 
of edrioasteroids, telescoped within the body, following the evo- 
lution of the introvert. 



1965 EARLY EVOLUTION OF THE ECHINOZOA 17 

EEFERENCES CITED 

Bassler, E. S. 

1935. The classification of the Edrioastcioidea. Smithson. Misc. Coll., 
93 (8): 1-11. 
Durham, J. W. and K. E. Caster 

19G3. Helicoplacoidea, a new class of echinoderms. Science, 140: 820- 
8-2-2. 
Fell, H. B. 

1962. A classification of echinoderms. Tuatara, 10 (3), 138-140. 
1963a. The phvlogeny of sea-stars. Phil. Trans. Eoy. Soc. London, B, 

246: 381-435, figs. 1-18. 
1963b. The evolution of the echinoderms. Ann. Eept. Smithson. Inst, 
for 1962 : 457-490, pis. 1-3. 
Fell, H. B. and E. C. Moore 

1965. General features and relationships (of echinozoans). Treatise 
on Invertebrate Paleontology, U: (in press). 
Jaekel, 0. 

1918. Phylogenie und System der Pelmatozoen. Pal. Zeitschr., Berlin, 
3: 1-128. 

KESLING, R. V. AND L. W. MiNTZ 

1960. Internal structure in two edrioasteroid species. Univ. Michigan 
Contrib. Paleont., 15: 313-348, figs. 1-4, pis. 1-9. 

Matsumoto, H. 

1929. Outline of a classification of the Echinodermata. Tohoku Imp. 
Univ., Sei. Eepts., ser. 2( Geol.), 13 (2): 27-33. 
Pawson, D. L. 

1963. The holothurian fauna of Cook Strait, New Zealand. Zool. 
Pubis. Victoria Univ., Wellington, 36: 1-38, pis. 1-7. 

1965. Phylogeny and evolution of holothuroids. Treatise on Inverte- 
brate Paleontology, U: (in press). 
Pawson, D. L. and H. B. Fell 

1965. A revised classification of the dendrochirote holothurians. Brevi- 
ora, Mus. Comp. Zool., No. 214: 1-7. 
Ubaghs, G. 

1961. Sur la nature de I'organe appeletige ou pedonele chez les car- 
poides Cornuta et Mitrata. C. E. Acad. Sci. Paris, 253: 2738- 
2740, figs. 1-5. 

Whitehouse, F. 

1941. The Cambrian faunas of north-eastern Australia. Mem. Queens- 
land Mus., Brisbane, 12: 1-28, figs. 1-9, pis. 1-4. 
Zittel, K. a. von 

1895. Grundziige der Palaeontologie (Palaeozoologie). Munich, 971 
pp. 

(Eeceivcd 18 January, 1965.) 



BREVIORA 

Mmseminti of Comnpsirative Zoology 



Cambridge, Mass. May 7, 1965 Number 220 

A NEW SPECIES OF ELEUTHERODACTYLUS 
FROM GUADELOUPE, WEST INDIES 

By John D. Lynch 

Department of Zoology and Museum of Natural History 
University of Illinois 
Urbana, Illinois 



The numerous Greater Antillean frogs of the genus Eleu- 
therodactylus have, for the most part, been studied as groups and 
in faunal studies. Cochran (1941) studied the forms occurring 
on Hispaniola, and Lynn and Grant (1940) studied the Ja- 
maican forms. In a series of papers, Schwartz (1957, 1958a-d, 
1959a, b, and 1960) has clarified the status of the forms 
occurring on Cuba. 

The frogs of this genus in the Lesser Antilles are less well 
understood. Five names have thus far been applied to the 
frogs occurring there. These are : E. martinicensis (Tschudi), E. 
johnstonei Barbour, E. lentus (Cope), E. aiitillensis Reinhardt 
and Lutken, and E. darludensis (Auffenberg). The last form 
was described as an extinct Hyla by Auffenberg (1958) but I 
have recently shown (Copeia, in press) that it is an Eleuthero- 
dactylus and is probably not extinct. My studies on the osteology 
of these frogs and the researches of Albert Schwartz, who is 
currently revising the Lesser Antillean Eleutherodactylus, show 
that in reality there are numerous forms of this genus occurring 
on the Lesser Antilles. 

James Lazell, Jr. and Patrice Barlagne collected two forms 
at Matouba, north of Basse Terre, La Guadeloupe, which they 
could distinguish by voice and habit of calling. On external 
features, however, they are nearly identical. But from Lazell's 
field notes and discussions with him it appeared that two species 
were involved. Inasmuch as I had had good fortune in separat- 
ing some of the other Eleutherodactylus on neighboring islands 
by the structure of their pelvic osteology, specimens of these 



BREVIORA 



No. 220 



forms were macerated, and strong differences were found be- 
tween the ilia of the two forms. These differences are as great 
or greater than those between any other of the forms of this 
genus occurring on Antigua, Barbuda, Martinique, St. Kitts or 
Grenada. In view of this, as well as the minor external differ- 
ences, and the call difference (which must be an important isolat- 
ing mechanism), it is evident that there are two species occurring 
in the vicinity of Matouba. 

One of these, the larger, also occurs on Martinique and is very 
probably E. martinicensis. The second species is apparently 
cryptic (although not sibling). It is here named for M. Patrice 
Barlagne, who collected the majority of the specimens and aided 
Lazell in collecting on the Souffriere-Saus Toucher massif of La 
Guadeloupe. 

Eleutherodacttlus barlagnei sp. nov. 

Holotype. Adult female, MCZ 35334, collected by Patrice Bar- 
lagne and James Lazell, Jr., at Matouba, La Guadeloupe, ca. 700 
meters elevation, on 17 August, 1961. 






FiGUEE 1. (Left) Eleutherodactylus martinicensis MCZ 35322; top, side 
of head; bottom, roof of mouth. (Right ( EleutTierodactylus harlagnei sp. 
nov., holotype MCZ 35334 ; top, side of head, bottom, roof of mouth. 



1965 NEW GUADELOUPEAN FROG 3 

Faratijpes. MCZ 35330-33 (4), same data as for holotype. MCZ 
35331 is a skeleton. 

Diagnosis. An Eleutherodactylus apparently related to E. 
martinicensis and separable from that species and all other 
Lesser Antillean species of the genus by the following combina- 
tion of characters : head narrow, not wider than body ; tympanum 
small, hidden dorsally, a small tubercle in the area of the hidden 
tympanic membrane ; tympanum separated from commissure of 
jaws by three-quarters to more than the horizontal diameter 
of the tympanum ; choanae completely visible from below ; vom- 
erine tooth bosses triangular in outline, within the borders of 
and posterior to the choanae ; the voice assumed to be that of the 
new form (since the species could not be separated on other 
bases in the field) is described by Lazell as "Teeen." 

Description of holotype. Adult female (see Fig. 1) : head 
very slightly broader than long; head not broader than body; 
eyes small, width of eyelid less than interorbital distance; can- 
thus rostralis distinct, not sharp ; loreal region slightly concave, 
sloping sharply to lip ; nostrils closer to tip of snout than to eye, 
area around them swollen; length of eye greater than distance 
from eye to nostril ; tympanum small, about one-third diameter 
of eye, upper portions hidden, not distinguishable ; tubercle pres- 
ent in area where upper rim of tympanum should be ; distance 
from tympanum to commissure of mouth slightly greater than hor- 
izontal length of tympanum ; anterior edge of tympanum from 
eye about one and one-half times horizontal width of tympanum ; 
no supra- or post-tympanic fold present. 

Tongue oval, free for about one-half its length; no vocal sac 
or slits; choanae not concealed by rim of upper jaws, small, 
round, slightly smaller than area of a vomerine tooth boss ; vom- 
erine tooth bosses between and posterior to choanae, triangular 
in outline, separated by a distance about equal to width of a 
single boss. 

No axillary membrane ; no tubercles or ridges on arm ; palmar 
tubercles small ; supernumerary tubercles on palms very faint or 
lacking; subarticular tubercles large, rounded, simple; lateral 
fringe present on fingers ; no webbing between fingers ; order of 
finger length, shortest to longest, 1, 2, 4, 3 ; circular disks on 
fingers, somewhat like pads of hylids in lateral view, notch pres- 
ent ; no tarsal fold or tubercles ; inner metatarsal tubercle small, 
elongate; outer metatarsal tubercle faint, but large; no super- 
numerary tubercles on soles ; subarticular tubercles large, round, 



BREVIORA 



No. 220 



simple ; lateral fringes on toes ; pads on toes like those on fingers, 
but smaller; faint webbing on toes except for web between toes 
3 and 4 which is clearly visible for about one-half of toe 3 ; legs 
short, heels do not overlap when flexed legs are held at right 
angles to body ; heel of adpressed hind limb extends to mid-eye. 





Figure 2. (Top) Eight ilium of Eleutherodactylus martinicensis, MCZ 
35321. (Bottom) Right ilium of Eleutherodactylus harlagnei sp. nov., para- 
type, MCZ 35331. 



1965 NEW GUADELOUPEAN FROG 5 

Skin of dorsum and sides smooth, that of belly and ventral 
and posterior surface of thighs granular. 

Color in alcohol. Dorsum nearly uniform red-brown with 
scattered faint reticulations of black. Bands are evident on the 
limbs. Those of thigh are broad, three on each side. Those of 
tibia are narrower and three on each side. Two bands are present 
on tarsus and foot and two on the forelimbs. The venter is a 
dusky brown with small lighter spots. The uudersurfaces of the 
forelimbs and the area across the chest lack the dusky brown 
pigment and are yellow. 

Measurements in mm. Snout to vent 33 ; width of head 10.2 ; 
length of head 9.8 ; horizontal length of tympanum 1.2 ; length 
of eye 3.6 ; eye to nostril 3.2 ; eyelid width 2.2 ; interorbital dis- 
tance 2.6; length of tibia 13.0. 

Variation. The four paratypes are quite similar in appearance. 
In all specimens the venter is grey-cream with cream spots on 
chin and throat. The legs are flecked with brown pigment giving 
the appearance of cream flecks. All specimens have a light tri- 
angular interorbital spot, although it is weak in the holotype. 
Two specimens have a dark chevron on the dorsum (MCZ 35330, 
35333). None of the type-series shows the wide or narrow ver- 
tebral stripe, although this variation is seen in examples of E. 
martinicensis collected with the type-series. 

This species has a narrower head than does E. martinicensis. 
The head width/body length ratio ranges from 0.27 to 0.32 with 
a mean of 0.30, whereas of twenty-four E. martinicensis taken at 
the type locality by Barlagne and Lazell, the ratio varies from 
0.32 to 0.41 with a mean of 0.33. 

Comparisons. While morphologically E. harlagnei is quite 
similar to E. martinicensis, there are several differences. E. 
harlagnei has a dark venter, the tympanum appears smaller and 
is farther from the mouth and the vomerine tooth bosses are 
triangular in outline, not elongate, and do not extend laterally 
beyond the inner borders of the choanae (in E. martinice'usis 
the bosses extend laterally as far as the outer borders of the 
choanae). 

E. urichi has less prominent vomerine tooth patches which 
are round. E. johnstonei has shorter limbs and the choanae are 
not completely visible when the roof of the mouth is viewed 
from directly below. E. harhudensis has elongate vomerine tooth 
patches. 

A further difference can be noted between these frogs. In 



6 



BREVIORA 



No. 220 



connection with a study regarding the identity of Hyla harbu- 
densis (= Eleutherodactylus harhudensis) , I prepared skeletons 
of the two species found at Matouba. E. harlagnei is very dis- 
similar to E. martinicensis with regards to the form of the 
ilium. These differences are readily apparent in Figure 2. E. 
harlagnei has a thinner ilial shaft, smaller angle of ventral ace- 
tabular expansion, less elevated ilial prominence and a very short 




ACTUAL HEAD WIDTH 

Figure 3. Head width iu per cent of snout-vent length vs. actual head 
width in mm. Crosses are E. martinicensis, circles are E. harlagnei sp. nov. 
Large cross is a small E. martinicensis with a narrow head ; in other fea- 
tures it is typical of its form. 



crest beginning at the anterior edge of the ilial prominence and 
extending anterad about one and one-half times the length of 
the prominence. There is also less of a ventral acetabular ex- 
pansion in E. harlagnei. 

Acknowledgment. The collections on which this study was 
based were made under the auspices of National Science Founda- 
tion Grant NSF-G 16066. 



1965 NEW GUADELOUPEAN FROG 7 

LITEEATUEE CITED 

AUFFENBEBG, W. A. 

1958. A small fossil herpetofauna from Barbuda, Leeward Islands, 
with the description of a new species of Hyla. Quart. J. Florida 
Acad. Sci., 21: 248-254. 

Cochran, D. M. 

1941. The herpetology of Hispaniola. Bull. U. S. Nat, Mus., 177: vii + 
1-398 pp. 

Lynch, J, D. 

In press. The status of the tree frog, Hyla iarhudensis Auffenberg, from 
Barbuda, BWI. Copeia. 

Lynn, W. G. and C. Grant 

1940. The herpetology of Jamaica. Bull. Inst. Jamaica, Sci. Ser., 
1: 1-148. 

Schwartz, A. 

1957. A new species of Eleutherodactylus (Amphibia: Leptodactyli- 

dae) from Cuba. Proc. Biol. Soc. Washington, 70: 209-212. 
1958a. Four new frogs of the genus Eleutherodactylus { Leptodactyli- 

dae) from Cuba. Anier. Mus. Novit. No. 1873: 1-20. 
1958b. Another new species of Eleutherodactylus (Amphibia: Lepto- 

dactylidae) from Western Cuba. J. Washington Acad. Sci., 

April, 1958: 127-131. 
1958c. Another new large Eleutherodactylus (Amphibia: Leptodactyli- 

dae) from Western Cuba. Proc. Biol. Soc. Washington, 71 : 

37-42. 
1958d. A new frog of the auriculatus group of the genus Eleuthero- 
dactylus from Western Cuba. Herpetologica, 14: 69-77. 
1959a. A new species of frog of the Eleutherodactylus ricordi group 

from Central Cuba. Amer. Mus. Novit., No. 1926: 1-16. 
1959b. The status of Eleutherodactylus pinarensis and a new species of 

the genus from Western Cuba. Herpetologica, 15(2) : 61-69. 
1960. Nine new Cuban frogs of the genus Eleutherodactylus. Eeading 

Pub. Mus. and Art Gallery, Sci. Pubis., 11: 1-50. 

(Eeceived 22 January, 1965.) 



■- \J 



BREVIORA 

Mmseium of Comparative Zoology 



Cambridge, Mass. May 7, 1965 Number 221 

NEW MELANESTAN ANTS OF THE GENERA 

SIMOPONE AND AMBLYOPONE (HYMENOPTERA- 

FORMICIDAE) OF ZOOGEOGRAPHIC SIGNIFICANCE ^-^ 

By Robert W. Taylor 

Biological Laboratories, Harvard University 

The two ants described below are of special zoogeographic 
interest. Simopone gressitti sp. n. (subfamily Cerapachyinae) is 
the second species of its genus recorded from the Indo-Australian 
area, and the first east of the Philippines. Amhlyopone noona- 
dan sp. n. (subfamily Ponerinae) is the first apparently endemic 
Amhlyopone to be described from Western Melanesia. 

Simopone Forel includes ten described Ethiopian and Mala- 
gasian species: 8. grandicUeri Forel, 1891 (in Grandidier, Hist. 
Nat. Phys. Madagascar, 20: 141, pi. 4, fig. 8, Imerina, Madagas- 
car) ; ^S*. emcnji Forel, 1891 {ihid. 247, Anoside, Madagascar) ; 
S. conradti Emery, 1899 (Ann. Soc. ent. Belg., 43: 475, Cam- 
eroon; 1911, Genera Insect., 118: 16, pi. 1, fig. 7) ; 8.{f) mayri 
(Emery), 1900 (Bull. Soc. ent. Ital., 31: 264 (Cerapachys) ; 
1911, Genera Insect., 118 : 16, Antongil Bay, Madagascar) ; S. 
marleyi Arnold, 1915 (Ann. S. Afr. Mus., 14 (1) : 20, Stella 
Bush, South Africa) ; S. grandis Santschi, 1923 (Rev. Zool. Afr., 
11 (3) : 259, Kungu, Congo) ; 8. schoutedeni Santschi, 1923 
(ibid. : 260, fig. 1 a-c, Kamaiembi, Congo) ; ^S*. fulvinodis Santschi, 
1923 (ihid.: 262, fig. Id, Kiclaba [Kitabola], Congo) ; ^^. wilhuri 
Weber, 1949 (Am. Mus. Novit., 1396 : 7, figs. 6, 7, N. of Beni, 
Congo) ; 8. laevissima Arnold, 1954 (Ann. Mus. Congo, n.s., 4°, 
Zool., 1:291, figs. 1, la, Zika Forest, Uganda). An eleventh 
species, 8. bakeri Menozzi, 1926, was described from Singapore 
(Atti Soc. Nat. Mat. Modena, (6) 4: 92 (1925)). All these 
species are known only from the worker, except 8. mayri, which 



1 Research supported by U.S. National Science Foundation Grant No. GB. lfi.34. 

2 The specimens discussed liere were provided by Drs. J. L. Gressitt (Bishop 
Museum, Honolulu) and B0rge Peterson (Universitetets Zoologiske Museum, Copen- 
hagen), whose assistance is gratefully acknowledged. 



2 BREVIORA No. 221 

is based on a unique male and may not really belong in Simo- 
pone. 

Simopone workers are small to medium sized slender ants 
(length about 5.0-8.5 mm), usually dark brown or black in 
color, with very weak to moderately intense sculpturation and 
pilosity. The head is elongate-subrectangular, prismatic behind, 
with a transverse occipital carina. Frontal carinae horizontal, 
diverging posteriorly and obscuring the antennal insertions in 
facial view; fused anteriorly with the median part of the cly- 
peus, and forming with it and the frontal area a continuous 
planar surface, thus producing an anterior cephalic structure 
much as in the aberrant Indo-Australian myrmicine genus Meta- 
pone. Eyes very large (maximum diameter about 0.3 to 0.5 x 
the head width), situated at or just behind the middle of the 
sides of the head. Ocelli present, usually minute and closely ap- 
proximated. The 11-segmented antennae have flattened stubby 
scapes (usually only about 3 x as long as broad) which lie at rest 
in well developed preocular antennal scrobes, each of which is 
enclosed dorsally by the frontal carina and ventrally by the 
characteristically cerapachyine genal carina. These carinae usu- 
ally reach the eye posteriorly and may become continuous with a 
very fine postorbital carina, so that the eye is essentially en- 
closed within the scrobal area. Mandibles obtusely triangular, 
strongly arched ventrally ; masticatory border with a number 
of small regular teeth. The palpal formula of a single African 
specimen (species evidently undescribed) in the MCZ collection 
is maxillary 6: laMal 2, possibly 3 (inspected). 

The structure of the mesosoma is generally like that of Phyra- 
caces niayri Forel, with its dorsolateral borders broadly or nar- 
rowly rounded, sometimes angled but never carinate. Pronotum 
prismatic anteriorly, with a transverse carina between the 
humeri. A similar carina may separate the dorsal and declivitous 
faces of the propodeum and the declivitous face may be laterally 
margined. Sutural traces on mesosomal dorsum weak or ves- 
tigial, the mesometanotal suture sometimes lacking. The leg 
segments, especially the femora, are often inflated, the fore and 
hind tibiae each bear a single pectinate spur, and the pretarsal 
claws are toothed or pectinate.^ The posterior flange of the hind 
coxa may be produced dorsally as a more or less raised lamella 



8 The characters of the tibial spurs and the pretaisiil claws have been selrlom 
mentioned in specific descriptions. All specimens which 1 have seen lack tibial 
spurs on the middle legs, and have a single median tooth on each tarsal claw. 



1965 MELANESIAN ANTS d 

(another character common to many cerapachyines). Petiole 
longer than broad, subrectangular-trapezoidal in dorsal view, 
the dorsolateral margins acarinate, though sometimes angled; 
there is usually a transverse anterior carina, and sometimes 
a posterior one. Postpetiole strongly constricted behind, sub- 
rectangular in dorsal view, about equal in size to the petiole or 
larger. Pygidium flattened at its apex, with a full or reduced 
complement of bristle-like marginal setae, the presence of which 
indubitably establishes the cerapachyine affinities of this genus 
(see Brown, 1954). 

Little is known of the biology of Simopone but its general 
habitus strongly implies that it is arboricolous ; several of the 
older types were collected on vegetation and one species (;Si. 
marleyi — see Arnold, 1915) has been taken in hollow twigs. 
Specimens in the MCZ collections are either labeled as having 
been swept from foliage, or else carry adherent moth wing 
scales, an almost sure sign that they were collected by sweep- 
net. The genus is apparently an aberrant arboricolous offshoot 
from Phyracaces-like stock. The feeding biology needs study, 
especially since many cerapachyines, including some Phyracaces 
species, are apparently specialized myrmecophagous feeders 
(Wilson, 1958). 

Amhlyopone Erichson is an almost cosmopolitan genus now 
containing 50 described and apparently valid species, including 
31 from the Indo-Australian area. The world fauna was exten- 
sively reviewed by Brown (1960), and one subsequently de- 
scribed species is known (Brown, 1962). A. noonadan sp. n. is a 
member of the Indo-Australian luzonica group, which includes 
the following species: luzo7iica (Wheeler and Chapman) 1925, 
Philippines {=williamsi (Wheeler and Chapman) 1925; synon- 
ymy by Brown, I960); silvcstrii (Wheeler) 1928, Japan; am- 
hlyops (Karawajew) 1935, Indo-China; and probably also celata 
(Mann) 1919, Solomon Islands (see Brown, 1960, for details and 
references). The features distinguishing noonadan from these 
species and others present in Melanesia are given below. 

Simopone gressitti Taylor, new species 

Type locality. NEW GUINEA (WEST) : HoUandia-Binnen, 
100 m. The unique holotype worker was collected on November 
1, 1958, by Dr. J. L. Gressitt for whom this species is named. 

Type deposition. Holotype deposited in the Bernice P. Bishop 
Museum, Honolulu, Hawaii. 



BREVIORA 



No. 221 







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1965 MELANESIAN ANTS 5 

Description. Dimensions (in mm): Total length (TL) 6.4; 
head length at midline (HL) 0.98; maximum head width (IIW) 
0.79; maximum pronotum width (PW) 0.70; Weber's length of 
mesosoma (WL) 1.4; petiolar node length at midline 0.72; maxi- 
mum petiolar node width 0.69 ; postpetiolar length at midline 
0.70 ; maximum width of postpetiole 0.75. General form as shown 
in Figures 1 and 2. Mandibles small, their outer borders sinuate, 
convex basally ; angle between masticatory and posterior borders 
broadly rounded; dentition worn, almost effaced, probably orig- 
inally like that of S. hakeri. Head 0.81 x as broad as long ; sides 
almost parallel, slightly concave before eyes, slightly convex 
behind ; occipital border feebly concave ; occipital carina well 
developed, ribbed along its anterior edge, enclosing the occiput 
laterally as well as dorsally, its ventral traces extended for- 
wards for a short distance on each side, along the sides of the 
postgenae. Eyes large, maximum diameter 0.30 mm (0.38 x the 
head width), separated by a distance of 0.46 mm (1.53 x their 
maximum diameter) ; ocelli minute. Scapes barely reaching 
anterior margins of eyes, flagellar proportions as in Figure 2. 
Anterior clypeal border feebly convex. Frontal carinae diverg- 
ing posteriorly, on each side meeting the postorbital carina, which 
is very fine and continuous below the eye with the genal carina. 

Mesosoma twice as long as broad in dorsal view, its dorsolateral 
borders approximately right-angled but acarinate; transverse 
pronotal carina well developed; angle between dorsal and de- 
clivitous propodeal faces abruptly rounded, acarinate. Sutura- 
tion of mesosomal walls as shown in Figure 1 ; promesonotal and 
mesometanotal sutures represented by transversely ribbed traces 
on mesosomal dorsum, mesometanotal suture weakest. Femora 
and tibiae moderately inflated (Fig. 1) ; posterior edge of hind 
coxa raised but not lamellate ; pretarsal claws each with a single 
median tooth. Petiole trapezoidal in dorsal view, broader behind 
than in front, with a distinct ribbed transverse anterior carina; 
lateral borders strongly angled at about 60 degrees, but not 
carinate ; profile as in Figure 1. Sides of postpetiole slightly con- 
vergent posteriorly in dorsal view. Pygidial spines reduced to a 
single minute pair on each side, at the extreme apex. 

Mandibles smooth and shining, with a few minute piligerous 
punctures. Entire body strongly shining, virtually lacking sculp- 
turation except for scattered minute piligerous punctures, and 
some effaced longitudinal rugosity along the sides of the pronotal 
dorsum and on the metepisternum, which is in part coarsely 
punctate-rugose. Pilosity reduced. A few moderately long 



6 BREVIORA No. 221 

suberect to reclinate hairs on mandibles, clypeus and underside 
of head, propleurae, coxae, and undersides of petiole and gaster ; 
hairs most abundant on the propleurae and coxae, and on the 
petiolar sternite where they form a peculiar brush-like series 
behind the subpetiolar process. Single, slightly longer erect 
hairs in the following positions : at the midlength of each frontal 
carina, above eyes, on pronotal humeri, on the anterior half of 
the sides of the node (2 pairs) and the anterodorsal corners of 
the postpetiole. Similar, but slightly less erect hairs increasingly 
long and abundant towards the gastric apex, which is surrounded 
by very long arched hairs. Pubescence virtually absent. Color 
very dark brown, almost black, the following areas weakly in- 
fuscated with reddish brown : mandibles and front of head, an- 
terior parts of each gastric tergite, gastric apex and legs, es- 
pecially the tibiae and tarsi; antennae medium dull reddish 
brown. 

Diagnosis. According to Menozzi's description (Menozzi, 1926) 
S. hakeri is smaller than gressitti, with a narrower head and 
petiolar node. I have tentatively identified as 8. hakeri a speci- 
men in the J. W. Chapman collection (MCZ) from the Philip- 
pine Island of Negros (Horns of Negros 3,600 ft., J. W. Chap- 
man). 

This individual agrees well with Menozzi's description, and 
resembles gressitti in color and habitus, but has very different 
cephalic, ocular and petiolar proportions as follows : TL c. 5 mm ; 
HL 0.91 mm; HW 0.68 mm (head 0.75 x as broad as long) ; PW 
0.55 mm; WL 1.2 mm; petiolar node length at midline 0.68 mm; 
maximum petiolar node width 0.56 mm (node 0.52 x as broad as 
long); postpetiolar length at midline 0.64 mm; maximum 
width of postpetiole 0.61 mm. The maximum diameter of the 
eyes is 0.30 mm (0.44 x head width) and they are separated 
by a distance of 0.34 mm (1.14 x their maximum diameter). 
Apart from these proportional differences the post-cephalic 
structure is similar to that of gressitti. The pilosity is similarly 
distributed but less abundant, and the subpetiolar "brush" is 
lacking. 

Amblyopone noonadan Taylor, new species 

Type locality. TERRITORY OF NEW GUINEA : New Brit- 
ain: Yalom, 1,000 m. May 19, 1962 (Danish Noona Dan Ex- 
pedition). The types were collected "in or on the ground in 
newly cleared secondar.v growth," no collector specified. 



1965 



MELANESIAN ANTS 



Type deposition. The holotype is deposited in the Universi- 
tetets Zoologiske Museum, Copenhagen, Denmark; the paratype 
is in the Museum of Comparative Zoology (Type No. 31148). 
The species is named, in apposition, for the Danish expedition 
vessel Noona Ban. 

Worker. The following description is based on the holotype 
and single paratype. 

Dimensions (in mm, holotype cited first). TL c. 5.5, 6 mm; 
HL (including clypeal denticles) 1.04, 1.12; HW (immediately 
behind genal teeth) 0.96, 1.04; maximum scape length (exclud- 
ing articular condyle) 0.64, 0.68; outside total length of man- 
dible 0.85, 0.90 ; PW 0.58, 0.65 ; WL 1.38, 1.42 ; midline length 
of petiolar dorsum 0.49, 0.55 ; maximum petiolar node width 
0.50, 0.57 ; postpetiolar length at midline 0.36, 0.40 ; maximum 
width of postpetiole 0.65, 0.73. 

General habitus as in Figures 3 and 4. Head with occipital 
border feebly concave, sides feebly convex, converging posteriad ; 




Figs. 3 and 4: Amblyopone noonadan sp. n. Fig. 3. Lateral view of meso- 
soma and node. Fig. 4. Frontal view of head, right antenna omitted. Scale 
line 1 mm. Holotype worker. 



8 BREVIORA No. 221 

anterior corners with strong acute genal teeth, the inner edges 
of which are about as long as the maximum width of the mandib- 
ular shafts. Frontal lobes approximate, separated by a deep 
linear groove. Clypeal apron strongly convex, with eight small 
denticles ; the four median ones closely approximate, less deeply 
separated from each other than from the more lateral denticles, 
their apices diverging from the midline ; the innermost of the 
two lateral denticles on each side moderately large, triangular, 
separated from the median quartet by a gap equal to its width 
at base ; the outer tooth large and blunt, its apex jagged, form- 
ing two or three indistinct cusps. Mandibles slender, their ex- 
ternal margins feebly concave, each bearing ten acute, slightly 
recurved teeth. The two basal teeth simple, conical, the basal- 
most blunt, the second acute ; the eight apical teeth arranged in 
four more or less separated pairs, in typical "stigmatommine" 
fashion ; the dorsalmost tooth of each pair lies slightly distal to 
its partner ; a distinct low reclinate tooth is present on the inner 
edge of the strong acute mandibular apex. The mandibular 
apices cross when the jaws are closed, leaving a triangular gap 
between the mandibular and clypeal teeth. Palpal formula 
maxillary 4: labial 3 (paratype dissected). Scapes slender, 
slightly incrassate ; funiculus with 11 segments proportioned as 
in Figure 4. Eyes small, variable in size, maximum diameter 0.02 
mm in holotype, 0.06 mm in paratype, with four and nine or ten 
indistinct facets, respectively. 

Mesosomal profile as in Figure 3. In dorsal view this tagma 
is widest at the pronotum and strongly narrowed at the base of 
the propodeum. Pronotal dorsum almost flat, with marginate 
lateral borders; inferior angles of pronotum broadly rounded. 
Promesonotal suture flexible ; mesonotum transverse ; mesometa- 
notal suture A^estigial, represented only by a sculptural break 
between the subopaque mesonotum and the moderately shining 
propodeum. Propodeal dorsum about as long as broad, its sides 
diverging posteriad; declivity feebly concave, rounding into 
dorsum, its lateral edges slightly raised, forming angles of a lit- 
tle more than 90 degrees in dorsal view. 

Petiole sessile, its profile as in Figure 3 ; subpetiolar process 
afenestrate; nodal dorsum slightly wider than long in dorsal 
view, the anterior border Avith a slight median emargination, the 
sides converging slightly anteriad. Postpetiole wider but shorter 
than node, and also shorter than the succeeding segment which 
is of about the same width. Gastric apex laterally compressed, 



1965 MELANESIAN ANTS 9 

sting stout. Tibial spurs vestigial on middle legs ; posterior tibiae 
each with a broad flat pectinate spur and a more slender simple 
conical one. 

Mandibles and frontal lobes obscurely longitudinally striate. 
Clypeal apron with somewhat radial longitudinal striae; lateral 
parts of clypeus similarly sculptured, the striae on each side 
radiating back from a focal point at about the level of the inner 
basal edge of the mandible. These lateral clypeal striae arch 
back over the cheeks, where those nearest the midline are 
almost longitudinal, reaching back to the base of the frontal 
carinae; the cheek striae become increasingly divergent towards 
the sides of the head and the most lateral ones gather apically, 
at the base of the genal tooth. Remainder of head coarsely and 
roughly punctate-rugose ; the sculptural trend faint, mainlj^ 
longitudinal, but transverse across a narrow posterior strip. 
Scapes finely shagreened. Postgenae somewhat obscurely and 
irregularly longitudinally striate, the striae diverging posteriad. 
Lateral parts of the dorsa of the pronotum and propodeum with 
scattered punctures, about 0.02 mm in diameter, separated by 
about twice this distance on pronotum and more widely spaced on 
propodeum ; a narrow longitudinal median strip on these scler- 
ites lacks punctures, the surface here is shining, with a very 
fine superficial scale-like surface pattern which is also present on 
the interpunctural areas of the lateral strips, and which has a 
transverse trend on the posterior propodeal dorsum. Mesonotum 
subopaque, coarsely and irregularly shagreened. 

Sides of mesosoma, except metepisternal area, subopaque, 
bearing somewhat effaced and polished, almost vertical fine 
striae, which are slightly curved (concave anteriorly) and slope 
posteriorly; sculptural intensity diminishing posteriad, with 
striae virtually absent behind the propodeal spiracle. Metepister- 
nal area longitudinally striate. Declivity of propodeum shin- 
ing, with very superficial, minutely scale-like transverse sculptur- 
ation. Petiolar dorsum subopaque, with scattered fine piliger- 
ous punctures ; gastric tergites similar, the punctures finer and 
more abundant. 

Pubescence adpressed and subadpressed, generally distributed 
over body except for the post-pronotal sides of the mesosoma and 
the sides of the petiolar node. Erect pilosity moderately 
abundant, especially on the dorsum of the body and towards 
the gastric apex, where the hairs are longest. Ground color dark 



10 BREVIORA No. 221 

chocolate browii, with the following areas infuscated with red- 
dish brown: clypeus, frontal lobes, anterior corners of head, 
including genal teeth, pronotal collar, area of mesosomal-petiolar 
junction, subpetiolar process, posterior edges of gastric tergites, 
and gastric apex. Mandibles, antennae, legs and sting rich 
golden brown. 

Diagnosis. A. noonadan is readily distinguished from the only 
other known western Melanesian Amhlyopone, A. australis 
Erichson, by the characters of its " stigmatommine habitus" (i.e., 
"double ranked" mandibular dentition and enlarged clypeal 
teeth, etc., — see Brown, 1960). In addition, australis is larger 
(minimum known HW on New Guinea about 1.5 mm), and has 
an acute tooth on each inferior pronotal angle. The third known 
Melanesian species, A. celata Mann (Solomon Islands), is much 
smaller (3 syntype workers in the MCZ collection have HL 
0.69-0.71 mm; HW 0.60-0.63 mm) with minute genal teeth (max- 
imally only about as large as the median clypeal denticles) and 
with the head evenly and rather finely shagreened. 

The other luzonica group species have reduced genal teeth, 
as in celata, and most specimens (MCZ collection) are somewhat 
smaller than the noonadan types : 3 silvestrii syntypes have HL 
0.87-0.92 mm; HW 0.76-0.81 mm; the luzonica female holo- 
type from Los Banos (Luzon) has HL 0.82 mm, HW 0.74 
mm; and six luzonica workers from Dumaguete (Negros) 
have HL 0.85-0.90 mm, HW 0.74-0.80 mm. Three williamsi 
syntype workers from Baguio (Luzon) are somewhat larger 
with HL 1.03-1.09 mm, HW 0.91-0.97 mm, and Karawajew 
(1935) gave similar measurements for his amhlyops type — 
HL 1.05 mm, HW 0.98 mm. These "species" differ among them- 
selves in the conformation of the clypeal denticles and in the 
sculpturation (Brown, 1960, p. 196), but they may ultimately 
prove to be geographical variants of a single species, especially 
considering the extensive variation known in other members of 
the genus. Notwithstanding, there can be little doubt that noona- 
dan and celata are good "species. 



5 J 



EEFEEENCES 

Arnold, G. 

1915. A monograph of the Formicidae of South Africa (Ponerinae; 
D.orylinae). Ann. S. Afr. Mus., 14 (1): 1-159. 
Brown, W. L., Jr. 

1954. Eemarks on the internal ph.ylogeny and subfamily classification 
of the family Formicidae. Insectes Soc, 1: 22-31. 



1965 MELANESIAN ANTS 11 

1960. Contributions toward a reclassification of the Formicidae. III. 
Tribe Amblyoponini (Hymenoptera). Bull. Mus. Comp. Zool., 
112 (4): 145-230. 
1962. A new ant of the genus Amblyoponc from Panama. Psyche, Cam- 
bridge, 69 (2): 73-76. 
Karawajew, W. 

1935. Neue Ameisen aus dem Indo-Australischen Gebiet. Treubia, 15 
(1): 57-117. 
Menozzi, C. 

1926. Nuove formiche delle isole Filippine e di Singapore. Atti Soe. 
Nat. Mat. Modena, 6 (4): 92-103 (1925). 
Wilson, E. O. 

1958. Observations on the behavior of the cerapachyine ants. Inseetes 
Soc, 5: 129-140. 



^v 



,iU 



BREVIORA 

MniseMiii of Compsirative Zoology 

Cambridge, Mass. May 28, 19G5 Number 222 

THE GENUS LEPTOTYPHLOPS IN THE WEST INDIES 

WITH DESCRIPTION OF A NEW SPECIES FROM HIS- 

PANIOLA (SERPENTES, LEPTOTYPHLOPIDAE) 

By Richard Thomas 

10,000 SW 84tli St., Miami, Florida 33143 



Three specimens of Leptotyphlops were collected in the summer 
of 1964 in the vicinity of the town of Pedernales, Dominican 
Republic, by Mr. David C. Leber and myself. These snakes ap- 
pear most closely related to L. hilineata Schlegel which they re- 
semble in the failure of the ocular to reach the labial border. 
No snakes of the genus Leptotyphlops have previously been 
definitely noted from the island of Hispaniola. 

Boulenger (1893), however, recorded a specimen of Lepto- 
typhlos alhifrons Wagler from Santo Domingo de Guzman col- 
lected by Dr. A. C. Buller. It so happens that the name of the 
capital city of the Dominican Republic, rarely referred to in its 
entirety, is Santo Domingo de Guzman. The question of the 
provenance and relationships of the Buller specimen might 
therefore be important. Thanks to Miss A. G. C. Grandison, I 
have been able to examine this specimen (BM 90.10.10.73) and 
can confirm its affinities with the alhifrons group of the genus. 
It appears to agree most closely with L. phenops hakewelli Oliver 
although the rostral-prefrontal fusion is apparently lacking. 
(Dunn and Saxe, 1950, regard phenops as a race of alhifrons; 
but I here follow more recent authors such as Peters, 1954, and 
Duellman, 1961, in giving phenops specific rank.) Miss A. G. C. 
Grandison writes (in litt.) that "Dr. Audley C. Buller . . . made 
quite extensive collections in Mexico in 1891 and 1892, travelling 
from L. [Lago] Chapala and Guadalajara ... to Bolafios and 
back to Ixtlan and later ... to an area west of Guadalajara." 
Examination of a recent map of Mexico shows that roughly 200 
km to the soutli of the city of Guadalajara in the state of Jalisco 
is a Ciudad Guziiuni, whicli may well he another case of the un- 
wieldy Santo Domingo de Guzman liaving been sliortened. Smith 



2 BREVIORA No. 222 

and Taylor (1945) list L. phenops hakewclli from the states of 
Colima, Guerrero, Jalisco, Michoacaii, and Oaxaea. That identity, 
collector, locality and time should all approximately coincide 
seems too remarkable for mere coincidence. This specimen (BM 
90.10.10.73)1 is here considered to be close to L. p. hakewclli 
from what is now Ciudad Guzman, Jalisco, Mexico, and hence 
irrelevant to the Hispaniolan problem. 

The specimens from Pedernales, however, are distinct from 
other species and apparently represent an endemic form, here 
named in allusion to the type locality - : 

Leptotyphlops pyrites new species 

Holotypc: MCZ 77239, collected at the southern outskirts of 
the town of Pedernales, approx. 1 km from the center of town, 
Pedernales Province, Dominican Republic, 3 July 1964, by 
Richard Thomas. 

Paratypes: USNM 152452, same locality as type, 26 June 1964, 
Richard Thomas; ASFS V2601, 9 km n' Pedernales, Pedernales 
Province, Dominican Republic, 26 June 1964, David C. Leber. 

Diagnosis: A species of Leptotyphlops of closest affinities to 
hilincata in that the second and third upper labial scales exclude 
the ocular from the labial border. It is further characterized by 
considerable attenuation, a high number of middorsal scales 
(from rostral to tail spine), 12 scale rows around the base of the 
tail, and 15 to 16 subcaudals, a trilineate dorsal pattern and 
unicolor, dark sides and venter. 

Distributio7i: Known presently only from the northwestern 
lowlands of the Barahona peninsular region of Hispaniola. 

Description of holotypc (Fig. 1) : Head rounded, of same width 
as neck. Rostral at widest point slightly less than width of head 
at eyes, broadly truncate posteriorly at contact with prefrontal. 
Nasals separated by a transverse suture proceeding from first 
labial diagonally upward across naris to rostral ; dorsal half of 
nasal also in contact with rostral, prefrontal, supraocular, ocular. 



IBM 90.10.10.7.1 : Total length ca. ITjI luin ; tiiil 7.(> iiiui : middorsal scales 
al)out 24() : siibcaiKhils ]<i: scale rows 14: 10 scale row.s around tail. Rostral e.\- 
tends posteriorly to slightly beyond level of eye, does not contact supraoculars. 
Suture coniplefely divides nasals : supiaoculars elongate, slanting sharply forward. 
One supralahial lielweeii ociilolahial and nasal, one behind ocuhd.ihial. Light spot 
on I'ostral scale and on underside of tail from l:!tli scale aiilcrinr tn caudal spine 
and including caudal spine. Dorsum darli, liglit scale edges form lines; anterior 
venter light, becoming darker posteriorly. 

2 "Pedernales" means "flints," in Si)anish. hence "pyrites," the Greek I'lpiiva- 
Icnt. 



1965 



WEST INDIAN LEI^TOTYPIILOPS 



and first and second supralabials. Four supralabials, first small- 
est, second and third of about equal size, fourth largest; second 
and third supralabials in contact with ocular, occluding it from 
contact with labial border. Prefrontal, frontal and interparietal 
(third middorsal scale) increase in size in that order; middorsal 





Fig. 1. Dorsal and lateral views of the head of the type specimen of Lepto- 
typhlops pyrites (MCZ 77239). 



scales posterior to the interparietal decrease gradually to stand- 
ard body scale size. Parietal and fourth upper labial in broad con- 
tact behind ocular. Parietal and postparietal distinctly enlarged, 
parietal the larger. Supraocular about equal in size to prefrontal. 
Middorsal scales 273 from rostral to caudal spine; 14 scales 
around body ; 12 around base of tail ; subcaudals 15. Anal scale 
enlarged and shield shaped, tri-lobate posteriorly. Body very 
slender, of nearly uniform diameter throughout ; total length 133 
mm; diameter at midbody (2.0 mm) into total length about 67 
times; length of tail into total length 24 times. 



BREVIORA 



No. 222 



Coloration (Fig. 2): Ground color of body chestnut; mid- 
dorsal scales from prefrontal back, chestnut with faint lighter 
(tannish brown) lateral corners. First paramedian dorsal rows 
(including supranasals, supraoculars, parietals and postparie- 
tals) plus dorsal third of second paramedian rows tannish 
brown. Central axis of first paramedian dorsal rows suffused 
with darker brown forming a thin darker line on these rows, 




A 



B 



Fig. 2. A, color pattern at midbody of L. pyrites; B, color pattern at 
midbody of L. iilineata. 



resulting pattern trilineate on light middorsal zone (Fig. 2A) ; 
middorsal stripe slightly club-shaped anteriorly because of en- 
largement of median cephalic scales. Upper and lower labial 
margins light, nearly continuous with light areas on top of head 
flanking median dark stripe ; paramedian dark lines commence 
on first scale behind second parietal; rostral faintly suffused 
with brown. 

Variation: Head scalation in the two paratypes is much the 
same as that of the type. Total length of each paratype 123 mm ; 
diameters at midbody (1.9 and 2.0 mm) go into total length 
65 and 61 times; tail lengths into total lengths go 21 times for 
each specimen. Middorsal scales 269 and 283; 15 and 16 sub- 
caudals; 14 scale rows and 12 scales around the base of tail in 
both paratypes. Color patterns the same as that of the type. 

Comparisons and discussion: As noted previously, L. pyrites 
is most closely related to hilineata, described by Schlegel from 
Martinique ; hilineata is additionally recorded from Guadeloupe 
(Dumeril and Bibron, fide Barbour, 1914), Barbados (Boulenger, 
1893), and St. Lucia. (See also below, specimens examined.) 



1965 WEST INDIAN LEPTOTYPHLOPS 5 

Underwood (1962) indicated that bilineata occurs on the main- 
land of South America, but he has informed me (m litt.) that this 
was a mistake. There appear to be no other records of Mlineata 
from the mainland. The Guadeloupe record has not been sub- 
stantiated in recent years. Dr. Guibe of the Museum National 
d'Histoire Naturelle, Paris, writes (tn litt.) that there are no 
specimens from Guadeloupe in that institution, and that he has 
not been able to find any indication of specimens from Guade- 
loupe. All of the specimens collected by Guyon and Plee have the 
locality indicated as Martinique. It seems best, therefore, to re- 
gard the Guadeloupe record as erroneous. ^ 

Of the eight specimens of bilineata examined from Martinique, 
St. Lucia and Barbados, middorsal scales range from 170 to 
189, in the neighborhood of one hundred scales lower than the 
counts for pyrites (269-283) ; scale rows are 14 in all cases, al- 
though two specimens reduce to 12 and 13 just anterior to the 
vent. Subcaudals range from 12 to 14 (15-16 in pyrites) ; scale 
rows around the base of the tail are 10 (11 in one) in all speci- 
mens from Martinique and St. Lucia versus 12 for pyrites. The 
Barbados specimen has 12 scale rows around the tail. Total 
lengths for the bilineata specimens 60-108 mm, for pyrites 123- 
133 mm. The ratio of midbody diameter into total length ranges 
from 34.6 to 43.2 in bilineata, 61.0-66.5 for pyrites. The ratio of 
tail length into total length varies from 15 to 18 for bilineata, 
21-22 for pyrites. It can be seen that the two species differ 
abundantly in body scalation and proportions. In head scala- 
tion there are no constant differences between the two. In colora- 
tion, once again, the differences are striking (Fig. 2B). The 
unicolor middorsal zone, dorsolateral light lines, dark sides and 
light venter of bilineata contrast strongly with the more complex 
dorsal pattern and uniformly dark sides and venter of pyrites. 
L. bilineata also has a rather large patch of light (cream or 
yellow) scales surrounding the cloaeal opening, while pyrites 
does not (the lips of the cloaca are light, but the scales surround- 
ing it are not). In addition to having 12 scales around the tail, 
the Barbados specimen also has the highest middorsal scale count 
(189). Klauber (1940) has found the number of scales around 
the tail to be a useful differential character ; it is possible that the 



3 Mr. James D. Lazell, Jr. states (pers. comm.) that the natives of Guadeloupe 
spoke of two kinds of "two-headed" snakes (i.e. Tmihlopa or Leptotyphlops) , of 
which one was said to have a lineate pattern and to inhabit the hotter, dryer 
parts of the island — a nather accurate and concise characterization of L. Mlineata. 



6 BREVIOBA No. 222 

Barbados snakes are distinct from those of Martinique and St. 
Lucia. 

Leptotyphlops alhifroiis Wagler has been recorded from An- 
tigua, Grenada (Boulenger, 1893), and from Swan Island (Dunn 
and Saxe, 1950) in the West Indies; it also occurs on the South 
American mainland. This species (and its relatives in the albi- 
frons group) is quickly distinguished from pyrites and hilineata 
by the extension of the ocular scale to the labial border; it is 
also characterized by a finely lineate dorsal and ventral pattern, 
and a light spot on the rostral and one on the tip of the tail. 
The name "alhifrons" is used here with some reservations. I 
have examined a number of Leptotyphlops from South America 
pertaining to this group, but unfortunately none from the type 
locality of albifrons (Para, Brazil). There is obviously more 
than one species involved in the material I have seen, but I have 
not tried to determine which of the profusion of available names 
applies to which forms except where West Indian specimens are 
concerned. 

Examination of the specimen of albifrons recorded by Boul- 
enger from Antigua shows it to agree closely, both in coloration 
and supraocular-first labial contact, with specimens from Trini- 
dad to which the name L. tenella "^ has been given (Klauber, 
1939). No further specimens of Leptotyphlops from Antigua 
have come to light, not even under the more intensive herpeto- 
logical collecting of the region in recent years. The locality for 
the specimen is possibly incorrect; it is probably best to regard 
the Antiguan record for Leptotyphlops as problematical until 
specimens are obtained or the negative evidence becomes 
stronger. Mr. Wayne King has informed me (pers. comm.) that 
the record of Leptotyphlops from the nearby island of Barbuda 
(Auffenberg, 1958) is in error and was due to an incorrectly 
identified Typhlops. 

The record of L. albifrons from Grenada (Boulenger, 1893) 
is based on two specimens in the British Museum. Miss Grandi- 
son has advised me that the correct datum for these specimens is 
New Granada. As New Granada is the old name for the South 



4 Subsequent to its proposal, the name tenella has been regarded variously as 
applying to a distinct species with sijeciniens reported from as far south as the 
Brazilian state of Mato Grosso (Bailey and Carvalho, 1940)) or as a race of alhijrons 
inhabiting northeastern South America (Roz<i, 19.52). It is my feeling that tenella 
represents either a subspecies of a wide-ranging South American form (presum- 
ably true albifronn) or a variant (supraoculars and lirst labials in contact) which 
occurs throughout mucli of the range of albifronR (as far south as Mato Grosso) 
but is of particularly high fre(inency in the northeast. However, my knowledge of 
the complex of forms in the albifrons group is too meager to venture a solution to 
the problem at this time. 



1965 WEST INDIAN LEPTOTYPIILOPS 7 

American country of Colombia, these specimens can no longer 
be regarded as pertaining to the Lesser Antillean island of 
Grenada. 

The New Granada specimens are indeed of the alhifrons group 
but do not pertain to that species. They have uniforml}' dark 
colored tails (with the exception of the terminal light spot) and 
relatively high middorsal scale counts (245, 253) ; they possibly 
pertain to a form or complex of forms including margaritae Roze 
(1952, described as a subspecies of alhifrons) from Isla Mar- 
garita, off of Venezuela, and melanoterma Cope (1862) described 
from Corrientes, Argentina. 

L. alhifrons magnamaculata Taylor is known from Swan 
Island in the West Indies and additionally from San Andres, 
Providencia, and the Bay Islands of Honduras, including Utila 
(type locality). It is supposedly distinguished by a larger 
white spot on the snout and a more extensive spot under the 
tail (Taylor, 1940), and more vivid markings (Dunn and Saxe, 
1950). 

Leptotyphlops columhi Klauber is known only from Watlings 
Island in the Bahamas, over 400 miles from its nearest congener. 
The Bahamas are as a whole poorly collected, and it may well be 
found to occur on other islands. L. columhi too can be distin- 
guished from pyrites in the possession of an oculo-labial contact ; 
it is further distinguished by a high subcaudal count (22-25 : 
Legler, 1959), and a nearly uniform dark dorsal coloration but 
paler venter {op. cit.). Klauber used the coloration and the 
high subcaudal count to distinguish it from forms of the alhi- 
frons group (however, Legler 's new data indicate an overlap 
with the highest "alhifrons" counts of 23 noted by Klauber, 
1939, and myself). 

Key to the West Indian Leptotyphlops 

1. Ocular excluded from labial border 2 

Ocular extends to labial border 3 

2. Middorsal scales (between rostral and terminal spine) 170-189, 

venter light Mlineata 

Middorsal scales 269-283, venter dark pyrites, n. sp. 

3. Light spot on snout and tip of tail 4 

No light spot on snout and tip of tail, coloration generally dark above, 
lighter below columhi 

4. Supraoculars contact first supralabials tenella 

Supraoculars not in contact with first 

supralabials alhifrons magnamaculata 



8 BREVIORA No. 222 

The strange apparent distribution of Leptotyphlops in the 
West Indies should be noted (Fig. 3). There has been a tendency 
to dismiss the erratic insular occurrence of these small snakes as 
due in large part to introduction by man, both pre- and post- 
Columbian. While it must be admitted that their small size and 
burrowing habits make them likely candidates for artificial 
transportation, we have no evidence that this has occurred. Such 
thinking is an easy way out of facing what might well be a real 
but complex zoogeographic problem. To begin with, the distri- 
bution of Leptotyphlops in the Lesser Antilles is perhaps no 
more unusual than the non-uniform distribution of several other 
forms in this region (cf. Leptodactylus, Ameiva, Gymnophthal- 
mus, Bothrops, Constrictor). The non-uniform distribution of 
other forms is attributable in part to the erratic nature of nat- 
ural dispersal across water and in part perhaps to selective 
extinction on some islands ; in the case of creatures like Lepto- 
typhlops some of the gaps may be more apparent than real 
due to incomplete collecting. 

Not all cases are, in any event, erratic. L. a. magnamaculata 
occurs on marginal islands (Swan Island) that are close to the 
mainland; it is closely allied to the mainland alhifrons repre- 
sentative (Dunn and Saxe, 1950) ; its distribution, therefore, 
is not particularly remarkable or unexpected. Although the 
specimen of Antiguan tenella is regarded as being of question- 
able provenance, the occurrence of a form of strong South 
American affinities so far up the chain would not be unprece- 
dented if this record is verified. Thus, the gecko Phyllodactylus 
occurs on Puerto Rico and the adjacent Caja de Muertos; the 
nearest records to the south for this genus are Grenada and 
Barbados. Leptotyphlops pyrites and L. columbi are the Antil- 
lean forms found farthest from the mainland and whose distri- 
butions are the most irregular. The fact that L. pyrites has an 
obvious relative in the Lesser Antilles and apparently none on 
the mainland strongly bespeaks a relict distribution in the West 
Indies. The exact relationships of L. columbi are uncertain ; 
although Klauber suggested a closer affinity with alhifrons than 
with other forms, it is apparently not close. L. columbi may 
either represent a fortuitous arrival in the Bahamas or a relict 
distribution. Its apparent distinctness would seem to speak 
against its having been artificially introduced, as has been sug- 
gested (Darlington, 1957:221). 

I wish to express my appreciation to Dr. Albert Schwartz who 
supported this study and the collecting that resulted in the 



1965 WEST INDIAN LEPTOTYPHLOPS 9 

acquisition of the form described. I also wish to thank the fol- 
lowing people for their help in various ways: Pere R. Pinchon 
of the Seminaire College (SC), Fort-de-France, Martinique, for 
the loan of specimens and for the donation of a specimen to the 
ASFS (Albert Schwartz Field Series) collection ; Miss A. G. C. 
Grandison of the British Museum (Natural History) (BM) 
for the loan of specimens and the biographical information on 
Dr. A. C. Buller; Dr. James A. Peters of the U. S. National 
Museum (USNM) for the loan of specimens and help in finding 
Leptotyphlops names and literature; Dr. Ernest E. Williams 
of the Museum of Comparative Zoology, Harvard (MCZ) for 
loan of specimens and help in obtaining literature ; Neil D. 
Richmond of the Carnegie Museum (CM) for the loan of speci- 
mens; and Mr. David C. Leber for his most able assistance in 
the field. RT designates specimens in the Richard Thomas 
private collection. 

Specimens Examined 

Leptotyphlops pyrites: As listed for the type and paratypes. 

Leptotyphlops hiliiieata: Martinique: BM 53.2.4.36, USNM 
119168; Fort-de-France, SC 1; Tartane, Morne Jesus, SC 2-3, 
ASFS V4150. St. Lucia: MCZ 10693. Barbados: BM 89.7.5.27. 

Leptotyphlops " albifrons" (no nomenclatural finality in- 
tended) : Specimens in the American Museum of Natural His- 
tory and the University of Michigan Museum of Zoology collec- 
tions, Brazil (1), Bolivia (7), Peru (16). 

Leptotyphlops tenella: Trinidad: St. George Co.: Mt. St. 
Benedict, CM 4888-89, 4892, 6612 (paratypes) ; El Dorado, CM 
4893 (paratype) ; Arima Ward, Santa Cruz Valley, 7.5 mi. N 
San Juan, RT 1186. ?Antigua: BM 50.4.29.3. 

Leptotyphlops cf. margaritae : New Granada : BM 80.2.26.4-5. 

Leptotyphlops phenops cf. hakewelli: Santo Domingo de 
Guzman, BM 90.10.10.73. 

LITERATUEE CITED 

AUTFENBERG, WALTER 

1958. A small fossil herpetofauna from Barbuda, Leeward Islands, 
with the description of a new species of Hyla. Quart. Jour. 
Florida Acad. Sci., vol. 21, no. 3, pp. 248-254. 
Bailett, Joseph R., and A. L. de Carvalho 

1946. A new Leptotyphlops from Mato Grosso, with notes on Lepto- 
typhlops tenella Klauber. Bol. Mus. Nac, Zool., no. 52, pp. 1-7, 
4 figs. 



10 BREVIORA No. 222 

Barbour, Thomas 

1914. A contribution to the zoogeography of the West Indies, with 
especial reference to amphibians and reptiles. Mem. Mus. Comp. 
Zool., vol. 44, no. 2, pp. 209-346. 

BOULKNGEB, GeORGE ALBERT 

1893. Catalogue of snakes in the British Museum (Natural History). 
London, vol. 1, xiii -|- 448 pp. 
Cope, E. D. 

1862. Catalogues of the reptiles obtained during the explorations of 
the Parana, Paraguay, Vermejo, and Uraguay rivers, by Capt. 
Thos. J. Page, U.S.N, and of those procured by Lieut. N. Michler, 
U. S. Trop. Eng., Commander of the expedition conducting the 
survey of the Atrato River. Proc. Acad. Nat. Sci. Philadelphia, 
pp. 346-359. 
Darlington, Philip J., Jr. 

1957. Zoogeography: the geographical distribution of animals. New 
York. John Wiley & Sons, Inc., pp. i-xi, 1-675, 80 figs. 
Duellman, William E. 

1961. The amphibians and reptiles of Michoacan, Mexico. Univ. Kansas 
Publ. Mus. Nat. Hist., vol. 10, pp. 1-148, 6 pis., 11 figs. 
Dunn, E. R. and L. H. Saxe, Jr. 

1950. Results of the Catherwood-Chaplin West Indies Expedition, 
1948. Part V. Amphibians and reptiles of San Andres and 
Providencia. Proc. Acad. Nat. Sci. Philadelphia, vol. 102, pp. 
141-165. 
Klaubek, Laurence M. 

1939. Three new worm snakes of the genus LeptotypMops. Trans. San 

Diego Soc. Nat. Hist., vol. 9, no. 14, pp. 59-66, 6 figs. 

1940. The worm snakes of the genus Leptotyphlops in the United 
States and northern Mexico. Trans. San Diego Soc. Nat. Hist., 
vol. 9, no. 18, pp. 87-162, 8 figs., 6 pis., 2 maps. 

Legler, John M. 

1959. Notes on the snake Leptotyphlops columhi Klauber. Herpeto- 
logica, vol. 15, p. 112. 
Peters, James A. 

1954. The amphibians and reptiles of the coast and coastal sierra of 
Michoacan, Mexico. Occ. Pap. Mus. Zool. Univ. Michigan, no. 
554, pp. 1-37. 
RozE, Janis a. 

1952. Contribucion al conocimiento de los ofidios de las familias Typhlo- 
pidae y Leptotyphlopidae en Venezuela. Mem. Soc. Cien. Nat. La 
Salle, vol. 12, no. 32, pp. 143-158. 
Smith, Hobart M. and Edvpard H. Taylor 

1945. An annotated checklist and key to the snakes of Mexico. Bull. 
U. S. Natl. Mus., vol. 187, iv + 239 pp. 



1965 



WEST INDIAN LEPTOTYPIILOPS 



11 



Taylor, Edward H. 

1940. Herpetologieal luisi-ellany no. I. Univ. Kansas Sci. Bull., vol. 26, 
no. 15, pp. 489-571, 5 pis., 7 figs. 
Underwood, Garth 

1962. Caribbean affairs No. 1. Reptiles of the eastern Caribbean. 
Dept. Extra-Mural Studies, Univ. West Indies., pp. 1-191. 

(Received 16 February, 1965.) 



Total 



Tail 



Mid- 



Sub- Rows, Midbody 



pyrites 

USNM 152452 
ASFS V2601 
MCZ 77239 
hilineata 
MCZ 10693 
BM 53.2.4.36 
SC 1 
SC 2 
SC 3 

ASFS V4150 
USNM 119168 
BM 89.7.5.27 

Table 1. Data for individual specimens of L. pyrites and L. hilineata 
examined. Measurements are in millimeters. 



lengtii 


lengtii 


dorsals 


caudals 


tail 


diameter 


123 


6 


283 


15 


12 


1.9 


123 


6 


269 


16 


12 


2.0 


133 


6 


273 


15 


12 


2.0 


74 


4 


182 


13 


10 


1.7 


108 


7 


183 


14 


11 


2.5 


107 


6 


174 


12 


10 


3.1 


93 


5 


181 


14 


10 


2.4 


60 


4 


187 


14 


10 


1.6 


108 


6 


175 


13 


10 


2.9 


102 


7 


170 


13 


10 




102 


6 


189 


13 


12 





12 



BREVIORA 



No. 222 




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BREVIORA 



Museum of Comparative Zoology 



Cambridge, Mass. May 28, 1965 Number 223 

A NEW SUBSPECIES OF CLE LI A CLE LI A (SERPENTES : 
COLUBRIDAE) FROM THE ISLAND OF GRENADA 

By Allen E. Greer 

The recent acquisition of two specimens of Clelia clelia from 
the island of Grenada, Lesser Antilles, draws attention to signifi- 
cant morphological and behavioral differences between the island 
and mainland populations of this species. The population of 
Clelia clelia from Grenada warrants taxonomic recognition and 
may be known as : 

Clelia clelia groomei^ new subspecies 
Fig. 1 



'&• 



Holotype. MCZ 79767, adult male, Beausejour, Grenada. Coll. James D. 

Lazell, Jr., 1 July, 1964. 
Paratypes. MCZ 79766, Du Quesne, Du Quesne River Valley, St. Marks, 

Grenada. Coll. James D. Lazell, Jr., 26 June, 1964. MCZ 4507, Grenada. 

Coll. Peter Gillinam. Received by the Museum, August, 1880. 
Diagnosis. Clelia clelia groomei is similar to the mainland 
populations of C. clelia except for the preocular which is either 
very much reduced in size or united with the supraocular. When 
present, the preocular does not reach dorsally to the level of the 
upper edge of the loreal. When the preocular is lacking, the 
supraocular extends ventrally in front of the eye to touch the 
third supralabial. 

Comments. In all three specimens the reduced preocular is 
present on the left side of the head and absent on the right. 
As only three specimens of Clelia are known from Grenada, 



1 Grenada is one of the few places in the world where snakes are protected by law. 
Dr. John R. Groome of Point Salines, Grenada, one of the few zoologists in the 
Lesser Antilles, has been the primary instigator of this reform. For this, and 
for his hospitality to Mr. Lazell during his stay on Grenada, the new subspecies is 
named in Dr. Groome's honor. 



BREVIORA 



No. 223 



it may be objected that the diagnostic head scale characters are 
nothing more than asymmetrical anomalies. Obviously, only 
an examination of a larger sample would test this objection satis- 
factorily. For the present, however, it seems significant to me 
that the diagnostic characters are found not only in two speci- 
mens (MCZ 79766-79767) recently collected in different areas 
of Grenada (Fig. 2) but also in a third specimen (MCZ 4507) 




B 





Fig. 1. Side of the head of Clelia clelia. A and B, C.c. groomei new 
subspecies, paratype, MCZ 79766, Du Quesiie, Grennda; C, C. clelia 
(Daudiii), MCZ 19933, Ulna River, Tela, Honduras. Preocular scale is 
cross-hatched. 



1965 NEW SUBSPECIES OF CLELIA 3 

collected at probably yet a tbird locality 83 years previously. 
Tbese admittedly slig'ht, but constant, morphological differences, 
together with the behavioral differences alluded to below, indi- 
cate that the Grenada population of Clelia should be given at 
least subspecific status. 

The holotype (MCZ 79767), a male, has 216 ventral scales, 
72 + subcaudal scales, and 19-19-19 longitudinal scale rows. The 
uniform slate grey color of the dorsum extends ventrally to the 
lateral quarter of the ventral scales. The chin, throat and entire 
midventer are white. 

MCZ 79766 is a female with 231 ventrals, 81 subcaudals and 
19-19-17 longitudinal scale rows. The color is essentially similar 
to that of the holotype. 

MCZ 4507 is a female with 232 ventrals, 44 + subcaudals and 
19-19-17 longitudinal scale rows. The distribution of the color 
is the same as in the holotype, but the dorsum is uniformly brown 
and the midventer is yellowish brown. The specimen has been 
long preserved in alcohol. 

All three specimens have well-defined paired apical pits on the 
dorsal scales. 

The hemipenes of the holotype (MCZ 79767) agree well with 
Cope's (1900, pi. 26, fig. 4) figure of Oxyrrhopus plumbeus 
(= Clelia clelia) from Brazil. 

Field Notes. Mr. James D. Lazell, Jr., who is currently study- 
ing the iguanid lizards of the Lesser Antilles, has kindly sum- 
marized his knowledge of C. clelia groomei for me. The account 
is presented in its colorful entirety below. 

"When I was first in Grenada, in 1959, I heard tales of a 
snake, supposedly huge and much feared by Grenadians, called 
'Cribo.' Underwood (1962, p. 164) having heard the same 
stories, recorded Coiistt'ictor constrictor from Grenada; he made 
the remarkable assumption (pers. comm.) that the stories could 
only apply to that species. As the distribution of Constrictor in 
the Lesser Antilles was of great interest to me (Lazell, 1964), 
when I returned to Grenada in June and July of 1964 I made a 
special effort to find out all I could about the famous 'Cribo.' 
Primary among my sources of information were Dr. John R. 
Groome, of Point Salines, Major Francis Power, of Tufton Hall, 
and Mr. Nick Neckles, of Beausejour. From these and manj^ 
others, I learned that the 'Cribo' is indeed a large snake (up 
to ten feet ?), much feared by the local people for its strength 
(not its venom), restricted in range to the %vet portions of the 



4 BBEVIORA No. 223 

island (though it is not known to what elevation it occurs), 
and uniform slate in dorsal color. 

"Obviously the 'Cribo' was not a Constrictor; I thought, 
however, that it might be Clelia clelia, a species supposedly re- 
corded from Grenada by one specimen (MCZ 4507). I knew 
from Drs. John Price and Jake Kenny (pers. comm.) and Mr. 
Michael Dix (pers. comm.) that in Trinidad and Central Amer- 
ica, respectively, C. clelia was a nocturnal, ophiophagous snake. 

Du Quesnev^AVvivi^v? " 



Beausejour. 



Fig. 2. Grenada. The stippled region indicates the wet portions of the 
island. Clelia is reported not to occur outside this region. X = localities 
represented by specimens. Question marks are centered on extremely wet, 
high, montane areas. 



I spent seven nights, from dark to dawn, driving north out of 
St. George's around the coast and back by the Grand Etang 
road in a fruitless effort to collect specimens. Throughout this 
time, I learned on the sixth day, people all along my route had 
been encountering 'Cribos' in the daytime. On 26 June, there- 
fore, I continued on around again after sunrise. At 9 :10 A.M., 
in the Du Quesne Valley in northern Grenada, the car in front 
of me swerved off onto the shoulder of the road and ran over a 
50-inch Clelia. This individual (MCZ 79766) was in the process 
of swallowing a rat (Rattus rattus) at the time, and the side 
of the neck was burst. I took the dead snake on around the island, 
stopping at villages to ask about it. To the question, 'What is 
this ? ' I got the invariable reply : ' a Cribo ' ; I then asked, ' How 



1965 NEW SUBSPECIES OF CLELIA 5 

old do you think it is ? ' and got always a variation on : 'It is 
very young.' I became famous on Grenada, needless to say. 

"On the morning of 1 July, I was coming south along the 
leeward coast road when I was stopped by a man who had just 
seen a 'Cribo' crossing the road. It was 9:30 A.M. I uncov- 
ered the specimen (MCZ 79767) in some bushes alongside the 
road, where my informant indicated it had gone. 

"This specimen was kept alive for several days. It seemed 
nervous, making abrupt darts with its head when handled, but it 
never attempted to bite. It was active in the morning, but re- 
mained quite dormant at night. It was a powerful constrictor. 
Fresh dead, it measured 62 inches. 

"The people of Grenada confirm my records of the 'Cribo' 
eating rats, and mention occasional hen house raids as well. This 
species appears to be strictly diurnal with an apparent activity 
peak between sunrise and noon. It does not occur in the ex- 
tremely dry area along the coast of Grenada from Point Salines 
to Telescope Point ; similarly, it is absent from the dry northeast 
part of the island (Fig. 2). Elsewhere, in the wet lowlands at 
least, it is apparently fairly common. 

"Several people in Carriacou maintain that a 'Cribo' (fit- 
ting the general description of the Grenada animal) occurs there ; 
they are not confusing it with the 'serpent' {Boa sp.), which is 
abundant in the Grenadines and Grenada. If Clelia does occur 
on Carriacou, it is remarkable, for that island is drier by far 
than any habitat occupied by Clelia on the Grenada mainland." 

Mr. Michael Dix of the Biological Laboratories, Harvard Col- 
lege, has studied the habits of C. clelia both in the wild and in 
captivity. He informs me that in the vicinity of Middlesex, 
British Honduras, the snake is an inhabitant of the "tropical 
wet forest" and is strictly nocturnal, being most active between 
8-10 P.M. It is both terrestrial and arboreal (one individual hav- 
ing been found 5 feet above the ground), an observation sup- 
ported by Beebe (1946, p. 23). 

Captive individuals from Central America will take birds the 
size of sparrows and a little larger but prefer their more tradi- 
tional bill of fare — snakes. Apparently food items are "trailed 
about the cage by olfaction. Mice, rats and frogs have been 
offered as food but never accepted. Several successful attempts 
to escape confinement also attest to the climbing abilities of the 
species." 

Large adults in the vicinity of Middlesex are black dorsally but 
slate grey when younger. 



6 BBEVIORA No. 223 

Discussion. Throughout its range on the mainland from Mexico 
to Argentina, and on the island of Trinidad, the "Mussurana" 
is known as a nocturnal, ophiophagous snake. On Grenada it 
appears as if the "Cribo" is primarily diurnal and not unwill- 
ing to make a meal of small rodents. Morphologically, the two 
populations are separated by what most taxonomists would con- 
sider to be subspecifie differences, while from the behavioral 
standpoint, the two populations are distinct to a degree suggest- 
ing specific separation. Assigning island populations to the 
species or subspecies category is largely a matter of personal 
preference and practicality. In this case our purpose is perhaps 
best served by recognizing the Grenada "Cribo" as a subspecies 
of the mainland C. clelia and thereby acknowledging its close 
relationship with the mainland population. 

Acknowledgments. My thanks go to Mr. James D. Lazell, Jr. 
for suggesting an examination of the Grenada specimens. The 
extensive knowledge of the Antillean herpetofauna possessed by 
both Dr. E. E. AVilliams and Mr. Lazell made their criticisms of 
the paper most valuable. 

Literature Cited 

Cope, Edward Drinker 

1900. The crocodilians, lizards and snakes of North America. Ann. 
Eept. United States Nat. Mus. for 1898: 153-1294, pis. 1-36, 
text-figs. 1-347. 
Beebe, William 

1946. Field notes on the snakes of Kartabo, British Guiana, and Cari- 
pito, Venezuela. Zoologica, 31 : 11-52, pis. 1-13, text-tigs. 1-4. 
Lazell, J. D., Jr. 

1964. The Lesser Antillean representatives of Botlirops and Constrictor. 
Bull. Mus. Comp. Zool., 132: 245-273. 
Underwood, Garth 

1962. Eeptiles of the eastern Caribbean. Dept. Extra-Mural Studies, 
Univ. West Indies (Trinidad). Caribbean Affairs, (N. S.) 1: 
1-192 + supplements. 

(Received 10 March, 1965.) 



L^{_y 



B R E V I O R A 

Mmseiiim of Contiparative Zoology 



Cambridge, Mass. July 15, 19G5 Xump.er 224 

NEW SPECIES OP LAND MOLLUSKS WTTTI NOTES ON 

OTHER SPECIES fro:m the soeomox islands 

By William J. Clench 



The three new species described in this paper were received 
from James R. Hood of Chattanooofa, Tennessee, who was sta- 
tioned in the Solomon Islands in 1949, and from the Rev. J. 
Vander Riet, a missionary in Ataa, Malaita. A tine series of 
land and freshwater shells from this region was received from 
the latter. 

AVhile working up this and other Solomon Island material sev- 
eral systematic problems were encountered and these are dis- 
cnssed in this study. 

CAMAENIDAE 

The following species, generally placed in the genus Papuina, 
are here placed in Solmopina Iredale. They have been confusi'd 
with one another and are in need of clarification. This confusion 
resulted from the fact that Solmopina macfarUnui (Cox) had 
not been figured, and the island in the Solomons from which it 
came was unknown until th(> locality was published by Rensch 
in 1934. 

Solmopina macfarlanei (Cox) 
PI. 1, figs. 1-2 

Helix (Geotrochus) macfarlanei Cox 1873, Proc. Zool. Soc. London, p. 567 

(Solomon Islands). 
Helix (Papuina) macfarlanei Cox. Pilsbry 1891, Manual of Conchology 

(2) 7:13 (Solomon Islands). 
Papuina macfarlanei (Cox). Eeiisch 1934, Amer. Miis. Novit. No. 763:7 

(Kiota, Bougainville, Solomon Islands). 



BREVIORA 



No. 224 



Height 


Grejiter dianiete 


nun 


mm 


23 


25 


23 


24.5 


21 


22 



Measurements 



near Buin, Bougainville 
near Buin, Bougainville 
near Buin, Bougainville 

Description : Shell troeliiforni, imperforate, smooth, and reach- 
ing 25 mm in greater diameter. Color a pale ivory with a dark 
brown band at the whorl periphery and a broad circular area of 
the same color on the base of the shell. In addition, there are 
numerous irregular spots of brown both above and below the 
periphery. These brown bands and spots are translucent. The 
somewhat lighter brown spots are a little more translucent than 
the bands. Whorls 51/0 and convex. Spire somewhat extended 
and produced at an angle of about 70°. Aperture subquadrate 
with the li]) slightly reflected and cast at an angle of about 42° 
from the base. Columella at an angle from the umbilical area to 
the base. Suture indented. Sculpture consisting of tine and 
irregular growth lines. Protocunch of IV^ whorls, white and 
smooth. 

Remarks: This species differs from S. coxiana (Angas) by 
being larger, having more convex whorls, and in having the 
peripheral band of brown. The shell of /S*. coxiana is more deli- 
cate in structure and the brown coloration is considerably lighter 
in color. The brown spots in coxiana are also translucent. 1 do 
not agree with Renscli that these two species may be only geo- 
graphic races. They appear quite distinct. In 101 specimens 
examined only a single specimen lacked the peripheral band. 
Rensch has recorded this species from Kieta, Bougainville. 

Spccinifns ( xaniini d : BorGAixvn.LE : Buin (MCZ). 

SOLMOPINA COXIANA (Augas) 

PI. l.tig. 3 

(ieotioiliu.s coxiana Angas 18(37, Pioe. Zool. Soi-. Lonchui, ji. SSii, pi. 4.'>, figs. 

7-8 (Ysabel Island, Solomon Islands). 
Helix (Papuina) coxiana (Angas). Pilslny isyl, Manual of ("oncholog.v 

(2) 7:13, pi. 3, figs. 36-37. 
Papuina coxiana (Angas). Renseh 1934, Amer. Mus. Xovit. No. 763:8. 

Measurements 



Height 


Greater Diameter 


mm 


mill 


17 


20 



Ysabel Island 



1965 LAM) MOLU'SKS FROM THE SOLOMONS 3 

Dcscripiion: Slu^ll trochifonn, imporforatc, smooth and 
r(»a('liiii<>- 20 niiu in <>-i'c'at('i" diaiiu'tcr. Color a j)al(' ivory with a 
broad ciiH-ular area of Ii»i'ht brown on tlic liase of tlie shell, and, 
in addition, a few brownish spots irre<;ularly disjiosed over the 
shell. Both the basal band and the spots are translucent. Whorls 
5, slightly eonvex. Spire somewhat extended and produced at an 
augle of about 70°. Aperture subcpiadrate with the li{) slightly 
reflected, colored brown and cast at an angle of about 40° from 
the base. Columella white and at an angle from the umbilical 
area to the base. Suture slightly indented. Sculpture consisting 
of very fine and irregular growth lines. Protoconch of IVij 
whorls, white and smooth. 

Remarks: See RcDiarks under S. mdcfarlanei. 

Spcci)ncns exa)}ii.)ic(1 : Ysabel. 

l^pecimcns recorded: Choiseul: Luti and Tauro (Kensch). 

ARIOPHANTIDAE 

Trociiomorpiia vanderrieti new species 
PI. 2, figs. 1-2 

Holoiiipc: MCZ 251176, from Ataa, Malaita Island, British 
Solomon Islands. Rev. J. Vander Riet, collector, 1964. 

Paratypes: Figured paratype, MCZ 251177; additional para- 
types, MCZ 247959, all from the same locality as the holotype. 





Greater 


Lesser 


Heiglit 


(lianu'tei- 


diameter 


mm 


mm 


mm 


11 


26.5 


22 


10 


27 


22 


11.5 


26 


20.5 


11.5 


25 


20.5 


10 


28 


21 



Holotype 

Paratype MCZ 251177 (figured) 
Paratype MCZ 251177 
Paratype MCZ 251177 
Paratype MCZ 247959 (figured) 

Description: Shell reaching 27 mm in width, subdepressed, 
finely carinated and umbilicated. Color a dark, yellowish brown, 
uuicolorous below the thread-like carina and flecked with nu- 
merous narrow, straw-colored patches in axial arrangement 
above the carina. The brown coloration is in the shell; the straw 
coloration is invested in the periostracum. Whorls 5 to 5iX>, con- 
vex, and the last whorl with a tine, thread-like carina. Spire 
depressed, dome-shaped and forming an angle of about 1:^5°. 
Aperture subovate, with the outer lip slightly depressed near its 
juncture with the whorl above. Outer liji narrowly reflected 



4 BREVIORA No. 224 

along- the basal area. Umbilicus rather narrow and very deep, 
and is Yq tlie width of the shell in greater diameter. Suture 
slightly indented and well defined. Sculpture consisting of nu- 
merous oblique and fine costae above the whorl periphery and 
nearly smooth below. 

Re))iarks: In relationship, this species is nearest to Trocho- 
morpha aukiensis Cla]ip, also from Malaita. Trochomorpha van- 
derricti differs from H. aukknsis by being: larger, not being 
spirally banded and in having a narrower umbilicus, being % of 
the greater diameter, while in aukiensis the umbilicus is I/4 of 
the greater diameter and there are no straw-yellow flecks above 
the whorl periphery. 

Including- T. vandcrrieti, there are seven species of Trocho- 
morpha now known from Malaita Island. These are : 

Trochomorpha aukiensis C'lapp 1923, Bull. Mus. Comp. 
ZooL, 65:361. j)l. 2, figs. 10-15 (Auki, Malaita). [Holotype, 
MCZ 32535]. 

TrochomorpJia^ hclmorci (Cox) 1871, Proc. Zool. Soc. Lon- 
don, J). 647, pi. 52, fig. 12 (Solomon Islands). 

Trochomorpha concava Clapp 1923, Bull. Mus. Comp. Zool., 
65:363, pi. 3, figs. 1-3 (Auki, Malaita). [Holotype, MCZ 
32523]. 

Trochomorpha crust idio)) (Cox) 1873, Proc. Zool. Soc. Lon- 
don, p. 15U (Solomon Islands). 

Trochomorpha flava Clapp 1923, Bull. Mus. Comi). Zool., 
65:366, pi. 3, figs. 4-6 (Auki, Malaita Island). [Holotype, 
MCZ 32521]. 

Trochomorpha mclcaiii Clench 1959, Natural History of 
Rennell Island, British Solomon Islands, I"^niv. Copenhagen, 
Denmark, Vol. 2:179, pi. 17, fig. 6 (10 miles inhuul from Sun, 
Malaita Island). [Holotype, Amer. Mus. Nat. Hist. 7!)016]. 

ASSIMINEIDAE 
Setaepoma Clench 

Setaepoma Clench 19.")0, Nautilus, 63:1. U (fy]K' species, Japonid ( .' ) hcdi fieri 
T. Miul R. Renscli). 

Setaepoma iioodi new species 
PI. 2, fig. 3 

Jlolotffpr: MCZ 25130!), from the west side of the Teiiarn 
River, about one-half mile above the Catholic Mission, Guadal- 
canal, Solomon Islands. James R. Hood, collector, 194i). 



1965 LAND MOLLISKS FROM TllK SOLOMONS 5 

Measurcinciits 

Height Width 

mm mm 

H:2 7.!) TTololypo 

Description: Shell tiii'binatc, thin, umbilicato and having' nu- 
merous, spiral rows of bristle-like processes of periostraeuiu. 
Color a o-oldeu brown. Whorls 6 and tubular. Spire moderately 
extended and produced at an angle of about 60 \ Aperture 
nearly circular, holostomatous and with a thin, simple lip. No 
columella ; umbilicus narrow and deep. Suture deeply indented. 
Periostracum thin, shining', and with numerous, spiral and axial 
ridges which support a bristle-like process at each intersection. 
Protoconch of about 2 whorls which are smooth and shining. 
Operculum calcareous, multispiral and dished. Inner surface 
with a papilliform central nucleus, smooth, shining, and with 
faint indications of the outer sculpture. 

Remarks: This species is distantly related to Sietoepoma 
hccligeri I. and B. Renscli (Bougainville Id.) and S. )iw}iri 
Clench (Ysabel and Choiseul Ids.). It differs from both by being 
more attenuated, having fewer spiral ridges and a much nar- 
rower umbilicus. 

This present new species is the third known from the Solomon 
Islands. The other two are : 

Sciacpoma Judifjcri (I. and B. Rensch) 1986, Revue Suisse 

de Zoologie, 43:678 (Bougainville Id., Solomon Ids.). 

SetacpoDia maijri Clench 1959, Natural History of Rennell 

Island, British Solomon Islands, Univ. Copenhagen, Denmark, 

Vol. 2:3 68, jil. 17, tig. 5, text fig. 1 (Fulakora, Ysabel Id., and 

the Wurulata River. Choiseul Id., Solomon Ids.). 

CYCLOPHORIDAE 

Palaina (Palaina) brazieri (Cox) 

PL 2, fig. 4 

Diplommaiina haizieri Cox 1870, Proe. Zool. Soc. London, p. 84 (Wanga, 
San Christoval Id., Solomon Islands). 
This species has not been figured previously. The specimen 
we figure came from Kira Kira, San Christoval Id., collected by 
the Whitney Expedition in 1929. 



6 BEEVIORA No. 224 

Palaixa (Palaina) delli new species 
PI. 2, fig-. 5 

Holotype: MCZ 258017, from one-half mile above the Seventh 
Day Adventist Mission Station on the west side of the Lunga 
River, Guadalcanal Island, Solomon Islands. James R. Hood, 
collector, li)4y. 

Paratypcs: Five paratypes, MCZ 2"'.8018, from the same lo- 
cality as the holotype. 

Measurements 

Height Width 

iiini mm 

4.5 2 Holotype 

4.7 2.2 Paratype 

4.5 2 Paratype 

4.5 2.1 Paratype 

4.3 1.8 Paratype 

Description: Shell reaching 4.7 mm in height, dextral, imper- 
forate and sculptured. Whorls Gi/o and convex. Color a dull 
grayish brown. Spire attenuate and straight. Aperture circular, 
with a I)roadly reflected lip, and attached at the parietal area. 
No columellar tooth. Suture impressed. Sculpture consisting of 
numerous, diagonal riblets which are very irregular as to number 
on the different whorls and between different specimens. These 
ril)lets may be singk' or in groups of 2 to 5 and varying from 
whorl to whorl. Protoconch of 2 whorls, smooth and rather dull. 
Operculum with a vertical, spiral lamella. 

Remarks: This species is nearest in relationship to P. hrazieri 
(Cox) from San Christoval. It diff'ers by being more than twice 
as large and in having more regular and more numerous axial 
riblets. Both are dextral and neither has a columellar tooth. 

Named for Dr. Richard K. Dell, Dominion Museum, Welling- 
ton, Xcw Zealand. 

DU'LOMMATINA AERARI Dell 

DiploiiDiKil inn (i<r(iri Dell 1955, Pat-ifie Scit'iicc, 9:4li5, i]^. 1 h (Mono Til., 
Treasury (iioup, Solomon Islands). 

Three s])ecimens of this species were collected by -James R. 
Hood, one-half mile above the SevcMith Day Adventist ^Mission 
Station on tlic west side of the Lnnga Kiver, (iuadalcanal Island. 



1965 



I.AM) MOLLrSKS FROM TlIK SOIiOMONS 



REFERKNCES CITED 

Clapp, W. F. 

1923. Sonu' Mollusc;! fi-inii tlic SoloiiKiii Islands. liiill. Mils, ('uiiiii. 
Zool., 65:351-418. 
Kobf.lt, Wilhelm 

19(ll\ Cyc-loplioridae. I)a.s Tierreii-h, Berlin, 16:393-412. 
PiLSBRY, H. A. 

1891. Manual of ConcholoRv, (2) 7:1-81, pis. 1-17. 
Rensch, Ilse 

1934. Studies on the Papuind and Dcndrot roclius, i)uhnonate niollusks 
from the Solomon Islands. Amer. ^lus. Novit., No. 763:1-24. 
SoLEM, Alan 

1960. Non-marine ]\Iollusca from the Florida Islands, Solomon Islands. 
Jour. Malaeologieal Soe. Australia, No. 4:39-56. 





PLATE 1 

Figs. 1-2. Solmopina macfarlanei (Cox). Buin, Bougainville Id. (about 
2X). 

Fig. 3. Snlniopiim coxiana (Angas). Ysabel Id. (2.5 X). 



8 



BREVIORA 



No. 224 







PLATE 2 

Fig. 1. Trochomorplia vanderrieti, n. sp. Ataa, Malaita Id., Solomon 
Islands. Holotype, MCZ 251176 (2.4 X). 

Fig. 2. Trochnmorpha vanderrieti, n. sp. Atan, Malaita Td., Solomon 
Islands. Paratype, MCZ 247959 (2.4 X). 

Fig. 3. Setaepoma hoodi, ii. sp. West side of the 'I'cnaru i\i\ci-, alunit 
one-lialf mile aliovc the Catholic Mission, Guadalcanal Id., Sdlcuiion Ids. 
Ilolotypc, MCZ l'.")i;i(l9 (4.(i X). 

Fig. 4. Palaina (Palaina) brcizicri (Coxj. Kira Kira, San Cliiistovai 
Id. (12.(j X). 

Fig. 5. PdUtiint (Palaina) dclli, n. sp. West side of the Lunga River, 
onedialf mile aijovc the Seventh Day Adventist Mission, Guadalcanal Td. 
Holotype, MCZ 258017 (18 X). 



BREVIORA 



semm of Comparative Zoology 



Cambridge, Mass. July 15, 19G5 Number 225 

THE ASIAN SPECIES OP GALERITULA STRAND 
(COLEOPTERA, CARABIDAE) 

By Hans Reichardt^ 

Very few attempts have been made to study the relatively few 
Asian species of Galeritula. All these species have been described 
individually, by separate authors, who usually had very few 
specimens of each. 

Heller (1923:65) was the first to attempt the characterization 
of species groups when he described szcfschwana from China. 
However, the characters he used to divide the Asian species of 
Galeritula into two groups are very variable (posterior constric- 
tion of pronotum more or less well developed, and margins of 
pronotum parallel or divergent behind the constriction), as will 
be seen below. The failure of this system is evidenced by Heller 's 
separation of peregrina Dohrn and hirmanica Bates into differ- 
ent groups, even though these species were considered synony- 
mous by Andrewes a few years earlier (1919:480). 

In 1949 Jeannel assigned the Asian species of Galeritula (to- 
gether with a few African forms) to a new genus, Galeritella. 

Finally, in 1963, Jedlicka presented a revision of the species 
found on the Asian Continent. According to this author (1963: 
474) it is very difficult to separate some species morphologically, 
since most of them are very similar ; Jedlicka separated the 
species, as did most earlier authors, solely on the basis of color. 
As pointed out later in this paper, the coloration is a very varia- 
ble character in G. orientalis and for this reason this species has 
been described several times (eight names are available for orien- 
talis). The fact that such variation occurs can, however, only be 
observed in large series of specimens. No author has yet had 
many specimens on which to base his studies. 



1 Departamento rte Zoologla, Secretaria da Agricultura, Sab Paulo, Brazil ; pres- 
pntly at Harvard University, Cambridge, JIass. 



2 BREVIORA No. 225 

In the course of my present work on the Neotropical species 
of Galeritula, I have been able to study many types as well as 
undetermined material of Asian species in the British Museum 
(Natural History), London, and the Museum National d'Histoire 
Naturelle, Paris. In addition, several other museums have pro- 
vided specimens for identification. Study of this relatively rich 
collection of 138 specimens has revealed some very interesting 
facts, and has produced many new distributional records. All 
this has led to a study of the Oriental species and their relation- 
ships, which is reported here. 

SOURCES OF MATERIAL, METHODS 
AND ACKNOWLEDGMENTS 

The present study is based on material from the following col- 
lections (abbreviations as used in the text) : 

British Museum (Natural History), London (BM) ; Mr. Jacques 
Negre's collection, Versailles (JN) ; Institute of Zoology, Polish 
Academy of Sciences, "Warsaw (IZ) ; Museum of Comparative 
Zoology, Cambridge, Mass. (MCZ) ; Museum National d'Histoire 
Naturelle, Paris (MNHN) ; Academy of Natural Sciences, Phila- 
delphia (ANSP), and Senckenberg Museum, Frankfurt am 
Main (SMF). 

The measurements of the specimens have been made with a 
microruler in the microscope ocular. Comparison is made be- 
tween measurements of parts of the beetles (expressed in the 
descriptions by ratios) as follows: Head — width/length (width 
taken at the widest point, including the eyes ; length taken from 
apex of clypeus to constriction behind the eyes, not including 
the neck) ; length of eyes/length of occiput (the measurement 
of the occiput is made parallel to the longitudinal axis of the 
body, from posterior margin of eye to beginning of neck). 
Pronotum — width/length (width taken at widest point; length 
along the median line). Elytra — width/length (width at widest 
point; length from base to apex, along suture). Total length of 
specimens has been measured from the tip of mandibles to apex 
of elytra, excluding the pygidium. 

I am greatly indebted to the curators of the above mentioned 
collections, without whom this study would have been impossible ; 
special thanks are due to the Evolutionary Biology Committee 
at Hai'vard University, which provided the funds for my Euro- 
pean studies in June 1964. I am also grateful to Professor P. J. 



1965 ASIAN GALERITULA 3 

Darlington, Jr., Dr. E. G. MacLeod and Dr. R. W. Taylor for 
having read and criticized the manuscript. 

Galebitula Strand, 1936 

Galerita Fabricius, 1801, Syst. Eleuth., i:214 [type, by subsequent designa- 
tion (Latreille, 1810, Consid. Gen. :426) Carahus americanus Linnaeus, 
1758] ; Jedlieka, 1941, Versueh einer Monograpliie der Truncatipennen 
. . . pp. 12, 25 [notes on Asian species]. Not Galerita Gouan, 1770. 
Galeritula Strand, 1936, Fol. Zool. Hydrobiol., 5:168 [new name for Galerita 
Fabricius, 1801] ; Basilewsky, 1963, Ann. Mus. Eoy. Afr. Centr., 8°, 
Sci. Zool., 120:5, 6, 7, 23 [genus restricted to Neotropical species]; 
Jedlieka, 1963, Ent. Abh. Mus. Dresden, ^5:474-475 [revision of con- 
tinental Asian species] . 
Galeritina Jeannel, 1949, Faun. Enip. Frang., ii:1058 [new name for 

Galerita Fabricius, 1801]. 
Galeritella Jeannel, 1949, Faun. Emp. Frang., li:1058 [type, by original 
designation, Galerita orientalis Schmidt-Goebel, 1846] ; Basilewsky, 
1963, Ann. Mus. Eoy. Afr. Centr., 8°, Sci. Zool., 1S0:8, 63-64 [revision 
of African species]. NEW SYNONYMY. 
Galericeps Jeannel, 1949, Faun. Emp. Fran?., 11:1058, 1062 [type, by 
original designation, Galericeps yheropsoplioides Jeannel, 1949] ; Basi- 
lewsky, 1963, Ann. Mus. Eoy. Afr. Centr., 8°, Sci. Zool., 120 -.QZ, 64 
[proposed synonymy with Galeritella] . 
In the revision of the Carabidae from Madagascar, Jeannel 
divided the pantropical genus Galerita into several genera. The 
new generic groups were based on the following characters: *'la 
forme de la dent labiale et . . . evolutions divergentes de la 
sculpture elytrale" (1949: 1057). As I have already mentioned 
in the introduction to this paper, only the Asian forms (which 
together with some species from Africa and Madagascar have 
been separated by Jeannel and subsequent authors as Galeritella) 
will be discussed here. 

The mouthparts of G. orientalis (Fig. 1), type species of 
Galeritella, have been compared with those of americana (Fig. 
2), the type species of Galeritella, and also with other Neotropical 
species of the genus. No important difference seems to be pres- 
ent, not even in the "dent labiale," the character stressed by 
previous authors. The same is true for all the other species 
found in Asia that I have been able to study in detail. 

The second character mentioned by Jeannel (1949 : 1058), the 
elytral structure, really seems to be of importance, but no con- 
sistent difference between American and Asian species could be 



BREVIORA 



No. 225 



found, as noted by Jeannel himself (1949: 1058, "... a de- 
veloppe la meme sculpture elytrale") ! 

More recently, Basilewsky (1963: 7-8, 63-64) added new char- 
acters, which should also be analyzed. He characterized Galeri- 
tella as having "... une membrane apicale transparente aux 
elytres, ' ' and a more voluminous aedeagus, the latter very often 
with "partie terminale individualisee" (the latter, however, only 
in the African species) . 




Fig. 1. Galeritula orientalw, 5 from Jabalpur (MCZ), inouthparts. 
Fig. 2. G. americana, $ from Trinidad (MCZ), mouthparts. 



As to the first character, Asian species have a membranous 
apical margin on the elytra (for a width of less than 0.1 mm). 
This membrane is much less developed in Neotropical species, but 
it can be found with high magnification and careful examination. 

As part of this paper, I have measured and studied the geni- 
talia of the Oriental species which were available (Figs. 4-8). 
Although the relative size of the aedeagus really seems to be 
larger in these than in most Neotropical species, it seems also to 
be true that the Oriental species are larger insects than the 
average Neotropical species, so that insofar as my investigation 
goes, the size of aedeagus seems to be directly related to the size 
of the species. I have not studied the genitalia of any African 
species of Galeritella; however, Basilewsky (1963: 64 and fig. 
28) indicates that the African species have a less well developed 
aedeagus, and this seems to agree with the average smaller size 
of the African species. 



1965 



ASIAN GALERITULA 



Figure 3 represents the linear regression for total length of 
aedeagus against total length of specimen in 25 species of 
Galcritula (21 Neotropical, 4 Oriental) and indicates clearly 
that the size of aedeagus is related to the size of species. Since 
material available for dissection is not always present in de- 
sirable quantities, I have represented in Figure 3 the size of 



_ 2- 
° mm 



1.5 




lOo on 



• Orienfol 
o Neotropicol 



15 18 21 24mm 

Tofol lenqfh of specimen 

Fig. 3. Galeritula species showing linear regression for total length of 
aedeagus against total length of specimen. 1, gracilis Brulle; 2, unicolor 
Latreille and Dejean; 3, palustris Liebke; 4, striala Klug; 5, microcostata 
Darlington; 6, lacordairci Dejean; 7, coeruleipennis Chaudoir; 8, collaris 
Dejean; 9, ruficoUis Dejean; 10, melanarthra Chaudoir; 11, nigra Chevrolat ; 
12, tucumana Liebke; 13, americana Linnaeus; 14, occidentalis Olivier; 15, 
orbignyi Lucas; 16, cliampioni Bates; 17, hrasiliensis Dejean; 18, hruchi 
Liebke; 19, carbonaria Mannerheini ; 20, corumhana Liebke; 21, ventricosa 
Lucas; 22, orientalis Sehmidt-Goebel, from India; 23, same species from 
Japan ; 24, carinifrons Schauf uss ; 25, feae Bates ; 26, toreuta Andrewes. 



aedeagus in relation to the specimen it comes from, rather than 
taking mean size of aedeagus against mean size of species in 
each case. Although variation of aedeagus is not large, in species 
where I have studied larger series (see for example "22" and 
" 23 " in Figure 3, which represent two different sized specimens 
of the same species, from different localities), it must be kept in 
mind that the points in Figure 3 do not represent the species, 
but only one specimen each. 



6 BREVIORA No. 225 

The linear regression for this character shows very clearly 
that there is no place to objectively draw a line between Oriental 
and Neotropical species. 

As to the apex of genitalia, the aedeagus of Neotropical species 
shows a large range of variation which includes forms very 
similar to those of the Asian and African ones. 

Thus, a careful study of the two important generic characters 
given by Jeannel for Galeritella, as well as characters added by 
Basilewsky, leads me to conclude that these are the same as in 
Galeritula and the two genera must be considered synonymous. 

The genus Galeritula, as considered here, occurs in the Neo- 
tropical (50 species more or less, of which several are as yet 
undescribed), Ethiopian (4 species recently revised by Basilew- 
sky, 1963 : 63-72), and Oriental regions (7 species). 

It seems unnecessary to give here a full description or even a 
diagnosis of the genus. The descriptions found in the literature 
seem to be more than sufficient. 

As in my current work on the Neotropical species of the genus, 
the Asian species of Galeritula are here separated into species 
groups, based on similarities which seem to indicate a phylogene- 
tic relationship. These groups are assemblages of related species, 
but are not to be considered subgenera. Each group is named 
after the oldest species. The Oriental species can easily be as- 
signed to two groups, as will be seen below. The African species, 
although related to Oriental and Neotropical ones, seem to con- 
stitute other, distinct groups, as do the Neotropical species. 

Key to species 

1. Legs black 2 

Legs red or yellowish-brown 3 

2. Head and pronotum more elongate ; carinulae-interstices much deeper 

than carinae-carinulae-interstices; carinae not very high; 21.8 mm; 

continental species feae Bates 

Head and pronotum less elongate; carinulae-interstices as deep as carinae- 
carinulae-interstices; carinulae very thin; carinae higher; 21.0-22.0 
mm ; Java toreuta Andrewes 

3. Legs completely red; 17.5 mm; India indica Chaudoir 

Legs with yellow femora and brownish tibiae; apices of femora dark- 
ened 4 

4. Head with shorter occiput (roughly as long as diameter of one eye) ; 

carinae usually less well developed; pubescence in carinae-carinulae- 
interstices dense, formed by two irregular rows of hairs; head red, 
clvtra usuallv bluish 5 



1965 ASIAN OAhERITULA 7 

Head with longer occiput (longer than diameter of one eye) ; carinae 
usually well developed; pubescence in carinae-carinulae-interstices less 
dense, formed by a single row of hairs; head, pronotum and elytra 
black (or sometimes very dark fuscous) 6 

5. Pronotum dark brown ; head completely red on superior face ; hind 

wings reduced ; 20.0 mm ; India ruficeps Chaudoir 

Pronotum varying from completely red, through red with dark margins, 
to completely dark brown or black; when completely dark brown, head 
with frontal ridge and sides of occiput much darkened; hind wings 
fully developed ; 18.5-22.0 mm ; India to Japan, Sumatra to Flores .... 
orientalis Schmidt-Goebel 

6. Antennae completely rufous; head, pronotum and elytra dark brownish; 

humeri very poorly developed, wingless species; 19.5-21.0 mm; Celebes 

carinifrons Schauf uss 

Antennae with apex of scape and segments 2-4 darkened ; head, pronotum 
and elytra black; humeri well developed (wings dimorphic?); 21.0 
mm ; Continental species batesi Andrewes 

''Orientalis" Group 

Characterized mainly by the shape of pronotum, with posterior 
constriction very basally placed, not well defined; basal angles 
very rounded; elytra usually with very conspicuous pilosity, 
formed by two more or less parallel rows of rufous hairs in each 
carinae-carinulae-interstice. 

Three species are known : orientalis Schmidt-Goebel, indica 
Chaudoir and ruficeps Chaudoir. 

Galeritula ORIENTALIS (Schmidt-Goebel, 1846) 
(Figs. 1,3-5, 9) 

Galerita orientalis Schmidt-Goebel, 1846, Faunula Coleopterorum Birmaniae, 

pp. 26-27 [types, $ and ?, "birmesische Provinzen, " Prague Museum; 

not examined] ; Andrewes, 1923, Trans. Ent. Soc. London : 8 [redescrip- 

tion]. 
Galerita nigripennis Chaudoir, 1861, Bull. Soc. Nat. Moscow, 34(1) -.557 

[types, $ and 2 , " Indes Orientales, ' ' Museum National d 'Histoire 

Naturelle, Paris ; not located]. NEW SYNONYMY. 
Galerita japonica Bates, 1873, Trans. Ent. Soc. London: 304 [types, Naga- 
saki, Yokohama; British Museum (Natural History), not located]; 

Chaudoir, 1877, Bull. Soc. Nat. Moscow, 5^(1) :255 [synonym of 

nigripennis?]. NEW SYNONYMY. 
Galerita peregrina Dohrn, 1880, Stett. Ent. Zeit., 41:291 [types. Hong 

Kong, Stettin Museum; not examined]. NEW SYNONYMY. 
Galerita ruficeps; Bates [nee Chaudoir], 1889, Ann. Mus. Civ. Geneva, 

f 7:109 [Bhanio, Burma]. 



8 



BREVIORA 



No. 225 





Fig. 4, Galeritula orientalis, from Tsusliiiiui (iSMF C13907), aedeagus; 
Fig. 5, G. orientalis from Jabalpur (MCZ), aedeagus; Fig. 6, G. carinifrons, 
from Bonthain (IZ), aedeagus; Fig. 7, G. feae, from Lambok (SMF 
C13910), aedeagus; Fig. 8, G. toreuta, from Eadeng (MCZ), aedeagus. 



1965 ASIAN OALKRITULA 9 

Galerita birmanica Bates, 1892, Aim. Mus. Civ. Geneva, 5^:385 [G. ruficeps; 

Bates, 1889; types, Bhamo, Burma, Geneva Museum; not examined]. 

Galerita ssetschwana Heller, 1923, Ent. Blaett., 19:65 [type, "Omisien, 

Szesc'huan, " Dresden Museum; not examined]. NEW S¥NON¥M¥. 
Galerita fonnosana Kano, 1930, Trans. Nat. Hist. Soc. Formosa, S0:29, 
fig. 3 [types, 2 9, Taihoku, Urai; Kano collection; not examined]. NEW 
SYNONYMY. 

Redescription: Head varying from completely blaeli to com- 
pletely red, with intermediates where head is red with black 
frontal ridge and blacli sides of occiput (typical peregrina) ; 
pronotum also varying from completely black or bluish (pere- 
grina) to completely red (szetschwana) , with intermediates 
which have more or less developed black margins (japonica, 
nigripennis and formosana) ; elytra blue, sometimes very dark, 
almost black ; antennae and mouthparts rufous ; femora yellow 
witli black apices; tibiae and tarsi rufous; inferior side dark 
brown, almost black on abdomen. 

Head longer than wide (1.07) with relatively small eyes; oc- 
ciput as long as eyes; surface punctate-rugose, with few yellow 
hairs, mainly in posterior half. 

Pronotum wider than head (1.25), slightly longer than wide 
(1.03) ; widest slightly in front of the middle; not much nar- 
rowed anteriorly or posteriorly ; posterior constriction very basal, 
sides varying from slightly convergent to slightly divergent 
behind the constriction ; surface convex, densely punctate-rugose. 

Elytra much wider than pronotum (1.7) ; 1.73 times as long 
as wide, almost parallel, slightly widened in posterior half; 
carinae strong, carinulae very thin, sometimes almost erased; 
carinulae-interstices with very shallow row of punctures ; carinae- 
carinulae-interstices with dense rufous pilosity, disposed in two 
somewhat irregular rows. Always fully winged. 

Measurements: length, 18.6-22.0 mm; width, 5.9-7.3 mm. 

Specimens examined (96) : India: Kerala, Wallardi, Travan- 
core (10 5 , 4 9 , MNHN) ; Madras, Nilgiri Hills (65,79, MCZ, 
BM) ; Madhya Pradesh, Jabalpur, 480 m (3 S , 1 9 , MCZ) ; 
Maharashtra, Nagpur (1 9 , MCZ); Assam, Patkai Mts. (1 S , 
BM). Burma: Bhamo (2^,79, BM, MNHN) ; Tenasserim (1 9 , 
BM) ; Kachin Hills, Malikha Valley, 300-750 m (1 9 , BM) ; no 
locality (1 9 , SMP C13899). Vietnam: Hoabinh (4^,29, BM, 
MNHN, IZ) ; Lactho (1 i , MNHN) ; Ha Ciang (1 9 , MNHN) ; 
Tuyenkwan (1 9 , MNHN); Hue (1 9 , BM) ; Pliuc-son (2 9, 
SMF C13900) ; Long Chuyan (1 9 , BM). Hong Kong (1 9 , 
BM). China: Kwangsi, Kweiling {1 $ , MCZ); Liuchow (1 $ , 



10 



BREVIORA 



No. 225 



MCZ) ; "Wuchow (15, MCZ) ; Kiangsu, Nanking (1 $ , MCZ) ; 
Tschekiang, Ning-Po (1$, BM) ; Szechuan, Chungking (1 $ , 
MCZ) ; Kwanshien (1 $ , SMF C13901) ; no locality {2 $ , 2 9 , 
BM). Japan: Kyoto (1^, SMF C13902, 1$, MNHN) ; Tokyo 
{1$, SMF C13903, 1$, MNHN); Saga, Kiushu {2$, SMF 
C13904, 1 5 , MNHN) ; Yuyama (1 $ , MCZ) ; Kobe (1 $ , SMF 
C13905, 15, ANSP); Tsushima (3 5, SMF C13906-13907) ; 
Hiogo (15, SMF C13908) ; Osaka (1 9 , SMF C13909) ; no local- 
ity (3 $ , MNHN). Indonesia: Sumatra, Manna (1 5 , 1 5 , BM) ; 
Java, Pelabuhan Ratu {2$, MCZ, MNHN); Buiteuzorg (1 5 , 
MNHN) ; no locality (1$, BM) ; Sunhawa {1$, MNHN) ; Flores 
(15, MNHN). 




Fig. 9. Galeritula orientalis, distribution. 



According to the literature, the species is also known from 
Korea and from Taiwan (Formosa: formosana) ; it was unknown 
from Indonesia. Its distribution is thus much Avider than that 
of any of the other Asian species (Fig. 9) . 



1965 ASIAN GALERITULA 11 

The large number of synonyms of this species is due to the 
variability of the color pattern, which, when only a few speci- 
mens are studied (as in all the previous work), seems quite 
constant. Study of larger series, however, shows that these 
characters have very little, if any, specific value. With respect 
to color and pattern, there seems to be no geographic variation 
in orienialis. 

The present study also included examination of the male 
genitalia (aedeagus), which seems to be quite constant in most 
populations of the species. A different aedeagal shape is, how- 
ever, found in Indian specimens (described originally as nigri- 
pennis — see Fig. 5). I have tried to correlate this character 
with others, but have failed. A series of "nigripennis" from 
Travancore (10 5, 4$, MNHN; the genitalia unfortunately 
could not be studied) shows the same color variation as orienialis 
from elsewhere. For this reason, I think it is not advisable to 
keep nigripennis separated from orienialis. There is a complete 
overlap of the two forms in external characters, mainly in color, 
and I feel that the difference in aedeagi of Indian orienialis is 
due simply to geographic variation. 

The possible synonymy of nigripennis and japonica was 
pointed out in 1877 by Chaudoir, who could not find any differ- 
ences between the two species. 

Galeritula indica (Chaudoir, 1861) 

Galerita indica Chaudoir, 1861, Bull. Soc. Nat. Moscow, 34(1) :557 [Holo- 
type, $ , "Nord de I'llindostan, " Museum National d'Histoire Natur- 
elle, Paris; examined]. 
Only the type specimen of this species seems to be known. 
G. indica is very close to orienialis but shows a few important 
differences, and for this reason I consider it separate, at least for 
the time being. G. indica has completely red legs, is a wingless 
species, and is much smaller than orienialis (17.5 mm). More 
material is essential for a final decision on its status. 

Galeritula RUFiCEPS (Chaudoir, 1861) 

Galerita ruficeps Chaudoir, 1861, Bull. Soc. Nat. Moscow, 5^(1) :556-557 

[Holotype, $, "Nord de I'Hindostan," Museum National d'Histoire 

Naturelle, Paris; examined]. 

Besides the type, I have seen only 1 $ from Harki Dun, India 

(BM). This species, like indica, is very close to orienialis, and 

I keep it separate mainly because of reduction of the wings. 

More material will eventually show its real status. 



12 BREVIORA No. 225 

' ' C ARINIFRONS ' ' GroUp 

Pronotum usually with better defined posterior constriction ; 
this is situated less basally than in orientalis group ; basal angles 
sharp; elytra with pilosity less visible, sometimes absent; species 
with more restricted distribution, some with reduced wings. 

Four species are included : hatesi Andrewes, carimfrons Schau- 
fuss, feae Bates and toreuta Andrewes. 

Galeeitula BATESi (Andrewcs, 1923) 

Galerita batesi Andrewes, 1923, Trans. Ent. Soc. London: 9 [Holotype, $, 
Karin Cheba, Burma, 900-1100 m; British Museum (Natural Historj-) ; 
examined]. 
Galerita orientalis; Bates [nee Schmidt-Goebel], 1889, Ann. Mus. Civ. 
Geneva, ^7:109 [pars]. 

Galeritula hatesi is very close to carinifrons, distinguished b.y 
very few characters, the main ones having been mentioned in 
the key. These two species are also close to feae and toreuta, 
distinguished mainly by the color of the legs. 

A superficial examination of the two types in the British 
Museum (Natural History) (2, Karin Cheba; S, Teinzo), 
seems to indicate that they have reduced posterior wings. All 
other examined specimens have normally developed wings. G. 
hatesi, however, does not show any reduction of the humeri. The 
species may be in the process of losing its wings, a process which 
has already been finished by its close relative, carinifrons, prob- 
ably independently. Cases of wing dimorphism are not common 
in the genus: the only cases knoAvn to me are those of africana 
Dejean from Africa and forreri Bates from Mexico. 

G. hatesi is known only from the mainland, and has a rather 
restricted area of distribution. 

Examined specimens (6) : Burma: Bhamo (1 5 , BM) ; Teinzo 
{Is, paratype, BM, l9, MNHN). India: Assatn, Shillong re- 
gion (1 S , MNHN) ; Sylhet, Chandkhira (1 9 , BM). 

Galeritula carinifrons (Schaufuss, 1887) 
(Figs. 3, 6) 

Galerita carinifrons Schaufuss, 1887, Ilor. Soc. Ent. Eoss., J'?: 103-104 
[types. Macassar, South Celebes; Zoologisches ^luseum dor Universitiit 
Berlin; not seen]. 
lie description: Black-brownisli, mouthparts and antennae ruf- 
ous ; legs as in hatesi. 



1965 ASIAN OALERITULA 13 

Head large, longer than wide (1.08), with relatively small 
eyes ; occiput slightly longer than the diameter of one eye ; 
surface moderately punctate-rugose, with j^ellow hairs. 

Pronotum wider than head (1.24), longer than wide (1.11) ; 
widest in anterior third, as narrowed anteriorly as posteriorly ; 
constriction well developed, sides parallel behind it ; surface con- 
vex, densely punctate ; covered with yellow hairs. 

Elytra 1.7 times as wide as pronotum; 1.77 times as long as 
wide ; carinae and carinulae strong, the latter thinner, slightly 
closer to each other than to next carinae; carinulae-interstices 
punctate ; carinae-carinulae-insterstices with single row of yellow 
hairs; scutellar carina not joining the first; humeri very much 
reduced ; "wingless species. 

Measurements : length, 19.5-21.0 mm ; width, 6.2-6.9 mm. 

Specimens examined (3): Celebes: Bonthain {S $ , IZ, 
MNHN). 

Notes: G. carinifrons seems to be restricted to the southern 
part of Celebes. As already mentioned, it is very close to hatesi. 
Both are probably derived from a common ancestor. The loss of 
Avings in carinifrons is interesting and may be correlated with 
the restricted distribution. 

Galeritula feae (Bates, 1892) 
(Figs. 3, 7) 

Galerita feae Bates, 1892, Ann. Mus. Civ. Geneva, 55:386 [types, Karin 
Cheba, Burma, 900-1100 m; Geneva Museum; not seen]. 

Re description: Completely black; antennal segments 5-11, 
brown. 

Head large, longer than wide (1.09), with relatively small 
eyes ; occiput slightly longer than the diameter of one eye ; 
posterior half almost impunctate, anterior half deeply punctate- 
rugose ; covered with few brownish hairs. 

Pronotum wider than head (1.21), longer than wide (1.12) ; 
form and sculpture as in hatesi and carinifrons, sides slightly 
divergent behind constriction. 

Elytra very similar to those of carinifrons, 1.78 times as wide 
as pronotum ; 1.7 times as long as wide ; carinae and carinulae 
well developed ; carinulae-interstices very deep, sulciform ; car- 
inae-earinulae-interstices with a row of hairs; scutellar carina not 
joining the fir.st ; humeri reduced ; wingless species. 

Measurements: {$ from Lambok, SMF) length, 21.8 mm; 
width, 7.15 mm. 



14 BEEVIORA No. 225 

Specimens examined (10): China: Yunnan (2 5, MNHN). 
Laos: Luang Prabang, Pang Bo {1 $ , BM) ; Ban Ban {1 $ , 
BM); Xieng Khouang {1 $ , BM). Vietnam: Tarn Dao (1 9 , 
BM). Localities which I have been unable to locate: "British 
Bootang" ( ? = Bhutan) (1 $ , MNHN) ; "Pedong" (2 $ , 
MNHN) ; Lambok, Sadjang {1 S , SMF C13910). 

Notes: G. feae is a species which seems to be confined to the 
Continent. (Lambok has not been located. It may, however, be 
Lombok. If the species really occurs on Lombok, it should also 
occur on Java, sympatrically with toreuta.) It is very similar to 
toreuta from Java. Both are wingless, with very reduced 
humeri. One of the best characters to distinguish the two seems 
to be the carinular interstice (see key). These two forms must 
also stem from a common ancestor, perhaps the same one from 
which 'batesi and carinifrons have been derived. 

Galeritula toreuta (Andrewes, 1933) 
(Figs. 3, 8) 

Galerita toreuta Andrewes, 1933, Treubia, 14:283-284 [holotype, $, Idjen 

Plateau, Java; British Museum (Natural History), examined]. 
Galerita insulindae Liebke, 1934, Arb. Morph. Tax. Ent., 1:280 [types, 
Pradjekan, east Java; Deutsches Entomologisches Institut, Berlin; not 
seen] ; Liebke, 1940, Fol. Zool. Hydrobiol., i(9:449 [proposed synonymy 
with toreuta], 
Galerita fortis Lowerens, 1952, Treubia, ^i:228 [type, central Java, Mt. 
Slamat, Kaligua; Lowerens collection (?); not seen]; Lowerens, 1955, 
Tijdschr. Ent., 98:5Q [proposed synonymy with toreuta]. 
It seems unnecessary to repeat a description of this well de- 
scribed species. 

Specimens examined (17) : Indonesia: Java, Soemberwringin, 
G. Raoeng (1 $ , BM, 3$ ,19 , MCZ) ; Tengger, Nongokodjadjar, 
1300 m (1 $ , BM) ; Idjes Plateau. Kenedeng Mts. (1 9 , holotype, 
is, BM) ; Kawie Mts. (4 5, 1$, MNHN); Toegoe (1 S , 
MNHN); Malang Romjer (1$, MNHN); Pradjekan (2 9, 
MCZ). 

RELATIONSHIPS AND ZOOGEOGRAPHY 
OF THE ASIAN SPECIES OP GALERITULA 

At present the Asian species of Galeritula are completely iso- 
lated from the other groups of the genus ; however, they must be 
considered as being relicts of what was probably a more wide- 
spread genus in the past, extending from Africa to South 
America. 



1965 ASIAN GALERITULA 15 

The relations of the species groups within the genus are very- 
obscure. According to Jeannel, Galeritula must have originated 
in Africa and from there it must have spread to South America 
and Asia when the continents were connected. No facts seem 
to support this view. The presence of two primitive genera of 
the tribe in the New World {Progaleritina Jeannel in North 
and Central America and Trichognathus Latreille in South 
America), and the immense diversification of the Neotropical 
species of GalcriUda, in my opinion suggest a New World origin. 

The Asian species have a relatively restricted distribution (see 
Fig. 9, which represents the distribution of orientalis. Inclusion 
of Celebes shows the limits of the distribution of the genus) . In 
the west these species reach India, in the northeast, Japan ; only 
a few forms have invaded the islands southeast of Asia, where 
they are found from Sumatra to southern Celebes and Flores. 
The genus is not known from Borneo, perhaps due to insufficient 
collecting there. Of the island forms, two are endemic to very 
restricted areas: toreuta is found only in Java, and carinifrons 
in southern Celebes. G. orientalis, the third species present on 
the islands, has the Avidest distribution of all Oriental species, 
occurring from India and Japan to Sumatra and Flores. It is a 
fully winged species, while toreuta and carinifrons are wingless. 
The two latter species are closely related to tatesi (winged) 
and feae (wingless) from the mainland. There has probably been 
one ancestral stock, which through isolation (especially on the 
Indonesian islands) has given origin to the 4 species. 

The other Oriental species form the second Asian group of 
species, that of G. orientalis, which is probably also derived from 
a single common ancestor. As seen above, the taxonomy of these 
three species is not yet well understood, since of two of them 
only the types are known. 

REFEEENCES 

Andrewes, H. E. 

1919. Papers on Oriental Carabidae. I. Ann. Mag. Nat. Hist., (9)3: 
469-483. 
Basilewsky, p. 

1963. Revision des Galeritininae d'Afrique et de Madagascar. Ann. 
Mus. Roy. Afr. Centr., ser. in 8°, Sci. Zool., No. 120, 93 pp., 
38 figs. 
Heller, K. M. 

1923. Die Coleopterenausbente der Stoetznerschen Sze-Tschwan-Expe- 
dition. Ent. Blaett., 19: 61-79, 3 figs. 



16 BREVIORA No. 225 

Jeannel, E. 

1949. Coleopteres Carabiques de la region malgache (Seme partie). 
Faune de 1 'Empire Frangais, 11: 767-1146, figs. 365-548. 
Jedlicka, a. 

1963. Monographie der Truncatipennen aus Ostasien. Ent. Abh. Mus. 
Dresden, 28 (7) : 269-579, 245 figs., 6 color plates. 

(Eeeeived 3 May 1965.) 



^Vl-UO' 



BREVIORA 

MiiseiLim of Connparative Zoology 



Cambridge, Mass. .Iily 15, 1965 Number 226 

THE LARVAL FORM OF THE HETEROMI (PLSCES) 

By Giles W. Mead 
Museum of Comparative Zoology, Harvard University 



The distinctive order Heteromi (Lypomi, Halosanriformes, or 
Notacantliiformes) includes about 25 species arrayed among 
about eight genera within the three families Notacanthidae, 
Halosauridae, and Lipogeiwidae. All appear to be bottom fishes 
which, as adults, live betAveen a fcAv hundred and about 3500 
meters. The placement of the order, which has been divided into 
two by many authors (Gill, 1889; Goode and Bean, 1896:129, 
162; Berg, 1940:453; and Lagler, Bardach and Miller, 1962:40), 
within the teleostean hierarchy has been problematical. The most 
penetrating study of possible relationships is that of Marshall 
(1962) who emphasized the unity of the order, and allied it with 
the true eels or Apodes, chiefly on the basis of significant resem- 
blances in swimbladder structure. The group was also recognized 
as a distinct order allied to the true eels within their superorder 
Elopomorpha by Greenwood, Rosen, Weitzman and Myers (1965) 
— a treatment in which I wholly concur. 

The reproductive biology of the group has been a complete 
enigma. Neither eggs nor larvae have been reported, although 
Marshall (1962) and Greenwood ct al. (1965) suggested that if 
the Heteromi shared common ancestry with the apodal fishes, a 
leptocephalus-like young should be expected. Similar speculation 
was included in the review of reproduction in the group provided 
by Mead, Bertelsen and Cohen (1964:583). It is thus most grati- 
fying to report here on a single, relatively large leptocephalus 
taken during the midwater trawling program of the International 
Indian Ocean Expedition. It is broken and badly damaged, but 
has the typical shape and transparency of a relatively large eel 
leptocephalus and head structure unmistakably that of a halosaur, 
probably of the genus Aldrovandia. This metamorphosing young 
was taken as follows : 



2 BREVIORA No. 226 

R/V Anton Bruun, Cruise VI, Sta. 351D, APB label 7354; 
29 June 1964; Southern Indian Ocean between 31°45'S, 65°08'E 
and 32°26'S, 65°05'E ; 0359 to 1507 hrs. ; 10-foot Isaacs-Kidd 
trawl equipped with Foxton Trousers (Foxton, 1963) set to trip 
at 350 m ; maximum depth of haul 1786 m ; probable depth of 
capture below 125 m; depth of bottom 4480 m. MCZ catalog 
number 43994. 

The body of this specimen Avhich lacks the terminal part is 
263 mm long and is composed of 250 somites. It lacks much of 
its skin, and the anal and the ventral fins are either unde- 
veloped or were lost during capture. The flanks are densely 
stippled with fine black pigment. The head is similarly but more 
darkly colored. Along the ventral profile are series of black spots, 
each continuous with its mate across the ventral midline and 
connected with the adjacent spots mid-ventrally by a fine line of 
dense pigment. These spots probably occur throughout the length 
of the fish, but both skin and pigment are missing in many areas 
(Fig. 1). Within or surmounting each ventral spot, at least 
anteriorly, is a pore which may be the precursor of the series of 
ventral luminous organs seen in Aldrovandia rostrata (Giinther, 
1887, pi. 59, fig. A). Spots and pores occur on every fourth or 
fifth somite, a relationship similar to that between light organs 
and rows of scales in certain species. Prior to preservation, the 
body was nearly as transparent as that of most eel leptocephali. 
The pectoral fin, which is set close behind the gill opening, is 
formed of twelve rays. The dorsal fin, badly damaged but includ- 
ing at least eight rays, is short-based and situated relatively far 
forward compared to that of other halosaurs, suggesting a sub- 
.stantial relative shortening of the post-dorsal part of the bodj^ 
with growth. The anus cannot be found. 

The head (Fig. 1) is about as broad as deep and terminates 
anterior to the gape in a prolonged fleshy snout. The gill open- 
ings are broad and continue anteriorly to near the symphysis of 
the lower jaw. Branchiostegal membranes are free from each 
other and from the isthmus. The gill membrane is thin and 
fragile. None of the osteological peculiarities noted by Marshall 
(1962:253) can be ascertained. Gill rakers are present and lath- 
like, are about as long as the opposite filaments, and number 4 + 
1 + 14 on the first arch. 

Jaws are poorly developed and bear teeth only at the tip of the 
mandible. Upper jaw bones are similar in structure to adult 
Aldrovandia, i.e. a somewhat flattened maxillary bone that 
abuts anteriorly on the end of the premaxillary which, with its 



1965 



LARVAL FORM OF THE HETEROMI 









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4 BEEVIORA No. 226 

opposite partner, courses transversel.v across the gape. Pores, 
presumably sensory, occur on the cheek (Fig. 1) and another pair 
is situated on the top of the head above the anterior edge of 
the orbit. 

The capture of a specimen, small but of adult appearance, 
which has been identified provisionally as Halosaurus nigerriynus 
Alcock, 1899, indicates that halosaurs can complete their meta- 
morphosis in midwater. This fish, 185 mm long, was also caught 
in the more southern latitudes of the Indian Ocean, as follows : 

R/V Anton Bruun, Cr. VI, Sta. 349B, APB label 7332; 26 
June 1964; 26°24'S, 65°02'E to 26°44'S, 65°05'E ; 0830 to 1425 
hrs. ; 10-foot Isaacs-Kidd trawl equipped with Foxton Trousers set 
to trip at 350 m ; maximum depth of haul 1470 m ; probable depth 
of capture below 125 m; depth of bottom 4571 m. MCZ catalog 
number 43993. 

These specimens thus clearly show that heteromous fishes have 
leptocephalus young so strikingly similar to those of the true 
eel that a close relationship between the two orders must be 
recognized, and that the j^oung of at least some of these benthic 
fishes are pelagic. Thus either a spawning migration of the adults 
must occur or, less likely, buoyant eggs spawned on the bottom 
must rise through a water column of substantial height. 

ACKNOWLEDGMENTS 

I wish to acknowledge, with my sincere thanks, the aid which 
the National Science Foundation has granted the author through 
GF 147 and its support of the field work aboard R/V Anion 
Bruun, International Indian Ocean Expedition, which resulted 
in the capture of the fishes here discussed. I am also grateful to 
Basil G. Nafpaktitis for the illustrations. 

LITERATURE CITED 

Berg, L. S. 

1940. Classific-ation of fishes both recont and fossil. Trav. Inst. Zooh 
Acad. Sei. U.R.S.S., 5:87-512. English trans., Ann Arbor, Michi- 
gan, 1947, 517 pp. 
Gill, T. X. 

1889. The halosauroid fishes typical of a peculiar order. Anier. Xat. 
23:1016-1017. 
Foxton, P. 

19C3. An automatic ()]ieiiins-closiiig device for larjjfe plankton nets and 
mid-water trawls. Jour. Mar. Biol. Assn. U.K., 43:295-308. 



1965 LARVAL FORM OF THE IIETEROMI 5 

GOODE, G. B., and T. H. Bean 

1896. Oceanic Ichthyology . . . U.S. Nat. Mus., Spec. Bull. 2, 553 pp., 
123 pis. 
Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers 

1965. Phyletie studies of teleostean fishes with a provisional classifica- 
tion of living forms. Bull. Amer. Mus. Nat. Hist., in press. 
GiJNTHER, A. 

1887. Eeport on the Deep-sea fishes collected by H.M.S. Challenger 
during the years 1873-76. Rep. Sci. Res. "Challenger," Zool., 
22. 335 pp. 
Lagler, K. F., J. E. Bardach, and R. R. Miller 

1962. Ichthyology. New York, 545 pp. 
^Marshall, N. B. 

1962. Observations on the Heteromi, an order of teleost fishes. Bull. 
British Mus. (Nat. Hist.), Zool., 9 (6) :251-270. 
Mead, G. W., E. Bertelsen and D. M. Cohen 

1964. Reproduction among deep-sea fishes. Deep-Sea Res., 11:569-596. 



^f^^LcJ 



BREVIORA 

MeseMim of Comparative Zoology 



Cambridge, Mass September 10, 1965 Number 227 

THE SPECIES OF HISPANIOLAN GREEN ANGLES 
(SAURIA, IGUANIDAE) 

By Ernest E. Williams 

INTRGDUCTIGN 

Certain Hispaniolan anoles of moderate to small size are 
dwellers in the crowns of trees, green in color, almost without 
other color or pattern, with short legs and long heads, and with 
relatively uniform squamation. These seem to be a natural 
species-group and not an assemblage of forms convergently 
adapted to the tree-crown niche. 

In many external details the Hispaniolan green anoles are 
strikingly similar; in most squamation characters the several 
species have overlapping variabilities, and there are only relatively 
few characters that assist in discriminating fully valid species. 

DEFINITION GF THE HISPANIOLAN 
GREEN ANGLE SPECIES-GROUP 

The nearly uniform green color and the tree-cro\\7i habitat 
provide the most immediate diagnostic features of this group. 
Coinciding with these are a number of morphological characters: 

1. Head long ( > 20% snout- vent length). 

2. Legs short (80 - 120% head length). 

3. Middorsal and flank scales smaller than ventrals, granular, 
subequal in size or very slightly and gradually enlarged in the 
middorsal line. 

4. Ventral scales hexagonal to square, subimbricate or juxta- 
posed, smooth or weakly keeled but never mucronate. 

5. Mental wider than long, 4 scales + 2 sublabials in contact 
with infralabials. Smallest anterior gular scales not smaller 
than 1 /6 first sublabial and usually larger. 

6. Dewlap small or vestigial. 

7. Lamellae under fourth toe always more than 20. 

8. Tail round or trigonal, never distinctly compressed; 4-5 
dorsal scales per verticil. 



2 BREVIORA No. 227 

SPECIES DIFFERENCES 

The green anoles of Hispaniola divide into two subgroups, 
each a superspecies with two species. The species of the first 
subgroup {Anolis chloroajanus Dumeril and Bibron 1837, and 
A. coelestinus Cope 1862) are widespread and very common in 
the lowlands and up to at least 1500 meters; those of the second 
subgroup (Anolis aliniger Mertens 1939, and A. singularis n. sp.) 
occur only at elevations well above sea level and appear always 
to be scarce. The species of subgroup 1 occur sympatrically with 
those of subgroup 2, but the species within each subgroup are 
primarily allopatric. The two wide-ranging species of the first 
subgroup show enough geographic variation to permit description 
of subspecies; the subspecies will not, however, be discussed in 
this paper, which is concerned solely with species differences. 

In a majority of the characters in which anoles are distinguished 
by taxonomists, there is conspicuous overlap between these two 
subgroups and even more overlap between the species of a single 
subgroup. It is significant that, lacking the distributional evi- 
dence that is now available, Mertens (1939) described aliniger 
(of the second subgroup of my terminology) as a subspecies of 
chlorocyanus (of the first subgroup) and was not very sure (p. 62) 
that coelestinus was a species distinct from chlorocyanus. 

Table 1 shows the extent of this overlap very strikingly. Though 
in the better represented species of the first subgroup there are 
evident differences in the means of the several numerical charac- 
ters, it is quite as evident that there are individuals it would not 
be possible to place on these characters alone. 

Tables 2 and 3 show the very few qualitative characters that 
seem useful in discriminating the species of Hispaniolan green 
anoles. Size also differs. A. chlorocyanus and A. coelestinus may 
exceed 70 mm in snout-vent length, A. aliniger and A. singularis 
do not reach much over 50 mm. 

SUBGROUP 1: A. CHLOROCYANUS - A. COELESTINUS 

These two species overlap very little geographically. One has 
a distribution north, the other south of the Cul de Sac trench 
(still partly below sea level) which formerly divided Hispaniola 
into northern and southern islands. At the southern edge of this 
trench, now dry and very arid land, there is contact and sporadic 
real sympatry, always it appears, with one species or the other 
predominating. The actual zone of contact has never been 
mapped and is known at the moment only from occasional tran- 
sects. A. coelestinus occurs in Port-au-Prince and extends to 
Damien, but A. chlorocyanus occurs in the area too. 



1965 



HISPANIOLAN GREEN ANGLES 



Even in tliia area of contact there is no indication of reinforce- 
ment of species differences in body squamation. There are only- 
average differences in the size of the head scales which are reflected 
in counts of loreal rows, scales across snout, scales between inter- 
parietal and supraorbital semicircles, etc. However, the best 
differences between the two species are in body color (the presence 
in codestinus of the labial white streak which is continued above 
the shoulder, and its absence in chlorocyanus) , and in dewlap 
color and squamation (very large dewlap scales and black dew- 
lap skin in chloroajanus, small dewlap scales and relatively unpig- 
mented skin in coelestinus) (Fig. 1). Thus, in color characters 
there is some evidence of reinforcement of species difference 
because of secondary contact. 





Fig. 1. Scales along edge of dewlap. Above: Anolis coelestinus, MCZ 
64883, Ca-ira near Leogane, Haiti; below: Anolis chlorocyanus, MCZ 80719, 
Nan Palmiste, Gonave Island, Haiti. Drawn to the same scale; the speci- 
mens have the same snout-vent length. 



Both body color and dewlap color differences, while they hold 
over most of the species ranges, are lost or weakened in popula- 
tions of these species remote from the zone of contact. Thus the 
Isle Vache population of coelestinus lacks the white labial to 
shoulder streak of other coelestinus populations. At least pre- 
served specimens of coelestinus remote from the Cul de Sac tend 
to show darker dewlap, occasionally almost to a degree that 



4 BREVIORA No. 227 

would be confusing were not the dewlap scales consistently small. 

In chlorocyanus also, geographic variation slightly weakens the 
diagnostic differences. The population described by Mertens 
from the vicinity of Santo Domingo City as A. chlorocyanus 
cyanostictus has reduced the black of the dewlap and exhibits a 
cadmium yellow basal spot. 

However, both the Isle Vache population of coelestinus and the 
Santo Domingo City population of chlorocyanus are very small 
segments of the total range of these species. The non-distinctive 
segment of coelestinus is isolated on an island far to the west; 
the exact range of chlorocyanus cyanostictus is very limited and on 
present evidence very uncertain: typical chlorocyanus have been 
collected in Santo Domingo itself. 

Both species are characteristic inhabitants of mesic lowland 
forests and appear not to extend to the highest peaks. They are, 
for example, unreported in the Foret des Pins, Massif de la Selle 
in Haiti, or at Valle Nuevo in the Cordillera Central in the 
Dominican Republic. Both these localities have been visited by 
several investigators and the apparent absences are quite probably 
real. 

SUBGROUP 2: A. ALINIGER-A. SINGULARIS N. SP. 

This pair of species is not known to overlap at all, but they 
both occur on the jMassif de la Selle. 

The previously described member of this pair, A. alinigcr, was 
until recently known only by the unique type, and was regarded by 
its describer jMertens as a subspecies of chlorocyanus. A. aliniger 
is, however, widely sympatric with chlorocyanus. A specimen of A. 
aliniger collected by Clayton Ray and A. S. Rand at 7 km N Car- 
pintero Prov., San Juan, Dominican Republic, was collected on a 
tree ca. 45 meters from a specimen of ^4. chlorocyanus collected on 
a pole fence. The head scales of these two specimens are shown 
in Figure 2. In the Constanza area. La Vega Province, Dominican 
Republic, natives collecting for Juan Rivero brought in A. aliniger 
and A. chlorocyanus at the same time. 

A. aliniger is one of the most peculiar of anoles, not indeed in 
general habitus which is that of a small and somewhat stockier 
version of chlorocyanus or coelestinus, but in the singular feature 
which gives it its name. This is the strange coloration of the axilla, 
bright orange or yellow followed by a larger or smaller spot of in- 
tense black. Thi.s is present in both sexes and very conspicuous 
in freshly preserved specimens, but the yellow or orange is, of 
course, (juickly bleached out by alcohol. There is, however, an 



1965 



HISPANIOLAN GREEN ANGLES 





JBC 



Fig. 2. Head squamation. Above: A. aliniger, ]^ICZ 57403; below: 
A. chlorociianus, MCZ 57473. Both specimens from 7 km N Carpiiitero, Prov. 
San Juan, Dominican Republic. 



area of scaleless skin which represents the area formerly occupied 
by the orange spot (Fig. 3). Since the dewlap is hardly developed, 
almost non-functional, it is very possible that the orange, made 
more conspicuous by the black behind it, is a flash pattern used 
in some fashion in high intensity agonistic behavior. This is at 
present a mere suspicion ; the only specimens of aliniger seen have 
been merely captured and preserved, or only very briefly ob- 
served. 



6 BREVIORA No. 227 

The best report of the ecology and behavior of A. aliniger is 
that by James Lazell (field notes, December, 1963) at Paraje 
la Palma near Constanza: "Up in largish trees along the edge of 
the woods by the stream. Since they retreat upwards, collect- 
ing them is merely a problem of having a long enough pole. Just 
like coelestinus-chlorocyanus, therefore, in habits — except for the 
vertical flattening in display. In display the whole body is ver- 
tically compressed — showing much of the venter. Extended, 
the throat fan is quite small." Lazell's observations were unfor- 
tunately terminated by rain which prevented him from seeing any 
further specimens during the remainder of his stay. 

The vertical compression, according to Lazell's sketch from life, 
emphasizes the yellow color of the belly. Lazell saw, however, no 
instance of exposure of the black and orange axillary pattern. 

A number of descriptions of color in life are available for aliniger. 
They appear to indicate not only that the green of the light phase 
is different in tone from that of the species of subgroup 1 but also 
that there is a greater play of patterns and tones in the darker 
state. (See also ^4. singularis below.) In view of our very inade- 
quate knowledge of the species, I quote these color descriptions 
in full. 

Mertens (1939, translated): "In life this Anolis when caught 
was a uniform chocolate-brown with a large pitch black spot in 
each axillary region. In the bag in which it was transported the 
lizard turned blue-green, the black axillary spot remaining unal- 
tered. The dewlap was bright green, the tip of the tail black." 

Rand (field notes, 1958). Female — 7 km N of Carpintero: 
"Gray green above, head grayer, upper lip white, rear of thigh 
with a dark line, axilla bright yellow with smaller black spot be- 
hind it." 

Lazell (field notes, 1963): "Duller and bluer green than coe- 
lestinus-chlorocyanus. Venter, throat fan and frosted spots on 
sides (especially shoulder region) bright saffron yellow. Orbital 
skin butter yellow. Axillary spot plain black. 

"Changes, when unhapjjy, to lichenate frosted grey-brown with 
white. A pattern of large dark bilaterally arranged spots emerges 
with transverse bands — especially posteriorly. Loreals and ir- 
regular stripe through eye emerge slate blackish." 

Schwartz (field notes, 1964). ASFS V 1625— 12 km S of Ca- 
brera de Loma: "Dorsal ground color blackish brown to olive with 
green sacral 'butterfly' marking and tail banded olive and cream. 
A white subocular mark. Ventral ground color dull greenish." 



1965 



HISPANIOLAN GREEN ANGLES 





Fig. 3. Axillary squamation. Above: A. chlorocyanus, MCZ 57473, 7 km 
N Carpintero, Prov. San Juan, Dominican Republic; below: A. aliniger, 
MCZ 79341, Paraje La Palma, Municipio Constanza, Prov. La Vega, 
Dominican Republic. 



8 BREVIORA No. 227 

The type locality of Anolis chlorocyanus aliniger Mertens was 
Paso Bajito on the northern rim of Valle Constanza in La Vega 
Province, Dominican Republic. Most of the recent specimens 
have been taken within that valley itself. However, a female has 
been taken in San Juan Province and a male in Dajabon Province. 
All specimens are from elevations near or above 600 meters. 

These Dominican Republic localities appeared to place A. 
aliniger as an anole of the Cordillera Central or its immediate 
foothills, present at moderate elevations — within the pine zone 
— and widely sympatric with A . chlorocyanus. 

It was with some surprise, therefore, that two specimens were 
found in a collection of .4. coelestinus made by the Whiteman 
brothers at Furcy in Haiti. Furcy is south of the Cul de Sac 
trough that separates Hispaniola into northern and southern faunal 
areas — formerly separate islands. 

It is not, of course, especially surprising that a form common in 
the northern faunal area should sometimes penetrate some dis- 
tance or even extensively into the southern area. This seems a 
frequent phenomenon. What is surprising about the discovery of 
aliniger at Furcy, Haiti at ca. 1500 meters at the north margin of 
the Massif de la Selle is that this is not a lowland species, which 
could without special difficulty cross the dry and hot Cul de Sac 
Plain, but a creature of higher elevations, cooler temperatures and 
of quite different floral associations from those which are now seen 
in either the desertic or mesic habitats of the lowlands. Essen- 
tially, the discovery of aliniger at Furcy is analogous to finding a 
species of one island present on one cape of a neighboring island. 

The ecology of aliniger appears to require that at some past 
time the Cul de Sac gap must have been climatically and floristi- 
cally passable for the species: the passage from one montane 
island to the other must have been made at a time of much cooler 
temperatures — presumably the last cool-wet period of the 
Pleistocene. 

The specimens from Furcy remain the only record of the 
species in Haiti. Doubtless in part this absence of record is an 
artifact of collecting. A spur of the Cordillera Central enters the 
northeast of Haiti, but little or no collecting has been done there. 

At present the known localities for aliniger are: Dominican 
Republic. La Vega Province. Below Paso Bajito at about 900 m 
(type locality), Senckenberg 25825. Valle Constanza, MCZ 56912. 
Tireo near Constanza, MCZ 56913-15. Paraje La Palma, east of 
Constanza, MCZ 75140-41, 79341-43. El Rio, AMNH 44852-53. 
San Juan Province. Seven kilometers north of Carpintero, 



1965 HISPAOTOLAN GREEN ANGLES 9 

MCZ 57463. Dajabon Province. Twelve kilometers south of 
Loma de Cabrera, ASFS V 1625. Haiti. Departement de VOuesl. 
Furcy, MCZ 63444-45. 

Over most of the southern island of Hispaniola — south of the 
Cul de Sac trough in both Haiti and Hispaniola — the absence of 
alinigcr is plausibly accounted for by the presence of a closely 
related species — essentially alinigcr without the axillary light 
and dark spots. This peculiar species is known from very few 
specimens, ever3^one of which is from a different locality. I have, 
therefore, called this new species by the Latin adjective which 
means "one at a time." 

Anolis singularis new species 

Holotypc: MCZ 72043, adult male, Pourcine, Massif de la Hotte, 
Haiti, collected by Frangois Vuilleumier, 31 December 1962. 

Parafypes: MCZ 13778, La Gonave Island, Haiti, G. AL Allen 
coll. 1919; YPAI 3229, Nan Cafe, La Gonave Island, P. Humphrey 
and Sarita Van Vleck coll., March 26, 1959; YPM 3194, Foret des 
Pins, Massif de la Selle, Haiti, P. Humphrey and Sarita Van Vleck 
coll., February 19, 1959; AMNH 51728, Valle de Polo, Dominican 
Republic, W. G. Hassler coll., September 14, 1932; ASFS V 2608, 
5 km NE Los Arroyos, 5800 feet {ca. 1750 meters) elevation, 
Pedernales, Dominican Republic, D. C. Leber coll., June 27, 1984; 
ASFS V 2985, 30 km N of Pedernales, 2680 feet {ca. 810 meters), 
Pedernales, Dominican Republic, hatched from an egg collected 
under a limestone rock in Cajetal, July 3, 1964, by R. Thomas; 
hatched July 16, 1964. 

Diagnosis: An anole most closely allied to ,4. alinigcr Mertens 
but differing in the absence of a scaleless, highly pigmented 
axillary area. Like A. alinigcr, the new species differs from 
A. chlorocyanus and A. coelestinus in the greater enlargement of 
scales around the interparietal (Fig. 4), and in reduction of the 
dewlap in males. 

Description: Head scales rather large, 6-8 scales across snout 
between second canthals. Frontal depression feeble, scales within 
it not smaller than those surrounding it. Five to seven scales 
border rostral posteriorly. Anterior nasal scale in contact with 
rostral. Three to four scales between supranasals. 

Supraorbital semicircles separated by 1-2 scales, separated 
by one row of scales from supraocular disk. Seven to eleven 
smooth scales in the poorly defined supraocular disk which is 
separated from supraciliary by 2-3 rows of granular or sub- 
granular scales. One rather short supraciliary, flanked medially 



10 



BREVIORA 



No. 227 



by smaller scales and continued posteriorly by granules. Six to 
eight canthals, second and third largest, decreasing thence for- 
ward, anteriormost under rostral. Three to five loreal rows. 





Fig. 4. Head squamation. Above: A. singularis Holotype, MCZ 72043; 
below: A. coelestinus, MCZ 74708. Both specimens from Pourcine, Massif 
de la Hotte, Haiti. 



Temporal scales subgranular, smallest in center. A poorly 
defined intertemporal line. Supratcmporals larger than temporals, 
increasing in size toward interparietal. Interparietal larger or 
slightly smaller than ear, separated from semicircles by 2-3 scales. 
Scales surrounding interparietal largest laterally but a very 
distinct zone of 5-6 rows of enlarged scales posterior to inter- 
parietal. 



1965 HISPANIOLAN GREEN ANGLES 11 

Suboculars broadly in contact with supralabials, anteriorly sepa- 
rated from canthal ridge by two scales, posteriorly merging into 
temporals. Seven to eight supralabials to center of eye. 

Mentals broader than long, in contact posteriorly with 6-7 scales 
between infralabials (2 sublabials and 4-5 smaller scales) ; 5-7 sub- 
labials in contact with infralabials. Central throat scales small, 
swollen, smooth. 

Dewlap: Hardly differentiated, only indicated as a longitudi- 
nal fold; scales larger than throat scales, smaller than ventrals. 

Trunk: Middorsals hardly enlarged, grading very gradually 
into flank granules. Axilla with normal granular scales. Ven- 
trals larger, smooth, quadrate, juxtaposed, transverse. Postanals 
enlarged in male. Scales posterior to vent smooth. 

Limbs: Largest forelimb scales smaller, largest hindlimb scales 
larger or smaller than ventrals, both weakly unicarinate. Supra- 
digital scales smooth; 21-23 lamellae (27 in one hatchling) under 
phalanges ii and iii of fourth toe. 

Tail : Verticils with four scales above, three below. Scales sub- 
equal. 

Color in life: YPM 3194, Foret des Pins, Haiti: Emerald green 
labials and limbs. Yellow eyelids, oUve head and back, yellow 
green venter, last quarter of tail black with yellow tip. 

YPM 3229, Nan Cafe, La Gonave Island, Haiti: Head silvery 
grey mottled with turquoise and brown, the latter extending to the 
shoulder. A brown spot just behind shoulder. Back silver gray 
mottled with turquoise. Tail and limbs very pale buff, tail banded 
with sky blue. Underparts pale turquoise becoming lemon on 
femora and at vent. 

ASFS V 2608. 5 km NE Los Arroyos, Dominican Republic: 
Dorsal ground color gray to brown, a pattern of dark chevron- 
shaped middorsal blotches and smaller, roughly triangular lateral 
blotches. Light lateral stripes. Venter faint rust, speckled with 
greenish. 

ASFS V 2985 (hatchhng). 30 km N Pedernales, Dominican 
Republic: Dorsal ground color green with longitudinal darker 
stripes, especially middorsally. 

Species status. On present evidence, A. aliniger and A. singu- 
laris are wholly allopatric. The question of species or subspecies 
status does therefore arise. My decision to describe singularis as 
a species is based upon two considerations: (1) a high valuation 
placed upon the axillary differences in pigmentation and squama- 
tion, which I assume to imply behavioral differences; (2) the 
presence of typical aliniger at Furcy appears to imply a capacity 



12 BREVIORA No. 227 

to invade the territory of an allied taxon — a feature more prob- 
able for a species than a subspecies. Both these points require 
further elaboration. 

(1) It is frequent in Anolis for the most conspicuous and taxo- 
nomically useful differences between very closely allied species to 
be in structures such as the dewlap that are involved in species 
recognition and intraspecies agonistic behavior. Thus, in sub- 
group 1 of the present paper, the southern island form A. coe- 
lestinus has a dewlap with pale skin and small scales while the 
northern island species A. chlorocyanus has a dewlap with pig- 
mented skin and greatly enlarged scales. A. semilineatus and A. 
olssoni differ in an exactly similar way, A. semilineatus having 
the dewlap skin pale, the dewlap scales small, A. olssoni having 
pigmented dewlap skin and greatly enlarged dewlap scales. (In 
the latter case there are other strong differences.) 

Dewlap color, however, does vary intraspecifically in Anolis 
(e.g. in A. distichus, A. hrevirostris, A. cybotes) and sometimes 
individually, as, of course, body pattern may do also. To what 
extent such color variation disturbs species recognition is quite un- 
known, and equally unknown are any differences in display be- 
havior between allied full species such as .4. coelestinus and .4. 
chlorocyanus or A. semilineatus and A. olssoni. 

In the present case, I have chosen to infer. that the orange 
axillary spot of aliniger, attended as it is by loss of normal squa- 
mation in that area and contrasted with the unmodified axilla of 
singularis, is comparable to the species recognition marks that 
distinguish species pairs and not to the simpler color variations 
that may occur intraspecifically. 

(2) The species pair A. coelestinus- A. chlorocyanus may be the 
only instance in which the geographic boundary between species 
is exactly that area of the island which at one time, as an open 
seaway, divided Hispaniola into northern and southern parts. It 
is certain that this seaway has been profoundly important for 
origin of many widel}^ distributed Hispaniolan species; Mertens 
(1939) cari^^ recognized its importance. (See also Williams, 1961.) 
But it is in fact very unusual for allied species which have origin- 
ated north and south of this important zoogcographic boundary to 
be precisely limited by it now that the seaway has become dry 
land. The situation which I described in the species pair .4. srmi- 
linvatus-A. olssoni is commoner: one or the other or perhaps ijoth 
members of the species pair interpenetrate the range of the other 
to a greater or lesser extent. Usually, ii would appear, there is, 



1965 HISPANIOLAN GREEN ANGLES 13 

as in .1. scmilincatus-A. olssoni, sufficient ecological difference be- 
tween the members of a species pair to permit this, but in the case 
of A. coelesfiniis-A . chlorocyanus the ecologies are too similar to 
permit anything but a stand-off, with a very narrow, perhaps 
fluctuating zone of sympatry. 

The case of A. aUnigcr-A. singnlaris, which is surely another 
example of a northern island-southern island pair, differs from that 
of ^4. coelestinus-A. chlorocyanus in that these are inhabitants of 
cooler upland areas and should now be quite separated by the 
whole extent of the hot Cul de Sac Plain and by much of the 
mesic woodlands on either side. That any A. aliniger occurs on 
the south side of the Cul de Sac Plain indicates, as I have suggested 
above, the former existence of a zone of passage right across the 
Cul de Sac for species now characteristic of cool upland climates. 
At such a period A. aliniger invaded the southern island and may 
well have met A. singnlaris. If contact occurred, certainly no 
merging of populations resulted. The Furcy aliniger are quite un- 
modified, as typical as any from Valle Constanza. Perhaps there 
was a slight ecological difference with A. singnlaris living at higher 
elevations than A. aliniger. (The known true south island records 
of A. singnlaris would fit this pattern but the Gonave records 
would not.) Or perhaps ^4. aliniger displaced A. singnlaris in part 
of the latter 's range. 

Too little is known about either taxon to permit any but the 
most provisional hypothesis. It does, however, seem plausible to 
regard the invasion of the southern island by aliniger, without any 
indication of compromise of its characters, as evidence that it and 
singnlaris have indeed achieved species status and to infer that 
A. aliniger and A. singnlaris are, like A. coelestinus-A . chlorocyanus, 
too similar ecologically to occur together. 

ACKNOWLEDGMENTS 

I am indebted to Dr. Albert Schwartz (Albert Schwartz Field 
Series, ASFS), Mr. C. M. Bogert, American ]Museum of Natural 
History (A]\1NH), and Dr. Charles Reed, Yale Peabody Museum 
(YP]\1) for the privilege of examining specimens under their care. 
This paper, as part of a general study of West Indian anoles, has 
been supported by National Science Foundation grants NSF- 
G 16066 and NSF-GB-2444. The illustrations were prepared by 
Mr. Joshua Clark. I have had the advantage of utilizing counts 
and tabulations made by Dr. A. S. Rand. 



14 



BREVIORA 



No. 227 




in 

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O 

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o 
o 

o 



a 

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a 

en 

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-1.3 



a 
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1965 



HISPANIOLAN GREEN ANGLES 



15 



REFERENCES 

COCHKAN, D. M. 

1941. The herpetology of Hispaniola. Bull. U. S. Nat. Mus., 177: 
1-398. 
Mertens, R. 

1939. Herpetologische Ergobnisse einer Roise nach der Insel Hispaniola, 
Westiudien. Abhandl. Senckenberg. Naturf. Ges., 449: 1-S4. 
Williams, E. E. 

1961. Notes on Hispaniolan herpetology. 3. The evolution and rela- 
tionships of the Anolis seniilinealus group. Breviora, Mus. Comp. 
Zool., No. 136: 1-S. 

(Received 10 May, 1965.) 

Table 1. 
Scale characters of the Hispaniolan green anoles 





coelestinus 


chlorocijanus 


aliniger 


singularis 


scales across snout 


6-10 


4-7 


6-9 


6-8 


loreal rows 


4-7 


3-5 


3-4 


3-5 


scales between 










supraorbital semicircles 


1-3 


0-2 


0-1 


1-2 


scales between inter- 










parietal and semicircles 


2-6 


1-5 


2-4 


2-3 


lamellae 


23-32 


27-34 


21-24 


21-23' 



27 in one hatchling, ASFS V 2985 



Scale charact 

coelestinus 

scales behind 
interparietal grad- 
ing rather gradu- 
ally into dorsal 
scales 



Table 2. 
ers differentiating the Hispaniolan green anoles 



chlorocyanus 
as in coelestinus 



aliniger 
Scales behind 
interparietal not 
grading into 
dorsals but 
several rows 
abruptly larger 



singularis 
as in aliniger 



size of scales of 
dewlap ca. = 
ventrals 



size of scales of 
dewlap > > 
ventrals 



dewlap vestigial, 
size of scales of 
dewlap area < 
ventrals 



as in aliniger 



supradigital scales 
multicarinate 



as in coelestinus supradigital scales as in aliniger 
smooth 



axillary area with 
normal granular 
squamation 



as in coelestinus axillary area with- as in coelestinus 
out granular scales 



16 



BREVIORA 



No. 227 



Table 3. 
Color characters differentiating the Hispaniolan green anoles 



coelestin us 
a white line on 
supralabials pro- 
duced posteriorly 
to above the 
shoulders (absent 
in He Vache popu- 
lation) 



chlorocyanus aliniger 

never any as in chlorocijanus 

white markings 

on side of head 

or neck 



singularis 

as in chlorocyanus 



axillary area with 
vertical black and 
light bars or 
unpatterned 



as in coelestinus 



axillary area with 
a light spot 
(orange in life) 
followed by an 
ink-black blotch 
of larger or 
smaller extent — 
both concealed by 
normal position 
of arm 



as in coelestinus 



dewlap scales blue, 
skin olive-gray 



dewlap scales 
white to blue, 
skin with black 
or (cyanostictus) 
cadmium 
yellow basal 
spot 



dewlap skin and 
scales greenish 



as in aliniger 



BREVIORA 

Mesemm of Coimpsirative Zoology 

Cambridge, Mass. September 15, 1965 Number 228 

RELATIONSHIPS AMONG INDO-AUSTRALIAN 
ZOSTEROPIDAE (AVES) 

By 

Ernst Mayr 



During- preparation of the maniiscriiDt of the Zosteropidae for 
Peters' Checklist of Birds of the World, I reviewed once more 
the relationships of the Indo-Anstralian -white-eyes. The first 
attempt to arrange these species in natural groups was made by 
Stresemann in a pioneering work in 1931. The revisionary 
studies of the ensuing 25 years were summarized by G. F. Mees 
in a very thorough systematic review of the Indo-Australian 
Zosteropidae (1957, 1961). Mees' work is based not only on an 
exhaustive analysis of the literature, but also on an examination 
of most of the available museum material. His fundamental 
monograph will remain for many decades the basis of all taxo- 
nomie research in Indo-Australian white-eyes. In a few cases 
my own interpretation of relationship differs from his, resulting 
in a somewhat different sequence of species. This paper may serve 
to explain the reason why, in these few instances, I have 
adopted a sequence different from that chosen by Mees in his 
systematic review. I am deeply indebted to Dr. Mees for much 
valuable information on Zosteropidae. 

Absolute criteria of relationship in the Zosteropidae do not 
exist at the present time, and neither Mees nor I can prove that 
the sequence adopted by one of us is "more correct" than the 
sequence adopted by the other. White-eyes have characteristic 
songs and call notes, and perhaps analysis of these and other 
behavioral characters may lead to a better understanding of rela- 
tionships. I know of no other group of birds in which close 
relatives, for example the subspecies of Zostcrops atrifrons or 
the semispecies of the superspecies griseotiitcta, may differ more 
from each other than do distantly related species. Indeed some 



2 BREVIORA No. 228 

Oriental species are almost indistinguishable from African forms, 
from which they must have been isolated since remote times. 

ZosTEROPS CEYLONENSis Holdsworth 

This species is often considered closely related to Z. palpcbrosa, 
and Mees (1957, p. 22) lists it immediately preceding palpchrosa. 
Yet, he states correctly, "morphologically Z. ceylonensis is un- 
doubtedly rather closer to several other species than to palpe- 
hrosa" (ibid., p. 26). For this reason I have placed it after 
palpchrosa and closer to some of the Indonesian species from 
which it might possibly have been derived. 

ZoSTEROPS ERYTHROPLEURA Swinhoe 

This species, with its rufous flanks, does not resemble any of 
the species (pajpehrosa and japonica) with which it is usually 
placed. Being also the only white-eye restricted to the Palearc- 
tic region, I prefer to indicate its distinctness by placing it 
first in the sequence, as had been previously done by Stresemann 
(1931, p. 206). 

ZoSTEROPS CONSPICILLATA Kittlitz 

Mees (1957) lists this Micronesian bird as the last species of 
the genus. To me this species does not seem nearly as aberrant 
as Z. cinerea. Indeed, in spite of its paleness, a frequent char- 
acter in island birds, conspiciUata resembles in some ways the 
japonica-palpehrosa assemblage. Since several Micronesian birds 
were derived from the west (e.g. Acroccphalus) , relationship of 
conspiciUata with japonica is a distinct possibility. It is hoped 
that placing the species earlier in the sequence will bring it 
closer to its real relatives. 

ZoSTEROPS WALLACEi Finsch 

As Mees has stated rightly, this is an old and peculiar en- 
demic. It seems distantly related to the western group of species 
(atricapilla, everetti, nigrorum, and others) and I have there- 
fore placed it earlier in the sequence. 

ZoSTEROPS FLAVA-CHLORIS-LUTEA grOUp 

I have adopted Mees' sequence for the sake of uniformity. I 
still feel, how^ever, that these species are more closely related 



1965 RELATIONSHIPS AMONO ZOSTEROPIDAE 3 

to each other than believed by Mees. To separate lutea from 
chloris by 14 other species does not seem to be the best possible 
arrangement. 

ZosTEROPS CONSOBRINORUM Meyer 

This species is so similar to Z. chloris citrmclla that in any 
other genus one M'ould consider them conspecific. Even though 
I have retained Z. consohrinorum as a full species, I have placed 
it next to chloris, while I now treat the very distinct peripheral 
forms grayi and uropygialis as full species (see also Mees, 
1953, 1961). 

ZoSTEROPS ATRIFRONS grOUp 

I agree, on the whole, with Mees' (1961) arrangement. How- 
ever, Z. atriceps is best listed before the atrifrons-minor-deli- 
catula series, because the latter is close to the forms on the 
islands east and northeast of New Guinea {meeki, hypoxantha) 
and should not be separated from them by atriceps. 

In this group of species close relatives may appear rather 
different (belly, yellow or white ; forehead, black or olive ; eye- 
ring, absent or broad; throat, orange, yellow or whitish). The 
delimitation of the species is therefore a difficult task. Two 
of the most distinct forms, minor and delicaUila of New Guinea 
are connected by the intermediate forms chrysolaema and 
rothschildi. Mees (1961) quite rightly combines atrifrons and 
minor in a single species, but if one goes that far one must 
also include meeki (close to delicatula) and hypoxantha. 

I have maintained Z. mysorensis as a separate species, because 
in its combination of characters (no yellow on throat, heavy 
gray wash on breast and flanks, olive forehead, absence of 
eye-ring, blackish loral region, and pale yellow under tail 
coverts) it resembles some other species (e.g. ugiensis) more 
closely than atrifrons. 

ZoSTEROPS NATALIS Listcr 

Mees (1957) notwithstanding, there is much to indicate that 
this species is closer to one of the east Indonesian or Australian 
species {chloris, lutea, etc.) than to any of the Malaysian 
species. I agree in this with Stresemann and Chasen. Mees 
(1957) makes the peculiar comment that my (Mayr, 1944) 



4 BREVIORA No. 228 

association of this species with lutca, instead of with citrinella, 
is an ' ' unfortunate choice, ' ' overlooking^ that I include citrinella 
in lutea in the cited paper. 

ZosTEROPS RENDOVAE Tristram 

The description of rendovac was based on a Rendova Island 
specimen misidentified as Tephras olivaceus Ramsay, but newly 
named rendovae since the name olivaceus Ramsay (nee Certhia 
olivacea Linnaeus 1766) was considered unavailable. That the 
original author, Tristram, considered rendovae a name for the 
Rendova bird far more than a replacement name for olivaceus 
is evident from his subsequent statement (1894, p. 30) : "I give 
a figure (PI. Ill, fig. 2) of Zosterops rendovac of Rendova 
Island . . . which I described in the Ibis for 1882, p. 135." 
Galbraith (1957) has well stated the reasons for retaining the 
name rendovae for the Rendova White-Eye to which it had 
been applied universally from 1882-1955, including general books 
in ornithology and evolutionary biology. 

Zosterops lateralis Latham 

The arrangement of the Australian races is largely based on 
unpublished research kindly made available to me by Drs. A. 
Keast and G. F. Mees. 

The resulting sequence of species of Indo-Australian Zos- 
terops is as follows : 

erythropleura uropygialis 

japonica anomala 

palpehrosa (ttric(ps 

ccylonensis atrifrons 

conspicillata mysorensis 

salvadorii fuscicapilla 

atricapilla h urucnsis 

everetti kuehni 

nigrorum novaeguincae 

montana metcalfii 



wallacei natalis 

flava lutea 

chloris griseotinctu 

consohrinorum renneUinna 

grayi vellaJarrlla 



1965 RELATIONSHIPS AMONG ZOSTEROPIDAE 5 

lutcirostris minuia 

rendovae xanthochroa 

murphyi lateralis 

ugiensis strenua 

stresemanni tenuirostris 

sanctaecrucis alhogularis 

samocnsis inornata 

cxplorator cincrea 
flavifrons 

The ABERRANT GENERA OF InDO- AUSTRALIA 

Some white-eyes differ from the normal structure or colora- 
tion of the genus Zosterops, as represented by a species like 
palpchrosa or lateralis, to such an extent that they have been 
separated generieally. If all the larger white-eyes, with some- 
what aberrant coloration and a longer or heavier bill, could 
be placed in a single genus, no one would mind. The fact 
of the matter is, how^ever, that 13 genera have been proposed 
to accommodate 18 species. Excluding Lophozosterops (with 
6 species), there are 11 generic names for 13 species. Some 
further simplification is possible by combining Sanfordia with 
Woodfordia, as well as the Micronesian Kuharyum, Megazos- 
terops, and Cinnyrorhyncha with Rukia. In the "Oreosterops 
group" of authors, Mees (1953, pp. 57-66) recognizes six 
genera, reduced in 1957 to five by combining Apoia with Lopho- 
zosterops. Of these five genera, three {Madanga, Tephrozos- 
terops, and Oculocincta) are monotypic, while Heleia has two 
species in one superspecies. The five genera seem to form a 
natural group and a renewed analysis may result in further 
lumping, perhaps of all five genera into Heleia. 

The sequence chosen by Mees (1953) does not seem quite 
natural. By starting with the species that is most like Zosterops 
and also keeping the pattern of geographic distribution in mind, 
we arrive at the following sequence: Tephrozosterops (staJhcri), 
Madanga (ruficollis), Lophozosterops {pinaiae, goodfellowi, 
squamiceps, javanica, superciliaris, dohertyi), Oculocincta 
(squamifrons) , and Heleia {miielleri, crassirostris) . 

I entirely agree with Mees {in litt.) that Hypocryptadius 
Hartert does not appear to be a white-eye. Fresh material is 
needed to determine whether it should go to the Monarcliini, 
Sylviinae, Timaliinae, or some other group. 



6 BREVIORA No. 228 

LITEBATURE CITED 

Galbraith, I. C. J. 

1957. On the application of the name Zosterops rendovae Tristram, 
1882. Bull. Brit. Orn. Club, 77:10-16. 
Mayr, Ernst 

1944. The birds of Timor and Sumba. Bull. Amer. Mus. Nat. Hist., 
83:123-194. 
Mees, G. F. 

1953. An attempt at a natural classification of certain Zosteropidae 
of the Indo-Australian archipelago. Zool. Meded., 32:57-68. 



1957. A systematic review of the Indo-Australian Zosteropidae (part 
I). Zool. Verh., No. 35: 1-204. 



1961. A systematic review of the Indo-Australian Zosteropidae (part 
II). Zool. Verh., No. 50: 1-168. 
Stresemann, E. 

1931. Die Zosteropiden der indo-australischen Region. Mitt. Zool. 
Mus. Berlin, 17:201-238. 
Tristram, H. B. 

1894. On some birds from Bugotu, Solomon Islands, and Santa Cruz. 
Ibis, (sixth series) 6:28-31. 



(Received 5 June, 1965.) 



JL-^t/ 



BREVIORA 

MmseiuiiM of Coniiparative Zoology 



Cambridge, Mass. September 15, 1965 Number 229 

THE GENUS DARLINGTONIA (SERPENTES) 

IN HISPANIOLA, 

INCLUDING A NEW SUBSPECIES FROM THE 

DOMINICAN REPUBLIC 

By Albert Schwartz 
AND Richard Thomas 

10,000 SW 84th St., Miami, Florida 33143 



The West Indian island of Hispaniola is outstanding for the rela- 
tively large number of endemic snake genera which inhabit it. 
These include laltris Cope (with two species), Uromacer Dumeril 
and Bibron (with apparently two or three species), Hypsirhynchus 
Giinther, and Darlingtonia Cochran. The latter is a small snake 
(snout-vent length to about 300 mm) which was described 
(Cochran, 1935) on the basis of a single Haitian individual col- 
lected by P. J. Darlington from Roche Croix, about 5000 feet 
(1515 meters) altitude, near Pic Macaya in the Massif de la Hotte. 
Since the genus was first described, the snake has been found to be 
not uncommon in Haiti in the vicinity of Furcy and Kenscoff on 
the Montague Noire. Additional specimens from the Massif de la 
Hotte remain rare, and we have seen only one other snake (in 
addition to the type) from Camp Perrin in that region. Considering 
that there is a distinct faunal resemblance between the Massif de 
la Hotte, Massif de la Selle, and Sierra de Baoruco, all serially 
arranged from west to east along the south island of Hispaniola, 
we expected that ultimately Darlingtonia might be taken in the 
last named range. Such indeed was the case when in the summer of 
1964 the junior author and David C. Leber succeeded in taking 
two females at a sawmill in the Baoruco. It was later learned that 
a single Darlingtonia had been collected in the Repiiblica Domini- 
cana by W. G. Hassler in 1935. 

In an attempt to compare these eastern specimens with the 
more western populations, we have borrowed Darlingtonia from 
the following collections: American Museum of Natural History 



BREVIORA 



No. 229 



(AMNH) ; Museum of Comparative Zoology, Harvard University 
(MCZ); United States National Museum (USNM); INIuseum of 
Zoology, University of Michigan (UMMZ). We wish to thank 
Charles M. Bogert, Grace M. Tilger, Ernest E. Williams, Doris M. 
Cochran, James A. Peters, Charles F. Walker, and George R. Zug 
for the loan of these small serpents; Mr. Leber and Ronald F. 
Klinikowski aided us in assembling material in the field (designated 
as ASFS), and Mr. Klinikowski has likewise made some of the 
illustrations for the present paper. We have examined 30 speci- 
mens in all, including the holotype (MCZ 38252) of the species, 
with all but five of these originating in the Furcy area. We are 
thus hampered in comparing these Furcy snakes with topotypical 
material from the east in the La Hotte, but for the moment there 
is no choice but to accept the agreement of the Montague Noire 
material with that from the La Hotte (admittedly an unlikely 
possibility, vide infra). The snakes from the Sierra de Baoruco, 
which are distinct from the more western populations, may be 
named : 




Fig. 1. Darlinglonia haedana haetiana, pattern at midbody, ASFS X20S0, 
Peneau, Bassin Bleu, 5000 feet, D^pt. de I'Ouest, Haiti. 



1965 



SNAKE GENUS DARLINGTONIA 



Darlingtonia haetiana perfector^ new subspecies 

Holotijpe: MCZ 77217, a female, from 24 km SW Barahona, 
3700 feet (1221 meters), Barahoua Province, Repuhlica Domini- 
cana, taken by David C. Leber on 6 July 19G4. Original number 
ASFS V 2897. 

Paratypes: ASFS V 2898, same data as type, but collected by 
Richard Thomas; AIMNH 49738, near Polo, 3000 feet (910 meters), 
Barahona Province, Republica Dominicana, W. G. Hassler, 19 
August 1935. 

Distribution: Known only from the Sierra de Baoruco in the 
Republica Dominicana, and probably occurring throughout 
moderate to higher elevations in that range. 

Diagnosis: A subspecies of Darlingtonia haetiana distinguished 
from the nominate race by a combination of a wide buffy nuchal 
collar (Fig. 3), a bolder and more contrasting longitudinally lined 
and dotted dorsal pattern (Fig. 2), and by less ventral and sub- 
caudal scales, giving a total underbody scale count from 174 to 
178 in contrast to 181 to 193 (both sexes) in D. h. haetiana. 




Fig. 2. Darlingtonia haetiana perfector, pattern at midbody, MCZ 77217, 
holotype, 24 km SW Barahona, .3700 feet, Barahona Province, Republica 
Dominicana. 



'From the Latin for "conipleter, finisher" in aUusion to the fact that with the specimens 
from Barahona the genus is known from the entire south island, whereas previously it had been 
known only from Morne La Selle and Massif de la Hotte. 



4 BREVIORA No. 229 

Description of holotype: A female with the following measure- 
ments and scale counts: snout-vent length, 190 mm, tail, 47 mm; 
ventral scales 133 (counted as suggested by Bowling, 1951), sub- 
caudal scales in 41 pairs; anal single; supralabials 7/7; infralabials 
8/8; loreal absent; preoculars 1/1; postoculars 2/2; temporals 
1 + 1 on right side, 1+2 on left side; dorsal scale rows 19-19-17. 
Coloration and pattern in life: dorsolateral area light brown with 
a middorsal dark reddish brown zone including a black middorsal 
stripe; a narrow dark brown conspicuous lateral stripe on scale 
rows 4 and 5 on each side; mid ventral region with a wide band al- 
most covering all of ventral scales including within it darker black 
blotches, the entire band with a blue iridescence; lowermost scale 
row on each side, and central part of second scale row and some- 
times third scale row and lateralmost ends of ventrals brick red; 
20 and 22 lateral dots on sixth to eighth scale rows on each side, 
each individual dot involving one, two, or three scales on adjacent 
rows, and yellowish orange with black margins in life. Tail brown 
above, black below. Head without a definitive pattern, but gen- 
erally blackish to reddish brown or orange on parietals, the brighter 
and paler color delimiting a vague, dark, T-shaped figure with its 
bar across the supraoculars and frontal, and the stem along the 
parietal suture, joining on the neck the middorsal black longi- 
tudinal band; a pale bar across the anterior half of the frontal, the 
prefrontals and supranasals variously marbled with brown and 
paler. A buffy collar, edged posteriorly with black, and two scales 
in width, goes from the angle of the jaws on either side, and is 
interrupted middorsally by the median black band. 

Variation: The female paratype (ASFS \'2898) has a snout- 
vent length of 180 mm and a tail length of 44 mm; there are 136 
ventral scales and 40 pairs of subcaudal scales. The supra- and 
infralabials, pre- and postoculars are as in the type. The temporals 
are + 2 on both sides, the single anterior temporal being fused 
with the parietal. The scale rows are 19-19-17, and the loreal is 
absent. In color and pattern the paratype is very like the tj^pe, 
except that, due to a smaller amount of pale color on the anterior 
half of the frontal, the transverse dark bar of the T-shaped cephalic 
figure is wider and more conspicuous. The buffy collar is present 
and readily visible. 

The male paratype (AAINH 49738) has a snout-vent length of 
216 mm and a tail length of 63 mm; there are 132 ventral scales 
and 46 pairs of subcaudal scales. The supralabials, pre- and 
postoculars, and temporals are like those of the type, except that 
there are 1 + 2 temporals on each side. The infralabials are 7 on 
each side. The scale rows are 19-19-17, and the loreal is absent. 



1965 



SNAKE GENUS DARLINGTONIA 



Although tlio male paratype is darker (possibly due to length of 
preservation) than the two smaller female pcrfector, the longitudi- 
nal zonation is still visible. The nuchal collar is bold and promi- 
nent. The head pattern is like that described for the type, except 
that the amount of pale coloration across the anterior portion of 
the frontal is less, thus making the transverse dark bar of the 
T-shaped figure broader. The ventral scales are entirely dark, 
with the lateralmost tips irregularly stippled light and dark. 





Fig. 3. Dorsal view of heads of Dariingionia haetiana: left, D. h. haeiiana, 
ASFS X2080; right, D. h. perfector, MCZ 77217. 



Comparisons and discussion of variation: Since there are more 
specimens available from the Furcy area than elsewhere, it is most 
profitable to first discuss the variation in snakes from this region, 
and then compare them both with the two western and three 
eastern serpents. At Furcy, the scalation shows the following 
variation: ventrals in males (13 specimens) range from 137 to 144 
(mean 139.7), ventrals in females (12 specimens) having the same 
range (mean 139.9). Subcaudals are 46 to 51 (mean 48.3) in 
males, and 41 to 50 (mean 45.3) in females. Ventrals + subcaudals 
are 185 to 193 (mean 188.0) in males, and 183 to 191 (mean 185.4) 
in females. Females have slightly less total underbody scales than 
do males. The largest male has a snout- vent length of 281 mm 
with a tail of 82 mm, whereas the largest female measures 305 in 
snout-vent length with a tail length of 83 mm. All specimens lack 
a loreal, and have 1 /I preoculars and 2/2 postoculars. The labials 
are normally 7/7 above and 8/8 below, although one snake has 
7/8 supralabials, three snakes have 7/7 infralabials, and four have 
7 /8 infralabials. The temporals are either 1 + 2 or 1 + 1 ; four 



6 BREVIORA No. 229 

snakes have 1 + 1 on each side, four have 1 + 1 and 1+2, and 
the balance have 1 + 2 on each side. The niimljer of lateral dots 
varies from (a single snake has no lateral dotting visible) and 4 
(one dot on one side and three on the other), to a maximum of 
54 (27 dots on each side) ; usually the number of dots on the two 
sides is not identical. The scale row formula is 19-19-17 with 
four exceptions; three of these are 20-19-17, and the other 21-19-17. 

The Furcy material shows an interesting ontogenetic pattern 
change. A series of six young snakes ranging in snout-vent lengths 
from 104 mm to 166 mm shows the development of the ventral 
dark band. In the smallest of these snakes (which still has evi- 
dence of the umbilicus at ventrals 115 to 117) the entire venter 
has a pale ground color with scattered dark brown or black blotches. 
The two snakes next in size (108 and 118 mm snout-vent) show 
the same condition, although the larger of these two shows the 
beginning of deposition of dark pigment ventrally. The next 
largest snake Avith a snout- vent length of 153 mm shows the 
obliterative effect of additional dark pigment so that the individual 
dark ventral blotches have become somewhat obscvn-cd. Finally, 
in two snakes with snout- vent lengths of 165 and 166 mm, the 
adult condition of a black or dark venter, with the original ju venal 
dark blotches barely discernible, is attained. In these young 
snakes the collar is pale and conspicuous; with increasing age the 
collar becomes fainter and less obvious so that in large adults it 
is much reduced and may be seen only as a small restricted pale 
nuchal area with some dark pigment deposited about it. Only 
two snakes from Furcy are as pale as are the two perfedor', and 
these two individuals have the lateral lines and dots as conspicuous 
as do the Baoruco snakes. In general, the Furcy snakes are dark 
brown: description in life of a series of D. h. haetiana from Bassin 
Bleu, Peneau, Dept. de FOuest, Haiti, noted the coloration as 
dark brown with a middorsal dark brown longitudinal band on a 
slightly lighter brown color (Fig. 1). The ventral ground color 
was dark brown with occasional whitish edges to the ventra's. 
The lateral margins of ventrals and first scale row on each side 
were brick red to orange-red, the extent of this color greatest 
anteriorly. The dorsal blotches were buffy-tan, outlined in brown. 
The collar (Fig. 3) was very much reduced or absent in this series 
which ranged in size from 159 to 245 mm in snout-vent length. 
Note also the lack of mention of the lateral stripe which was not 
obvious because of the dark lateral coloration. 

From the abo\e details in both scalation and coloration, it i.s 
apparent that 1). h. pcrfcclor differs not only in having fewer 



1965 SNAKE GENUS DARLINGTONIA 7 

uiiderbody scales than docs D. h. haetiana, but also in having a 
prominent nuchal collar and prominent lateral lines. 

There are two specimens of Darlimjtonia from the JNIassif de la 
Hotte, the type and another from Camp Perrin. These two indi- 
viduals are both females, the type having a snout-vent length of 
294 mm and the second specimen a snout-vent length of 247 mm. 
The tail of the type (88 mm) is unusually long, longer than any 
specimen of either sex, regardless of snout- vent length, from the 
Furc}^ area. The ventrals in these two snakes are 186 and 137, 
and the subcaudals 49 and 51, with total underbody scales 185 
and 188. There are no other scale differences, although the Camp 
Perrin specimen has 1 /2 preoculars and the type has 2+1 and 
2 + 2 temporals. Both are pallid snakes with conspicuous lateral 
lines and without collars; the amount of ventral darkening is 
variable, involving almost the entire width of the ventrals in the 
type, and with a clear reddish area on the lateral ends of the ven- 
trals in the second specimen. Using ventrals minus subcaudals 
as an index, the two western snakes both (with 86) lie just outside 
the range of the series of females from Furcy (with a range of 87 
to 102). Whether this hints at a basic difference between these 
two populations can be determined only by the acquisition of 
more material from the Massif de la Hotte. The extremely long 
tail of the type and, in the two La Hotte females, the relatively 
high number of subcaudals (49 and 51 versus 41 to 50 in Furcy 
females), and the tail /snout-vent ratio of 29.9 and 31.2 (versus 
23.7 to 27.2 in Furcy females) are all likewise suggestive of differ- 
entiation. 

Altitudinally, DarUngtonia haetiana ranges from 1000 feet 
(303 meters) at Camp Perrin to a know^i maximum of 5000 feet 
(1515 meters) at Roche Croix. Specimens from the Furcy area, 
from localities which can be mapped, show an altitudinal range 
of from 5000 feet (1515 meters) at Kenscoff and Peneau to 5600 
feet (1697 meters) at Furcy. The narrow altitudinal range is 
doubtless an artifact of collection, since the wider altitudinal 
limits to the west indicate that the snake may occur much lower. 
Once again, the extreme deforestation of the accessible mountains 
near Port-au-Prince may have been a crucial factor in limiting 
this snake to more favored higher elevations. The elevations for 
D. h. perfector (3000 and 3700 feet = 910 and 1221 meters) are 
intermediate and within the known limits of the jNIassif de la Hotte 
distribution of DarUngtonia. 



8 BREVIORA No. 229 

The series of five specimens from Peneau was collected in a 
montane ravine with dense growth of bamboo; the specimen from 
Kenscoff was secured under a pile of drying vetiver. The Camp 
Perrin snake was received from a native and there are no habitat 
data available for it. Of the two recent specimens of D. h. per- 
fector, the type was taken under a rock in a small weedy clearing 
surrounded by broad-leaf forest and cafetales, and the paratype 
was taken within a mat of cut vines in the same clearing. 

The absence of records of Darlingtonia from the area between 
Furcy and the eastern end of the Sierra de Baoruco is puzzling. 
If Darlingtonia has the altitudinal limits indicated by the La 
Hotte specimens, it is strange that it has not been taken at such 
localities as Foret des Pins in e.xtreme eastern Haiti or along the 
Dominico-Haitian border. It may be significant, however, that 
the junior author and Mr. Leber visited the type locality of 
D. h. perfedor in the summer of 1963, and although they secured 
a long series of Wetmorena, no Darlingtonia were taken. The two 
specimens were secured at the same locality a year later. One 
would expect that Darlingtonia occurs throughout much of the 
south island montane massifs, but, on the other hand, it is possible 
that the populations are in reality disjunct. Only further collect- 
ing may pro^"e which is the case. 

Hemipenis 

The everted hemipenis of Darlingtonia haetiana is relatively 
small (extending to the level of the 6th or 8th sul)caudal), and 
bilobed; the sulcus spermaticus is deeply forked: and the sulcate 
side (we depart from the usage of Dowling and Savage, 1960, and 
use sulcate and non-sulcate instead of medial and lateral for the 
surfaces of the everted organ) is strongly differentiated from the 
non-sulcate side in being covered with papillae from the region 
of the sulcus spermaticus to the apices. The sulcus spermaticus 
proceeds through the papiilate region to the apices. This papillate 
zone is sharply set off from the non-sulcate side which is uni- 
formly adorned with very small spines. The or-gan has the ap- 
pearance of having an elongate cordate shield (papillate zone) 
affixed to the sulcate side (Fig. 4). A row of enlarged but pro- 
gressively smaller spines begins on each edge of the non-sulcate 
side about midway the length of the organ (!e\-el of tiiird sub- 
caudal) and proceeds basally and diagonally, becoming lost in the 
profusion of smaller spines on the basal sulcate side. Papillate 
calyces cover the papillate sides of each lobe; the papillae become 



1965 



SNAKE GENUS DARLIxNGTONIA 





Fig. 4. Hemipenis of Darlingtonia haetiana; left, sulcate surface; right, 
non-sulcate surface; ]\ICZ 65100, near Palmiste, Furcy, Dept. de I'Ouest, 
Haiti. 



smaller and more sparse on the more basal portions of the hemi- 
penis. Several larger, isolated papillae are present on the non- 
sulcate surfaces of the lobes or in the crotch between them. One 
specimen, ASFS X2078, appears abnormal in having more sparse 
and less prominent calyces, and accessory sulci partially di\dding 
the papillate zone, making the organ almost capitate. 



Specimens Examined 

Darlingtonia haetiana haetiana: Haiti, Dept. du SucI, Roche 
Croix, northeastern foothills. Massif de la Hotte (= Pic Macaya), 
about 5000 feet (1515 meters) altitude, MCZ 38252 (type): Camp 
Perrin, 1000 feet (303 meters), ASFS X3058; Dept. de VOuest, 
Peneau, Bassin Bleu, 5000 feet (1515 meters), ASFS X2077-81 ; 
Furcy, AICZ 60060-61, MCZ 66996-98, UMMZ 123097 (4 speci- 
mens), USNM 123803; Vendome near Furcy, MCZ 65104-07; 
near Palmiste, Furcy, MCZ 65098-101 ; Morne Bourette, USNM 
117286; Kenscoff, 5000 feet (1515 meters), ASFS X2254. 

Darlingtonia haetiana perfector: Republica Dominicana, Bara- 
hona Prov., 24 km SW Barahona, 3700 feet (1221 meters), INICZ 
77217, ASFS V2898 (type and paratype); AMNH 49738, near 
Polo, 3000 feet (910 meters) (paratype). 



10 BREVIORA No. 229 

LITERATURE CITED 

Cochran, Doris M. 

1935. New reptiles and amphibians collected in Haiti by P. J. Darlington. 
Proc. Boston Soc. Nat. Hist., Vol. 40, no. 6, pp. 367-376. 
DowLiNG, Herndon G. 

1951. A proposed standard system of counting ventrals in snakes. 
Brit. Jour. Herpetology, Vol. 1, no. 5, pp. 97-99, 1 fig. 
DowLiNG, Herndon G., and Jay M. Savage 

1960. A guide to the snake hemipenis: a survey of basic structure and 
systematic characteristics. Zoologica, Vol. 45, pt. 1, pp. 17-28. 

(Received 2 June, 1965.) 



BREVIORA 

MiiseiiitM of Comparsitive Zoology 



Cambuiuge, Mass. Skptkmhek 15, 19(55 Number 2:^0 



NOTES ON SOME NON-PASSERTNE BIRDS FROM 
EASTERN ECUADOR 

By 
David W. Norton 



A new collection of birds from eastern Ecuador contains a 
number of significant specimens. The Ecuadorian segment of 
the upper Amazon basin has been generally neglected by orni- 
thologists since Chapman's study of 1926, while the adjacent 
areas of Colombia and Peru have been more recently investi- 
gated. The latter studies have raised questions about the 
continuity of populations along the eastern base of the Andes 
and this new collection answers some of these questions by 
documenting the presence of certain forms in eastern Ecuador. 

In 1963 and 1964, during the months June-September, I made 
collecting trips to eastern Ecuador, working at altitudes be- 
tween 300 and 1,500 meters. In 1963, I spent a month and a 
half at Limoncocha, and a month on Mount Sumaco. Accom- 
panied by Richard D. Chandler in 1964, I returned to collect 
on the slopes of Sumaco for two months. A collection of 1,900 
specimens from these two expeditions is deposited in the Mu- 
seum of Comparative Zoology. The present paper contains the 
most noteworthy records from among the 145 forms of non- 
passerines represented in this collection. 

COLLECTING LOCALITIES 

CoTAPiNO (often labeled on maps as ''Concepcion") : 0°45'S, 
77°25'W; alt. 700 m^ 25 June-12 July 1964; 250 specimens. 

At the junction of Rio Pucuno and Rio Cotapino, this hacienda 
serves as a departure point for Sumaco trips. Besides an airstrip 

1 Altitudes are based on readings taken with a pocket altimeter at each locality. 



2 - BREVIORA No. 230 

and some 60 acres of cultivated land, Cotapino offers shelter and 
a small number of Quechuan Indian laborers. Most of the speci- 
mens were collected here with mist nets. . . 

CuYUJA: 0°25'S. 78°08'W; alt. 2,400 m; 19 June 1904; 9 
specimens. 

Chandler and I visited this town 10 km ea.st of Papallaeta, 
hoping to find conditions comparable to those on the slopes of 
Sumaco. Collecting here was incidental, but promising, because 
the deforestation characterizing Papallaeta has not yet reached 
Cuyuja. 

EuGENio: 0°46'S, 77°24'AY; alt. 700 m; 16-22 July 1964; 111 
specimens. 

A day's foot travel northwest toward Sumaco from Cotapino, 
the locality takes its name from the lone Quechuan inhabitant of 
this last settlement on the way to Sumaco. 

LiMONCOCHA : P°25'S, 7d°38'W ; alt. 300 m ; 25 June-2 August 
1963 ; 300 sj^ecimens. 

Limoncocha is the site of a large missionary base camp, in- 
cluding an airstrip, housing for United States missionary fami- 
lies, and a Quechuan village, all of which has been established 
since 1955 on the shores of a lake in the midst of tropical forest. 
The large lake whose lemonade-colored water inspired the name, 
Limoncocha, lies two kilometers inland from the mouth of Rio 
Jivino on the Rio Napo. There are few such lakes in eastern 
Ecuador, and none as unspoiled as Limoncocha. Consequently, 
the lake and its outlet have yielded some unique specimens, sev- 
eral forms new to Ecuador, and at least one new race. 

Lower Rio Pucuno: 0°46'S, 77°28'W; alt. 500 m; 19-31 Au- 
gust 1963 ; 135 specimens. 

Halfway between Cotapino and Eugenio, a traveller must ford 
the Rio Pucuno afoot. In 1963, high water made the river im- 
passable for a few days, during which delay 1 collected along 
the banks above and below the ford. 

MoNTALVO: 02°05'S, 76°57'W; alt. 250 m; 5 specimens. 

I purchased several specimens of apparent interest from this 
locality from R. Olalla, who worked here as an independent 
collector in 1964. 

Rio Negro: 01°25'S, 78°03'W; alt. 1,200 m; 12-14 September 
1964; 35 specimens. 

Along 'the road near this village, a few specimens were col- 
lected for comparison with those from similar altitiules on 
Sunii^po. 



1965 BIRDS FROAl EASTERN ECUADOR 3 

Mount Sumaco : Mount Siimaco, an isolated massif rising to 
about 4,000 ni, lies about 40 km southeast of Baeza, and about 
100 km southeast of Quito. Pour collecting cami)s were estab- 
lished on the southeast slojies. I have arbitrarily prefixed the 
name Sumaeo to all localities above Eugenio, the highest point 
settled by Quechuan Indians. 

Sumaco, Guaticocha: 0°45'S, 77°24'W; alt. 750 m; 16-24 
August 1964; 123 specimens. 

Only a few hours due west of Eugenio lies this tiny, perfectly 
round, and very deep lake, where Chandler and I established a 
camp. 

Sumaco, head of Kio Guataraco : 0°40'S, 77°35'W; alt. 
1,350 m ; 24 July-5 August 1964 ; 350 specimens. 

The junction of several brooks in a series of waterfalls a few 
hundred meters below this camp, marks the beginning of the 
Rio Guataraco. 

Sumaco, Palm Peak (translated from local Quechuan hunt- 
ers' designation, Ramus-Urcu) -. 0°39'S, 77°36'W; alt. 1,500 m; 
6-14 August 1964 ; 209 specimens. 

Palm Peak is the rim of the altiplano which slopes gradually 
upward for two days' travel to the final steep slope of the moun- 
tain peak. Most of the collecting was done on the altiplano. 

Sumaco, Upper Rio Pucuno: 0°36'S, 77°35'W; alt. 1,200 m; 
11-16 August 1963; 135 specimens. 

The highest camp in 1963 was on the narrow ridge separating 
the two principal rivers of this face of Sumaco, the Pucuno and 
the Guataraco. 

SYSTEMATIC NOTES 
Nycticorax pileatus (Boddaert) 

Limoncocha, 1 £ . 

This wide-ranging form has not been recorded before from 
Ecuador, where its occurrence was to be expected. This indi- 
vidual was shot in the cow pasture at Limoncocha, a habitat 
somewhat unusual in eastern Ecuador, as cattle in the country 
are confined mostly to the highlands. 

Ixobrychus exilis limoncochae subsp. nov. 

Type: Adult male, No. 285,860, Museum of Comparative 
Zoology, collected at Limoncocha, alt. 300 m, Rio Napo, eastern 
Ecuador, by D. W. Norton, on 6 July 1963. 



BREVIORA 



No. 230 



Diagnosis: Similar to nominate exilis, but cheeks and auricu- 
lars shading to chestnut, ventrum less streaked. Similar to 
erythromelas in color of cheeks, but wing and bill somewhat 
longer, ventrum less richly colored, throat less streaked ; dorsum 
of female grayish brown, not reddish brown as in erythromelas. 
Distinguished from hogotensis by chestnut rather than ochra- 
ceous cheeks, by lighter ventrum, with less streaking on throat, 
and by a somewhat shorter bill. Generally smaller than pe- 
ruvianus, with ventrum, head, and wing coverts tawny rather 
than ochraceous. 

Range: Known only from the type locality. 









Measurements 










Wing 


Culmen (exposed) 


limoncochae 


(1$ - 


-type) 


120 mm 


45 mm 




(19) 




120 


46 


exilis 


(45) 




112-121 (117.8 ± 1.7) 


44-49 (46.3 ± .09) 




(1$) 




112 


43 


erythromelas 


(35) 




106-113 (109.3 ± 1.7) 


42-44 (42.7 rt 0.5) 




(2$) 




109-113 (111.0 ± 1.4) 


40.0 


hogotensis 


(2 5) 




116-122 (119.0 ± 2.1) 


40.0 




(5$) 




113-125 (120.8 ± 2.1) 


37-40 (39.0 ± 0.6) 


peruvianus 


(15) 




125 


53 



Remarks: The two specimens of limoncochae were collected 
with a single shot and, therefore, are presumed to have been 
mated, although their gonads were not enlarged. 

While examining comparative material, I found that a female 
(AMNH No. 151,639) from Antioquia, Colombia, is referable 
to hogotensis, which extends the known range of that race con- 
siderably to the northwest. 

I tentatively identified as peruvianus a mounted specimen in 
adult male plumage at the Colegio San Bolivar in Ambato, 
Ecuador. The wing measures 127 mm and the culmen 50 mm. 
Although there were no data, the curator of the collection, who 
is the widow of the collector, claimed that her husband shot all 
his specimens in Ecuador. Corroboration of this doubtful record 
would extend the known range of peruvianus north from coastal 
Peru (Dept. Libertad) to western Ecuador. 

The origins of the races in northern South America are deserv- 
ing of speculation. Significantly, I believe, the southern limits 
of migration of the two North American races are just to the 
north (eastern Panama). Ixohrychus exilis may originally have 
been a temperate and entirely migratory species. In this case. 



1965 BIRDS FROM EASTERN ECUADOR 5 

the South American races have arisen at the southern limits of 
the wintering populations, and may represent descendants of a 
few birds which gave up the migratory habit to breed near the 
equator. The South American forms must then be more recent 
developments than the splitting of the migratory population 
into eastern and western subspecies, because the four southern 
forms are neatly divisible morphologically into eastern and west- 
ern types. The easternmost form, erythromelas, seemingly shows 
the extreme in characteristics of nominate exilis of eastern 
United States, in its small size and generally reddish brown 
coloring. The westernmost, peruvianus, is similarly related to 
hesperis of the western United States, as it shows the extreme 
of large size and grayish brown coloring. In between, limon- 
cochac is a less extreme eastern type, and hogotensis a less 
extreme western type. 

Specimens examined: limoncochae : Ecuador, Limoncocha, 
1 (? , 1 9 ; erythromelas : Surinam, Paramaibo, Z $ , 1 9 ; hogoten- 
sis: Colombia, Savana de Bogota, 2$, 49 ; Antioquia, l9 ; 
peruvianus: Peru, Dept. Lima, Vegueta, 1^,1 imm. ; "Ecua- 
dor", 1[5 ]. 

Aramides calopterus Sclater and Salvin 

Sumaco, Guaticocha, 1 i ; Montalvo, 1 $ . 

A specimen from Montalvo, collected by R. Olalla in April 
1 964, is significantly lighter than any in a series from Rio Suno 
and Sumaco. However, I believe Olalla dries skins in direct sun- 
light, which Avould cause fading, and explain this disparity. 

PoRPHYRULA MARTiNiCA (Liunaeus) 

Limoncocha, 1 6 , 1 9 . 

Although it is recorded from western Ecuador, I find no pre- 
vious record of this wide-ranging species in eastern Ecuador, 
where its occurrence was to be expected. 

PoRPHYRULA FLAViROSTRis (Boddaert) 

Limoncocha, 1 $ . 

The species, new to Ecuador, is apparently uncommon in up- 
per Amazonia, for de Schauensee (1949: 432) includes it in the 
faunal list of Colombia, also on the basis of a single specimen 
(Florencia). Individuals occur regularly at Limoncocha, but I 
have not seen any elsewhere in eastern Ecuador. 



6 BREVIORA No. 230 

Jacana spinosa intermedia (Sclater) 

Limoncocha, 2 5 . 

Specimens from Limoncocha are clearly referable to inter- 
media. This first record of the species from eastern Ecuador 
extends the known range of intermedia south from eastern Co- 
lombia to at least the Rio Napo in Ecuador. Hellmayr and 
Conover (1948 :9) questioned the validity of peruviana, of north- 
eastern Peru but comparison of 14 specimens from northeastern 
Peru with a series of intermedia from Venezuela, Colombia, and 
Ecuador, shows Peruvian specimens to be much the largest and 
darkest in the group, and easily separable from the reasonably 
uniform intermedia specimens. 

Vanellus resplendens (Tschudi) 

Limoncocha, 2 i ; Mt. Cotopaxi, Laguna de Limpio, 2 9 . 

The Andean lapwing seems never to have been recorded from 
below 2,000 m anywhere in its range (Ecuador to northern 
Chile). The Limoncocha specimens were observed daily (per- 
sonal communication) on the grassy airfield of the mission sta- 
tion (alt. 300 m) from February to June 1963, before I collected 
them there in July. This unusual pair, frequenting an artificial 
habitat cleared only recently of tropical forest, proved to be two 
males coming into adult plumage and having small gonads. 
These birds were much tamer than individuals observed and 
collected on Mt. Cotopaxi. The unique record probably repre- 
sents young strays lost during the seasonal altitudinal migrations 
of this species in the Andes. Aside from the airstrip, the only 
other sizable unforested areas near Limoncocha are the gravel 
bars of the nearby Rio Napo. This pair was indeed traced twice 
to the gravel bars during the daylight hours. Significantly, the 
Quechuan Indians of Limoncoclia could give no local name for 
this bird, whereas all regularly occurring species on the Napo 
receive specific and descriptive Quechuan names. 

Genus Eutoxeres 

Eiitoxeres condamini condamini (Bourcicr) : Cotapino, 3,5, 

59. 

Eutoxeres aquila aquila (Boureier) : Eugenio, 2 c^ ; Sumaco, 
head of Rio Guataraeo, 1 S ; Sumaco, Guaticocha, 1 9 . 

The ranges of these sibling si)eeies overlap in eastern Ecuador 
and adjacent areas of Colombia and Peru. Since Chandler and I 



1965 



BIRDS FROM EASTERN ECUADOR 




Fig. 1. Distribution of Euioxeres aquila and Eutoxcres condamini. 



never found both species in the same locality, it is i)ossible that 
there is a difference in the habitat preferences of the two species. 
It seems that E. condamini occurs in open cultivated areas, at 
low elevations, whereas E. aquila occurs in dense forest at any 
altitude up to at least 1,500 m. 



8 BREVIORA No. 230 

The distribution of the forms of Eutoxeres may be related to 
the distribution of a principal food source. Greenewalt (1960: 
legend, PL 34) observed E. aquila feeding on "platanillos, " or 
plantains of the genus Hcliconia, using its highly specialized, 
downcurved bill to draw nectar from the deep, upright bracts 
of the flowers. Chandler and I had best results netting both 
species of Eutoxeres when the nets were placed close to Hcliconia 
plants, which further suggests the dependence of Eutoxeres 
upon the plantains. The genus Eutoxeres occurs in northwestern 
South America, roughly where the greatest concentrations of the 
35-odd species of Heliconia also occur. The flowering periods of 
the various species of Heliconia doubtless span the year, prob- 
ably providing a steady supph^ of food for the hummingbirds. 

Observations made in eastern Ecuador suggest that the critical 
factor in determining local distribution of the sibling species is 
the abundance of Heliconia plants. When land in eastern Ecua- 
dor is cleared, and particularly when bananas are planted, many 
"platanillos" invade the clearing. Apparentlj^ E. c. condamini 
establishes itself in the midst of this abundance, while E. a. 
aquila retreats to the forest. Although both forms have been 
recorded sympatrically at La Morelia, Colombia (de Schauensee, 
1949 : 541), and at the mouth of Rio Curaray (Zimmer, 1950 : 1), 
I suspect that condamini is found in the settlements while aquila 
is found farther afield. These records reflect either inexactness 
in recording the locality, or possibly the passage of time be- 
tween collectors' visits, during which a locality supporting aquila 
was cultivated, thereby attracting coiulamini . 

The origin of these sibling species merits some speculations. 
The simplest explanation of the present distribution of the spe- 
cies of Eutoxeres is that an original South American population 
was split by the Andean uplift, giving rise to condamini to the 
east and aquila to the Avest of the cordillera. Subsequently, 
aquila has colonized the isthmus of Panama, and has spilled east- 
ward over the Andes to invade the range of condamini. 

Genus Topaza 

Topaza pyra (Gould) : Cotapino, 1 S ; Sumaco, Guaticocha, 
19. 

These specimens are the first definitely to extend the known 
range of pyra so far up the Rio Napo, contradicting Oberholser's 
( 1902 : 322) assertion that the species does not occur above Coca 
on the upper Napo. Of greater interest is the proof that T. pyra 



1965 BIRDS FROM EASTERN ECUADOR 9 

occurs west of T. pella pamprepta, an endemic form known only 
from Boca Suno on the Napo in eastern Ecuador. The popula- 
tion of T. p. pamprepta seems to be surrounded by the morjilio- 
logically similar T. pijra, isolated in Boca Suno, 2,000 km from 
the other races of pella, which occur in the Guianas and in Para, 
Brazil. The validity of pamprepta, and the present range exten- 
sion of pyra make it impossible to accept Peters' (1945: 92) 
suggestion that pella and pyra might be conspecific. I have 
examined the type of pamprepta, finding Oberholser's (1902: 
322) description and the data given by the collectors, both 
accurate. The type is similar in pattern to nominate pella, 
but it is distinguished by a much longer tail and somewhat 
shorter bill. One can conclude that the curiously spotty distri- 
butions of the taxa of Topaza are artifacts due to the rarity of 
these equatorial hummingbirds. Future records of pella may 
come from northern Brazil, between Ecuador and the Guianas. 

Trogon rufus sulphureus Spix 

Sumaco, Guaticocha, 1 $ ; Sumaco, Guaticocha, 1 9 (preserved 
as a skeleton). 

Although this subspecies was to be expected in eastern Ecua- 
dor, Peters (1945: 157) omits both eastern Ecuador and eastern 
Colombia from its range. Apparently, the only previous records 
were specimens labeled with doubtful accuracy as being from 
Coca, Rio Napo, and "Equateur" (Zimmer, 1948: 29). The 
present record shows sulphureus to occur almost to the foot of 
the Andes and up to nearly 800 m in this part of its range. 

CaPITO NIGER PUNCTATUS (LcSSOn) 

Limoncocha, 3 S , 4 $ ; Lower Rio Pucuno, 1 S ; Cotapino, 

1$, 1$. 

The subspecific status of this species in upper Amazonia is in 
confusion. Several authors have disputed Brodkorb's (1939) 
two races, macintyrei of eastern Ecuador and conjunctus of 
northeastern Peru. Examining a large series from Colombia, 
Ecuador, and Peru, I fail to find any of the consistent geo- 
graphic variations mentioned by Brodkorb. Furthermore, the 
12 birds from Limoncocha show all extremes in pileum color 
and streaking of the ventrum. Bond (1954: 49) synonymized 
conjunctus with macintyrei, but distinguished macintyrei from 
pnnctatus on the basis of the amount of ventral streaking in one 
female from Villavicencio, Colombia. Two of four Limoncocha 



10 BREVIORA No. 230 

females have the ventrum as lightly streaked as this female from 
Villavicencio. I therefore agree with Peters (1948: 25) and 
ascribe to punctatus birds of upper Amazonia from Colombia to 
northeastern Peru. 

PicuMNUs RUFR'^NTRis RUFiVENTRis (Bonaparte) 

Limoncocha, 1 £ ; Cotapino, 1 S ; Eugenio, 1 S ; Sumaco, Palm 
Peak, 1$ , 19. 

Chandler and I found these birds quite common up to at least 
1,500 m on Mt. Sumaco, although the species is usually consid- 
ered an inhabitant of the Tropical Zone (de Schauensee, 1949: 
641). At Palm Peak, specimens were taken simultaneously with 
such typically subtropical forms as Cyanocorax yncas. 

ACKNOWLEDGMENTS 

I thank Dean Amadou of the American Museum of Natural 
History, James Bond of the Philadelphia Academy of Natural 
Sciences, and George E. Watson of the U.S. National Museum, 
all of whom lent comparative material for this study. Richard 
D. Chandler was a valuable companion and collector on the trip 
in 1964. I am especially grateful to Raymond A. Paynter, Jr. 
for his guidance during the preparation of this paper. 

LITERATUEE CITED 

Bond, James 

1954. Notes on Peruvian Pieiformes. Proc. Acad. Nat. Sci. Phila- 
delphia, 106: 45-61. 

Bbodkorb, Pierce 

1939. Two undeseribed South American barbets. Proc. Biol. Soc. 
Washington, 52: 135-136. 
Chapman, F. M. 

1926. Distribution of bird-life in Ecuador. Bull. Am. Mus. Nat. Hist., 
55: xiii -f 784 pp. 
DE Schauensee, R. M. 

1949. The birds of the Republic of Colombia. Caldasia, 5 (23j: 381- 
644. 
Grebinewalt, H. C. 

1960. Hummingbirds. New York, Doubleday. 250 pp. 
Hellmayr, C. E. and B. Conover 

1948. Catalogue of l)irds of the Americas. Field Mus. Nat. Hist., 
Zool. Ser., 13: ]it. 1, no. 3, vi -f 383 pp. 



1965 BIRDS KHOM EASTERN ECUADOR 11 

Oberholsek, H. C. 

1902. Catalogue of a collection of huniniinsliinls from Ecuador and 
Colombia. Proc. V. S. Nat. Mus., 24: .309-342. 
Peters, J. L. 

1945. Check-li.st of birds of the world. Vol. 5. Cambridge, Mass., 

Harvard Univ. Press, xi + 2.38 pp. 
1948. Check-list of birds of the Avorld. Vol. (5. Cambridge, Mass., 
Harvard Univ. Press, xi + 306 pp. 
ZiMMER, J. T . 

1948. The family Trogonidae. Am. Mus. Novit., No. 1380, 56 pp. 
1950. The genera Eutoxeres, Campylopterus, Eupetomcna, and Flnri- 
.suga. Am. Mus. Novit., No. 1450, 14 pp. 

(Received 15 June 1965.) 



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